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.

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.

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.

Dependence of hydropower energy generation on forests in the Amazon Basin at local and regional scales.

Tropical rainforest regions have large hydropower generation potential that figures prominently in many nations' energy growth strategies. Feasibility studies of hydropower plants typically ignore the effect of future deforestation or assume that deforestation will have a positive effect on river discharge and energy generation resulting from declines in evapotranspiration (ET) associated with forest conversion. Forest loss can also reduce river discharge, however, by inhibiting rainfall. We used land use, hydrological, and climate models to examine the local "direct" effects (through changes in ET within the watershed) and the potential regional "indirect" effects (through changes in rainfall) of deforestation on river discharge and energy generation potential for the Belo Monte energy complex, one of the world's largest hydropower plants that is currently under construction on the Xingu River in the eastern Amazon. In the absence of indirect effects of deforestation, simulated deforestation of 20% and 40% within the Xingu River basin increased discharge by 4-8% and 10-12%, with similar increases in energy generation. When indirect effects were considered, deforestation of the Amazon region inhibited rainfall within the Xingu Basin, counterbalancing declines in ET and decreasing discharge by 6-36%. Under business-as-usual projections of forest loss for 2050 (40%), simulated power generation declined to only 25% of maximum plant output and 60% of the industry's own projections. Like other energy sources, hydropower plants present large social and environmental costs. Their reliability as energy sources, however, must take into account their dependence on forests.

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.

Soil moisture depletion under simulated drought in the Amazon: impacts on deep root uptake.

*Deep root water uptake in tropical Amazonian forests has been a major discovery during the last 15 yr. However, the effects of extended droughts, which may increase with climate change, on deep soil moisture utilization remain uncertain. *The current study utilized a 1999-2005 record of volumetric water content (VWC) under a throughfall exclusion experiment to calibrate a one-dimensional model of the hydrologic system to estimate VWC, and to quantify the rate of root uptake through 11.5 m of soil. *Simulations with root uptake compensation had a relative root mean square error (RRMSE) of 11% at 0-40 cm and < 5% at 350-1150 cm. The simulated contribution of deep root uptake under the control was c. 20% of water demand from 250 to 550 cm and c. 10% from 550 to 1150 cm. Furthermore, in years 2 (2001) and 3 (2002) of throughfall exclusion, deep root uptake increased as soil moisture was available but then declined to near zero in deep layers in 2003 and 2004. *Deep root uptake was limited despite high VWC (i.e. > 0.30 cm(3) cm(-3)). This limitation may partly be attributable to high residual water contents (theta(r)) in these high-clay (70-90%) soils or due to high soil-to-root resistance. The ability of deep roots and soils to contribute increasing amounts of water with extended drought will be limited.

Mortality of large trees and lianas following experimental drought in an Amazon forest.

Severe drought episodes such as those associated with El Niño Southern Oscillation (ENSO) events influence large areas of tropical forest and may become more frequent in the future. One of the most important forest responses to severe drought is tree mortality, which alters forest structure, composition, carbon content, and flammability, and which varies widely. This study tests the hypothesis that tree mortality increases abruptly during drought episodes when plant-available soil water (PAW) declines below a critical minimum threshold. It also examines the effect of tree size, plant life form (palm, liana, tree) and potential canopy position (understory, midcanopy, overstory) on drought-induced plant mortality. A severe, four-year drought episode was simulated by excluding 60% of incoming throughfall during each wet season using plastic panels installed in the understory of a 1-ha forest treatment plot, while a 1-ha control plot received normal rainfall. After 3.2 years, the treatment resulted in a 38% increase in mortality rates across all stems >2 cm dbh. Mortality rates increased 4.5-fold among large trees (>30 cm dbh) and twofold among medium trees (10-30 cm dbh) in response to the treatment, whereas the smallest stems were less responsive. Recruitment rates did not compensate for the elevated mortality of larger-diameter stems in the treatment plot. Overall, lianas proved more susceptible to drought-induced mortality than trees or palms, and potential overstory tree species were more vulnerable than midcanopy and understory species. Large stems contributed to 90% of the pretreatment live aboveground biomass in both plots. Large-tree mortality resulting from the treatment generated 3.4 times more dead biomass than the control plot. The dramatic mortality response suggests significant, adverse impacts on the global carbon cycle if climatic changes follow current trends.

