How deregulation, drought and increasing fire impact Amazonian biodiversity.

Biodiversity contributes to the ecological and climatic stability of the Amazon Basin1,2, but is increasingly threatened by deforestation and fire3,4. Here we quantify these impacts over the past two decades using remote-sensing estimates of fire and deforestation and comprehensive range estimates of 11,514 plant species and 3,079 vertebrate species in the Amazon. Deforestation has led to large amounts of habitat loss, and fires further exacerbate this already substantial impact on Amazonian biodiversity. Since 2001, 103,079-189,755 km2 of Amazon rainforest has been impacted by fires, potentially impacting the ranges of 77.3-85.2% of species that are listed as threatened in this region5. The impacts of fire on the ranges of species in Amazonia could be as high as 64%, and greater impacts are typically associated with species that have restricted ranges. We find close associations between forest policy, fire-impacted forest area and their potential impacts on biodiversity. In Brazil, forest policies that were initiated in the mid-2000s corresponded to reduced rates of burning. However, relaxed enforcement of these policies in 2019 has seemingly begun to reverse this trend: approximately 4,253-10,343 km2 of forest has been impacted by fire, leading to some of the most severe potential impacts on biodiversity since 2009. These results highlight the critical role of policy enforcement in the preservation of biodiversity in the Amazon.

Cerrado deforestation threatens regional climate and water availability for agriculture and ecosystems.

The Brazilian Cerrado is one of the most biodiverse savannas in the world, yet 46% of its original cover has been cleared to make way for crops and pastures. These extensive land-use transitions (LUTs) are expected to influence regional climate by reducing evapotranspiration (ET), increasing land surface temperature (LST), and ultimately reducing precipitation. Here, we quantify the impacts of LUTs on ET and LST in the Cerrado by combining MODIS satellite data with annual land use and land cover maps from 2006 to 2019. We performed regression analyses to quantify the effects of six common LUTs on ET and LST across the entire gradient of Cerrado landscapes. Results indicate that clearing forests for cropland or pasture increased average LST by ~3.5°C and reduced mean annual ET by 44% and 39%, respectively. Transitions from woody savannas to cropland or pasture increased average LST by 1.9°C and reduced mean annual ET by 27% and 21%, respectively. Converting native grasslands to cropland or pasture increased average LST by 0.9 and 0.6°C, respectively. Conversely, grassland-to-pasture transitions increased mean annual ET by 15%. To date, land changes have caused a 10% reduction in water recycled to the atmosphere annually and a 0.9°C increase in average LST across the biome, compared to the historic baseline under native vegetation. Global climate changes from increased atmospheric greenhouse gas concentrations will only exacerbate these effects. Considering potential future scenarios, we found that abandoning deforestation control policies or allowing legal deforestation to continue (at least 28.4 Mha) would further reduce yearly ET (by -9% and -3%, respectively) and increase average LST (by +0.7 and +0.3°C, respectively) by 2050. In contrast, policies encouraging zero deforestation and restoration of the 5.2 Mha of illegally deforested areas would partially offset the warming and drying impacts of land-use change.

