Asymmetric response of Amazon forest water and energy fluxes to wet and dry hydrological extremes reveals onset of a local drought-induced tipping point.

Understanding the effects of intensification of Amazon basin hydrological cycling-manifest as increasingly frequent floods and droughts-on water and energy cycles of tropical forests is essential to meeting the challenge of predicting ecosystem responses to climate change, including forest "tipping points". Here, we investigated the impacts of hydrological extremes on forest function using 12+ years of observations (between 2001-2020) of water and energy fluxes from eddy covariance, along with associated ecological dynamics from biometry, at the Tapajós National Forest. Measurements encompass the strong 2015-2016 El Niño drought and La Niña 2008-2009 wet events. We found that the forest responded strongly to El Niño-Southern Oscillation (ENSO): Drought reduced water availability for evapotranspiration (ET) leading to large increases in sensible heat fluxes (H). Partitioning ET by an approach that assumes transpiration (T) is proportional to photosynthesis, we found that water stress-induced reductions in canopy conductance (Gs ) drove T declines partly compensated by higher evaporation (E). By contrast, the abnormally wet La Niña period gave higher T and lower E, with little change in seasonal ET. Both El Niño-Southern Oscillation (ENSO) events resulted in changes in forest structure, manifested as lower wet-season leaf area index. However, only during El Niño 2015-2016, we observed a breakdown in the strong meteorological control of transpiration fluxes (via energy availability and atmospheric demand) because of slowing vegetation functions (via shutdown of Gs and significant leaf shedding). Drought-reduced T and Gs , higher H and E, amplified by feedbacks with higher temperatures and vapor pressure deficits, signaled that forest function had crossed a threshold, from which it recovered slowly, with delay, post-drought. Identifying such tipping point onsets (beyond which future irreversible processes may occur) at local scale is crucial for predicting basin-scale threshold-crossing changes in forest energy and water cycling, leading to slow-down in forest function, potentially resulting in Amazon forests shifting into alternate degraded states.

Understanding water and energy fluxes in the Amazonia: Lessons from an observation-model intercomparison.

Tropical forests are an important part of global water and energy cycles, but the mechanisms that drive seasonality of their land-atmosphere exchanges have proven challenging to capture in models. Here, we (1) report the seasonality of fluxes of latent heat (LE), sensible heat (H), and outgoing short and longwave radiation at four diverse tropical forest sites across Amazonia-along the equator from the Caxiuanã and Tapajós National Forests in the eastern Amazon to a forest near Manaus, and from the equatorial zone to the southern forest in Reserva Jaru; (2) investigate how vegetation and climate influence these fluxes; and (3) evaluate land surface model performance by comparing simulations to observations. We found that previously identified failure of models to capture observed dry-season increases in evapotranspiration (ET) was associated with model overestimations of (1) magnitude and seasonality of Bowen ratios (relative to aseasonal observations in which sensible was only 20%-30% of the latent heat flux) indicating model exaggerated water limitation, (2) canopy emissivity and reflectance (albedo was only 10%-15% of incoming solar radiation, compared to 0.15%-0.22% simulated), and (3) vegetation temperatures (due to underestimation of dry-season ET and associated cooling). These partially compensating model-observation discrepancies (e.g., higher temperatures expected from excess Bowen ratios were partially ameliorated by brighter leaves and more interception/evaporation) significantly biased seasonal model estimates of net radiation (Rn ), the key driver of water and energy fluxes (LE ~ 0.6 Rn and H ~ 0.15 Rn ), though these biases varied among sites and models. A better representation of energy-related parameters associated with dynamic phenology (e.g., leaf optical properties, canopy interception, and skin temperature) could improve simulations and benchmarking of current vegetation-atmosphere exchange and reduce uncertainty of regional and global biogeochemical models.

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.

The Amazon basin in transition.

Agricultural expansion and climate variability have become important agents of disturbance in the Amazon basin. Recent studies have demonstrated considerable resilience of Amazonian forests to moderate annual drought, but they also show that interactions between deforestation, fire and drought potentially lead to losses of carbon storage and changes in regional precipitation patterns and river discharge. Although the basin-wide impacts of land use and drought may not yet surpass the magnitude of natural variability of hydrologic and biogeochemical cycles, there are some signs of a transition to a disturbance-dominated regime. These signs include changing energy and water cycles in the southern and eastern portions of the Amazon basin.

