17Drought threatens tropical rainforests over seasonal to decadal timescales [1][2][3][4] , but the drivers 18 of tree mortality following drought remain poorly understood 5,6 . It has been suggested that 19 reduced availability of non-structural carbohydrates (NSC) critically increases mortality risk 20 through insufficient carbon supply to metabolism ('carbon starvation') 7,8 . However little is 21 known about how NSC stores are affected by drought, especially over the long term, and 22 whether they are more important than hydraulic processes in determining drought-induced 23 mortality. Using data from the world's longest-running experimental drought study in tropical 24 rainforest (in the Brazilian Amazon), we test whether carbon starvation or deterioration of the 25 water-conducting pathways from soil to leaf trigger tree mortality. Biomass loss from 26 mortality in the experimentally-droughted forest increased substantially after >10 years of 27 reduced soil moisture availability. The mortality signal was dominated by the death of large 28 trees, which were at a much greater risk of hydraulic deterioration than smaller trees. 29However, we find no evidence that the droughted trees suffered carbon starvation, as their 30 NSC concentrations were similar to those of un-droughted trees, and growth rates did not 31 decline in either living or dying individuals. Our results indicate that hydraulics, rather than 32 carbon starvation, triggers tree death from drought in tropical rainforest. 34Drought-response observations from both field-scale experiments and natural droughts have 35 demonstrated increased mortality over the short-term (1-3 years), with notably higher 36 vulnerability for some taxa, and for larger trees 6,9,10 . After several years of drought, 37 recovering growth rates in smaller trees, dbh (diameter at breast height) <40 cm, and reduced 38 mortality have been recorded at different locations 6,11,12 . However, the long-term (>10 yr) 39 sensitivity of tropical forests to predicted prolonged and repeated water deficit [1][2][3] we synthesise these data to test whether long-term soil moisture deficit alters NSC storage 64 and use in tropical rainforest trees, and if this, or hydraulic processes, are most strongly 65 associated with increased mortality rates. 66By 2014, following 13 years of the TFE treatment, cumulative biomass loss through mortality 67 was 41.0±2.7% relative to pre-treatment values (Fig. 1a), and the rate of loss had increased 68 substantially since the previous reported value of 17.2±0.8%, after 7 years of TFE 6 . 69Accelerating biomass loss and failure to recover substantially, or to reach a new 70 equilibrium 13 , has led to a committed flux to the atmosphere from decomposing necromass of 71 101.9±19.1 Mg C ha -1 (Fig. 1a). This biomass loss has been driven by elevated mortality in 72 the largest trees (Fig. 1b), as previously observed over shorter timescales 6 , and has created a 73 canopy that has had a persistently lower average leaf area index during 2010-2014 74 (12.0±1...

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The allocation and cycling of carbon (C) within forests is an important component of the biospheric C cycle, but is particularly understudied within tropical forests. We synthesise reported and unpublished results from three lowland rainforest sites in Amazonia (in the regions of Manaus, Tapajos and Caxiuana), all major sites of the Large-Scale Biosphere-Atmosphere Programme (LBA). We attempt a comprehensive synthesis of the C stocks, nutrient status and, particularly, the allocation and internal C dynamics of all three sites. The calculated net primary productivities (NPP) are 10.1 +/- 1.4 Mg C ha(-1) yr(-1) (Manaus), 14.4 +/- 1.3 Mg C ha(-1) yr(-1) (Tapajos) and 10.0 +/- 1.2 Mg C ha(-1) yr(-1) (Caxiuana). All errors bars report standard errors. Soil and leaf nutrient analyses indicate that Tapajos has significantly more plant-available phosphorus and calcium. Autotrophic respiration at all three sites (14.9-21.4 Mg C ha yr(-1)) is more challenging to measure, with the largest component and greatest source of uncertainty being leaf dark respiration. Comparison of measured soil respiration with that predicted from C cycling measurements provides an independent constraint. It shows general good agreement at all three sites, with perhaps some evidence for measured soil respiration being less than expected. Twenty to thirty percent of fixed C is allocated belowground. Comparison of gross primary productivity (GPP), derived from ecosystem flux measurements with that derived from component studies (NPP plus autotrophic respiration) provides an additional crosscheck. The two approaches are in good agreement, giving increased confidence in both approaches to estimating GPP. The ecosystem carbon-use efficiency (CUEs), the ratio of NPP to GPP, is similar at Manaus (0.34 +/- 0.10) and Caxiuana (0.32 +/- 0.07), but may be higher at Tapajos (0.49 +/- 0.16), although the difference is not significant. Old growth or infertile tropical forests may have low CUE compared with recently disturbed and/or fertile forests