O Cerrado brasileiro é uma das savanas mais biodiversas do mundo. Apesar disso, 46% da sua cobertura original foi desmatada para dar lugar a cultivos agrícolas e pastos. Estas extensas transições de uso do solo (LUT) têm o potencial de influenciar o clima regional, reduzindo a evapotranspiração (ET), aumentando a temperatura da superfície terrestre (LST) e por fim reduzindo a precipitação. O objetivo deste estudo foi quantificar os impactos de LUTs sobre ET e LST no Cerrado, combinando dados do satélite MODIS com mapas anuais de uso e cobertura do solo de 2006-2019. Foram realizadas análises de regressão para quantificar os efeitos de seis LUTs usuais sobre ET e LST, ao longo de todo o gradiente de paisagens do Cerrado. Os resultados indicaram que a retirada de florestas para dar lugar à agricultura ou pastagem aumentou a LST média em ~3.5°C e reduziu a ET média anual em 44% e 39%, respectivamente. Transições de formações savânicas para agricultura ou pastagem aumentaram a LST média em 1.9°C e reduziram a ET média anual em 27% e 21%, respectivamente. A conversão de campos nativos para agricultura ou pastagem aumentou a LST média em 0.9 e 0.6°C, respectivamente. Em contrapartida, transições de formações campestres nativas para pastagens aumentaram a ET média anual em 15%. Até o momento, as mudanças de uso do solo causaram redução de 10% da água reciclada para a atmosfera anualmente e aumento de 0.9°C da LST média ao longo do bioma, em comparação com a linha de base histórica sob vegetação nativa. As mudanças climáticas globais decorrentes do aumento das concentrações atmosféricas de gases do efeito estufa irão exacerbar esses efeitos. Considerando potenciais cenários futuros, observou-se que o abandono das políticas de controle do desmatamento ou o avanço do desmatamento legal (ao menos 28.4 Mha) reduziriam a ET anual (em −9% e −3%, respectivamente) e aumentariam a LST média (em +0.7 e +0.3ºC, respectivamente) até 2050. Por outro lado, políticas que promovam desmatamento zero e restauração dos 5.2 Mha de áreas ilegalmente desmatadas compensariam parte dos impactos de aquecimento e seca causados por alterações de uso do solo.

Starch and lipid storage strategies in tropical trees relate to growth and mortality.

Non-structural carbon (NSC) storage (i.e. starch, soluble sugras and lipids) in tree stems play important roles in metabolism and growth. Their spatial distribution in wood may explain species-specific differences in carbon storage dynamics, growth and survival. However, quantitative information on the spatial distribution of starch and lipids in wood is sparse due to methodological limitations. Here we assessed differences in wood NSC and lipid storage between tropical tree species with different growth and mortality rates and contrasting functional types. We measured starch and soluble sugars in wood cores up to 4 cm deep into the stem using standard chemical quantification methods and histological slices stained with Lugol's iodine. We also detected neutral lipids using histological slices stained with Oil-Red-O. The histological method allowed us to group individuals into two categories according to their starch storage strategy: fiber-storing trees and parenchyma-storing trees. The first group had a bigger starch pool, slower growth and lower mortality rates than the second group. Lipid storage was found in wood parenchyma in five species and was related to low mortality rates. The quantification of the spatial distribution of starch and lipids in wood improves our understanding of NSC dynamics in trees and reveals additional dimensions of tree growth and survival strategies.

Burning in southwestern Brazilian Amazonia, 2016-2019.

Fire is one of the most powerful modifiers of the Amazonian landscape and knowledge about its drivers is needed for planning control and suppression. A plethora of factors may play a role in the annual dynamics of fire frequency, spanning the biophysical, climatic, socioeconomic and institutional dimensions. To uncover the main forces currently at play, we investigated the area burned in both forested and deforested areas in the outstanding case of Brazil's state of Acre, in southwestern Amazonia. We mapped burn scars in already-deforested areas and intact forest based on satellite images from the Landsat series analyzed between 2016 and 2019. The mapped burnings in already-deforested areas totalled 550,251 ha. In addition, we mapped three forest fires totaling 34,084 ha. Fire and deforestation were highly correlated, and the latter occurred mainly in federal government lands, with protected areas showing unprecedented forest fire levels in 2019. These results indicate that Acre state is under increased fire risk even during average rainfall years. The record fires of 2019 may continue if Brazil's ongoing softening of environmental regulations and enforcement is maintained. Acre and other Amazonian states must act quickly to avoid an upsurge of social and economic losses in the coming years.

Thinner bark increases sensitivity of wetter Amazonian tropical forests to fire.

Understory fires represent an accelerating threat to Amazonian tropical forests and can, during drought, affect larger areas than deforestation itself. These fires kill trees at rates varying from < 10 to c. 90% depending on fire intensity, forest disturbance history and tree functional traits. Here, we examine variation in bark thickness across the Amazon. Bark can protect trees from fires, but it is often assumed to be consistently thin across tropical forests. Here, we show that investment in bark varies, with thicker bark in dry forests and thinner in wetter forests. We also show that thinner bark translated into higher fire-driven tree mortality in wetter forests, with between 0.67 and 5.86 gigatonnes CO2 lost in Amazon understory fires between 2001 and 2010. Trait-enabled global vegetation models that explicitly include variation in bark thickness are likely to improve the predictions of fire effects on carbon cycling in tropical forests.

