Revealing legacy effects of extreme droughts on tree growth of oaks across the Northern Hemisphere.

Forests are undergoing increasing risks of drought-induced tree mortality. Species replacement patterns following mortality may have a significant impact on the global carbon cycle. Among major hardwoods, deciduous oaks (Quercus spp.) are increasingly reported as replacing dying conifers across the Northern Hemisphere. Yet, our knowledge on the growth responses of these oaks to drought is incomplete, especially regarding post-drought legacy effects. The objectives of this study were to determine the occurrence, duration, and magnitude of legacy effects of extreme droughts and how that vary across species, sites, and drought characteristics. The legacy effects were quantified by the deviation of observed from expected radial growth indices in the period 1940-2016. We used stand-level chronologies from 458 sites and 21 oak species primarily from Europe, north-eastern America, and eastern Asia. We found that legacy effects of droughts could last from 1 to 5 years after the drought and were more prolonged in dry sites. Negative legacy effects (i.e., lower growth than expected) were more prevalent after repetitive droughts in dry sites. The effect of repetitive drought was stronger in Mediterranean oaks especially in Quercus faginea. Species-specific analyses revealed that Q. petraea and Q. macrocarpa from dry sites were more negatively affected by the droughts while growth of several oak species from mesic sites increased during post-drought years. Sites showing positive correlations to winter temperature showed little to no growth depression after drought, whereas sites with a positive correlation to previous summer water balance showed decreased growth. This may indicate that although winter warming favors tree growth during droughts, previous-year summer precipitation may predispose oak trees to current-year extreme droughts. Our results revealed a massive role of repetitive droughts in determining legacy effects and highlighted how growth sensitivity to climate, drought seasonality and species-specific traits drive the legacy effects in deciduous oak species.

Contrasting impact of extreme soil and atmospheric dryness on the functioning of trees and forests.

Recent studies indicate an increase in the frequency of extreme compound dryness days (days with both extreme soil AND air dryness) across central Europe in the future, with little information on their impact on the functioning of trees and forests. This study aims to quantify and assess the impact of extreme soil dryness, extreme air dryness, and extreme compound dryness on the functioning of trees and forests. For this, >15 years of ecosystem-level (carbon dioxide and water vapor fluxes) and 6-10 years of tree-level measurements (transpiration and growth) each from a montane mixed deciduous forest (CH-Lae) and a subalpine evergreen coniferous forest (CH-Dav) in Switzerland, is used. The results showed extreme air dryness limitation on CO2 fluxes and extreme soil dryness limitations on water vapor fluxes. Additionally, CH-Dav was mainly affected by extreme air dryness whereas CH-Lae was affected by both extreme soil dryness and extreme air dryness. The impact of extreme compound dryness on net CO2 uptake (about 75 % decrease) was more due to higher increased ecosystem respiration (40 % and 70 % increase at CH-Dav and CH-Lae, respectively) than decreased gross primary productivity (10 % and 40 % decrease at CH-Dav and CH-Lae, respectively). A significant negative impact on evapotranspiration and transpiration was only observed at CH-Lae during extreme soil and compound dryness (about 25 % decrease). Furthermore, with some differences, the tree-level impact on tree water deficit, transpiration, and growth were consistent with the ecosystem-level impact on carbon uptake and evapotranspiration. Finally, the impact of extreme dryness showed no significant relationship with tree allometry (diameter and height) but across different tree species. The projected future is likely to expose these forest areas to more extreme and frequent dryness conditions, thus compromising the functioning of trees and forests, thereby calling for management interventions to increase the adaptive capacity and resistance of these forests.

Networking the forest infrastructure towards near real-time monitoring - A white paper.

