The carbon sink of secondary and degraded humid tropical forests.

The globally important carbon sink of intact, old-growth tropical humid forests is declining because of climate change, deforestation and degradation from fire and logging1-3. Recovering tropical secondary and degraded forests now cover about 10% of the tropical forest area4, but how much carbon they accumulate remains uncertain. Here we quantify the aboveground carbon (AGC) sink of recovering forests across three main continuous tropical humid regions: the Amazon, Borneo and Central Africa5,6. On the basis of satellite data products4,7, our analysis encompasses the heterogeneous spatial and temporal patterns of growth in degraded and secondary forests, influenced by key environmental and anthropogenic drivers. In the first 20 years of recovery, regrowth rates in Borneo were up to 45% and 58% higher than in Central Africa and the Amazon, respectively. This is due to variables such as temperature, water deficit and disturbance regimes. We find that regrowing degraded and secondary forests accumulated 107 Tg C year-1 (90-130 Tg C year-1) between 1984 and 2018, counterbalancing 26% (21-34%) of carbon emissions from humid tropical forest loss during the same period. Protecting old-growth forests is therefore a priority. Furthermore, we estimate that conserving recovering degraded and secondary forests can have a feasible future carbon sink potential of 53 Tg C year-1 (44-62 Tg C year-1) across the main tropical regions studied.

Increased Amazon carbon emissions mainly from decline in law enforcement.

The Amazon forest carbon sink is declining, mainly as a result of land-use and climate change1-4. Here we investigate how changes in law enforcement of environmental protection policies may have affected the Amazonian carbon balance between 2010 and 2018 compared with 2019 and 2020, based on atmospheric CO2 vertical profiles5,6, deforestation7 and fire data8, as well as infraction notices related to illegal deforestation9. We estimate that Amazonia carbon emissions increased from a mean of 0.24 ± 0.08 PgC year-1 in 2010-2018 to 0.44 ± 0.10 PgC year-1 in 2019 and 0.52 ± 0.10 PgC year-1 in 2020 (± uncertainty). The observed increases in deforestation were 82% and 77% (94% accuracy) and burned area were 14% and 42% in 2019 and 2020 compared with the 2010-2018 mean, respectively. We find that the numbers of notifications of infractions against flora decreased by 30% and 54% and fines paid by 74% and 89% in 2019 and 2020, respectively. Carbon losses during 2019-2020 were comparable with those of the record warm El Niño (2015-2016) without an extreme drought event. Statistical tests show that the observed differences between the 2010-2018 mean and 2019-2020 are unlikely to have arisen by chance. The changes in the carbon budget of Amazonia during 2019-2020 were mainly because of western Amazonia becoming a carbon source. Our results indicate that a decline in law enforcement led to increases in deforestation, biomass burning and forest degradation, which increased carbon emissions and enhanced drying and warming of the Amazon forests.

Direct evidence for phosphorus limitation on Amazon forest productivity.

The productivity of rainforests growing on highly weathered tropical soils is expected to be limited by phosphorus availability1. Yet, controlled fertilization experiments have been unable to demonstrate a dominant role for phosphorus in controlling tropical forest net primary productivity. Recent syntheses have demonstrated that responses to nitrogen addition are as large as to phosphorus2, and adaptations to low phosphorus availability appear to enable net primary productivity to be maintained across major soil phosphorus gradients3. Thus, the extent to which phosphorus availability limits tropical forest productivity is highly uncertain. The majority of the Amazonia, however, is characterized by soils that are more depleted in phosphorus than those in which most tropical fertilization experiments have taken place2. Thus, we established a phosphorus, nitrogen and base cation addition experiment in an old growth Amazon rainforest, with a low soil phosphorus content that is representative of approximately 60% of the Amazon basin. Here we show that net primary productivity increased exclusively with phosphorus addition. After 2 years, strong responses were observed in fine root (+29%) and canopy productivity (+19%), but not stem growth. The direct evidence of phosphorus limitation of net primary productivity suggests that phosphorus availability may restrict Amazon forest responses to CO2 fertilization4, with major implications for future carbon sequestration and forest resilience to climate change.