En los bosques tropicales de la Amazonia, los incendios de sotobosque representan una amenaza que se está acelerando. Durante la sequía, pueden afectar un área mayor que la deforestación misma. Estos incendios pueden matan árboles a tasas que varían desde <10 hasta cerca de 90% dependiendo de la intensidad del fuego, la historia de perturbaciones forestales y los rasgos funcionales de los árboles. En este estudio, examinamos la variación en el grosor de la corteza en la Amazonía. La corteza puede proteger los árboles de los incendios, pero normalmente se supone que es uniformemente delgada en los bosques tropicales. Aquí, mostramos que el grosor de la corteza varía bastante, con una corteza más gruesa en los bosques secos y más delgada en los bosques húmedos. También, mostramos que cortezas más delgadas resultan en tasas de mortalidad más altas en bosques más húmedos. En total, estimamos que los incendios en el sotobosque de la Amazonía han añadido entre 0,67 y 5,86 gigatoneladas de CO2 atmosférico entre 2001-2010. Los modelos globales de vegetación que predicen los efectos de los incendios sobre el reciclaje de carbono en los bosques tropicales deberían incluir explícitamente la variación en el grosor de la corteza.

Os incêndios rasteiros de sub-bosque representam uma ameaça cada vez maior às florestas tropicais da Amazônia. Durante secas, eles podem afetar áreas maiores do que àquelas desmatadas. Esses incêndios matam árvores a taxas que variam de <10 a c. 90%, dependendo da intensidade do fogo, da história de distúrbios florestais e das características funcionais das árvores. Neste estudo, examinamos a variação na espessura da casca na Amazônia. A casca pode proteger árvores do fogo, mas geralmente é considerada uniformemente fina para diversas florestas tropicais. Aqui, mostramos que a espessura da casca varia, com cascas mais espessas ocorrendo em florestas secas e mais finas ocorrendo em florestas mais úmidas. Mostramos também que a casca mais fina resulta em taxas de mortalidade mais altas em florestas úmidas. No total, estimamos que os incêndios de sub-bosque adicionaram entre 0,67 e 5,86 gigatoneladas de CO2 atmosférico entre 2001-2010. Os modelos globais de vegetação devem incluir explicitamente a variação na espessura da casca ao prever os efeitos do fogo no ciclo do carbono de florestas tropicais.

Prolonged tropical forest degradation due to compounding disturbances: Implications for CO2 and H2 O fluxes.

Drought, fire, and windstorms can interact to degrade tropical forests and the ecosystem services they provide, but how these forests recover after catastrophic disturbance events remains relatively unknown. Here, we analyze multi-year measurements of vegetation dynamics and function (fluxes of CO2 and H2 O) in forests recovering from 7 years of controlled burns, followed by wind disturbance. Located in southeast Amazonia, the experimental forest consists of three 50-ha plots burned annually, triennially, or not at all from 2004 to 2010. During the subsequent 6-year recovery period, postfire tree survivorship and biomass sharply declined, with aboveground C stocks decreasing by 70%-94% along forest edges (0-200 m into the forest) and 36%-40% in the forest interior. Vegetation regrowth in the forest understory triggered partial canopy closure (70%-80%) from 2010 to 2015. The composition and spatial distribution of grasses invading degraded forest evolved rapidly, likely because of the delayed mortality. Four years after the experimental fires ended (2014), the burned plots assimilated 36% less carbon than the Control, but net CO2 exchange and evapotranspiration (ET) had fully recovered 7 years after the experimental fires ended (2017). Carbon uptake recovery occurred largely in response to increased light-use efficiency and reduced postfire respiration, whereas increased water use associated with postfire growth of new recruits and remaining trees explained the recovery in ET. Although the effects of interacting disturbances (e.g., fires, forest fragmentation, and blowdown events) on mortality and biomass persist over many years, the rapid recovery of carbon and water fluxes can help stabilize local climate.