Effects of high-frequency understorey fires on woody plant regeneration in southeastern Amazonian forests.

Anthropogenic understorey fires affect large areas of tropical forest, yet their effects on woody plant regeneration post-fire remain poorly understood. We examined the effects of repeated experimental fires on woody stem (less than 1 cm at base) mortality, recruitment, species diversity, community similarity and regeneration mode (seed versus sprout) in Mato Grosso, Brazil. From 2004 to 2010, forest plots (50 ha) were burned twice (B2) or five times (B5), and compared with an unburned control (B0). Stem density recovered within a year after the first burn (initial density: 12.4-13.2 stems m(-2)), but after 6 years, increased mortality and decreased regeneration--primarily of seedlings--led to a 63 per cent and 85 per cent reduction in stem density in B2 and B5, respectively. Seedlings and sprouts across plots in 2010 displayed remarkable community similarity owing to shared abundant species. Although the dominant surviving species were similar across plots, a major increase in sprouting occurred--almost three- and fourfold greater in B2 and B5 than in B0. In B5, 29 species disappeared and were replaced by 11 new species often present along fragmented forest edges. By 2010, the annual burn regime created substantial divergence between the seedling community and the initial adult tree community (greater than or equal to 20 cm dbh). Increased droughts and continued anthropogenic ignitions associated with frontier land uses may promote high-frequency fire regimes that may substantially alter regeneration and therefore successional processes.

Defending public interests in private lands: compliance, costs and potential environmental consequences of the Brazilian Forest Code in Mato Grosso.

Land-use regulations are a critical component of forest governance and conservation strategies, but their effectiveness in shaping landholder behaviour is poorly understood. We conducted a spatial and temporal analysis of the Brazilian Forest Code (BFC) to understand the patterns of regulatory compliance over time and across changes in the policy, and the implications of these compliance patterns for the perceived costs to landholders and environmental performance of agricultural landscapes in the southern Amazon state of Mato Grosso. Landholdings tended to remain in compliance or not according to their status at the beginning of the study period. The perceived economic burden of BFC compliance on soya bean and beef producers (US$3-5.6 billion in net present value of the land) may in part explain the massive, successful campaign launched by the farm lobby to change the BFC. The ecological benefits of compliance (e.g. greater connectivity and carbon) with the BFC are diffuse and do not compete effectively with the economic benefits of non-compliance that are perceived by landholders. Volatile regulation of land-use decisions that affect billions in economic rent that could be captured is an inadequate forest governance instrument; effectiveness of such regulations may increase when implemented in tandem with positive incentives for forest conservation.

Responding to climate change and the global land crisis: REDD+, market transformation and low-emissions rural development.

Climate change and rapidly escalating global demand for food, fuel, fibre and feed present seemingly contradictory challenges to humanity. Can greenhouse gas (GHG) emissions from land-use, more than one-fourth of the global total, decline as growth in land-based production accelerates? This review examines the status of two major international initiatives that are designed to address different aspects of this challenge. REDD+ is an emerging policy framework for providing incentives to tropical nations and states that reduce their GHG emissions from deforestation and forest degradation. Market transformation, best represented by agricultural commodity roundtables, seeks to exclude unsustainable farmers from commodity markets through international social and environmental standards for farmers and processors. These global initiatives could potentially become synergistically integrated through (i) a shared approach for measuring and favouring high environmental and social performance of land use across entire jurisdictions and (ii) stronger links with the domestic policies, finance and laws in the jurisdictions where agricultural expansion is moving into forests. To achieve scale, the principles of REDD+ and sustainable farming systems must be embedded in domestic low-emission rural development models capable of garnering support across multiple constituencies. We illustrate this potential with the case of Mato Grosso State in the Brazilian Amazon.

Testing the Amazon savannization hypothesis: fire effects on invasion of a neotropical forest by native cerrado and exotic pasture grasses.