Amazonia as a carbon source linked to deforestation and climate change.

Amazonia hosts the Earth's largest tropical forests and has been shown to be an important carbon sink over recent decades1-3. This carbon sink seems to be in decline, however, as a result of factors such as deforestation and climate change1-3. Here we investigate Amazonia's carbon budget and the main drivers responsible for its change into a carbon source. We performed 590 aircraft vertical profiling measurements of lower-tropospheric concentrations of carbon dioxide and carbon monoxide at four sites in Amazonia from 2010 to 20184. We find that total carbon emissions are greater in eastern Amazonia than in the western part, mostly as a result of spatial differences in carbon-monoxide-derived fire emissions. Southeastern Amazonia, in particular, acts as a net carbon source (total carbon flux minus fire emissions) to the atmosphere. Over the past 40 years, eastern Amazonia has been subjected to more deforestation, warming and moisture stress than the western part, especially during the dry season, with the southeast experiencing the strongest trends5-9. We explore the effect of climate change and deforestation trends on carbon emissions at our study sites, and find that the intensification of the dry season and an increase in deforestation seem to promote ecosystem stress, increase in fire occurrence, and higher carbon emissions in the eastern Amazon. This is in line with recent studies that indicate an increase in tree mortality and a reduction in photosynthesis as a result of climatic changes across Amazonia1,10.

Linking land-use and land-cover transitions to their ecological impact in the Amazon.

Human activities pose a major threat to tropical forest biodiversity and ecosystem services. Although the impacts of deforestation are well studied, multiple land-use and land-cover transitions (LULCTs) occur in tropical landscapes, and we do not know how LULCTs differ in their rates or impacts on key ecosystem components. Here, we quantified the impacts of 18 LULCTs on three ecosystem components (biodiversity, carbon, and soil), based on 18 variables collected from 310 sites in the Brazilian Amazon. Across all LULCTs, biodiversity was the most affected ecosystem component, followed by carbon stocks, but the magnitude of change differed widely among LULCTs and individual variables. Forest clearance for pasture was the most prevalent and high-impact transition, but we also identified other LULCTs with high impact but lower prevalence (e.g., forest to agriculture). Our study demonstrates the importance of considering multiple ecosystem components and LULCTs to understand the consequences of human activities in tropical landscapes.

Large-scale commodity agriculture exacerbates the climatic impacts of Amazonian deforestation.

In the Amazon rainforest, land use following deforestation is diverse and dynamic. Mounting evidence indicates that the climatic impacts of forest loss can also vary considerably, depending on specific features of the affected areas. The size of the deforested patches, for instance, was shown to modulate the characteristics of local climatic impacts. Nonetheless, the influence of different types of land use and management strategies on the magnitude of local climatic changes remains uncertain. Here, we evaluated the impacts of large-scale commodity farming and rural settlements on surface temperature, rainfall patterns, and energy fluxes. Our results reveal that changes in land-atmosphere coupling are induced not only by deforestation size but also, by land use type and management patterns inside the deforested areas. We provide evidence that, in comparison with rural settlements, deforestation caused by large-scale commodity agriculture is more likely to reduce convective rainfall and increase land surface temperature. We demonstrate that these differences are mainly caused by a more intensive management of the land, resulting in significantly lower vegetation cover throughout the year, which reduces latent heat flux. Our findings indicate an urgent need for alternative agricultural practices, as well as forest restoration, for maintaining ecosystem processes and mitigating change in the local climates across the Amazon basin.

Tracking the impacts of El Niño drought and fire in human-modified Amazonian forests.

