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  1. Abstract Increases in hydrological extremes, including drought, are expected for Amazon forests. A fundamental challenge for predicting forest responses lies in identifying ecological strategies which underlie such responses. Characterization of species-specific hydraulic strategies for regulating water-use, thought to be arrayed along an ‘isohydric–anisohydric’ spectrum, is a widely used approach. However, recent studies have questioned the usefulness of this classification scheme, because its metrics are strongly influenced by environments, and hence can lead to divergent classifications even within the same species. Here, we propose an alternative approach positing that individual hydraulic regulation strategies emerge from the interaction of environments with traits. Specifically, we hypothesize that the vertical forest profile represents a key gradient in drought-related environments (atmospheric vapor pressure deficit, soil water availability) that drives divergent tree water-use strategies for coordinated regulation of stomatal conductance (gs) and leaf water potentials (ΨL) with tree rooting depth, a proxy for water availability. Testing this hypothesis in a seasonal eastern Amazon forest in Brazil, we found that hydraulic strategies indeed depend on height-associated environments. Upper canopy trees, experiencing high vapor pressure deficit (VPD), but stable soil water access through deep rooting, exhibited isohydric strategies, defined by little seasonal change in the diurnal pattern of gs and steady seasonal minimum ΨL. In contrast, understory trees, exposed to less variable VPD but highly variable soil water availability, exhibited anisohydric strategies, with fluctuations in diurnal gs that increased in the dry season along with increasing variation in ΨL. Our finding that canopy height structures the coordination between drought-related environmental stressors and hydraulic traits provides a basis for preserving the applicability of the isohydric-to-anisohydric spectrum, which we show here may consistently emerge from environmental context. Our work highlights the importance of understanding how environmental heterogeneity structures forest responses to climate change, providing a mechanistic basis for improving models of tropical ecosystems. 
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  2. Abstract 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). PartitioningETby an approach that assumes transpiration (T) is proportional to photosynthesis, we found that water stress‐induced reductions in canopy conductance (Gs) droveTdeclines partly compensated by higher evaporation (E). By contrast, the abnormally wet La Niña period gave higherTand lowerE, with little change in seasonalET. 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 ofGsand significant leaf shedding). Drought‐reducedTandGs, higherHandE, 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. 
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  3. Summary 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 structuredLAIalong axes of both canopy height and light environments.Upper canopyLAIincreased during the dry season, whereas lower canopyLAIdecreased. The low canopy decrease was driven by highly illuminated leaves of smaller trees in gaps. By contrast, understoryLAIincreased 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. 
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  4. null (Ed.)
    Plant ecophysiological trade-offs between different strategies for tolerating stresses are widely theorized to shape forest functional diversity and vulnerability to climate change. However, trade-offs between hydraulic and stomatal regulation during natural droughts remain under-studied, especially in tropical forests. We investigated eleven mature forest canopy trees in central Amazonia during the strong 2015 El Niño. We found greater xylem embolism resistance (P50 = − 3.3 ± 0.8 MPa) and hydraulic safety margin (HSM = 2.12 ± 0.57 MPa) than previously observed in more precipitation-seasonal rainforests of eastern Amazonia and central America. We also discovered that taller trees exhibited lower embolism resistance and greater stomatal sensitivity, a height-structured trade-off between hydraulic resistance and active stomatal regulation. Such active regulation of tree water status, triggered by the onset of stem embolism, acted as a feedback to avoid further increases in embolism, and also explained declines in photosynthesis and transpiration. These results suggest that canopy trees exhibit a conservative hydraulic strategy to endure drought, with trade-offs between investment in xylem to reduce vulnerability to hydraulic failure, and active stomatal regulation to protect against low water potentials. These findings improve our understanding of strategies in tropical forest canopies and contribute to more accurate prediction of drought responses. 
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    Abstract. Canopy stomatal conductance is commonly estimated fromeddy covariance measurements of the latent heat flux (LE) by inverting thePenman–Monteith equation. That method ignores eddy covariance measurementsof the sensible heat flux (H) and instead calculates H implicitly as theresidual of all other terms in the site energy budget. Here we show thatcanopy stomatal conductance is more accurately calculated from eddy covariance (EC)measurements of both H and LE using the flux–gradient equations that defineconductance and underlie the Penman–Monteith equation, especially when thesite energy budget fails to close due to pervasive biases in the eddy fluxesand/or the available energy. The flux–gradient formulation dispenses withunnecessary assumptions, is conceptually simpler, and is as or more accuratein all plausible scenarios. The inverted Penman–Monteith equation, on theother hand, contributes substantial biases and erroneous spatial andtemporal patterns to canopy stomatal conductance, skewing its relationshipswith drivers such as light and vapor pressure deficit. 
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  7. Canopy stomatal conductance (gsV) is commonly estimated from eddy covariance (EC) measurements of latent heat flux (LE) by inverting the Penman-Monteith (PM) equation. That method implicitly represents the sensible heat flux (H) as the residual of all other terms in the site energy budget – even though H is measured at least as accurately as LE at every EC site while the rest of the energy budget almost never is. We argue that gsV should instead be calculated from EC measurements of both H and LE, using the flux-gradient formulation that defines conductance and underlies the PM equation. The flux-gradient formulation dispenses with unnecessary assumptions, is conceptually simpler, and provides more accurate values of gsV for all plausible scenarios in which the measured energy budget fails to close, as is common at EC sites. The PM equation, on the other hand, contributes biases and erroneous spatial and temporal patterns to gsV, skewing its relationships with drivers such as light and vapor pressure deficit. To minimize the impact of the energy budget closure problem on the PM equation, it was previously proposed that the eddy fluxes should be corrected to close the long-term energy budget while preserving the Bowen ratio (B = H/LE). We show that such a flux correction does not fully remedy the PM equation but should produce accurate values of gsV when combined with the flux-gradient formulation. 
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