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.

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.

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.

Competition influences tree growth, but not mortality, across environmental gradients in Amazonia and tropical Africa.

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

Evolutionary diversity is associated with wood productivity in Amazonian forests.

Higher levels of taxonomic and evolutionary diversity are expected to maximize ecosystem function, yet their relative importance in driving variation in ecosystem function at large scales in diverse forests is unknown. Using 90 inventory plots across intact, lowland, terra firme, Amazonian forests and a new phylogeny including 526 angiosperm genera, we investigated the association between taxonomic and evolutionary metrics of diversity and two key measures of ecosystem function: aboveground wood productivity and biomass storage. While taxonomic and phylogenetic diversity were not important predictors of variation in biomass, both emerged as independent predictors of wood productivity. Amazon forests that contain greater evolutionary diversity and a higher proportion of rare species have higher productivity. While climatic and edaphic variables are together the strongest predictors of productivity, our results show that the evolutionary diversity of tree species in diverse forest stands also influences productivity. As our models accounted for wood density and tree size, they also suggest that additional, unstudied, evolutionarily correlated traits have significant effects on ecosystem function in tropical forests. Overall, our pan-Amazonian analysis shows that greater phylogenetic diversity translates into higher levels of ecosystem function: tropical forest communities with more distantly related taxa have greater wood productivity.

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.

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.

Evolutionary heritage influences Amazon tree ecology.

Lineages tend to retain ecological characteristics of their ancestors through time. However, for some traits, selection during evolutionary history may have also played a role in determining trait values. To address the relative importance of these processes requires large-scale quantification of traits and evolutionary relationships among species. The Amazonian tree flora comprises a high diversity of angiosperm lineages and species with widely differing life-history characteristics, providing an excellent system to investigate the combined influences of evolutionary heritage and selection in determining trait variation. We used trait data related to the major axes of life-history variation among tropical trees (e.g. growth and mortality rates) from 577 inventory plots in closed-canopy forest, mapped onto a phylogenetic hypothesis spanning more than 300 genera including all major angiosperm clades to test for evolutionary constraints on traits. We found significant phylogenetic signal (PS) for all traits, consistent with evolutionarily related genera having more similar characteristics than expected by chance. Although there is also evidence for repeated evolution of pioneer and shade tolerant life-history strategies within independent lineages, the existence of significant PS allows clearer predictions of the links between evolutionary diversity, ecosystem function and the response of tropical forests to global change.

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 Amazonian forest carbon cycling.

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

The flood pulse as the underlying driver of vegetation in the largest wetland and fishery of the Mekong Basin.

The Tonle Sap is the largest wetland in Southeast Asia and one of the world's most productive inland fisheries. The Mekong River inundates the Tonle Sap every year, shaping a mosaic of natural and agricultural habitats. Ongoing hydropower development, however, will dampen the flood pulse that maintains the Tonle Sap. This study established the current underlying relationship among hydrology, vegetation, and human use. We found that vegetation is strongly influenced by flood duration; however, this relationship was heavily distorted by fire, grazing, and rice cultivation. The expected flood pulse alteration will result in higher water levels during the dry season, permanently inundating existing forests. The reduction of the maximum flood extent will facilitate agricultural expansion into natural habitats. This study is the most comprehensive field survey of the Tonle Sap to date, and it provides fundamental knowledge needed to understand the underlying processes that maintain this important wetland.

Quantifying changes in flooding and habitats in the Tonle Sap Lake (Cambodia) caused by water infrastructure development and climate change in the Mekong Basin.

The economic value of the Tonle Sap Lake Floodplain to Cambodia is arguably among the highest provided to a nation by a single ecosystem around the world. Nonetheless, the Mekong River Basin is changing rapidly due to accelerating water infrastructure development (hydropower, irrigation, flood control, and water supply) and climate change, bringing considerable modifications to the flood pulse of the Tonle Sap Lake in the foreseeable future. This paper presents research conducted to determine how the historical flooding regime, together with human action, influenced landscape patterns of habitats in the Tonle Sap Lake, and how these habitats might shift as a result of hydrological changes. Maps of water depth, annual flood duration, and flood frequency were created for recent historical hydrological conditions and for simulated future scenarios of water infrastructure development and climate change. Relationships were then established between the historical flood maps and land cover, and these were subsequently applied to assess potential changes to habitat cover in future decades. Five habitat groups were clearly distinguishable based on flood regime, physiognomic patterns, and human activity: (1) Open water, flooded for 12 months in an average hydrological year; (2) Gallery forest, with flood duration of 9 months annually; (3) Seasonally flooded habitats, flooded 5-8 months and dominated by shrublands and grasslands; (4) transitional habitats, flooded 1-5 months and dominated by abandoned agricultural fields, receding rice/floating rice, and lowland grasslands; and (5) Rainfed habitats, flooded up to 1 month and consisting mainly of wet season rice fields and village crops. It was found that water infrastructure development could increase the area of open water (+18 to +21%) and the area of rainfed habitats (+10 to +14%), while reducing the area covered with seasonally flooded habitats (-13 to -22%) and gallery forest (-75 to -83%). Habitat cover shifts as a result of climate change include a net increase of open water (2-21%), as well as a reduction of rainfed habitats by 2-5% and seasonally flooded habitats by 5-11%. Findings from this study will help guide on-going and future conservation and restoration efforts throughout this unique and critical ecosystem.