With humanity facing an unprecedented climate crisis, the conservation of tropical forests has never been so important - their vast terrestrial carbon stocks can be turned into emissions by climatic and human disturbances. However, the duration of these effects is poorly understood, and it is unclear whether impacts are amplified in forests with a history of previous human disturbance. Here, we focus on the Amazonian epicenter of the 2015-16 El Niño, a region that encompasses 1.2% of the Brazilian Amazon. We quantify, at high temporal resolution, the impacts of an extreme El Niño (EN) drought and extensive forest fires on plant mortality and carbon loss in undisturbed and human-modified forests. Mortality remained higher than pre-El Niño levels for 36 mo in EN-drought-affected forests and for 30 mo in EN-fire-affected forests. In EN-fire-affected forests, human disturbance significantly increased plant mortality. Our investigation of the ecological and physiological predictors of tree mortality showed that trees with lower wood density, bark thickness and leaf nitrogen content, as well as those that experienced greater fire intensity, were more vulnerable. Across the region, the 2015-16 El Niño led to the death of an estimated 2.5 ± 0.3 billion stems, resulting in emissions of 495 ± 94 Tg CO2 Three years after the El Niño, plant growth and recruitment had offset only 37% of emissions. Our results show that limiting forest disturbance will not only help maintain carbon stocks, but will also maximize the resistance of Amazonian forests if fires do occur.

Declining Amazon biomass due to deforestation and subsequent degradation losses exceeding gains.

In the Amazon, deforestation and climate change lead to increased vulnerability to forest degradation, threatening its existing carbon stocks and its capacity as a carbon sink. We use satellite L-Band Vegetation Optical Depth (L-VOD) data that provide an integrated (top-down) estimate of biomass carbon to track changes over 2011-2019. Because the spatial resolution of L-VOD is coarse (0.25°), it allows limited attribution of the observed changes. We therefore combined high-resolution annual maps of forest cover and disturbances with biomass maps to model carbon losses (bottom-up) from deforestation and degradation, and gains from regrowing secondary forests. We show an increase of deforestation and associated degradation losses since 2012 which greatly outweigh secondary forest gains. Degradation accounted for 40% of gross losses. After an increase in 2011, old-growth forests show a net loss of above-ground carbon between 2012 and 2019. The sum of component carbon fluxes in our model is consistent with the total biomass change from L-VOD of 1.3 Pg C over 2012-2019. Across nine Amazon countries, we found that while Brazil contains the majority of biomass stocks (64%), its losses from disturbances were disproportionately high (79% of gross losses). Our multi-source analysis provides a pessimistic assessment of the Amazon carbon balance and highlights the urgent need to stop the recent rise of deforestation and degradation, particularly in the Brazilian Amazon.

Potential aboveground biomass increase in Brazilian Atlantic Forest fragments with climate change.

Fragmented tropical forest landscapes preserve much of the remaining biodiversity and carbon stocks. Climate change is expected to intensify droughts and increase fire hazard and fire intensities, thereby causing habitat deterioration, and losses of biodiversity and carbon stock losses. Understanding the trajectories that these landscapes may follow under increased climate pressure is imperative for establishing strategies for conservation of biodiversity and ecosystem services. Here, we used a quantitative predictive modelling approach to project the spatial distribution of the aboveground biomass density (AGB) by the end of the 21st century across the Brazilian Atlantic Forest (AF) domain. To develop the models, we used the maximum entropy method with projected climate data to 2100, based on the Intergovernmental Panel on Climate Change Representative Concentration Pathway (RCP) 4.5 from the fifth Assessment Report. Our AGB models had a satisfactory performance (area under the curve > 0.75 and p value < .05). The models projected a significant increase of 8.5% in the total carbon stock. Overall, the projections indicated that 76.9% of the AF domain would have suitable climatic conditions for increasing biomass by 2100 considering the RCP 4.5 scenario, in the absence of deforestation. Of the existing forest fragments, 34.7% are projected to increase their AGB, while 2.6% are projected to have their AGB reduced by 2100. The regions likely to lose most AGB-up to 40% compared to the baseline-are found between latitudes 13° and 20° south. Overall, although climate change effects on AGB vary latitudinally for the 2071-2100 period under the RCP 4.5 scenario, our model indicates that AGB stocks can potentially increase across a large fraction of the AF. The patterns found here are recommended to be taken into consideration during the planning of restoration efforts, as part of climate change mitigation strategies in the AF and elsewhere in Brazil.