Partitioning controls on Amazon forest photosynthesis between environmental and biotic factors at hourly to interannual timescales.

Gross ecosystem productivity (GEP) in tropical forests varies both with the environment and with biotic changes in photosynthetic infrastructure, but our understanding of the relative effects of these factors across timescales is limited. Here, we used a statistical model to partition the variability of seven years of eddy covariance-derived GEP in a central Amazon evergreen forest into two main causes: variation in environmental drivers (solar radiation, diffuse light fraction, and vapor pressure deficit) that interact with model parameters that govern photosynthesis and biotic variation in canopy photosynthetic light-use efficiency associated with changes in the parameters themselves. Our fitted model was able to explain most of the variability in GEP at hourly (R2  = 0.77) to interannual (R2  = 0.80) timescales. At hourly timescales, we found that 75% of observed GEP variability could be attributed to environmental variability. When aggregating GEP to the longer timescales (daily, monthly, and yearly), however, environmental variation explained progressively less GEP variability: At monthly timescales, it explained only 3%, much less than biotic variation in canopy photosynthetic light-use efficiency, which accounted for 63%. These results challenge modeling approaches that assume GEP is primarily controlled by the environment at both short and long timescales. Our approach distinguishing biotic from environmental variability can help to resolve debates about environmental limitations to tropical forest photosynthesis. For example, we found that biotically regulated canopy photosynthetic light-use efficiency (associated with leaf phenology) increased with sunlight during dry seasons (consistent with light but not water limitation of canopy development) but that realized GEP was nonetheless lower relative to its potential efficiency during dry than wet seasons (consistent with water limitation of photosynthesis in given assemblages of leaves). This work highlights the importance of accounting for differential regulation of GEP at different timescales and of identifying the underlying feedbacks and adaptive mechanisms.

Do dynamic global vegetation models capture the seasonality of carbon fluxes in the Amazon basin? A data-model intercomparison.

To predict forest response to long-term climate change with high confidence requires that dynamic global vegetation models (DGVMs) be successfully tested against ecosystem response to short-term variations in environmental drivers, including regular seasonal patterns. Here, we used an integrated dataset from four forests in the Brasil flux network, spanning a range of dry-season intensities and lengths, to determine how well four state-of-the-art models (IBIS, ED2, JULES, and CLM3.5) simulated the seasonality of carbon exchanges in Amazonian tropical forests. We found that most DGVMs poorly represented the annual cycle of gross primary productivity (GPP), of photosynthetic capacity (Pc), and of other fluxes and pools. Models simulated consistent dry-season declines in GPP in the equatorial Amazon (Manaus K34, Santarem K67, and Caxiuanã CAX); a contrast to observed GPP increases. Model simulated dry-season GPP reductions were driven by an external environmental factor, 'soil water stress' and consequently by a constant or decreasing photosynthetic infrastructure (Pc), while observed dry-season GPP resulted from a combination of internal biological (leaf-flush and abscission and increased Pc) and environmental (incoming radiation) causes. Moreover, we found models generally overestimated observed seasonal net ecosystem exchange (NEE) and respiration (Re ) at equatorial locations. In contrast, a southern Amazon forest (Jarú RJA) exhibited dry-season declines in GPP and Re consistent with most DGVMs simulations. While water limitation was represented in models and the primary driver of seasonal photosynthesis in southern Amazonia, changes in internal biophysical processes, light-harvesting adaptations (e.g., variations in leaf area index (LAI) and increasing leaf-level assimilation rate related to leaf demography), and allocation lags between leaf and wood, dominated equatorial Amazon carbon flux dynamics and were deficient or absent from current model formulations. Correctly simulating flux seasonality at tropical forests requires a greater understanding and the incorporation of internal biophysical mechanisms in future model developments.

Empirical evidence for resilience of tropical forest photosynthesis in a warmer world.

