Soil moisture supply and atmospheric demand for water independently limit-and profoundly a ect-vegetation productivity and water use during periods of hydrologic stress [1][2][3][4] . Disentangling the impact of these two drivers on ecosystem carbon and water cycling is di cult because they are often correlated, and experimental tools for manipulating atmospheric demand in the field are lacking. Consequently, the role of atmospheric demand is often not adequately factored into experiments or represented in models 5-7 . Here we show that atmospheric demand limits surface conductance and evapotranspiration to a greater extent than soil moisture in many biomes, including mesic forests that are of particular importance to the terrestrial carbon sink 8,9 . Further, using projections from ten general circulation models, we show that climate change will increase the importance of atmospheric constraints to carbon and water fluxes in all ecosystems. Consequently, atmospheric demand will become increasingly important for vegetation function, accounting for >70% of growing season limitation to surface conductance in mesic temperate forests. Our results suggest that failure to consider the limiting role of atmospheric demand in experimental designs, simulation models and land management strategies will lead to incorrect projections of ecosystem responses to future climate conditions. Ecosystem moisture stress is often characterized by changes in soil water availability 10,11 . Declining soil moisture impedes the movement of water to evaporating sites at the soil or leaf surface 12 , reducing the surface conductance to water vapour (G S )-a key determinant of carbon and water cycling-and thereby evapotranspiration (ET). However, atmospheric demand for water, which is directly related to the atmospheric vapour pressure deficit (VPD), also affects G S and ET. Plants close their stomata to prevent excessive water loss when VPD is high [13][14][15][16] and thus, increases in VPD during periods of hydrologic stress represent an independent constraint on plant carbon uptake and water use in ecosystems.While the plant physiological community has long recognized the critical role of VPD in determining plant functioning, VPD is often overlooked in many fields of hydrologic and climate science. For example, precipitation manipulation experiments are frequently used to draw conclusions about ecosystem response to drought stress, even though VPD is unaffected by precipitation manipulation 10 . Some terrestrial ecosystem and ecohydrological models do not permit stomatal conductance to vary with atmospheric demand 5,11 . Many models designed to capture these impacts rely on empirical parameterizations for soil moisture and VPD stress that promote compensating effects and model equifinality 5 , and/or use relative humidity instead of VPD as the primary driver, with significant consequences for projections of

The FLUXNET2015 dataset provides ecosystem-scale data on CO 2 , water, and energy exchange between the biosphere and the atmosphere, and other meteorological and biological measurements, from 212 sites around the globe (over 1500 site-years, up to and including year 2014). These sites, independently managed and operated, voluntarily contributed their data to create global datasets. Data were quality controlled and processed using uniform methods, to improve consistency and intercomparability across sites. The dataset is already being used in a number of applications, including ecophysiology studies, remote sensing studies, and development of ecosystem and Earth system models. FLUXNET2015 includes derived-data products, such as gap-filled time series, ecosystem respiration and photosynthetic uptake estimates, estimation of uncertainties, and metadata about the measurements, presented for the first time in this paper. In addition, 206 of these sites are for the first time distributed under a Creative Commons (CC-BY 4.0) license. This paper details this enhanced dataset and the processing methods, now made available as open-source codes, making the dataset more accessible, transparent, and reproducible.

Abstract. Trees, the most successful biological power plants on earth, build and plumb the critical zone (CZ) in ways that we do not yet understand. To encourage exploration of the character and implications of interactions between trees and soil in the CZ, we propose nine hypotheses that can be tested at diverse settings. The hypotheses are roughly divided into those about the architecture (building) and those about the water (plumbing) in the CZ, but the two functions are intertwined. Depending upon one's disciplinary background, many of the nine hypotheses listed below may appear obviously true or obviously false. (1) Tree roots can only physically penetrate and biogeochemically comminute the immobile substrate underlying mobile soil where that underlying substrate is fractured or pre-weathered. (2) In settings where the thickness of weathered material, H , is large, trees primarily shape the CZ through biogeochemical reactions within the rooting zone. (3) In forested uplands, the thickness of mobile soil, h, can evolve toward a steady state because of feedbacks related to root disruption and tree throw. (4) In settings where h H and the rates of uplift and erosion are low, the uptake of phosphorus into trees is buffered by the fine-grained fraction of the soil, and the ultimate source of this phosphorus is dust. (5) In settings of limited water availability, trees maintain the highest length density of functional roots at depths where water can be extracted over most of the growing season with the least amount of energy expenditure. (6) Trees grow the majority of their roots in the zone where the most growth-limiting resource is abundant, but they also grow roots at other depths to forage for other resources and to hydraulically redistribute those resources to depths where they can be taken up more efficiently. (7) Trees rely on matrix water in the unsaturated zone that at times may have anPublished by Copernicus Publications on behalf of the European Geosciences Union. 5116 S. L. Brantley et al.: Reviews and syntheses: on the roles trees play in building and plumbing the critical zone isotopic composition distinct from the gravity-drained water that transits from the hillslope to groundwater and streamflow. (8) Mycorrhizal fungi can use matrix water directly, but trees can only use this water by accessing it indirectly through the fungi. (9) Even trees growing well above the valley floor of a catchment can directly affect stream chemistry where changes in permeability near the rooting zone promote intermittent zones of water saturation and downslope flow of water to the stream. By testing these nine hypotheses, we will generate important new cross-disciplinary insights that advance CZ science.

