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The original version of this Article contained an error in Table 2, where the number of individuals in the “All Amazonia” row was reported as 11,6431 instead of 116,431. Also, the original version of this Article contained an error in the Methods, where the R2 for the proportion of broken/uprooted dead trees increase per year was reported as 0.12, the correct value being 0.06. The original version of this Article contained errors in the author affiliations. The affiliation of Gerardo A. Aymard C. with UNELLEZGuanare, Herbario Universitario (PORT), Portuguesa, Venezuela Compensation International Progress S.A. Ciprogress–Greenlife.

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 similar to 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.

Forest biomass is an essential indicator for monitoring the Earth's ecosystems and climate. It is a critical input to greenhouse gas accounting, estimation of carbon losses and forest degradation, assessment of renewable energy potential, and for developing climate change mitigation policies such as REDD+, among others. Wall-to-wall mapping of aboveground biomass (AGB) is now possible with satellite remote sensing (RS). However, RS methods require extant, up-to-date, reliable, representative and comparable in situ data for calibration and validation. Here, we present the Forest Observation System (FOS) initiative, an international cooperation to establish and maintain a global in situ forest biomass database. AGB and canopy height estimates with their associated uncertainties are derived at a 0.25 ha scale from field measurements made in permanent research plots across the world's forests. All plot estimates are geolocated and have a size that allows for direct comparison with many RS measurements. The FOS offers the potential to improve the accuracy of RS- based biomass products while developing new synergies between the RS and ground-based ecosystem research communities.

To provide an empirical foundation for estimates of the Amazonian tree diversity, we recently published a checklist of 11,675 tree species recorded to date in the region (ter Steege H, et al. (2016) The discovery of the Amazonian tree flora with an updated checklist of all known tree taxa. Scientific Reports 6:29549). From this total of plant records compiled from public databases and literature, widely used in studies on the Amazonian plant diversity, only 6,727 tree species belong to the first taxonomically-vetted checklist published for the region (Cardoso D, et al. (2017) Amazon plant diversity revealed by a taxonomically verified species list. PNAS 114:10695-10700). The striking difference in these two numbers spurred us to evaluate both lists, in order to release an improved Amazonian tree list; to discuss species inclusion criteria; and to highlight the ecological importance of verifying the occurrence of "non-Amazonian" trees in the region through the localization and identification of specimens. A number of species in the 2016 checklist that are not trees, non-native, synonyms, or misspellings were removed and corresponded to about 23% of the names. Species not included in the taxonomicallyvetted checklist but verified by taxonomists to occur in Amazonia as trees were retained. Further, the inclusion of recently recorded/new species (after 2016), and recent taxonomic changes added up to an updated checklist including 10,071 species recorded for the Amazon region and shows the dynamic nature of establishing an authoritative checklist of Amazonian tree species. Completing and improving this list is a long-term, high-value commitment that will require a collaborative approach involving ecologists, taxonomists, and practitioners.

Many plant traits covary in a non-random manner reflecting interdependencies associated with "ecological strategy" dimensions. To understand how plants integrate their structural and physiological investments, data on leaf and leaflet size and the ratio of leaf area to sapwood area (ΦLS) obtained for 1020 individual trees (encompassing 661 species) located in 52 tropical forest plots across the Amazon Basin were incorporated into an analysis utilising existing data on species maximum height (Hmax), seed size, leaf mass per unit area (MA), foliar nutrients and δ13C, and branch xylem density (ρx). Utilising a common principal components approach allowing eigenvalues to vary between two soil fertility dependent species groups, five taxonomically controlled trait dimensions were identified. The first involves primarily cations, foliar carbon and MA and is associated with differences in foliar construction costs. The second relates to some components of the classic "leaf economic spectrum", but with increased individual leaf areas and a higher ΦLS newly identified components for tropical tree species. The third relates primarily to increasing Hmax and hence variations in light acquisition strategy involving greater MA, reductions in ΦLS and less negative δ13C. Although these first three dimensions were more important for species from high fertility sites the final two dimensions were more important for low fertility species and were associated with variations linked to reproductive and shade tolerance strategies. Environmental conditions influenced structural traits with ρx of individual species decreasing with increased soil fertility and higher temperatures. This soil fertility response appears to be synchronised with increases in foliar nutrient concentrations and reductions in foliar [C]. Leaf and leaflet area and ΦLS were less responsive to the environment than ρx. Thus, although genetically determined foliar traits such as those associated with leaf construction costs coordinate independently of structural characteristics such as maximum height, others such as the classical "leaf economic spectrum" covary with structural traits such as leaf size and ΦLS. Coordinated structural and physiological adaptions are also associated with light acquisition/shade tolerance strategies with several traits such as MA and [C] being significant components of more than one ecological strategy dimension. This is argued to be a consequence of a range of different potential underlying causes for any observed variation in such "ambiguous" traits. Environmental effects on structural and physiological characteristics are also coordinated but in a different way to the gamut of linkages associated with genotypic differences.