Drivers and mechanisms of tree mortality in moist tropical forests.

Tree mortality rates appear to be increasing in moist tropical forests (MTFs) with significant carbon cycle consequences. Here, we review the state of knowledge regarding MTF tree mortality, create a conceptual framework with testable hypotheses regarding the drivers, mechanisms and interactions that may underlie increasing MTF mortality rates, and identify the next steps for improved understanding and reduced prediction. Increasing mortality rates are associated with rising temperature and vapor pressure deficit, liana abundance, drought, wind events, fire and, possibly, CO2 fertilization-induced increases in stand thinning or acceleration of trees reaching larger, more vulnerable heights. The majority of these mortality drivers may kill trees in part through carbon starvation and hydraulic failure. The relative importance of each driver is unknown. High species diversity may buffer MTFs against large-scale mortality events, but recent and expected trends in mortality drivers give reason for concern regarding increasing mortality within MTFs. Models of tropical tree mortality are advancing the representation of hydraulics, carbon and demography, but require more empirical knowledge regarding the most common drivers and their subsequent mechanisms. We outline critical datasets and model developments required to test hypotheses regarding the underlying causes of increasing MTF mortality rates, and improve prediction of future mortality under climate change.

Impacts of fire on sources of soil CO2 efflux in a dry Amazon rain forest.

Fire at the dry southern margin of the Amazon rainforest could have major consequences for regional soil carbon (C) storage and ecosystem carbon dioxide (CO2 ) emissions, but relatively little information exists about impacts of fire on soil C cycling within this sensitive ecotone. We measured CO2 effluxes from different soil components (ground surface litter, roots, mycorrhizae, soil organic matter) at a large-scale burn experiment designed to simulate a severe but realistic potential future scenario for the region (Fire plot) in Mato Grosso, Brazil, over 1 year, and compared these measurements to replicated data from a nearby, unmodified Control plot. After four burns over 5 years, soil CO2 efflux (Rs ) was ~5.5 t C ha-1  year-1 lower on the Fire plot compared to the Control. Most of the Fire plot Rs reduction was specifically due to lower ground surface litter and root respiration. Mycorrhizal respiration on both plots was around ~20% of Rs . Soil surface temperature appeared to be more important than moisture as a driver of seasonal patterns in Rs at the site. Regular fire events decreased the seasonality of Rs at the study site, due to apparent differences in environmental sensitivities among biotic and abiotic soil components. These findings may contribute toward improved predictions of the amount and temporal pattern of C emissions across the large areas of tropical forest facing increasing fire disturbances associated with climate change and human activities.

Ecophysiological plasticity of Amazonian trees to long-term drought.

Episodic multi-year droughts fundamentally alter the dynamics, functioning, and structure of Amazonian forests. However, the capacity of individual plant species to withstand intense drought regimes remains unclear. Here, we evaluated ecophysiological responses from a forest community where we sampled 83 woody plant species during 5 years of experimental drought (throughfall exclusion) in an eastern Amazonian terra firme forest. Overall, the experimental drought resulted in shifts of some, but not all, leaf traits related to photosynthetic carbon uptake and intrinsic water-use efficiency. Leaf δ13C values increased by 2-3‰ within the canopy, consistent with increased diffusional constraints on photosynthesis. Decreased leaf C:N ratios were also observed, consistent with lower investments in leaf structure. However, no statistically significant treatment effects on leaf nitrogen content were observed, consistent with a lack of acclimation in photosynthetic capacity or increased production of nitrogen-based secondary metabolites. The results of our study provide evidence of robust acclimation potential to drought intensification in the diverse flora of an Amazonian forest community. The results reveals considerable ability of several species to respond to intense drought and challenge commonly held perspectives that this flora has attained limited adaptive plasticity because of a long evolutionary history in a favorable and stable climate.

Projections of future meteorological drought and wet periods in the Amazon.