Forests account for nearly 90 % of the world's terrestrial biomass in the form of carbon and they support 80 % of the global biodiversity. To understand the underlying forest dynamics, we need a long-term but also relatively high-frequency, networked monitoring system, as traditionally used in meteorology or hydrology. While there are numerous existing forest monitoring sites, particularly in temperate regions, the resulting data streams are rarely connected and do not provide information promptly, which hampers real-time assessments of forest responses to extreme climate events. The technology to build a better global forest monitoring network now exists. This white paper addresses the key structural components needed to achieve a novel meta-network. We propose to complement - rather than replace or unify - the existing heterogeneous infrastructure with standardized, quality-assured linking methods and interacting data processing centers to create an integrated forest monitoring network. These automated (research topic-dependent) linking methods in atmosphere, biosphere, and pedosphere play a key role in scaling site-specific results and processing them in a timely manner. To ensure broad participation from existing monitoring sites and to establish new sites, these linking methods must be as informative, reliable, affordable, and maintainable as possible, and should be supplemented by near real-time remote sensing data. The proposed novel meta-network will enable the detection of emergent patterns that would not be visible from isolated analyses of individual sites. In addition, the near real-time availability of data will facilitate predictions of current forest conditions (nowcasts), which are urgently needed for research and decision making in the face of rapid climate change. We call for international and interdisciplinary efforts in this direction.

Number of growth days and not length of the growth period determines radial stem growth of temperate trees.

Radial stem growth dynamics at seasonal resolution are essential to understand how forests respond to climate change. We studied daily radial growth of 160 individuals of seven temperate tree species at 47 sites across Switzerland over 8 years. Growth of all species peaked in the early part of the growth season and commenced shortly before the summer solstice, but with species-specific seasonal patterns. Day length set a window of opportunity for radial growth. Within this window, the probability of daily growth was constrained particularly by air and soil moisture, resulting in intermittent growth to occur only on 29 to 77 days (30% to 80%) within the growth period. The number of days with growth largely determined annual growth, whereas the growth period length contributed less. We call for accounting for these non-linear intra-annual and species-specific growth dynamics in tree and forest models to reduce uncertainties in predictions under climate change.

Climate sensitivity and drought seasonality determine post-drought growth recovery of Quercus petraea and Quercus robur in Europe.

Recent studies have identified strong relationships between delayed recovery of tree growth after drought and tree mortality caused by subsequent droughts. These observations raise concerns about forest ecosystem services and post-drought growth recovery given the projected increase in drought frequency and extremes. For quantifying the impact of extreme droughts on tree radial growth, we used a network of tree-ring width data of 1689 trees from 100 sites representing most of the distribution of two drought tolerant, deciduous oak species (Quercus petraea and Quercus robur). We first examined which climatic factors and seasons control growth of the two species and if there is any latitudinal, longitudinal or elevational trend. We then quantified the relative departure from pre-drought growth during droughts, and how fast trees were able to recover the pre-drought growth level. Our results showed that growth was more related to precipitation and climatic water balance (precipitation minus potential evapotranspiration) than to temperature. However, we did not detect any clear latitudinal, longitudinal or elevational trends except a decreasing influence of summer water balance on growth of Q. petraea with latitude. Neither species was able to maintain the pre-drought growth level during droughts. However, both species showed rapid recovery or even growth compensation after summer droughts but displayed slow recovery in response to spring droughts where none of the two species was able to fully recover the pre-drought growth-level over the three post-drought years. Collectively, our results indicate that oaks which are considered resilient to extreme droughts have also shown vulnerability when droughts occurred in spring especially at sites where long-term growth is not significantly correlated with climatic factors. This improved understanding of the role of drought seasonality and climate sensitivity of sites is key to better predict trajectories of post-drought growth recovery in response to the drier climate projected for Europe.

Why trees grow at night.

The timing of diel stem growth of mature forest trees is still largely unknown, as empirical data with high temporal resolution have not been available so far. Consequently, the effects of day-night conditions on tree growth remained uncertain. Here we present the first comprehensive field study of hourly-resolved radial stem growth of seven temperate tree species, based on 57 million underlying data points over a period of up to 8 yr. We show that trees grow mainly at night, with a peak after midnight, when the vapour pressure deficit (VPD) is among the lowest. A high VPD strictly limits radial stem growth and allows little growth during daylight hours, except in the early morning. Surprisingly, trees also grow in moderately dry soil when the VPD is low. Species-specific differences in diel growth dynamics show that species able to grow earlier during the night are associated with the highest number of hours with growth per year and the largest annual growth increment. We conclude that species with the ability to overcome daily water deficits faster have greater growth potential. Furthermore, we conclude that growth is more sensitive than carbon uptake to dry air, as growth stops before stomata are known to close.