Tropical forests may be vulnerable to climate change1-3 if photosynthetic carbon uptake currently operates near a high temperature limit4-6. Predicting tropical forest function requires understanding the relative contributions of two mechanisms of high-temperature photosynthetic declines: stomatal limitation (H1), an indirect response due to temperature-associated changes in atmospheric vapour pressure deficit (VPD)7, and biochemical restrictions (H2), a direct temperature response8,9. Their relative control predicts different outcomes-H1 is expected to diminish with stomatal responses to future co-occurring elevated atmospheric [CO2], whereas H2 portends declining photosynthesis with increasing temperatures. Distinguishing the two mechanisms at high temperatures is therefore critical, but difficult because VPD is highly correlated with temperature in natural settings. We used a forest mesocosm to quantify the sensitivity of tropical gross ecosystem productivity (GEP) to future temperature regimes while constraining VPD by controlling humidity. We then analytically decoupled temperature and VPD effects under current climate with flux-tower-derived GEP trends in situ from four tropical forest sites. Both approaches showed consistent, negative sensitivity of GEP to VPD but little direct response to temperature. Importantly, in the mesocosm at low VPD, GEP persisted up to 38 °C, a temperature exceeding projections for tropical forests in 2100 (ref. 10). If elevated [CO2] mitigates VPD-induced stomatal limitation through enhanced water-use efficiency as hypothesized9,11, tropical forest photosynthesis may have a margin of resilience to future warming.

Species-Specific Shifts in Diurnal Sap Velocity Dynamics and Hysteretic Behavior of Ecophysiological Variables During the 2015-2016 El Niño Event in the Amazon Forest.

Current climate change scenarios indicate warmer temperatures and the potential for more extreme droughts in the tropics, such that a mechanistic understanding of the water cycle from individual trees to landscapes is needed to adequately predict future changes in forest structure and function. In this study, we contrasted physiological responses of tropical trees during a normal dry season with the extreme dry season due to the 2015-2016 El Niño-Southern Oscillation (ENSO) event. We quantified high resolution temporal dynamics of sap velocity (Vs), stomatal conductance (gs) and leaf water potential (ΨL) of multiple canopy trees, and their correlations with leaf temperature (Tleaf) and environmental conditions [direct solar radiation, air temperature (Tair) and vapor pressure deficit (VPD)]. The experiment leveraged canopy access towers to measure adjacent trees at the ZF2 and Tapajós tropical forest research (near the cities of Manaus and Santarém). The temporal difference between the peak of gs (late morning) and the peak of VPD (early afternoon) is one of the major regulators of sap velocity hysteresis patterns. Sap velocity displayed species-specific diurnal hysteresis patterns reflected by changes in Tleaf. In the morning, Tleaf and sap velocity displayed a sigmoidal relationship. In the afternoon, stomatal conductance declined as Tleaf approached a daily peak, allowing ΨL to begin recovery, while sap velocity declined with an exponential relationship with Tleaf. In Manaus, hysteresis indices of the variables Tleaf-Tair and ΨL-Tleaf were calculated for different species and a significant difference (p < 0.01, α = 0.05) was observed when the 2015 dry season (ENSO period) was compared with the 2017 dry season ("control scenario"). In some days during the 2015 ENSO event, Tleaf approached 40°C for all studied species and the differences between Tleaf and Tair reached as high at 8°C (average difference: 1.65 ± 1.07°C). Generally, Tleaf was higher than Tair during the middle morning to early afternoon, and lower than Tair during the early morning, late afternoon and night. Our results support the hypothesis that partial stomatal closure allows for a recovery in ΨL during the afternoon period giving an observed counterclockwise hysteresis pattern between ΨL and Tleaf.

Nocturnal accumulation of CO2 underneath a tropical forest canopy along a topographical gradient.

Flux measurements of carbon dioxide and water vapor above tropical rain forests are often difficult to interpret because the terrain is usually complex. This complexity induces heterogeneity in the surface but also affects lateral movement of carbon dioxide (CO2) not readily detected by the eddy covariance systems. This study describes such variability using measurements of CO2 along vertical profiles and along a toposequence in a tropical rain forest near Manaus, Brazil. Seasonal and diurnal variation was recorded, with atmospheric CO2 concentration maxima around dawn, generally higher CO2 build-up in the dry season and stronger daytime CO2 drawdown in the wet season. This variation was reflected all along the toposequence, but the slope and valley bottom accumulated clearly more CO2 than the plateaus, depending on atmospheric stability. Particularly during stable nights, accumulation was along lines of equal altitude, suggesting that large amounts of CO2 are stored in the valleys of the landscape. Flushing of this store only occurs during mid-morning, when stored CO2 may well be partly transported back to the plateaus. It is clear that, for proper interpretation of tower fluxes in such complex and actively respiring terrain, the horizontal variability of storage needs to be taken into account not only during the night but also during the mornings.