Future intensification of Amazon drought resulting from climate change may cause increased fire activity, tree mortality, and emissions of carbon to the atmosphere across large areas of Amazonia. To provide a basis for addressing these issues, we examine properties of recent and future meteorological droughts in the Amazon in 35 climate models participating in the Coupled Model Intercomparison Project Phase 5 (CMIP5). We find that the CMIP5 climate models, as a group, simulate important properties of historical meteorological droughts in the Amazon. In addition, this group of models reproduces observed relationships between Amazon precipitation and regional sea surface temperature anomalies in the tropical Pacific and the North Atlantic oceans. Assuming the Representative Concentration Pathway 8.5 scenario for future drivers of climate change, the models project increases in the frequency and geographic extent of meteorological drought in the eastern Amazon, and the opposite in the West. For the region as a whole, the CMIP5 models suggest that the area affected by mild and severe meteorological drought will nearly double and triple, respectively, by 2100. Extremes of wetness are also projected to increase after 2040. Specifically, the frequency of periods of unusual wetness and the area affected by unusual wetness are projected to increase after 2040 in the Amazon as a whole, including in locations where annual mean precipitation is projected to decrease. Our analyses suggest that continued emissions of greenhouse gases will increase the likelihood of extreme events that have been shown to alter and degrade Amazonian forests.

Abrupt increases in Amazonian tree mortality due to drought-fire interactions.

Interactions between climate and land-use change may drive widespread degradation of Amazonian forests. High-intensity fires associated with extreme weather events could accelerate this degradation by abruptly increasing tree mortality, but this process remains poorly understood. Here we present, to our knowledge, the first field-based evidence of a tipping point in Amazon forests due to altered fire regimes. Based on results of a large-scale, long-term experiment with annual and triennial burn regimes (B1yr and B3yr, respectively) in the Amazon, we found abrupt increases in fire-induced tree mortality (226 and 462%) during a severe drought event, when fuel loads and air temperatures were substantially higher and relative humidity was lower than long-term averages. This threshold mortality response had a cascading effect, causing sharp declines in canopy cover (23 and 31%) and aboveground live biomass (12 and 30%) and favoring widespread invasion by flammable grasses across the forest edge area (80 and 63%), where fires were most intense (e.g., 220 and 820 kW ⋅ m(-1)). During the droughts of 2007 and 2010, regional forest fires burned 12 and 5% of southeastern Amazon forests, respectively, compared with <1% in nondrought years. These results show that a few extreme drought events, coupled with forest fragmentation and anthropogenic ignition sources, are already causing widespread fire-induced tree mortality and forest degradation across southeastern Amazon forests. Future projections of vegetation responses to climate change across drier portions of the Amazon require more than simulation of global climate forcing alone and must also include interactions of extreme weather events, fire, and land-use change.

Effects of experimental fuel additions on fire intensity and severity: unexpected carbon resilience of a neotropical forest.

Global changes and associated droughts, heat waves, logging activities, and forest fragmentation may intensify fires in Amazonia by altering forest microclimate and fuel dynamics. To isolate the effects of fuel loads on fire behavior and fire-induced changes in forest carbon cycling, we manipulated fine fuel loads in a fire experiment located in southeast Amazonia. We predicted that a 50% increase in fine fuel loads would disproportionally increase fire intensity and severity (i.e., tree mortality and losses in carbon stocks) due to multiplicative effects of fine fuel loads on the rate of fire spread, fuel consumption, and burned area. The experiment followed a fully replicated randomized block design (N = 6) comprised of unburned control plots and burned plots that were treated with and without fine fuel additions. The fuel addition treatment significantly increased burned area (+22%) and consequently canopy openness (+10%), fine fuel combustion (+5%), and mortality of individuals ≥5 cm in diameter at breast height (dbh; +37%). Surprisingly, we observed nonsignificant effects of the fuel addition treatment on fireline intensity, and no significant differences among the three treatments for (i) mortality of large trees (≥30 cm dbh), (ii) aboveground forest carbon stocks, and (iii) soil respiration. It was also surprising that postfire tree growth and wood increment were higher in the burned plots treated with fuels than in the unburned control. These results suggest that (i) fine fuel load accumulation increases the likelihood of larger understory fires and (ii) single, low-intensity fires weakly influence carbon cycling of this primary neotropical forest, although delayed postfire mortality of large trees may lower carbon stocks over the long term. Overall, our findings indicate that increased fine fuel loads alone are unlikely to create threshold conditions for high-intensity, catastrophic fires during nondrought years.

