One sixth of Amazonian tree diversity is dependent on river floodplains.

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.

Mapping density, diversity and species-richness of the Amazon tree flora.

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.

Long-term Ecological Research: Chasing fashions or being prepared for fashion changes?

Long-term-ecological-research (LTER) faces many challenges, including the difficulty of obtaining long-term funding, changes in research questions and sampling designs, keeping researchers collecting standardized data for many years, impediments to interactions with local people, and the difficulty of integrating the needs of local decision makers with "big science". These issues result in a lack of universally accepted guidelines as to how research should be done and integrated among LTER sites. Here we discuss how the RAPELD (standardized field infrastructure system), can help deal with these issues as a complementary technique in LTER studies, allowing comparisons across landscapes and ecosystems and reducing sampling costs. RAPELD uses local surveys to understand broad spatial and temporal patterns while enhancing decision-making and training of researchers, local indigenous groups and traditional communities. Sampling of ecological data can be carried out by different researchers through standardized protocols, resulting in spatial data that can be used to answer temporal questions, and allow new questions to be investigated. Results can also be integrated into existing biodiversity networks. Integrated systems are the most efficient way to save resources, maximize results, and accumulate information that can be used in the face of the unknown unknowns upon which our future depends.

Unraveling Amazon tree community assembly using Maximum Information Entropy: a quantitative analysis of tropical forest ecology.

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.

Amazon tree dominance across forest strata.

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.

The other side of droughts: wet extremes and topography as buffers of negative drought effects in an Amazonian forest.

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.

Biased-corrected richness estimates for the Amazonian tree flora.

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.

Rarity of monodominance in hyperdiverse Amazonian forests.

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.

Can traits predict individual growth performance? A test in a hyperdiverse tropical forest.

The functional trait approach has, as a central tenet, that plant traits are functional and shape individual performance, but this has rarely been tested in the field. Here, we tested the individual-based trait approach in a hyperdiverse Amazonian tropical rainforest and evaluated intraspecific variation in trait values, plant strategies at the individual level, and whether traits are functional and predict individual performance. We evaluated > 1300 tree saplings belonging to > 383 species, measured 25 traits related to growth and defense, and evaluated the effects of environmental conditions, plant size, and traits on stem growth. A total of 44% of the trait variation was observed within species, indicating a strong potential for acclimation. Individuals showed two strategy spectra, related to tissue toughness and organ size vs leaf display. In this nutrient- and light-limited forest, traits measured at the individual level were surprisingly poor predictors of individual growth performance because of convergence of traits and growth rates. Functional trait approaches based on individuals or species are conceptually fundamentally different: the species-based approach focuses on the potential and the individual-based approach on the realized traits and growth rates. Counterintuitively, the individual approach leads to a poor prediction of individual performance, although it provides a more realistic view on community dynamics.

Near-infrared spectrometry allows fast and extensive predictions of functional traits from dry leaves and branches.

The linking of individual functional traits to ecosystem processes is the basis for making generalizations in ecology, but the measurement of individual values is laborious and time consuming, preventing large-scale trait mapping. Also, in hyper-diverse systems, errors occur because identification is difficult, and species level values ignore intra-specific variation. To allow extensive trait mapping at the individual level, we evaluated the potential of Fourrier-Transformed Near Infra-Red Spectrometry (FT-NIR) to adequately describe 14 traits that are key for plant carbon, water, and nutrient balance. FT-NIR absorption spectra (1,000-2,500 nm) were obtained from dry leaves and branches of 1,324 trees of 432 species from a hyper-diverse Amazonian forest. FT-NIR spectra were related to measured traits for the same plants using partial least squares regressions. A further 80 plants were collected from a different site to evaluate model applicability across sites. Relative prediction error (RMSErel ) was calculated as the percentage of the trait value range represented by the final model RMSE. The key traits used in most functional trait studies; specific leaf area, leaf dry matter content, wood density and wood dry matter content can be well predicted by the model (R2  = 0.69-0.78, RMSErel  = 9-11%), while leaf density, xylem proportion, bark density and bark dry matter content can be moderately well predicted (R2  = 0.53-0.61, RMSErel  = 14-17%). Community-weighted means of all traits were well estimated with NIR, as did the shape of the frequency distribution of the community values for the above key traits. The model developed at the core site provided good estimations of the key traits of a different site. An evaluation of the sampling effort indicated that 400 or less individuals may be sufficient for establishing a good local model. We conclude that FT-NIR is an easy, fast and cheap method for the large-scale estimation of individual plant traits that was previously impossible. The ability to use dry intact leaves and branches unlocks the potential for using herbarium material to estimate functional traits; thus advancing our knowledge of community and ecosystem functioning from local to global scales.

Species Distribution Modelling: Contrasting presence-only models with plot abundance data.

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.

Estimating the global conservation status of more than 15,000 Amazonian tree species.

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.

Hyperdominance in the Amazonian tree flora.

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.

Amazon forest carbon dynamics predicted by profiles of canopy leaf area and light environment.

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