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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.
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Biodiversidad , Inundaciones , Ríos , Árboles , Brasil , BosquesRESUMEN
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.
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ARN Largo no Codificante , Árboles , Bosques , Suelo , TemperaturaRESUMEN
Biodiversity loss is one of the main challenges of our time,1,2 and attempts to address it require a clear understanding of how ecological communities respond to environmental change across time and space.3,4 While the increasing availability of global databases on ecological communities has advanced our knowledge of biodiversity sensitivity to environmental changes,5,6,7 vast areas of the tropics remain understudied.8,9,10,11 In the American tropics, Amazonia stands out as the world's most diverse rainforest and the primary source of Neotropical biodiversity,12 but it remains among the least known forests in America and is often underrepresented in biodiversity databases.13,14,15 To worsen this situation, human-induced modifications16,17 may eliminate pieces of the Amazon's biodiversity puzzle before we can use them to understand how ecological communities are responding. To increase generalization and applicability of biodiversity knowledge,18,19 it is thus crucial to reduce biases in ecological research, particularly in regions projected to face the most pronounced environmental changes. We integrate ecological community metadata of 7,694 sampling sites for multiple organism groups in a machine learning model framework to map the research probability across the Brazilian Amazonia, while identifying the region's vulnerability to environmental change. 15%-18% of the most neglected areas in ecological research are expected to experience severe climate or land use changes by 2050. This means that unless we take immediate action, we will not be able to establish their current status, much less monitor how it is changing and what is being lost.
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Biodiversidad , Bosques , Humanos , Bosque Lluvioso , Brasil , Clima Tropical , Conservación de los Recursos Naturales , EcosistemaRESUMEN
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.
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Biodiversidad , Ecosistema , Entropía , Bosques , Plantas , Ecología , Clima TropicalRESUMEN
A new Leucauge species, widespread in the Amazon rainforest, is described and illustrated. Leucauge behemoth n. sp. lives in colonies, usually positioned above turbulent waters. Both males and females have a unique color pattern, especially on the abdomen, which makes this species easily identifiable. Males are distinguished from most congeners by the long hook of the cymbium and from their closest relatives by the lack of a cymbial dorsobasal process. Females of this species differ from their congeners by the presence of a rounded ventral process in the epigynum. Leucauge argyroaffins Soares & Camargo, 1948 is proposed as a junior synonym of Leucauge argyra (Walckenaer, 1841).
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Gastrópodos , Arañas , Masculino , Femenino , Animales , Bosque LluviosoRESUMEN
A new Leucauge species, widespread in the Amazon rainforest, is described and illustrated. Leucauge behemoth n. sp. lives in colonies, usually positioned above turbulent waters. Both males and females have a unique color pattern, especially on the abdomen, which makes this species easily identifiable. Males are distinguished from most congeners by the long hook of the cymbium and from their closest relatives by the lack of a cymbial dorsobasal process. Females of this species differ from their congeners by the presence of a rounded ventral process in the epigynum. Leucauge argyroaffins Soares & Camargo, 1948 is proposed as a junior synonym of Leucauge argyra (Walckenaer, 1841).
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Migratory species are the most important commercial fishes in the Amazon. They are also now the most threatened directly by some combination of overfishing, floodplain deforestation, and dam construction. Limited governmental monitoring and implemented regulations impede adequate management of the fisheries at adequate scale. We summarize the current stock status of the three most heavily exploited long-distance migratory species, which are two goliath catfishes (Brachyplatystoma rousseauxii and B. vaillantii) and the characiform Colossoma macropomum. In addition, we analyze impacts beyond overfishing on these species. Our results indicate: (i) the overfishing trends for these important species are either ominous or indicate the verge of collapse of the commercial fisheries based on them, and (ii) a dangerous synergy between overfishing, hydroelectric dams, and floodplain deforestation further challenge fisheries management of migratory species in the Amazon. We propose eight direct governmental actions as a proactive approach that addresses the main impacts on the fisheries. We consider that the most practical way to assess and manage overfishing of migratory species in the short run in an area as large as the main commercial fishing area in the Amazon is at market sites where enforced regulations can control fish catch. The management of the three species considered here has implications beyond just their sustainability. Their management would represent a paradigm shift where the governments assume their legal responsibilities in fishery management. These responsibilities include regulation enforcement, data collecting, inter-jurisdictional cooperation to protect migratory species at realistic life history scales, mitigation of the Madeira dams to assure goliath catfish passage to the largest western headwater region, and recognition of monitoring and managing wetland deforestation for the protection of fish and other aquatic and terrestrial biodiversity.
