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The increasing frequency of global change-type droughts has created a need for fast, accurate and widely applicable techniques for estimating xylem embolism resistance to improve forecasts of future forest changes. We used data from 12 diffuse-porous temperate tree species covering a wide range of xylem safety to compare the pneumatic and flow-centrifuge method, two rapid methods used for constructing xylem vulnerability curves. We evaluated the agreement between parameters estimated with both methods and the sensitivity of pneumatic measurements to the duration of air discharge (AD) measurements. There was close agreement between xylem water potentials at 50% air discharged (PAD), estimated with the Pneumatron, and 50% loss of hydraulic conductivity (PLC), estimated with the flow-centrifuge method (mean signed deviation: 0.12 MPa, Pearson correlation: 0.96 after 15 s of gas extraction). However, the relationship between the estimated slopes was more variable, resulting in lower agreement in the xylem water potential at 12% and 88% PAD/PLC. The agreement between the two methods was not affected by species-specific vessel length distributions. All pneumatic parameters were sensitive to AD time. Overall agreement was highest at relatively short AD times, with an optimum at 16 s. Our results highlight the value of the Pneumatron as an easy and reliable tool to estimate 50% embolism thresholds for a wide range of diffuse-porous temperate angiosperms. Further, our study provides a set of useful metrics for methodological comparisons of vulnerability curves in terms of systematic and random deviations, as well as overall agreement.

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We took advantage of the European 2018 drought and assessed the mechanisms causing differences in drought vulnerability among mature individuals of nine co-occurring tree species at the Swiss Canopy Crane II site in Switzerland. Throughout the drought we monitored leaf water status and determined native embolism formation in the canopy of the trees as indicators of drought vulnerability. We also determined hydraulic vulnerability thresholds (Ψ12 -, Ψ50 - and Ψ88 -values), corresponding hydraulic safety margins (HSMs) and carbohydrate reserves for all species as well as total average leaf area per tree, and used stable isotopes to assess differences in root water uptake depth among the nine species as variables predicting differences in drought vulnerability among species. Marked differences in drought vulnerability were observed among the nine tree species. Six species maintained their water potentials above hydraulic thresholds, while three species, Fagus sylvatica, Carpinus betulus and Picea abies, were pushed beyond their hydraulic thresholds and showed loss of hydraulic conductivity in their canopies at the end of the drought. Embolism resistance thresholds and associated HSMs did not explain why the co-existing species differed in their drought vulnerability, neither did their degree of isohydry, nor their regulation of carbohydrate reserves. Instead, differences in structural-morphological traits, in particular root water uptake depth, were associated with the risk of reaching hydraulic vulnerability thresholds and embolism formation among the nine species. Our study shows that structural-morphological traits, such as root water uptake depth, determine how quickly different species approach hydraulic vulnerability thresholds during a drought event and can thus explain species differences in drought vulnerability among mature field-grown trees.

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Leaf morphological and physiological traits control the carbon and water relations of mature trees and are determinants of drought tolerance, but it is not well understood how they are modified in response to water deficits. We analysed five sun-canopy leaf traits (mean leaf size (LS), specific leaf area (SLA), Huber value (HV), water potential at turgor loss point (Ψtlp ) and foliar carbon isotope signature (δ13 C)) in European beech (Fagus sylvatica L.) across three precipitation gradients sampled in moist (2010), dry (2019) and very dry (2018) summers, and tested their response to short-term water deficits (climatic water balance (CWB) preceding sample collection) and long-term water availability (mean annual precipitation (MAP), plant-available soil water capacity (AWC) and neighbourhood competition). Across the 34 sites, LS varied seven-fold (3.9-27.0 cm2 ), SLA four-fold (77.1-306.9 cm²·g-1 ) and HV six-fold (1.0-6.65 cm2 ·m-2 ). In the 2018 dataset, LS showed a negative and HV a positive relationship to MAP, which contradicts relations found in multi-species samples. Average Ψtlp ranged from -1.90 to -2.62 MPa and decreased across the sites with decreasing CWB in the month prior to measurement, as well as with decreasing MAP and AWC in 2019. Studied leaf traits varied considerably between years, suggesting that mast fruiting and the severe 2018 drought caused the formation of smaller leaves. We conclude that sun-canopy leaf traits of European beech exhibit considerable plasticity in response to climatic and edaphic aridity, and that osmotic adjustment may be an important element in the drought response strategy of this anisohydric tree species.

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