The linkages between photosynthesis, productivity, growth and biomass in lowland Amazonian forests.

Understanding the relationship between photosynthesis, net primary productivity and growth in forest ecosystems is key to understanding how these ecosystems will respond to global anthropogenic change, yet the linkages among these components are rarely explored in detail. We provide the first comprehensive description of the productivity, respiration and carbon allocation of contrasting lowland Amazonian forests spanning gradients in seasonal water deficit and soil fertility. Using the largest data set assembled to date, ten sites in three countries all studied with a standardized methodology, we find that (i) gross primary productivity (GPP) has a simple relationship with seasonal water deficit, but that (ii) site-to-site variations in GPP have little power in explaining site-to-site spatial variations in net primary productivity (NPP) or growth because of concomitant changes in carbon use efficiency (CUE), and conversely, the woody growth rate of a tropical forest is a very poor proxy for its productivity. Moreover, (iii) spatial patterns of biomass are much more driven by patterns of residence times (i.e. tree mortality rates) than by spatial variation in productivity or tree growth. Current theory and models of tropical forest carbon cycling under projected scenarios of global atmospheric change can benefit from advancing beyond a focus on GPP. By improving our understanding of poorly understood processes such as CUE, NPP allocation and biomass turnover times, we can provide more complete and mechanistic approaches to linking climate and tropical forest carbon cycling.

Threshold Responses to Soil Moisture Deficit by Trees and Soil in Tropical Rain Forests: Insights from Field Experiments.

Many tropical rain forest regions are at risk of increased future drought. The net effects of drought on forest ecosystem functioning will be substantial if important ecological thresholds are passed. However, understanding and predicting these effects is challenging using observational studies alone. Field-based rainfall exclusion (canopy throughfall exclusion; TFE) experiments can offer mechanistic insight into the response to extended or severe drought and can be used to help improve model-based simulations, which are currently inadequate. Only eight TFE experiments have been reported for tropical rain forests. We examine them, synthesizing key results and focusing on two processes that have shown threshold behavior in response to drought: (1) tree mortality and (2) the efflux of carbon dioxdie from soil, soil respiration. We show that: (a) where tested using large-scale field experiments, tropical rain forest tree mortality is resistant to long-term soil moisture deficit up to a threshold of 50% of the water that is extractable by vegetation from the soil, but high mortality occurs beyond this value, with evidence from one site of increased autotrophic respiration, and (b) soil respiration reaches its peak value in response to soil moisture at significantly higher soil moisture content for clay-rich soils than for clay-poor soils. This first synthesis of tropical TFE experiments offers the hypothesis that low soil moisture-related thresholds for key stress responses in soil and vegetation may prove to be widely applicable across tropical rain forests despite the diversity of these forests.

Landscape fragmentation, severe drought, and the new Amazon forest fire regime.

Changes in weather and land use are transforming the spatial and temporal characteristics of fire regimes in Amazonia, with important effects on the functioning of dense (i.e., closed-canopy), open-canopy, and transitional forests across the Basin. To quantify, document, and describe the characteristics and recent changes in forest fire regimes, we sampled 6 million ha of these three representative forests of the eastern and southern edges of the Amazon using 24 years (1983-2007) of satellite-derived annual forest fire scar maps and 16 years of monthly hot pixel information (1992-2007). Our results reveal that changes in forest fire regime properties differentially affected these three forest types in terms of area burned and fire scar size, frequency, and seasonality. During the study period, forest fires burned 15% (0.3 million ha), 44% (1 million ha), and 46% (0.6 million ha) of dense, open, and transitional forests, respectively. Total forest area burned and fire scar size tended to increase over time (even in years of average rainfall in open canopy and transitional forests). In dense forests, most of the temporal variability in fire regime properties was linked to El Nino Southern Oscillation (ENSO)-related droughts. Compared with dense forests, transitional and open forests experienced fires twice as frequently, with at least 20% of these forests' areas burning two or more times during the 24-year study period. Open and transitional forests also experienced higher deforestation rates than dense forests. During drier years, the end of the dry season was delayed by about a month, which resulted in larger burn scars and increases in overall area burned later in the season. These observations suggest that climate-mediated forest flammability is enhanced by landscape fragmentation caused by deforestation, as observed for open and transitional forests in the Eastern portion of the Amazon Basin.