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Bagres , Explotaciones Pesqueras , Animales , Biodiversidad , Bagres/fisiología , Conservación de los Recursos Naturales , PecesRESUMEN
Beta diversity is usually high along elevational gradients. We studied a spider community at the Pico da Neblina (Brazil), an Amazonian mountain which is one of the southern components of the Guayana region. We sampled six elevations and investigated if beta diversity patterns correspond to the elevational division proposed for the region, between lowlands (up to 500 m), uplands (500 m to 1500 m), and highlands (>1500 m). Patterns of dominance increased with elevation along the gradient, especially at the two highest elevations, indicating that changes in composition may be accompanied by changes in species abundance distribution. Beta diversity recorded was very high, but the pattern observed was not in accordance with the elevationaldivision proposed for the region. While the highlands indeed harbored different fauna, the three lowest elevationshad similar species compositions, indicating that the lowlands spider community extends into the uplands zone. Other measures of compositional change, such as similarity indices and species indicator analysis, also support this pattern. Our results, in addition to a revision of the literature, confirm the high diversity and endemism rates of montane spider communities, and we stress the importance of protecting those environments, especially considering the climate crisis.
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The Mid-Domain Effect (MDE) and the Rapoport (RE) effect are two biogeographical theories that make predictions about biogeogaphic patterns. MDE predicts higher richness in the central portions of a gradient if it is within a bounded domain. RE predicts a positive relation between altitude and species range size along an altitudinal gradient. Our aim was to document the distribution of spider species richness along an altitudinal gradient in the Brazilian Amazon, and to test the influence of MDE and RE on the diversity patterns. Our study was conducted at the Pico da Neblina (Amazonas state, Brazil), and we sampled spiders at six different altitudes using two methods: nocturnal hand sampling and a beating tray. We obtained 3,140 adult spiders from 39 families, sorted to 529 species/morphospecies. Richness declined continuously with an altitude increase, but the fit with the MDE richness estimates was very weak and was not significant. Range size was not related to altitude, i. e., no RE. Finally, the abundance distribution within each species range varied more specifically, which prevented the occurrence of a RE at the community level. The influence of MDE was extremely low, a consequence of our community characteristics, formed mostly by small range size species. Short and medium range species were located at all altitudes, preventing a significant relation between range size and altitude. The distribution of abundance within a species range varied specifically and do not support a RE hypothesis.
O Efeito do Domínio Central (MDE) e o Efeito Rapoport (ER) são duas teorias biogeografias que fazem previsões sobre a distribuição da diversidade ao longo de gradientes. O MDE prevê maior riqueza nas porções centrais de um gradiente, se este estiver dentro de um domínio fechado. O ER prevê uma relação positiva entre altitude e tamanho da distribuição ao longo do gradiente altitudinal. Nosso objetivo foi o de registrar a distribuição de uma comunidade de aranhas ao longo de um gradiente altitudinal na Amazônia Brasileira, e testar se há uma influência do EDC e do ER sobre os padrões de diversidade da comunidade. Nosso estudo foi feito no Parque Nacional do Pico da Neblina (AM, Brasil), e nós amostramos aranhas em seis altitudes diferentes. Nós coletamos 3.140 exemplares adultos de 39 famílias, que foram divididos em 529 espécies/morfoespécies. A riqueza declinou com o aumento de altitude, mas o padrão não mostrou ajuste com as previsões feitas pelo EDC. O tamanho da distribuição altitudinal também não esteve relacionado ao previsto pelo ER. Por fim, a distribuição de abundância ao longo da distribuição altitudinal das espécies variou de maneira específica, o que impediu a ocorrência de um ER nos padrões da comunidade. A influência do EDC sobre os padrões observados foi baixíssima, uma consequência de características de nossa comunidade, já que esta é formada por espécies com pequena distribuição altitudinal. Espécies de distribuição altitudinal médias e grandes ocorreram em todas as partes do gradiente o que impediu a ocorrência de um ER. Por fim, o ER também não foi observado na distribuição de abundância das espécies ao longo do gradiente, já que essa variou de maneira específica.