Anthropogenic disturbance in tropical forests can double biodiversity loss from deforestation.

Concerted political attention has focused on reducing deforestation, and this remains the cornerstone of most biodiversity conservation strategies. However, maintaining forest cover may not reduce anthropogenic forest disturbances, which are rarely considered in conservation programmes. These disturbances occur both within forests, including selective logging and wildfires, and at the landscape level, through edge, area and isolation effects. Until now, the combined effect of anthropogenic disturbance on the conservation value of remnant primary forests has remained unknown, making it impossible to assess the relative importance of forest disturbance and forest loss. Here we address these knowledge gaps using a large data set of plants, birds and dung beetles (1,538, 460 and 156 species, respectively) sampled in 36 catchments in the Brazilian state of Pará. Catchments retaining more than 69­80% forest cover lost more conservation value from disturbance than from forest loss. For example, a 20% loss of primary forest, the maximum level of deforestation allowed on Amazonian properties under Brazil's Forest Code, resulted in a 39­54% loss of conservation value: 96­171% more than expected without considering disturbance effects. We extrapolated the disturbance-mediated loss of conservation value throughout Pará, which covers 25% of the Brazilian Amazon. Although disturbed forests retained considerable conservation value compared with deforested areas, the toll of disturbance outside Pará's strictly protected areas is equivalent to the loss of 92,000­139,000 km2 of primary forest. Even this lowest estimate is greater than the area deforested across the entire Brazilian Amazon between 2006 and 2015 (ref. 10). Species distribution models showed that both landscape and within-forest disturbances contributed to biodiversity loss, with the greatest negative effects on species of high conservation and functional value. These results demonstrate an urgent need for policy interventions that go beyond the maintenance of forest cover to safeguard the hyper-diversity of tropical forest ecosystems.

Drought-driven wildfire impacts on structure and dynamics in a wet Central Amazonian forest.

While the climate and human-induced forest degradation is increasing in the Amazon, fire impacts on forest dynamics remain understudied in the wetter regions of the basin, which are susceptible to large wildfires only during extreme droughts. To address this gap, we installed burned and unburned plots immediately after a wildfire in the northern Purus-Madeira (Central Amazon) during the 2015 El-Niño. We measured all individuals with diameter of 10 cm or more at breast height and conducted recensuses to track the demographic drivers of biomass change over 3 years. We also assessed how stem-level growth and mortality were influenced by fire intensity (proxied by char height) and tree morphological traits (size and wood density). Overall, the burned forest lost 27.3% of stem density and 12.8% of biomass, concentrated in small and medium trees. Mortality drove these losses in the first 2 years and recruitment decreased in the third year. The fire increased growth in lower wood density and larger sized trees, while char height had transitory strong effects increasing tree mortality. Our findings suggest that fire impacts are weaker in the wetter Amazon. Here, trees of greater sizes and higher wood densities may confer a margin of fire resistance; however, this may not extend to higher intensity fires arising from climate change.

Rapid responses of root traits and productivity to phosphorus and cation additions in a tropical lowland forest in Amazonia.

Soil nutrient availability can strongly affect root traits. In tropical forests, phosphorus (P) is often considered the main limiting nutrient for plants. However, support for the P paradigm is limited, and N and cations might also control tropical forests functioning. We used a large-scale experiment to determine how the factorial addition of nitrogen (N), P and cations affected root productivity and traits related to nutrient acquisition strategies (morphological traits, phosphatase activity, arbuscular mycorrhizal colonisation and nutrient contents) in a primary rainforest growing on low-fertility soils in Central Amazonia after 1 yr of fertilisation. Multiple root traits and productivity were affected. Phosphorus additions increased annual root productivity and root diameter, but decreased root phosphatase activity. Cation additions increased root productivity at certain times of year, also increasing root diameter and mycorrhizal colonisation. P and cation additions increased their element concentrations in root tissues. No responses were detected with N addition. Here we showed that rock-derived nutrients determined root functioning in low-fertility Amazonian soils, demonstrating not only the hypothesised importance of P, but also highlighting the role of cations. The changes in fine root traits and productivity indicated that even slow-growing tropical rainforests can respond rapidly to changes in resource availability.