Changes in climate and land use that interact synergistically to increase fire frequencies and intensities in tropical regions are predicted to drive forests to new grass-dominated stable states. To reveal the mechanisms for such a transition, we established 50 ha plots in a transitional forest in the southwestern Brazilian Amazon to different fire treatments (unburned, burned annually (B1yr) or at 3-year intervals (B3yr)). Over an 8-year period since the commencement of these treatments, we documented: (i) the annual rate of pasture and native grass invasion in response to increasing fire frequency; (ii) the establishment of Brachiaria decumbens (an African C4 grass) as a function of decreasing canopy cover and (iii) the effects of grass fine fuel on fire intensity. Grasses invaded approximately 200 m from the edge into the interiors of burned plots (B1yr: 4.31 ha; B3yr: 4.96 ha) but invaded less than 10 m into the unburned plot (0.33 ha). The probability of B. decumbens establishment increased with seed availability and decreased with leaf area index. Fine fuel loads along the forest edge were more than three times higher in grass-dominated areas, which resulted in especially intense fires. Our results indicate that synergies between fires and invasive C4 grasses jeopardize the future of tropical forests.

Comment on "The incidence of fire in Amazonian forests with implications for REDD".

Aragão and Shimabukuro (Reports, 4 June 2010, p. 1275) reported that fires increase in agricultural frontiers even as deforestation decreases and concluded that these fires lead to unaccounted carbon emissions under the United Nations climate treaty's tropical deforestation and forest degradation component. Emissions from post-deforestation management activities are, in fact, included in these estimates--but burning of standing forests is not.

Interactions among Amazon land use, forests and climate: prospects for a near-term forest tipping point.

Some model experiments predict a large-scale substitution of Amazon forest by savannah-like vegetation by the end of the twenty-first century. Expanding global demands for biofuels and grains, positive feedbacks in the Amazon forest fire regime and drought may drive a faster process of forest degradation that could lead to a near-term forest dieback. Rising worldwide demands for biofuel and meat are creating powerful new incentives for agro-industrial expansion into Amazon forest regions. Forest fires, drought and logging increase susceptibility to further burning while deforestation and smoke can inhibit rainfall, exacerbating fire risk. If sea surface temperature anomalies (such as El Niño episodes) and associated Amazon droughts of the last decade continue into the future, approximately 55% of the forests of the Amazon will be cleared, logged, damaged by drought or burned over the next 20 years, emitting 15-26Pg of carbon to the atmosphere. Several important trends could prevent a near-term dieback. As fire-sensitive investments accumulate in the landscape, property holders use less fire and invest more in fire control. Commodity markets are demanding higher environmental performance from farmers and cattle ranchers. Protected areas have been established in the pathway of expanding agricultural frontiers. Finally, emerging carbon market incentives for reductions in deforestation could support these trends.

Drought effects on litterfall, wood production and belowground carbon cycling in an Amazon forest: results of a throughfall reduction experiment.

The Amazon Basin experiences severe droughts that may become more common in the future. Little is known of the effects of such droughts on Amazon forest productivity and carbon allocation. We tested the prediction that severe drought decreases litterfall and wood production but potentially has multiple cancelling effects on belowground production within a 7-year partial throughfall exclusion experiment. We simulated an approximately 35-41% reduction in effective rainfall from 2000 through 2004 in a 1ha plot and compared forest response with a similar control plot. Wood production was the most sensitive component of above-ground net primary productivity (ANPP) to drought, declining by 13% the first year and up to 62% thereafter. Litterfall declined only in the third year of drought, with a maximum difference of 23% below the control plot. Soil CO2 efflux and its 14C signature showed no significant treatment response, suggesting similar amounts and sources of belowground production. ANPP was similar between plots in 2000 and declined to a low of 41% below the control plot during the subsequent treatment years, rebounding to only a 10% difference during the first post-treatment year. Live aboveground carbon declined by 32.5Mgha-1 through the effects of drought on ANPP and tree mortality. Results of this unreplicated, long-term, large-scale ecosystem manipulation experiment demonstrate that multi-year severe drought can substantially reduce Amazon forest carbon stocks.

Globalization of the Amazon soy and beef industries: opportunities for conservation.