The effectiveness of contrasting protected areas in preventing deforestation in Madre de Dios, Peru.

Accurate monitoring of the effectiveness of protected areas (PAs) in decreasing deforestation is increasingly important given the vital role of forest protection in climate change mitigation. Recent studies on PA effectiveness have used remote-sensing imagery to compare deforestation rates within PAs to surrounding areas. However, remote-sensing data used in isolation provides limited information on the factors contributing to effectiveness. We used landscape-modelling techniques to estimate the effectiveness of ten PAs in Madre de Dios, Peru. Factors influencing PA effectiveness were investigated using in situ key-informant interviews. Although all of the PAs studied had positive effectiveness scores, those with the highest scores were ecotourism and conservation concessions, where monitoring and surveillance activities and good relations with surrounding communities were reported as possible factors in decreasing deforestation rates. Native community areas had the lowest scores, with deforestation mainly driven by internal resource use and population growth. Weak local governance and immigration were identified as underlying factors decreasing the effectiveness of protection, whereas good relations with surrounding communities and monitoring activity increased effectiveness. The results highlight the need to combine remote sensing with in situ information on PA management because identification of drivers and deterrents of deforestation is vital for improving the effectiveness of protection.

Conservation strategies to mitigate impacts from climate change in Amazonia.

Protected area systems and conservation corridors can help mitigate the impacts of climate change on Amazonian biodiversity. We propose conservation design criteria that will help species survive in situ or adjust range distributions in response to increased drought. The first priority is to protect the western Amazon, identified as the 'Core Amazon', due to stable rainfall regimes and macro-ecological phenomena that have led to the evolution of high levels of biodiversity. Ecotones can buffer the impact from climate change because populations are genetically adapted to climate extremes, particularly seasonality, because high levels of habitat diversity are associated with edaphic variability. Future climatic tension zones should be surveyed for geomorphological features that capture rain or conserve soil moisture to identify potential refugia for humid forest species. Conservation corridors should span environmental gradients to ensure that species can shift range distributions. Riparian corridors provide protection to both terrestrial and aquatic ecosystems. Multiple potential altitudinal corridors exist in the Andes, but natural and anthropogenic bottlenecks will constrain the ability of species to shift their ranges and adapt to climate change. Planned infrastructure investments are a serious threat to the potential to consolidate corridors over the short and medium term.

Climate change, deforestation, and the fate of the Amazon.

The forest biome of Amazonia is one of Earth's greatest biological treasures and a major component of the Earth system. This century, it faces the dual threats of deforestation and stress from climate change. Here, we summarize some of the latest findings and thinking on these threats, explore the consequences for the forest ecosystem and its human residents, and outline options for the future of Amazonia. We also discuss the implications of new proposals to finance preservation of Amazonian forests.

Thirty years of land-cover change in Bolivia.

Land-cover change in eastern lowland Bolivia was documented using Landsat images from five epochs for all landscapes situated below the montane tree line at approximately 3000 m, including humid forest, inundated forest, seasonally dry forest, and cloud forest, as well as scrublands and grasslands. Deforestation in eastern Bolivia in 2004 covered 45,411 km2, representing approximately 9% of the original forest cover, with an additional conversion of 9042 km2 of scrub and savanna habitats representing 17% of total historical land-cover change. Annual rates of land-cover change increased from approximately 400 km2 y(-1) in the 1960s to approximately 2900 km2 y(-1) in the last epoch spanning 2001 to 2004. This study provides Bolivia with a spatially explicit information resource to monitor future land-cover change, a prerequisite for proposed mechanisms to compensate countries for reducing carbon emissions as a result of deforestation. A comparison of the most recent epoch with previous periods shows that policies enacted in the late 1990s to promote forest conservation had no observable impact on reducing deforestation and that deforestation actually increased in some protected areas. The rate of land-cover change continues to increase linearly nationwide, but is growing faster in the Santa Cruz department because of the expansion of mechanized agriculture and cattle farms.

Increasing biomass in Amazonian forest plots.

A previous study by Phillips et al. of changes in the biomass of permanent sample plots in Amazonian forests was used to infer the presence of a regional carbon sink. However, these results generated a vigorous debate about sampling and methodological issues. Therefore we present a new analysis of biomass change in old-growth Amazonian forest plots using updated inventory data. We find that across 59 sites, the above-ground dry biomass in trees that are more than 10 cm in diameter (AGB) has increased since plot establishment by 1.22 +/- 0.43 Mg per hectare per year (ha(-1) yr(-1), where 1 ha = 10(4) m2), or 0.98 +/- 0.38 Mg ha(-1) yr(-1) if individual plot values are weighted by the number of hectare years of monitoring. This significant increase is neither confounded by spatial or temporal variation in wood specific gravity, nor dependent on the allometric equation used to estimate AGB. The conclusion is also robust to uncertainty about diameter measurements for problematic trees: for 34 plots in western Amazon forests a significant increase in AGB is found even with a conservative assumption of zero growth for all trees where diameter measurements were made using optical methods and/or growth rates needed to be estimated following fieldwork. Overall, our results suggest a slightly greater rate of net stand-level change than was reported by Phillips et al. Considering the spatial and temporal scale of sampling and associated studies showing increases in forest growth and stem turnover, the results presented here suggest that the total biomass of these plots has on average increased and that there has been a regional-scale carbon sink in old-growth Amazonian forests during the previous two decades.