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.

Evaluation of the management potential of timber resources in clearwater floodplain forests in the Amazon using growth models.

The Amazonian clearwater igapós are poorly studied floodplain ecosystems that are mainly covered by forests and are undergoing massive threats due to changes in land use and climate. Their hydrochemical characteristics and edaphic conditions fall between those of the eutrophic várzea floodplains on whitewater rivers and those of the oligotrophic igapós on blackwater rivers. Previous studies have indicated the management potential of timber species in the highly dynamic várzea floodplains due to the fast tree growth and high forest productivity. Timber resource management, however, is not recommended for the blackwater ecosystem because of its slow dynamics and high vulnerability to disturbances. For clearwater igapós, information on the potential for sustainable management of timber resources is lacking. In this study, we modeled the growth in diameter, height, and volume to derive species-specific minimum logging diameters (MLD) and felling cycles (FC) for eight merchantable species in the clearwater igapós of the Branco and Tapajós rivers in the northern and southern Amazon Basin, respectively. Diameter growth was modeled by analyzing the tree rings that are annually formed in the Amazonian floodplains as a consequence of the regular and predicable long-term flooding. Growth modeling followed the guidelines of the Growth-Oriented Logging (GOL) concept, with the adjustment of diameter growth improved by applying nonlinear mixed-effects regression. MLDs varied from 36 to 90 cm and FCs ranged from 6 to 21 years, which diverges from the standards of Brazilian logging regulations (MLD: 50 cm; FC: 25-35 years). This indicates the potential for timber resource management, which should be tested and introduced at small scales, integrated in protected areas to stepwise promote the sustainable management of these natural resources by traditional communities to increase their income and the conservation of this ecosystem.

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.

More than 10,000 pre-Columbian earthworks are still hidden throughout Amazonia.

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.

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.

Convergent evolution of tree hydraulic traits in Amazonian habitats: implications for community assemblage and vulnerability to drought.

Amazonian droughts are increasing in frequency and severity. However, little is known about how this may influence species-specific vulnerability to drought across different ecosystem types. We measured 16 functional traits for 16 congeneric species from six families and eight genera restricted to floodplain, swamp, white-sand or plateau forests of Central Amazonia. We investigated whether habitat distributions can be explained by species hydraulic strategies, and if habitat specialists differ in their vulnerability to embolism that would make water transport difficult during drought periods. We found strong functional differences among species. Nonflooded species had higher wood specific gravity and lower stomatal density, whereas flooded species had wider vessels, and higher leaf and xylem hydraulic conductivity. The P50 values (water potential at 50% loss of hydraulic conductivity) of nonflooded species were significantly more negative than flooded species. However, we found no differences in hydraulic safety margin among species, suggesting that all trees may be equally likely to experience hydraulic failure during severe droughts. Water availability imposes a strong selection leading to differentiation of plant hydraulic strategies among species and may underlie patterns of adaptive radiation in many tropical tree genera. Our results have important implications for modeling species distribution and resilience under future climate scenarios.

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.

Floristic composition, structure and soil-vegetation relations in three white-sand soil patches in central Amazonia / Composição florística, estrutura e relação solo-vegetação em três áreas de campinarana na Amazônia central

ABSTRACT The Amazonian white-sand vegetation presents a set of unique features, such as the dominance of a few species, high endemism and low species richness, which differentiate it from other Amazonian forests. Soil parameters have long been recognized as the main drivers of white-sand vegetation (WSV) characteristics. However, how they influence the composition, richness and structure of this vegetation type is still poorly understood. In this study we investigated the variation in floristic composition between patches and the soil-vegetation relations in three central Amazonian WSV patches. We tested whether slight differences in soil properties are linked with differences in floristic composition, species richness and forest structure in adjacent patches. In each patch three plots of 50 x 50 m were sampled (a total of 2.25 ha). Soil samples were collected for each plot. The sampling cutoff for arboreal individuals was DBH ≥ 5 cm. We sampled a total of 3956 individuals belonging to 40 families and 140 species. In each patch only a few species were dominant, but the dominant species varied among patches. Differences among patches were significant, but plots in the same patch tended to have similar species composition. The variable sum of bases (SB) was directly related to species composition, however, species richness and forest structure were not related to soil parameters. Even small variations in soil parameters can change species composition in WSV, although these variations do not necessarily influence the richness and other structural parameters.