Structure and composition of altered riparian forests in an agricultural Amazonian landscape.

Deforestation and fragmentation influence the microclimate, vegetation structure, and composition of remaining patches of tropical forest. In the southern Amazon, at the frontier of cropland expansion, forests are converted and fragmented in a pattern that leaves standing riparian forests whose dimensions are mandated by the Brazilian National Forest Code. These altered riparian forests share many characteristics of well-studied upland forest fragments, but differ because they remain connected to larger areas of forest downstream, and because they may experience wetter soil conditions because reduction of forest cover in the surrounding watershed raises groundwater levels and increases stream runoff. We compared forest regeneration, structure, composition, and diversity in four areas of intact riparian forest and four areas each of narrow, medium, and wide altered riparian forests that have been surrounded by agriculture since the early 1980s. We found that seedling abundance was reduced by as much as 64% and sapling abundance was reduced by as much as 67% in altered compared to intact riparian forests. The most pronounced differences between altered and intact forest occurred near forest edges and within the narrowest sections of altered riparian forests. Woody plant species composition differed and diversity was reduced in altered forests compared to intact riparian forests. However, despite being fragmented for several decades, large woody plant biomass and carbon storage, the number of live or dead large woody plants, mortality rates, and the size distribution of woody plants did not differ significantly between altered and intact riparian forests. Thus, even in these relatively narrow forests with high edge: area ratios, we saw no evidence of the increases in mortality and declines in biomass that have been found in other tropical forest fragment studies. However, because of the changes in both species community and reduced regeneration, it is unclear how long this relative lack of change will be sustained. Additionally, Brazil recently passed a law in their National Forest Code allowing narrower riparian buffers than those studied here in restored areas, which could affect their long-term sustainability.

Confronting model predictions of carbon fluxes with measurements of Amazon forests subjected to experimental drought.

Considerable uncertainty surrounds the fate of Amazon rainforests in response to climate change. Here, carbon (C) flux predictions of five terrestrial biosphere models (Community Land Model version 3.5 (CLM3.5), Ecosystem Demography model version 2.1 (ED2), Integrated BIosphere Simulator version 2.6.4 (IBIS), Joint UK Land Environment Simulator version 2.1 (JULES) and Simple Biosphere model version 3 (SiB3)) and a hydrodynamic terrestrial ecosystem model (the Soil-Plant-Atmosphere (SPA) model) were evaluated against measurements from two large-scale Amazon drought experiments. Model predictions agreed with the observed C fluxes in the control plots of both experiments, but poorly replicated the responses to the drought treatments. Most notably, with the exception of ED2, the models predicted negligible reductions in aboveground biomass in response to the drought treatments, which was in contrast to an observed c. 20% reduction at both sites. For ED2, the timing of the decline in aboveground biomass was accurate, but the magnitude was too high for one site and too low for the other. Three key findings indicate critical areas for future research and model development. First, the models predicted declines in autotrophic respiration under prolonged drought in contrast to measured increases at one of the sites. Secondly, models lacking a phenological response to drought introduced bias in the sensitivity of canopy productivity and respiration to drought. Thirdly, the phenomenological water-stress functions used by the terrestrial biosphere models to represent the effects of soil moisture on stomatal conductance yielded unrealistic diurnal and seasonal responses to drought.

Interactions between repeated fire, nutrients, and insect herbivores affect the recovery of diversity in the southern Amazon.