Dry and hot: the hydraulic consequences of a climate change-type drought for Amazonian trees.

How plants respond physiologically to leaf warming and low water availability may determine how they will perform under future climate change. In 2015-2016, an unprecedented drought occurred across Amazonia with record-breaking high temperatures and low soil moisture, offering a unique opportunity to evaluate the performances of Amazonian trees to a severe climatic event. We quantified the responses of leaf water potential, sap velocity, whole-tree hydraulic conductance (Kwt), turgor loss and xylem embolism, during and after the 2015-2016 El Niño for five canopy-tree species. Leaf/xylem safety margins (SMs), sap velocity and Kwt showed a sharp drop during warm periods. SMs were negatively correlated with vapour pressure deficit, but had no significant relationship with soil water storage. Based on our calculations of canopy stomatal and xylem resistances, the decrease in sap velocity and Kwt was due to a combination of xylem cavitation and stomatal closure. Our results suggest that warm droughts greatly amplify the degree of trees' physiological stress and can lead to mortality. Given the extreme nature of the 2015-2016 El Niño and that temperatures are predicted to increase, this work can serve as a case study of the possible impact climate warming can have on tropical trees.This article is part of a discussion meeting issue 'The impact of the 2015/2016 El Niño on the terrestrial tropical carbon cycle: patterns, mechanisms and implications'.

Widespread reduction in sun-induced fluorescence from the Amazon during the 2015/2016 El Niño.

The tropical carbon balance dominates year-to-year variations in the CO2 exchange with the atmosphere through photosynthesis, respiration and fires. Because of its high correlation with gross primary productivity (GPP), observations of sun-induced fluorescence (SIF) are of great interest. We developed a new remotely sensed SIF product with improved signal-to-noise in the tropics, and use it here to quantify the impact of the 2015/2016 El Niño Amazon drought. We find that SIF was strongly suppressed over areas with anomalously high temperatures and decreased levels of water in the soil. SIF went below its climatological range starting from the end of the 2015 dry season (October) and returned to normal levels by February 2016 when atmospheric conditions returned to normal, but well before the end of anomalously low precipitation that persisted through June 2016. Impacts were not uniform across the Amazon basin, with the eastern part experiencing much larger (10-15%) SIF reductions than the western part of the basin (2-5%). We estimate the integrated loss of GPP relative to eight previous years to be 0.34-0.48 PgC in the three-month period October-November-December 2015.This article is part of a discussion meeting issue 'The impact of the 2015/2016 El Niño on the terrestrial tropical carbon cycle: patterns, mechanisms and implications'.

Variability of carbon and water fluxes following climate extremes over a tropical forest in southwestern Amazonia.

The carbon and water cycles for a southwestern Amazonian forest site were investigated using the longest time series of fluxes of CO2 and water vapor ever reported for this site. The period from 2004 to 2010 included two severe droughts (2005 and 2010) and a flooding year (2009). The effects of such climate extremes were detected in annual sums of fluxes as well as in other components of the carbon and water cycles, such as gross primary production and water use efficiency. Gap-filling and flux-partitioning were applied in order to fill gaps due to missing data, and errors analysis made it possible to infer the uncertainty on the carbon balance. Overall, the site was found to have a net carbon uptake of ≈5 t C ha(-1) year(-1), but the effects of the drought of 2005 were still noticed in 2006, when the climate disturbance caused the site to become a net source of carbon to the atmosphere. Different regions of the Amazon forest might respond differently to climate extremes due to differences in dry season length, annual precipitation, species compositions, albedo and soil type. Longer time series of fluxes measured over several locations are required to better characterize the effects of climate anomalies on the carbon and water balances for the whole Amazon region. Such valuable datasets can also be used to calibrate biogeochemical models and infer on future scenarios of the Amazon forest carbon balance under the influence of climate change.