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Abstract: The Mid-Domain Effect (MDE) and the Rapoport (RE) effect are two biogeographical theories that make predictions about biogeogaphic patterns. MDE predicts higher richness in the central portions of a gradient if it is within a bounded domain. RE predicts a positive relation between altitude and species range size along an altitudinal gradient. Our aim was to document the distribution of spider species richness along an altitudinal gradient in the Brazilian Amazon, and to test the influence of MDE and RE on the diversity patterns. Our study was conducted at the Pico da Neblina (Amazonas state, Brazil), and we sampled spiders at six different altitudes using two methods: nocturnal hand sampling and a beating tray. We obtained 3,140 adult spiders from 39 families, sorted to 529 species/morphospecies. Richness declined continuously with an altitude increase, but the fit with the MDE richness estimates was very weak and was not significant. Range size was not related to altitude, i. e., no RE. Finally, the abundance distribution within each species range varied more specifically, which prevented the occurrence of a RE at the community level. The influence of MDE was extremely low, a consequence of our community characteristics, formed mostly by small range size species. Short and medium range species were located at all altitudes, preventing a significant relation between range size and altitude. The distribution of abundance within a species range varied specifically and do not support a RE hypothesis.
Resumo: O Efeito do Domínio Central (MDE) e o Efeito Rapoport (ER) são duas teorias biogeografias que fazem previsões sobre a distribuição da diversidade ao longo de gradientes. O MDE prevê maior riqueza nas porções centrais de um gradiente, se este estiver dentro de um domínio fechado. O ER prevê uma relação positiva entre altitude e tamanho da distribuição ao longo do gradiente altitudinal. Nosso objetivo foi o de registrar a distribuição de uma comunidade de aranhas ao longo de um gradiente altitudinal na Amazônia Brasileira, e testar se há uma influência do EDC e do ER sobre os padrões de diversidade da comunidade. Nosso estudo foi feito no Parque Nacional do Pico da Neblina (AM, Brasil), e nós amostramos aranhas em seis altitudes diferentes. Nós coletamos 3.140 exemplares adultos de 39 famílias, que foram divididos em 529 espécies/morfoespécies. A riqueza declinou com o aumento de altitude, mas o padrão não mostrou ajuste com as previsões feitas pelo EDC. O tamanho da distribuição altitudinal também não esteve relacionado ao previsto pelo ER. Por fim, a distribuição de abundância ao longo da distribuição altitudinal das espécies variou de maneira específica, o que impediu a ocorrência de um ER nos padrões da comunidade. A influência do EDC sobre os padrões observados foi baixíssima, uma consequência de características de nossa comunidade, já que esta é formada por espécies com pequena distribuição altitudinal. Espécies de distribuição altitudinal médias e grandes ocorreram em todas as partes do gradiente o que impediu a ocorrência de um ER. Por fim, o ER também não foi observado na distribuição de abundância das espécies ao longo do gradiente, já que essa variou de maneira específica.
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Abstract: Florivory can alter plant reproductive success by damaging sexual reproductive structures and disrupting plant-pollinator interactions through decreased flower attractiveness. Here, we report new records of the cactus Tacinga inamoena flower consumption by the Tropidurus hispidus lizard in the Brazilian Caatinga. We monitored 53 flowers from 11 T. inamoena individuals with camera traps over a 1-year period, totalling 450 camera-days of sample effort. We detected four florivory events. In three of these events, flowers were entirely consumed or had their reproductive structures severely damaged, leading to no fruit formation. Florivory events occurred in the morning, right after anthesis, in flowers near the ground, and lizards did not climb the cactus. Our results suggest that T. hispidus florivory on T. inamoena could have a negative impact on fruit set, since the consumed flowers were entirely destroyed. However, the long-term effects of florivory by lizards on T. inamoena reproductive success in the Caatinga still needs to be elucidated.