Amazon beef and soybean industries, the primary drivers of Amazon deforestation, are increasingly responsive to economic signals emanating from around the world, such as those associated with bovine spongiform encephalopathy (BSE, "mad cow disease") outbreaks and China's economic growth. The expanding role of these economic "teleconnections" (coupled phenomena that take place in distant places on the planet) led to a 3-year period (2002-2004) of historically high deforestation rates. But it also increases the potential for large-scale conservation in the region as markets and finance institutions demand better environmental and social performance of beef and soy producers. Cattle ranchers and soy farmers who have generally opposed ambitious government regulations that require forest reserves on private property are realizing that good land stewardship-including compliance with legislation-may increase their access to expanding domestic and international markets and to credit and lower the risk of "losing" their land to agrarian reform. The realization of this potential depends on the successful negotiation of social and environmental performance criteria and an associated system of certification that are acceptable to both the industries and civil society. The foot-and-mouth eradication system, in which geographic zones win permission to export beef, may provide an important model for the design of a low-cost, peer-enforced, socioenvironmental certification system that becomes the mechanism by which beef and soy industries gain access to markets outside the Amazon.

Rainfall exclusion in an eastern Amazonian forest alters soil water movement and depth of water uptake.

Deuterium-labeled water was used to study the effect of the Tapajós Throughfall Exclusion Experiment (TTEE) on soil moisture movement and on depth of water uptake by trees of Coussarea racemosa, Sclerolobium chrysophyllum, and Eschweilera pedicellata. The TTEE simulates an extended dry season in an eastern Amazonian rainforest, a plausible scenario if the El Niño phenomenon changes with climate change. The TTEE excludes 60% of the wet season throughfall from a 1-ha plot (treatment), while the control 1-ha plot receives precipitation year-round. Mean percolation rate of the label peak in the control plot was greater than in the treatment plot during the wet season (0.75 vs. 0.07 m/mo). The rate was similar for both plots during the dry season (ca. 0.15 m/mo), indicative that both plots have similar topsoil structure. Interestingly, the label peak in the control plot during the dry season migrated upward an average distance of 64 cm. We show that water probably moved upward through soil pores-i.e., it did not involve roots (hydraulic lift)-most likely because of a favorable gradient of total (matric + gravitational) potential coupled with sufficient unsaturated hydraulic conductivity. Water probably also moved upward in the treatment plot, but was not detectable; the label in this plot did not percolate below 1 m or beyond the depth of plant water uptake. During the dry season, trees in the rainfall exclusion plot, regardless of species, consistently absorbed water significantly deeper, but never below 1.5-2 m, than trees in the control plot, and therefore may represent expected root function of this understory/subcanopy tree community during extended dry periods.

Hydraulic redistribution in three Amazonian trees.

About half of the Amazon rainforest is subject to seasonal droughts of 3 months or more. Despite this drought, several studies have shown that these forests, under a strongly seasonal climate, do not exhibit significant water stress during the dry season. In addition to deep soil water uptake, another contributing explanation for the absence of plant water stress during drought is the process of hydraulic redistribution; the nocturnal transfer of water by roots from moist to dry regions of the soil profile. Here, we present data on patterns of soil moisture and sap flow in roots of three dimorphic-rooted species in the Tapajós Forest, Amazônia, which demonstrate both upward (hydraulic lift) and downward hydraulic redistribution. We measured sap flow in lateral and tap roots of our three study species over a 2-year period using the heat ratio method, a sap-flow technique that allows bi-directional measurement of water flow. On certain nights during the dry season, reverse or acropetal flow (i.e.,in the direction of the soil) in the lateral roots and positive or basipetal sap flow (toward the plant) in the tap roots of Coussarea racemosa (caferana), Manilkara huberi (maçaranduba) and Protium robustum (breu) were observed, a pattern consistent with upward hydraulic redistribution (hydraulic lift). With the onset of heavy rains, this pattern reversed, with continuous night-time acropetal sap flow in the tap root and basipetal sap flow in lateral roots, indicating water movement from wet top soil to dry deeper soils (downward hydraulic redistribution). Both patterns were present in trees within a rainfall exclusion plot (Seca Floresta) and to a more limited extent in the control plot. Although hydraulic redistribution has traditionally been associated with arid or strongly seasonal environments, our findings now suggest that it is important in ameliorating water stress and improving rain infiltration in Amazonian rainforests. This has broad implications for understanding and modeling ecosystem process and forest function in this important biome.