Physiological response of Swiss ecosystems to 2018 drought across plant types and elevation.

Using five eddy covariance flux sites (two forests and three grasslands), we investigated ecosystem physiological responses to the 2018 drought across elevational gradients in Switzerland. Flux measurements showed that at lower elevation sites (below 1000 m.a.s.l.; grassland and mixed forest) annual ecosystem productivity (GPP) declined by approximately 20% compared to the previous 2 years (2016 and 2017), which led to a reduced annual net ecosystem productivity (NEP). At the high elevation sites, however, GPP increased by approximately 14% and as a result NEP increased in the alpine and montane grasslands, but not in the subalpine coniferous forest. There, increased ecosystem respiration led to a reduced annual NEP, despite increased GPP and lengthening of the growing period. Among all ecosystems, the coniferous forest showed the most pronounced negative stomatal response to atmospheric dryness (i.e. vapour pressure deficit, VPD) that resulted in a decline in surface conductance and an increased water-use efficiency during drought. While increased temperature enhanced the water-use efficiency of both forests, de-coupling of GPP from evapotranspiration at the low-elevation grassland site negatively affected water-use efficiency due to non-stomatal reductions in photosynthesis. Our results show that hot droughts (such as in 2018) lead to different responses across plants types, and thus ecosystems. Particularly grasslands at lower elevations are the most vulnerable ecosystems to negative impacts of future drought in Switzerland. This article is part of the theme issue 'Impacts of the 2018 severe drought and heatwave in Europe: from site to continental scale'.

Determinants of legacy effects in pine trees - implications from an irrigation-stop experiment.

Tree responses to altered water availability range from immediate (e.g. stomatal regulation) to delayed (e.g. crown size adjustment). The interplay of the different response times and processes, and their effects on long-term whole-tree performance, however, is hardly understood. Here we investigated legacy effects on structures and functions of mature Scots pine in a dry inner-Alpine Swiss valley after stopping an 11-yr lasting irrigation treatment. Measured ecophysiological time series were analysed and interpreted with a system-analytic tree model. We found that the irrigation stop led to a cascade of downregulations of physiological and morphological processes with different response times. Biophysical processes responded within days, whereas needle and shoot lengths, crown transparency, and radial stem growth reached control levels after up to 4 yr only. Modelling suggested that organ and carbon reserve turnover rates play a key role for a tree's responsiveness to environmental changes. Needle turnover rate was found to be most important to accurately model stem growth dynamics. We conclude that leaf area and its adjustment time to new conditions is the main determinant for radial stem growth of pine trees as the transpiring area needs to be supported by a proportional amount of sapwood, despite the growth-inhibiting environmental conditions.

Towards comparable assessment of the soil nutrient status across scales-Review and development of nutrient metrics.

Nutrient availability influences virtually every aspect of an ecosystem, and is a critical modifier of ecosystem responses to global change. Although this crucial role of nutrient availability in regulating ecosystem structure and functioning has been widely acknowledged, nutrients are still often neglected in observational and experimental synthesis studies due to difficulties in comparing the nutrient status across sites. In the current study, we explain different nutrient-related concepts and discuss the potential of soil-, plant- and remote sensing-based metrics to compare the nutrient status across space. Based on our review and additional analyses on a dataset of European, managed temperate and boreal forests (ICP [International Co-operative Programme on Assessment and Monitoring of Air Pollution Effects on Forests] Forests dataset), we conclude that the use of plant- and remote sensing-based metrics that rely on tissue stoichiometry is limited due to their strong dependence on species identity. The potential use of other plant-based metrics such as Ellenberg indicator values and plant-functional traits is also discussed. We conclude from our analyses and review that soil-based metrics have the highest potential for successful intersite comparison of the nutrient status. As an example, we used and adjusted a soil-based metric, previously developed for conifer forests across Sweden, against the same ICP Forests data. We suggest that this adjusted and further adaptable metric, which included the organic carbon concentration in the upper 20 cm of the soil (including the organic fermentation-humus [FH] layer), the C:N ratio and pHCaCl2 of the FH layer, can be used as a complementary tool along with other indicators of nutrient availability, to compare the background nutrient status across temperate and boreal forests dominated by spruce, pine or beech. Future collection and provision of harmonized soil data from observational and experimental sites is crucial for further testing and adjusting the metric.