Resumo: A florivoria pode alterar o sucesso reprodutivo da planta através de danos às estruturas reprodutivas sexuais e interrupção das interações planta-polinizador por meio da diminuição da atratividade das flores. Aqui, relatamos novos registros de consumo de flores da cactácea opuntióide Tacinga inamoena pelo lagarto Tropidurus hispidus na Caatinga brasileira. Nós monitoramos 53 flores de 11 indivíduos de T. inamoena com armadilhas fotográficas por um período de um ano, totalizando 450 dias-câmera de esforço amostral. Detectamos quatro eventos de florivoria. Em três desses eventos, as flores foram totalmente consumidas ou tiveram suas estruturas reprodutivas severamente danificadas, não levando à formação de frutos. Os eventos de florivoria ocorreram pela manhã, logo após a antese, em flores próximas ao solo, e os lagartos não escalaram o cacto. Nossos resultados sugerem que a florivoria de T. hispidus em T. inamoena pode ter um impacto negativo na frutificação, uma vez que as flores consumidas foram totalmente destruídas. No entanto, os efeitos a longo prazo da florivoria por lagartos sobre o sucesso reprodutivo de T. inamoena na Caatinga ainda precisam ser elucidados.
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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.
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Biodiversidad , Clasificación/métodos , Bosques , Ríos , Árboles/clasificación , BrasilRESUMEN
The persistent high deforestation rate and fragmentation of the Amazon forests are the main threats to their biodiversity. To anticipate and mitigate these threats, it is important to understand and predict how species respond to the rapidly changing landscape. The short-eared dog Atelocynus microtis is the only Amazon-endemic canid and one of the most understudied wild dogs worldwide. We investigated short-eared dog habitat associations on two spatial scales. First, we used the largest record database ever compiled for short-eared dogs in combination with species distribution models to map species habitat suitability, estimate its distribution range and predict shifts in species distribution in response to predicted deforestation across the entire Amazon (regional scale). Second, we used systematic camera trap surveys and occupancy models to investigate how forest cover and forest fragmentation affect the space use of this species in the Southern Brazilian Amazon (local scale). Species distribution models suggested that the short-eared dog potentially occurs over an extensive and continuous area, through most of the Amazon region south of the Amazon River. However, approximately 30% of the short-eared dog's current distribution is expected to be lost or suffer sharp declines in habitat suitability by 2027 (within three generations) due to forest loss. This proportion might reach 40% of the species distribution in unprotected areas and exceed 60% in some interfluves (i.e. portions of land separated by large rivers) of the Amazon basin. Our local-scale analysis indicated that the presence of forest positively affected short-eared dog space use, while the density of forest edges had a negative effect. Beyond shedding light on the ecology of the short-eared dog and refining its distribution range, our results stress that forest loss poses a serious threat to the conservation of the species in a short time frame. Hence, we propose a re-assessment of the short-eared dog's current IUCN Red List status (Near Threatened) based on findings presented here. Our study exemplifies how data can be integrated across sources and modelling procedures to improve our knowledge of relatively understudied species.