Surface fires burn extensive areas of tropical forests each year, altering resource availability, biotic interactions, and, ultimately, plant diversity. In transitional forest between the Brazilian cerrado (savanna) and high stature Amazon forest, we took advantage of a long-term fire experiment to establish a factorial study of the interactions between fire, nutrient availability, and herbivory on early plant regeneration. Overall, five annual burns reduced the number and diversity of regenerating stems. Community composition changed substantially after repeated fires, and species common in the cerrado became more abundant. The number of recruits and their diversity were reduced in the burned area, but burned plots closed to herbivores with nitrogen additions had a 14 % increase in recruitment. Diversity of recruits also increased up to 50 % in burned plots when nitrogen was added. Phosphorus additions were related to an increase in species evenness in burned plots open to herbivores. Herbivory reduced seedling survival overall and increased diversity in burned plots when nutrients were added. This last result supports our hypothesis that positive relationships between herbivore presence and diversity would be strongest in treatments that favor herbivory--in this case herbivory was higher in burned plots which were initially lower in diversity. Regenerating seedlings in less diverse plots were likely more apparent to herbivores, enabling increased herbivory and a stronger signal of negative density dependence. In contrast, herbivores generally decreased diversity in more species rich unburned plots. Although this study documents complex interactions between repeated burns, nutrients, and herbivory, it is clear that fire initiates a shift in the factors that are most important in determining the diversity and number of recruits. This change may have long-lasting effects as the forest progresses through succession.

Seasonal and interannual variability of climate and vegetation indices across the Amazon.

Drought exerts a strong influence on tropical forest metabolism, carbon stocks, and ultimately the flux of carbon to the atmosphere. Satellite-based studies have suggested that Amazon forests green up during droughts because of increased sunlight, whereas field studies have reported increased tree mortality during severe droughts. In an effort to reconcile these apparently conflicting findings, we conducted an analysis of climate data, field measurements, and improved satellite-based measures of forest photosynthetic activity. Wet-season precipitation and plant-available water (PAW) decreased over the Amazon Basin from 1996-2005, and photosynthetically active radiation (PAR) and air dryness (expressed as vapor pressure deficit, VPD) increased from 2002-2005. Using improved enhanced vegetation index (EVI) measurements (2000-2008), we show that gross primary productivity (expressed as EVI) declined with VPD and PAW in regions of sparse canopy cover across a wide range of environments for each year of the study. In densely forested areas, no climatic variable adequately explained the Basin-wide interannual variability of EVI. Based on a site-specific study, we show that monthly EVI was relatively insensitive to leaf area index (LAI) but correlated positively with leaf flushing and PAR measured in the field. These findings suggest that production of new leaves, even when unaccompanied by associated changes in LAI, could play an important role in Basin-wide interannual EVI variability. Because EVI variability was greatest in regions of lower PAW, we hypothesize that drought could increase EVI by synchronizing leaf flushing via its effects on leaf bud development.

Intraspecific trait variability facilitates tree species persistence along riparian forest edges in Southern Amazonia.

Tropical forest fragmentation from agricultural expansion alters the microclimatic conditions of the remaining forests, with effects on vegetation structure and function. However, little is known about how the functional trait variability within and among tree species in fragmented landscapes influence and facilitate species' persistence in these new environmental conditions. Here, we assessed potential changes in tree species' functional traits in riparian forests within six riparian forests in cropland catchments (Cropland) and four riparian forests in forested catchments (Forest) in southern Amazonia. We sampled 12 common functional traits of 123 species across all sites: 64 common to both croplands and forests, 33 restricted to croplands, and 26 restricted to forests. We found that forest-restricted species had leaves that were thinner, larger, and with higher phosphorus (P) content, compared to cropland-restricted ones. Tree species common to both environments showed higher intraspecific variability in functional traits, with leaf thickness and leaf P concentration varying the most. Species turnover contributed more to differences between forest and cropland environments only for the stem-specific density trait. We conclude that the intraspecific variability of functional traits (leaf thickness, leaf P, and specific leaf area) facilitates species persistence in riparian forests occurring within catchments cleared for agricultural expansion in Amazonia.