Improving the spatial-temporal analysis of Amazonian fires.

There is a growing interest in Amazonian fires, accompanied by a substantial increase in research in the subject. Here, we list five common misunderstandings about Amazonian climate, vegetation, fires and the deforestation process to help to support future research.

Fine root dynamics across pantropical rainforest ecosystems.

Fine roots constitute a significant component of the net primary productivity (NPP) of forest ecosystems but are much less studied than aboveground NPP. Comparisons across sites and regions are also hampered by inconsistent methodologies, especially in tropical areas. Here, we present a novel dataset of fine root biomass, productivity, residence time, and allocation in tropical old-growth rainforest sites worldwide, measured using consistent methods, and examine how these variables are related to consistently determined soil and climatic characteristics. Our pantropical dataset spans intensive monitoring plots in lowland (wet, semi-deciduous, and deciduous) and montane tropical forests in South America, Africa, and Southeast Asia (n = 47). Large spatial variation in fine root dynamics was observed across montane and lowland forest types. In lowland forests, we found a strong positive linear relationship between fine root productivity and sand content, this relationship was even stronger when we considered the fractional allocation of total NPP to fine roots, demonstrating that understanding allocation adds explanatory power to understanding fine root productivity and total NPP. Fine root residence time was a function of multiple factors: soil sand content, soil pH, and maximum water deficit, with longest residence times in acidic, sandy, and water-stressed soils. In tropical montane forests, on the other hand, a different set of relationships prevailed, highlighting the very different nature of montane and lowland forest biomes. Root productivity was a strong positive linear function of mean annual temperature, root residence time was a strong positive function of soil nitrogen content in montane forests, and lastly decreasing soil P content increased allocation of productivity to fine roots. In contrast to the lowlands, environmental conditions were a better predictor for fine root productivity than for fractional allocation of total NPP to fine roots, suggesting that root productivity is a particularly strong driver of NPP allocation in tropical mountain regions.

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.

Seasonal and drought-related changes in leaf area profiles depend on height and light environment in an Amazon forest.

Seasonal dynamics in the vertical distribution of leaf area index (LAI) may impact the seasonality of forest productivity in Amazonian forests. However, until recently, fine-scale observations critical to revealing ecological mechanisms underlying these changes have been lacking. To investigate fine-scale variation in leaf area with seasonality and drought we conducted monthly ground-based LiDAR surveys over 4 yr at an Amazon forest site. We analysed temporal changes in vertically structured LAI along axes of both canopy height and light environments. Upper canopy LAI increased during the dry season, whereas lower canopy LAI decreased. The low canopy decrease was driven by highly illuminated leaves of smaller trees in gaps. By contrast, understory LAI increased concurrently with the upper canopy. Hence, tree phenological strategies were stratified by height and light environments. Trends were amplified during a 2015-2016 severe El Niño drought. Leaf area low in the canopy exhibited behaviour consistent with water limitation. Leaf loss from short trees in high light during drought may be associated with strategies to tolerate limited access to deep soil water and stressful leaf environments. Vertically and environmentally structured phenological processes suggest a critical role of canopy structural heterogeneity in seasonal changes in Amazon ecosystem function.

Hydraulic traits explain differential responses of Amazonian forests to the 2015 El Niño-induced drought.