RESUMO As campinaranas amazônicas apresentam uma série de características únicas, como a dominância de poucas espécies, alto grau de endemismos e baixa riqueza de espécies, que as diferenciam de outras formações florestais amazônicas. Parâmetros edáficos têm sido apontados como os principais responsáveis pelas características das campinaranas. Contudo, como estes parâmetros influenciam a composição, riqueza e estrutura deste tipo de vegetação ainda é pouco entendido. Neste estudo investigamos a variação estrutural, a composição florística e a relação solo-vegetação em três áreas de campinarana na Amazônia central, com intuito de testar se pequenas diferenças nos parâmetros edáficos do solo estão relacionados com diferenças na composição, riqueza e estrutura do componente arbóreo em áreas de campinarana adjacentes. Em cada área foram amostradas três parcelas de 50 x 50 m (totalizando 2.25 ha), com o critério de inclusão para os indivíduos de DAP ≥ 5 cm. Amostras de solo foram coletadas em cada parcela. O número total de indivíduos amostrados foi 3956, pertencendo a 40 famílias e 140 espécies. Em cada área poucas espécies foram dominantes, mas estas variaram entre as áreas. Diferenças entre as áreas foram significativas, porém parcelas da mesma área tenderam a ter composição florística similar. A variável soma de bases (SB) foi diretamente relacionada à composição de espécies; contudo, riqueza de espécies e estrutura florestal não foram relacionadas a nenhum dos parâmetros do solo amostrados. Concluimos que mesmo pequenas variações nos parâmetros edáficos do solo podem mudar a composição de espécies em campinaranas, embora esta variação não necessariamente influencie a riqueza e outros parâmetros estruturais da vegetação.

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.

Damming the rivers of the Amazon basin.

More than a hundred hydropower dams have already been built in the Amazon basin and numerous proposals for further dam constructions are under consideration. The accumulated negative environmental effects of existing dams and proposed dams, if constructed, will trigger massive hydrophysical and biotic disturbances that will affect the Amazon basin's floodplains, estuary and sediment plume. We introduce a Dam Environmental Vulnerability Index to quantify the current and potential impacts of dams in the basin. The scale of foreseeable environmental degradation indicates the need for collective action among nations and states to avoid cumulative, far-reaching impacts. We suggest institutional innovations to assess and avoid the likely impoverishment of Amazon rivers.

The Brazilian freshwater wetscape: Changes in tree community diversity and composition on climatic and geographic gradients.

Wetlands harbor an important compliment of regional plant diversity, but in many regions data on wetland diversity and composition is still lacking, thus hindering our understanding of the processes that control it. While patterns of broad-scale terrestrial diversity and composition typically correlate with contemporary climate it is not clear to what extent patterns in wetlands are complimentary, or conflicting. To elucidate this, we consolidate data from wetland forest inventories in Brazil and examine patterns of diversity and composition along temperature and rainfall gradients spanning five biomes. We collated 196 floristic inventories covering an area >220 ha and including >260,000 woody individuals. We detected a total of 2,453 tree species, with the Amazon alone accounting for nearly half. Compositional patterns indicated differences in freshwater wetland floras among Brazilian biomes, although biomes with drier, more seasonal climates tended to have a larger proportion of more widely distributed species. Maximal alpha diversity increased with annual temperature, rainfall, and decreasing seasonality, patterns broadly consistent with upland vegetation communities. However, alpha diversity-climate relationships were only revealed at higher diversity values associated with the uppermost quantiles, and in most sites diversity varied irrespective of climate. Likewise, mean biome-level differences in alpha-diversity were unexpectedly modest, even in comparisons of savanna-area wetlands to those of nearby forested regions. We describe attenuated wetland climate-diversity relationships as a shifting balance of local and regional effects on species recruitment. Locally, excessive waterlogging strongly filters species able to colonize from regional pools. On the other hand, increased water availability can accommodate a rich community of drought-sensitive immigrant species that are able to track buffered wetland microclimates. We argue that environmental conditions in many wetlands are not homogeneous with respect to regional climate, and that responses of wetland tree communities to future climate change may lag behind that of non-wetland, terrestrial habitat.