Global-scale studies suggest that dryland ecosystems dominate an increasing trend in the magnitude and interannual variability of the land CO sink. However, such analyses are poorly constrained by measured CO exchange in drylands. Here we address this observation gap with eddy covariance data from 25 sites in the water-limited Southwest region of North America with observed ranges in annual precipitation of 100-1000 mm, annual temperatures of 2-25°C, and records of 3-10 years (150 site-years in total). Annual fluxes were integrated using site-specific ecohydrologic years to group precipitation with resulting ecosystem exchanges. We found a wide range of carbon sink/source function, with mean annual net ecosystem production (NEP) varying from -350 to +330 gCm across sites with diverse vegetation types, contrasting with the more constant sink typically measured in mesic ecosystems. In this region, only forest-dominated sites were consistent carbon sinks. Interannual variability of NEP, gross ecosystem production (GEP), and ecosystem respiration (R ) was larger than for mesic regions, and half the sites switched between functioning as C sinks/C sources in wet/dry years. The sites demonstrated coherent responses of GEP and NEP to anomalies in annual evapotranspiration (ET), used here as a proxy for annually available water after hydrologic losses. Notably, GEP and R were negatively related to temperature, both interannually within site and spatially across sites, in contrast to positive temperature effects commonly reported for mesic ecosystems. Models based on MODIS satellite observations matched the cross-site spatial pattern in mean annual GEP but consistently underestimated mean annual ET by ~50%. Importantly, the MODIS-based models captured only 20-30% of interannual variation magnitude. These results suggest the contribution of this dryland region to variability of regional to global CO exchange may be up to 3-5 times larger than current estimates.

A North American epidemic of mountain pine beetle (MPB) has disturbed over 5 million ha of forest containing headwater catchments crucial to water resources. However, there are limited observations of MPB effects on partitioning of precipitation between vapor loss and streamflow, and to our knowledge these fluxes have not been observed simultaneously following disturbance. We combined eddy covariance vapor loss (V), catchment streamflow (Q), and stable isotope indicators of evaporation (E) to quantify hydrologic partitioning over 3 years in MPB-impacted and control sites. Annual control V was conservative, varying only from 573 to 623 mm, while MPB site V varied more widely from 570 to 700 mm. During wet periods, MPB site V was greater than control V in spite of similar above-canopy potential evapotranspiration (PET). During a wet year, annual MPB V was greater and annual Q was lower as compared to an average year, while in a dry year, essentially all water was partitioned to V. Ratios of 2 H and 18 O in stream and soil water showed no kinetic evaporation at the control site, while MPB isotope ratios fell below the local meteoric water line, indicating greater E and snowpack sublimation (S s ) counteracted reductions in transpiration (T) and sublimation of canopy-intercepted snow (S c ). Increased E was possibly driven by reduced canopy shading of shortwave radiation, which averaged 21 W m 22 during summer under control forest as compared to 66 W m 22 under MPB forest. These results show that abiotic vapor losses may limit widely expected streamflow increases.

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[1] Feedbacks among vegetation dynamics, pedogenesis, and topographic development affect the "critical zone"-the living filter for Earth's hydrologic, biogeochemical, and rock/ sediment cycles. Assessing the importance of such feedbacks, which may be particularly pronounced in water-limited systems, remains a fundamental interdisciplinary challenge. The sky islands of southern Arizona offer an unusually well-defined natural experiment involving such feedbacks because mean annual precipitation varies by a factor of five over distances of approximately 10 km in areas of similar rock type (granite) and tectonic history. Here we compile high-resolution, spatially distributed data for Effective Energy and Mass Transfer (EEMT: the energy available to drive bedrock weathering), above-ground biomass, soil thickness, hillslope-scale topographic relief, and drainage density in two such mountain ranges (Santa Catalina: SCM; Pinaleño: PM). Strong correlations exist among vegetation-soil-topography variables, which vary nonlinearly with elevation, such that warm, dry, low-elevation portions of these ranges are characterized by relatively low above-ground biomass, thin soils, minimal soil organic matter, steep slopes, and high drainage densities; conversely, cooler, wetter, higher elevations have systematically higher biomass, thicker organic-rich soils, gentler slopes, and lower drainage densities. To test if eco-pedo-geomorphic feedbacks drive this pattern, we developed a landscape evolution model that couples pedogenesis and topographic development over geologic time scales, with rates explicitly dependent on vegetation density. The model self-organizes into states similar to those observed in SCM and PM. Our results highlight the potential importance of eco-pedo-geomorphic feedbacks, mediated by soil thickness, in water-limited systems.

HighlightWe studied for the first time the temporal and spatial limits within which active and passive chlorophyll fluorescence measurements are comparable.