Reducing uncertainties in the response of tropical forests to global change requires understanding how intra- and interannual climatic variability selects for different species, community functional composition and ecosystem functioning, so that the response to climatic events of differing frequency and severity can be predicted. Here we present an extensive dataset of hydraulic traits of dominant species in two tropical Amazon forests with contrasting precipitation regimes - low seasonality forest (LSF) and high seasonality forest (HSF) - and relate them to community and ecosystem response to the El Niño-Southern Oscillation (ENSO) of 2015. Hydraulic traits indicated higher drought tolerance in the HSF than in the LSF. Despite more intense drought and lower plant water potentials in HSF during the 2015-ENSO, greater xylem embolism resistance maintained similar hydraulic safety margin as in LSF. This likely explains how ecosystem-scale whole-forest canopy conductance at HSF maintained a similar response to atmospheric drought as at LSF, despite their water transport systems operating at different water potentials. Our results indicate that contrasting precipitation regimes (at seasonal and interannual time scales) select for assemblies of hydraulic traits and taxa at the community level, which may have a significant role in modulating forest drought response at ecosystem scales.

Effects of climate and land-use change scenarios on fire probability during the 21st century in the Brazilian Amazon.

The joint and relative effects of future land-use and climate change on fire occurrence in the Amazon, as well its seasonal variation, are still poorly understood, despite its recognized importance. Using the maximum entropy method (MaxEnt), we combined regional land-use projections and climatic data from the CMIP5 multimodel ensemble to investigate the monthly probability of fire occurrence in the mid (2041-2070) and late (2071-2100) 21st century in the Brazilian Amazon. We found striking spatial variation in the fire relative probability (FRP) change along the months, with October showing the highest overall change. Considering climate only, the area with FRP ≥ 0.3 (a threshold chosen based on the literature) in October increases 6.9% by 2071-2100 compared to the baseline period under the representative concentration pathway (RCP) 4.5 and 27.7% under the RCP 8.5. The best-case land-use scenario ("Sustainability") alone causes a 10.6% increase in the area with FRP ≥ 0.3, while the worse-case land-use scenario ("Fragmentation") causes a 73.2% increase. The optimistic climate-land-use projection (Sustainability and RCP 4.5) causes a 21.3% increase in the area with FRP ≥ 0.3 in October by 2071-2100 compared to the baseline period. In contrast, the most pessimistic climate-land-use projection (Fragmentation and RCP 8.5) causes a widespread increase in FRP (113.5% increase in the area with FRP ≥ 0.3), and prolongs the fire season, displacing its peak. Combining the Sustainability land-use and RCP 8.5 scenarios causes a 39.1% increase in the area with FRP ≥ 0.3. We conclude that avoiding the regress on land-use governance in the Brazilian Amazon (i.e., decrease in the extension and level of conservation of the protected areas, reduced environmental laws enforcement, extensive road paving, and increased deforestation) would substantially mitigate the effects of climate change on fire probability, even under the most pessimistic RCP 8.5 scenario.

Compositional response of Amazon forests to climate change.

Most of the planet's diversity is concentrated in the tropics, which includes many regions undergoing rapid climate change. Yet, while climate-induced biodiversity changes are widely documented elsewhere, few studies have addressed this issue for lowland tropical ecosystems. Here we investigate whether the floristic and functional composition of intact lowland Amazonian forests have been changing by evaluating records from 106 long-term inventory plots spanning 30 years. We analyse three traits that have been hypothesized to respond to different environmental drivers (increase in moisture stress and atmospheric CO2 concentrations): maximum tree size, biogeographic water-deficit affiliation and wood density. Tree communities have become increasingly dominated by large-statured taxa, but to date there has been no detectable change in mean wood density or water deficit affiliation at the community level, despite most forest plots having experienced an intensification of the dry season. However, among newly recruited trees, dry-affiliated genera have become more abundant, while the mortality of wet-affiliated genera has increased in those plots where the dry season has intensified most. Thus, a slow shift to a more dry-affiliated Amazonia is underway, with changes in compositional dynamics (recruits and mortality) consistent with climate-change drivers, but yet to significantly impact whole-community composition. The Amazon observational record suggests that the increase in atmospheric CO2 is driving a shift within tree communities to large-statured species and that climate changes to date will impact forest composition, but long generation times of tropical trees mean that biodiversity change is lagging behind climate change.