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Globally, conversion of pristine areas to anthropogenic landscapes is one of the main causes of ecosystem service losses. Land uses associated with urbanization and farming can be major sources of pollution to freshwaters promoting artificial inputs of several elements, leading to impaired water quality. However, how the effects of land use on freshwater quality are contingent on properties of the local landscape and climate is still poorly understood. The aim of this study was to evaluate the effects of landscape properties (morphometric measurements of lakes and their catchments), precipitation patterns, and land use properties (extent and proximity of the land use to freshwaters) on water quality of 98 natural lakes and reservoirs in northeast Brazil. Water quality impairment (WQI) was expressed as a composite variable incorporating parameters correlated with eutrophication including nitrogen (N), phosphorus (P) and Chlorophyll-a concentration. Regression tree analysis showed that WQI is mainly related to highly impacted "buffer areas". However, the effects of land use in these adjacent lands were contingent on precipitation variability for 13% of waterbodies and on surface area of the buffer in relation to the volume of waterbody (BA:Vol) for 87% of waterbodies. Overall, effects on WQI originating from the land use in the adjacent portion of the lake were amplified by high precipitation variability for ecosystems with highly impacted buffer areas and by high BA:Vol for ecosystems with less impacted buffer areas, indicating that ecosystems subjected to intense episodic rainfall events (e.g. storms) and higher buffer areas relative to aquatic ecosystem size (i.e. small waterbodies) are more susceptible to impacts of land use. Land use at the catchment scale was important for the largest ecosystems. Thus, our findings point toward the need for considering a holistic approach to managing water quality, which includes watershed management within the context of climate change.
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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.
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The role of tropical lakes and reservoirs in the global carbon cycle has received increasing attention in the past decade, but our understanding of its variability is still limited. The metabolism of tropical systems may differ profoundly from temperate systems due to the higher temperatures and wider variations in precipitation. Here, we investigated the spatial and temporal patterns of the variability in the partial pressure of carbon dioxide (pCO2) and its drivers in a set of 102 low-latitude lakes and reservoirs that encompass wide gradients of precipitation, productivity and landscape properties (lake area, perimeter-to-area ratio, catchment size, catchment area-to-lake area ratio, and types of catchment land use). We used multiple regressions and structural equation modeling (SEM) to determine the direct and indirect effects of the main in-lake variables and landscape properties on the water pCO2 variance. We found that these systems were mostly supersaturated with CO2 (92% spatially and 72% seasonally) regardless of their trophic status and landscape properties. The pCO2 values (9-40,020⯵atm) were within the range found in tropical ecosystems, and higher (pâ¯<â¯0.005) than pCO2 values recorded from high-latitude ecosystems. Water volume had a negative effect on the trophic state (râ¯=â¯-0.63), which mediated a positive indirect effect on pCO2 (râ¯=â¯0.4), representing an important negative feedback in the context of climate change-driven reduction in precipitation. Our results demonstrated that precipitation drives the pCO2 seasonal variability, with significantly higher pCO2 during the rainy season (Fâ¯=â¯16.67; pâ¯<â¯0.001), due to two potential main mechanisms: (1) phytoplankton dilution and (2) increasing inputs of terrestrial CO2 from the catchment. We conclude that at low latitudes, precipitation is a major climatic driver of pCO2 variability by influencing volume variations and linking lentic ecosystems to their catchments.
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We determined the filtered tree species pool of Amazonian wetland forests, based on confirmed occurrence records, to better understand how tree diversity in wetland environments compares to tree diversity in the entire Amazon region. The tree species pool was determined using data from two main sources: 1) a compilation of published tree species lists plus one unpublished list of our own, derived from tree plot inventories and floristic surveys; 2) queries on botanical collections that include Amazonian flora, curated by herbaria and available through the SpeciesLink digital biodiversity database. We applied taxonomic name resolution and determined sample-based species accumulation curves for both datasets, to estimate sampling effort and predict the expected species richness using Chao's analytical estimators. We report a total of 3 615 valid tree species occurring in Amazonian wetland forests. After surveying almost 70 years of research efforts to inventory the diversity of Amazonian wetland trees, we found that 74% these records were registered in published species lists (2 688 tree species). Tree species richness estimates predicted from either single dataset underestimated the total pooled species richness recorded as occurring in Amazonian wetlands, with only 41% of the species shared by both datasets. The filtered tree species pool of Amazonian wetland forests comprises 53% of the 6 727 tree species taxonomically confirmed for the Amazonian tree flora to date. This large proportion is likely to be the result of significant species interchange among forest habitats within the Amazon region, as well as in situ speciation processes due to strong ecological filtering. The provided tree species pool raises the number of tree species previously reported as occurring in Amazonian wetlands by a factor of 3.2.
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Bosques , Árboles/clasificación , Humedales , Evolución Biológica , Análisis de SupervivenciaRESUMEN
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.