One Century of Forest Monitoring Data in Switzerland Reveals Species- and Site-Specific Trends of Climate-Induced Tree Mortality.

Climate-induced tree mortality became a global phenomenon during the last century and it is expected to increase in many regions in the future along with a further increase in the frequency of drought and heat events. However, tree mortality at the ecosystem level remains challenging to quantify since long-term, tree-individual, reliable observations are scarce. Here, we present a unique data set of monitoring records from 276 permanent plots located in 95 forest stands across Switzerland, which include five major European tree species (Norway spruce, Scots pine, silver fir, European beech, and sessile and common oak) and cover a time span of over one century (1898-2013), with inventory periods of 5-10 years. The long-term average annual mortality rate of the investigated forest stands was 1.5%. In general, species-specific annual mortality rates did not consistently increase over the last decades, except for Scots pine forests at lower altitudes, which exhibited a clear increase of mortality since the 1960s. Temporal trends of tree mortality varied also depending on diameter at breast height (DBH), with large trees generally experiencing an increase in mortality, while mortality of small trees tended to decrease. Normalized mortality rates were remarkably similar between species and a modest, but a consistent and steady increasing trend was apparent throughout the study period. Mixed effects models revealed that gradually changing stand parameters (stand basal area and stand age) had the strongest impact on mortality rates, modulated by climate, which had increasing importance during the last decades. Hereby, recent climatic changes had highly variable effects on tree mortality rates, depending on the species in combination with abiotic and biotic stand and site conditions. This suggests that forest species composition and species ranges may change under future climate conditions. Our data set highlights the complexity of forest dynamical processes such as long-term, gradual changes of forest structure, demography and species composition, which together with climate determine mortality rates.

Long-term stem CO2 concentration measurements in Norway spruce in relation to biotic and abiotic factors.

Stem CO(2) concentrations (stem [CO(2)]) undergo large temporal variations that need to be understood to better link tree physiological processes to biosphere-atmosphere CO(2) exchange. During 19 months, stem [CO(2)] was continuously measured in mature subalpine Norway spruce trees (Picea abies) and jointly analysed with stem, soil and air temperatures, sap flow rates, stem radius changes and CO(2) efflux rates from stem and soil on different time scales. Stem [CO(2)] exhibited a strong seasonality, of which over 80% could be explained with stem and soil temperatures. Both physical equilibrium processes of CO(2) between water and air according to Henry's law as well as physiological effects, including sap flow and local respiration, concurrently contributed to these temporal variations. Moreover, the explanatory power of potential biological drivers (stem radius changes, sap flow and soil respiration) varied strongly with season and temporal resolution. We conclude that seasonal and daily courses of stem [CO(2)] in spruce trees are a combined effect of physical equilibrium and tree physiological processes. Furthermore, we emphasize the relevance of axial diffusion of CO(2) along air-filled spaces in the wood, and potential wound response processes owing to sensor installation.

Thermal optimality of net ecosystem exchange of carbon dioxide and underlying mechanisms.

• It is well established that individual organisms can acclimate and adapt to temperature to optimize their functioning. However, thermal optimization of ecosystems, as an assemblage of organisms, has not been examined at broad spatial and temporal scales. • Here, we compiled data from 169 globally distributed sites of eddy covariance and quantified the temperature response functions of net ecosystem exchange (NEE), an ecosystem-level property, to determine whether NEE shows thermal optimality and to explore the underlying mechanisms. • We found that the temperature response of NEE followed a peak curve, with the optimum temperature (corresponding to the maximum magnitude of NEE) being positively correlated with annual mean temperature over years and across sites. Shifts of the optimum temperature of NEE were mostly a result of temperature acclimation of gross primary productivity (upward shift of optimum temperature) rather than changes in the temperature sensitivity of ecosystem respiration. • Ecosystem-level thermal optimality is a newly revealed ecosystem property, presumably reflecting associated evolutionary adaptation of organisms within ecosystems, and has the potential to significantly regulate ecosystem-climate change feedbacks. The thermal optimality of NEE has implications for understanding fundamental properties of ecosystems in changing environments and benchmarking global models.