Summary At least one climate model predicts severe reductions of rainfall over Amazonia during this century. Long‐term throughfall exclusion (TFE) experiments represent the best available means to investigate the resilience of the Amazon rainforest to such droughts. Results are presented from a 7 yr TFE study at Caxiuanã National Forest, eastern Amazonia. We focus on the impacts of the drought on tree mortality, wood production and above‐ground biomass. Tree mortality in the TFE plot over the experimental period was 2.5% yr−1, compared with 1.25% yr−1 in a nearby control plot experiencing normal rainfall. Differences in stem mortality between plots were greatest in the largest (> 40 cm diameter at breast height (dbh)) size class (4.1% yr−1 in the TFE and 1.4% yr−1 in the control). Wood production in the TFE plot was c. 30% lower than in the control plot. Together, these changes resulted in a loss of 37.8 ± 2.0 Mg carbon (C) ha−1 in the TFE plot (2002–2008), compared with no change in the control. These results are remarkably consistent with those from another TFE (at Tapajós National Forest), suggesting that eastern Amazonian forests may respond to prolonged drought in a predictable manner.

SummaryConsiderable 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.

a b s t r a c tWe investigated the seasonal patterns of Amazonian forest photosynthetic activity, and the effects thereon of variations in climate and land-use, by integrating data from a network of ground-based eddy flux towers in Brazil established as part of the 'Large-Scale Biosphere Atmosphere Experiment in Amazonia' project. We found that degree of water limitation, as indicated by the seasonality of the ratio of sensible to latent heat flux (Bowen ratio) predicts seasonal patterns of photosynthesis. In equatorial Amazonian forests (5• N-5 • S), water limitation is absent, and photosynthetic fluxes (or gross ecosystem productivity, GEP) exhibit high or increasing levels of photosynthetic activity as the dry season progresses, likely a consequence of allocation to growth of new leaves. In contrast, forests along the southern flank of the Amazon, pastures converted from forest, and mixed forest-grass savanna, exhibit dry-season declines in GEP, consistent with increasing degrees of water limitation. Although previous work showed tropical ecosystem evapotranspiration (ET) is driven by incoming radiation, GEP observations reported here surprisingly show no or negative relationships with photosynthetically active radiation (PAR). Instead, GEP fluxes largely followed the phenology of canopy photosynthetic capacity (Pc), with only deviations from this primary pattern driven by variations in PAR. Estimates of leaf flush at three * Corresponding author. Tel.: +1 520 6261500; fax: +1 520 621 9190. 182-183 (2013) 128-144 129 non-water limited equatorial forest sites peak in the dry season, in correlation with high dry season light levels. The higher photosynthetic capacity that follows persists into the wet season, driving high GEP that is out of phase with sunlight, explaining the negative observed relationship with sunlight. Overall, these patterns suggest that at sites where water is not limiting, light interacts with adaptive mechanisms to determine photosynthetic capacity indirectly through leaf flush and litterfall seasonality. These mechanisms are poorly represented in ecosystem models, and represent an important challenge to efforts to predict tropical forest responses to climatic variations.

Version 5 of the Community Land Model (CLM5) introduces the plant hydraulic stress (PHS) configuration of vegetation water use, which is described and compared with the corresponding parameterization from CLM4.5. PHS updates vegetation water stress and root water uptake to better reflect plant hydraulic theory, advancing the physical basis of the model. The new configuration introduces prognostic vegetation water potential, modeled at the root, stem, and leaf levels. Leaf water potential replaces soil potential as the basis for stomatal conductance water stress, and root water potential is used to implement hydraulic root water uptake, replacing a transpiration partitioning function. Point simulations of a tropical forest site (Caxiuanã, Brazil) under ambient conditions and partial precipitation exclusion highlight the differences between PHS and the previous CLM implementation. Model description and simulation results are contextualized with a list of benefits and limitations of the new model formulation, including hypotheses that were not testable in previous versions of the model. Key results include reductions in transpiration and soil moisture biases relative to a control model under both ambient and exclusion conditions, correcting excessive dry season soil moisture stress in the control model. PHS implements hydraulic gradient root water uptake, which allows hydraulic redistribution and compensatory root water uptake and results in PHS utilizing a larger portion of the soil column to buffer shortfalls in precipitation. The new model structure, which bases water stress on leaf water potential, could have significant implications for vegetation-climate feedbacks, including increased sensitivity of photosynthesis to atmospheric vapor pressure deficit.