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.
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.
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.
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.
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.
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.
Trees structure the Earth's most biodiverse ecosystem, tropical forests. The vast number of tree species presents a formidable challenge to understanding these forests, including their response to environmental change, as very little is known about most tropical tree species. A focus on the common species may circumvent this challenge. Here we investigate abundance patterns of common tree species using inventory data on 1,003,805 trees with trunk diameters of at least 10 cm across 1,568 locations1-6 in closed-canopy, structurally intact old-growth tropical forests in Africa, Amazonia and Southeast Asia. We estimate that 2.2%, 2.2% and 2.3% of species comprise 50% of the tropical trees in these regions, respectively. Extrapolating across all closed-canopy tropical forests, we estimate that just 1,053 species comprise half of Earth's 800 billion tropical trees with trunk diameters of at least 10 cm. Despite differing biogeographic, climatic and anthropogenic histories7, we find notably consistent patterns of common species and species abundance distributions across the continents. This suggests that fundamental mechanisms of tree community assembly may apply to all tropical forests. Resampling analyses show that the most common species are likely to belong to a manageable list of known species, enabling targeted efforts to understand their ecology. Although they do not detract from the importance of rare species, our results open new opportunities to understand the world's most diverse forests, including modelling their response to environmental change, by focusing on the common species that constitute the majority of their trees.
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.
We describe the geographical variation in tree species composition across Amazonian forests and show how environmental conditions are associated with species turnover. Our analyses are based on 2023 forest inventory plots (1 ha) that provide abundance data for a total of 5188 tree species. Within-plot species composition reflected both local environmental conditions (especially soil nutrients and hydrology) and geographical regions. A broader-scale view of species turnover was obtained by interpolating the relative tree species abundances over Amazonia into 47,441 0.1-degree grid cells. Two main dimensions of spatial change in tree species composition were identified. The first was a gradient between western Amazonia at the Andean forelands (with young geology and relatively nutrient-rich soils) and central-eastern Amazonia associated with the Guiana and Brazilian Shields (with more ancient geology and poor soils). The second gradient was between the wet forests of the northwest and the drier forests in southern Amazonia. Isolines linking cells of similar composition crossed major Amazonian rivers, suggesting that tree species distributions are not limited by rivers. Even though some areas of relatively sharp species turnover were identified, mostly the tree species composition changed gradually over large extents, which does not support delimiting clear discrete biogeographic regions within Amazonia.
Aim: Amazonia hosts more tree species from numerous evolutionary lineages, both young and ancient, than any other biogeographic region. Previous studies have shown that tree lineages colonized multiple edaphic environments and dispersed widely across Amazonia, leading to a hypothesis, which we test, that lineages should not be strongly associated with either geographic regions or edaphic forest types. Location: Amazonia. Taxon: Angiosperms (Magnoliids; Monocots; Eudicots). Methods: Data for the abundance of 5082 tree species in 1989 plots were combined with a mega-phylogeny. We applied evolutionary ordination to assess how phylogenetic composition varies across Amazonia. We used variation partitioning and Moran's eigenvector maps (MEM) to test and quantify the separate and joint contributions of spatial and environmental variables to explain the phylogenetic composition of plots. We tested the indicator value of lineages for geographic regions and edaphic forest types and mapped associations onto the phylogeny. Results: In the terra firme and v & aacute;rzea forest types, the phylogenetic composition varies by geographic region, but the igap & oacute; and white-sand forest types retain a unique evolutionary signature regardless of region. Overall, we find that soil chemistry, climate and topography explain 24% of the variation in phylogenetic composition, with 79% of that variation being spatially structured (R-2 = 19% overall for combined spatial/environmental effects). The phylogenetic composition also shows substantial spatial patterns not related to the environmental variables we quantified (R-2 = 28%). A greater number of lineages were significant indicators of geographic regions than forest types. Main Conclusion: Numerous tree lineages, including some ancient ones (>66 Ma), show strong associations with geographic regions and edaphic forest types of Amazonia. This shows that specialization in specific edaphic environments has played a long-standing role in the evolutionary assembly of Amazonian forests. Furthermore, many lineages, even those that have dispersed across Amazonia, dominate within a specific region, likely because of phylogenetically conserved niches for environmental conditions that are prevalent within regions.
Human activities exert a pronounced influence on forest ecosystems, impacting both biodiversity and function across multiple scales. Despite this, the consequences of low-intensity human activities on tropical forest ecosystems are difficult to assess and, therefore, remain poorly explored. Here, the influence of human activities and other site-specific variables on forest tree assemblages in central-west Africa was investigated. By dividing forest tree species into edible (from the perspective of humans) and inedible species, we aimed to assess the differential impacts of human resource use on different species; in particular, the greatest impact of human activity was expected to be seen on edible tree species. Tree data from 66 plots across Nigeria and Cameroon collected between 2002 and 2019 and Generalized Dissimilarity Models (GDMs) were used to assess pairwise beta-diversity between plots, accounting for candidate factors including proximity to human presence, elevation, and stem density. The analysis revealed that human activity significantly affects beta-diversity within the Nigeria-Cameroon forest region. The key variables that shape total beta-diversity included geographical distance between plots, plot elevation, stem density, proximity to human presence, and forest species composition. The forest species composition (monodominant or mixed forest) appeared to influence dissimilarity in beta-diversity, specifically for edible tree species. This pattern was not observed for inedible species, likely linked to the cultural practices in the region. While stem density contributed to the edible species models, elevation was more relevant for inedible species. These findings underscore the critical role of human influence in shaping tree species assemblages in African tropical forests and stress the necessity for further research in this area.
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.
Aim: To investigate the geographic patterns and ecological correlates in the geographic distribution of the most common tree dispersal modes in Amazonia (endozoo-chory, 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
Indigenous societies are known to have occupied the Amazon basin for more than 12,000 years, but the scale of their influence on Amazonian forests remains uncertain. We report the discovery, using LIDAR (light detection and ranging) information from across the basin, of 24 previously undetected pre-Columbian earthworks beneath the forest canopy. Modeled distribution and abundance of large-scale archaeological sites across Amazonia suggest that between 10,272 and 23,648 sites remain to be discovered and that most will be found in the southwest. We also identified 53 domesticated tree species significantly associated with earthwork occurrence probability, likely suggesting past management practices. Closed-canopy forests across Amazonia are likely to contain thousands of undiscovered archaeological sites around which pre-Columbian societies actively modified forests, a discovery that opens opportunities for better understanding the magnitude of ancient human influence on Amazonia and its current state.