An expert system model for mapping tropical wetlands and peatlands reveals South America as the largest contributor.

Wetlands are important providers of ecosystem services and key regulators of climate change. They positively contribute to global warming through their greenhouse gas emissions, and negatively through the accumulation of organic material in histosols, particularly in peatlands. Our understanding of wetlands' services is currently constrained by limited knowledge on their distribution, extent, volume, interannual flood variability and disturbance levels. We present an expert system approach to estimate wetland and peatland areas, depths and volumes, which relies on three biophysical indices related to wetland and peat formation: (1) long-term water supply exceeding atmospheric water demand; (2) annually or seasonally water-logged soils; and (3) a geomorphological position where water is supplied and retained. Tropical and subtropical wetlands estimates reach 4.7 million km2 (Mkm2 ). In line with current understanding, the American continent is the major contributor (45%), and Brazil, with its Amazonian interfluvial region, contains the largest tropical wetland area (800,720 km2 ). Our model suggests, however, unprecedented extents and volumes of peatland in the tropics (1.7 Mkm2 and 7,268 (6,076-7,368) km3 ), which more than threefold current estimates. Unlike current understanding, our estimates suggest that South America and not Asia contributes the most to tropical peatland area and volume (ca. 44% for both) partly related to some yet unaccounted extended deep deposits but mainly to extended but shallow peat in the Amazon Basin. Brazil leads the peatland area and volume contribution. Asia hosts 38% of both tropical peat area and volume with Indonesia as the main regional contributor and still the holder of the deepest and most extended peat areas in the tropics. Africa hosts more peat than previously reported but climatic and topographic contexts leave it as the least peat-forming continent. Our results suggest large biases in our current understanding of the distribution, area and volumes of tropical peat and their continental contributions.

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.

Composition, structure and floristic diversity in dense rain forest in the Eastern Amazon, Amapá, Brazil / Composição, estrutura e diversidade florística em floresta ombrófila densa na Amazônia Oriental, Amapá, Brasil

This study aims to evaluate the phytosociology and floristic composition of tree species in the eastern Amazon, at the Iratapuru River Sustainable Development Reserve (RDS), State of Amapá. Fouteen quarters with dimensions of 100 m x 100 m were randomly inventoried, and 50 sub-plots of 10 m x 20 m were established. In each sub-plot all living individuals were sampled, being taken from the height data and DAP (breast height diameter) for tree species ≥ 10 cm. A total of 5,233 individuals belonging to 33 families and 184 species were registered. The families with the largest number of species were Fabaceae (32), Lauraceae (17), Sapotaceae (12), Moraceae (10), Lecythidaceae (8) and Annonaceae (8). The six most abundant families (18.18% of total families) in the present study were responsible for more than half (57.92%) of the total number of species. The floristic structure of the area studied was diverse, with species of varied interests, including: medicinal, timber and oil-producing.