Amazonia is the most biodiverse rainforest on Earth, and the debate over how many tree species grow there remains contentious. Here we provide a checklist of all tree species collected to date, and describe spatial and temporal trends in data accumulation. We report 530,025 unique collections of trees in Amazonia, dating between 1707 and 2015, for a total of 11,676 species in 1225 genera and 140 families. These figures support recent estimates of 16,000 total Amazonian tree species based on ecological plot data from the Amazonian Tree Diversity Network. Botanical collection in Amazonia is characterized by three major peaks, centred around 1840, 1920, and 1980, which are associated with flora projects and the establishment of inventory plots. Most collections were made in the 20th century. The number of collections has increased exponentially, but shows a slowdown in the last two decades. We find that a species' range size is a better predictor of the number of times it has been collected than the species' estimated basin-wide population size. Finding, describing, and documenting the distribution of the remaining species will require coordinated efforts at under-collected sites.

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 thatmost 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.

Background: Trees and arborescent palms adopt different rooting strategies and responses to physical limitations imposed by soil structure, depth and anoxia. However, the implications of these differences for understanding variation in the relative abundance of these groups have not been explored. Aims: We analysed the relationship between soil physical constraints and tree and palm basal area to understand how the physical properties of soil are directly or indirectly related to the structure and physiognomy of lowland Amazonian forests. Methods: We analysed inventory data from 74 forest plots across Amazonia, from the RAINFOR and PPBio networks for which basal area, stand turnover rates and soil data were available.We related patterns of basal area to environmental variables in ordinary least squares and quantile regression models. Results: Soil physical properties predicted the upper limit for basal area of both trees and palms. This relationship was direct for palms but mediated by forest turnover rates for trees. Soil physical constraints alone explained up to 24% of palm basal area and, together with rainfall, up to 18% of tree basal area. Tree basal area was greatest in forests with lower turnover rates on well-structured soils, while palm basal area was high in weakly structured soils. Conclusions: Our results show that palms and trees are associated with different soil physical conditions. We suggest that adaptations of these life-forms drive their responses to soil structure, and thus shape the overall forest physiognomy of Amazonian forest vegetation.

Floristic variation is high in the Neotropics, but little is known about the factors shaping this variation at the mesoscale. We examined floristic composition and its relationship with environmental factors across 220 1-ha permanent plots in tropical lowland Bolivia. For each plot, abundance of 100 species (93 tree and 7 palm species =10 cm diam) was obtained. Climatic data, related to rainfall seasonality and temperature, were interpolated from all available weather stations in the region, and soil properties, related to texture and fertility, were obtained for each plot. Floristic variation was strongly associated with differences in water availability and temperature, and therefore the climatic gradient shaped floristic variation more strongly than the edaphic gradient. Detrended correspondence analysis ordination divided lowland Bolivia primarily into two major groups (Southern Chiquitano region vs. the Amazon region) and a multiple response permutation procedure distinguished five floristic regions. Overall, the tested environmental variables differed significantly among the five regions. Using indicator species analysis, we distinguished 82 strong indicator species, which had significant environmental preferences for one floristic region. These species can be used as indicators of environmental conditions or to determine which floristic region a certain forest belongs. Given the predicted decreases in rainfall and increases in temperature for tropical lowland forests, our gradient approach suggests that species composition may shift drastically with climate change

A soil charcoal survey was undertaken across 60,000 ha of closed-canopy tropical forest in central Guyana to determine the occurrence, ubiquity, and age of past forest fires across a range of terra firme soil types. Samples were clustered around six centers consisting of spatially nested sample stations. Most charcoal was found between 40 and 60 cm depth with fewest samples yielding material at 0-20 cm depth. The first core yielded charcoal at most stations. Charcoal ages of a random subsample ranged from less than 200 YBP to 9500 YBP with a noticeable peak between 1000 and 1250 YBP. Results reinforce a view that most closed-canopy tropical forests in eastern Amazonia have been subject to palaeo-fire events of unknown severity with a peak in charcoal age consistently appearing between 1000 and 2000 YBP. The two samples dated to the early Holocene represent some of the oldest indicators of paleo-fire known from upland Neotropical forest soils. Ubiquitous soil charcoal in central Guyana further indicate both forest resilience to fire and the widespread propensity for regional forests to burn, particularly during anomalous periods of drought.

Large-scale patterns of Amazonian biodiversity have until now been obscured by a sparse and scattered inventory record. Here we present the first comprehensive spatial model of tree α-diversity and tree density in Amazonian rainforests, based on the largest-yet compilation of forest inventories and bolstered by a spatial interpolation technique that allows us to estimate diversity and density in areas that have never been inventoried. These data were then compared to continent-wide patterns of rainfall seasonality. We find that dry season length, while only weakly correlated with average tree α-diversity, is a strong predictor of tree density and of maximum tree α-diversity. The most diverse forests for any given DSL are concentrated in a narrow latitudinal band just south of the equator, while the least diverse forests for any given DSL are found in the Guayana Shield and Amazonian Bolivia. Denser forests are more diverse than sparser forests, even when we used a measure of diversity that corrects for sample size. We propose that rainfall seasonality regulates tree α-diversity and tree density by affecting shade tolerance and subsequently the number of different functional types of trees that can persist in an area.