Este trabalho objetiva avaliar a composição florística e a fitossociologia de espécies arbóreas na Amazônia Oriental, Reserva de Desenvolvimento Sustentável do Rio Iratapuru, Estado do Amapá. Foram inventariados 14 quadrantes de 100 m x 100 m distribuídos aleatoriamente, onde se estabeleceram 50 sub-parcelas de 10 m x 20 m. Em cada sub-parcela foram amostrados todos os indivíduos vivos, sendo tomados dados de altura e DAP (diâmetro a altura do peito) para espécies arbóreas ≥10 cm. Foram registrados 5.233 indivíduos distribuídos em 33 famílias e 184 espécies. As famílias com maior número de espécies foram Fabaceae (17), Lauraceae (17), Sapotaceae (12), Moraceae (10), Lecythidaceae (8) e Annonaceae (8). Essas seis famílias mais abundantes (18,18% total das famílias) presentes no estudo foram responsáveis por mais da metade (57,92%) do número total de espécies. A estrutura florística da área estudada mostrou-se diversificada, apresentando espécies de interesses variados, como medicinal, madeireira e oleífera.

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.

Drought responses of flood-tolerant trees in Amazonian floodplains.

BACKGROUND: Flood-tolerant tree species of the Amazonian floodplain forests are subjected to an annual dry period of variable severity imposed when low river-water levels coincide with minimal precipitation. Although the responses of these species to flooding have been examined extensively, their responses to drought, in terms of phenology, growth and physiology, have been neglected hitherto, although some information is found in publications that focus on flooding. SCOPE: The present review examines the dry phase of the annual flooding cycle. It consolidates existing knowledge regarding responses to drought among adult trees and seedlings of many Amazonian floodplain species. MAIN FINDINGS: Flood-tolerant species display variable physiological responses to dry periods and drought that indicate desiccation avoidance, such as reduced photosynthetic activity and reduced root respiration. However, tolerance and avoidance strategies for drought vary markedly among species. Drought can substantially decrease growth, biomass and photosynthetic activity among seedlings in field and laboratory studies. When compared with the responses to flooding, drought can impose higher seedling mortality and slower growth rates, especially among evergreen species. Results indicate that tolerance and avoidance strategies for drought vary markedly between species. Both seedling recruitment and photosynthetic activity are affected by drought, CONCLUSIONS: For many species, the effects of drought can be as important as flooding for survival and growth, particularly at the seedling phase of establishment, ultimately influencing species composition. In the context of climate change and predicted decreases in precipitation in the Amazon Basin, the effects of drought on plant physiology and species distribution in tropical floodplain forest ecosystems should not be overlooked.

Struggle in the flood: tree responses to flooding stress in four tropical floodplain systems.

BACKGROUND AND AIMS: In the context of the 200th anniversary of Charles Darwin's birth in 1809, this study discusses the variation in structure and adaptation associated with survival and reproductive success in the face of environmental stresses in the trees of tropical floodplains. SCOPE: We provide a comparative review on the responses to flooding stress in the trees of freshwater wetlands in tropical environments. The four large wetlands we evaluate are: (i) Central Amazonian floodplains in South America, (ii) the Okavango Delta in Africa, (iii) the Mekong floodplains of Asia and (iv) the floodplains of Northern Australia. They each have a predictable 'flood pulse'. Although flooding height varies between the ecosystems, the annual pulse is a major driving force influencing all living organisms and a source of stress for which specialized adaptations for survival are required. MAIN POINTS: The need for trees to survive an annual flood pulse has given rise to a large variety of adaptations. However, phenological responses to the flood are similar in the four ecosystems. Deciduous and evergreen species respond with leaf shedding, although sap flow remains active for most of the year. Growth depends on adequate carbohydrate supply. Physiological adaptations (anaerobic metabolism, starch accumulation) are also required. CONCLUSIONS: Data concerning the ecophysiology and adaptations of trees in floodplain forests worldwide are extremely scarce. For successful floodplain conservation, more information is needed, ideally through a globally co-ordinated study using reproducible comparative methods. In the light of climatic change, with increasing drought, decreased groundwater availability and flooding periodicities, this knowledge is needed ever more urgently to facilitate fast and appropriate management responses to large-scale environmental change.