Dr. Roman Zweifel

Increasing outbreaks of invasive insect pests pose a substantial threat to the functioning and viability of cultivated and wild woody perennial species worldwide. In the eastern U.S., the spotted lanternfly (Lycorma delicatula White; SLF), an invasive phloem-feeding planthopper, was reported to be able to negatively impact late-season plant carbon dynamics in various woody perennial species following repeated or prolonged infestation and feeding events. However, it remains unclear if SLF infestations also impact plant water relations and if SLF-mediated effects persist when populations are controlled and feeding stops. This study investigated how late-season exposure to SLF impacts whole-plant water relations by assessing diurnal sap flow and trunk radius changes in grapevines subjected to varying infestation densities of adult SLF. In two seasons, vines exposed to high infestation densities (an average of 180 SLFs per vine) for up to 31–32 days of cumulative SLF exposure had significantly lower sap flow rate than those with no SLF, resulting in up to 38 % less daily total water use. Trunk diurnal amplitudes increased under SLF infestation, but impacts were less dependent on infestation density, suggesting that grapevines may be utilizing trunk water storage during infestation to meet both grapevine water use and SLF sap ingestion. In both cases, SLF-mediated effects persisted following removal of SLFs, suggesting that exposure to high populations of SLF can alter patterns of late-season grapevine water use, at least when populations are not effectively controlled. These results indicate that SLF can modify both whole-plant water relations and carbon dynamics concurrently, further defining the implications that intensive infestations by high populations of SLF have for woody perennial whole-plant physiology.

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.

Climate change is predicted to increase atmospheric vapor pressure deficit, exacerbating soil drought, and thus enhancing tree evaporative demand and mortality. Yet, few studies have addressed the longer-term drought acclimation strategy of trees, particularly the importance of morphological versus hydraulic plasticity. Using a long-term (20 years) irrigation experiment in a natural forest, we investigated the acclimation of Scots pine (Pinus sylvestris) morpho-anatomical traits (stomatal anatomy and crown density) and hydraulic traits (leaf water potential, vulnerability to cavitation (ψ₅₀), specific hydraulic conductivity (K_s), and tree water deficit) to prolonged changes in soil moisture. We found that low water availability reduced twig water potential and increased tree water deficit during the growing season. Still, the trees showed limited adjustments in most branch-level hydraulic traits (ψ₅₀ and K_s) and needle anatomy. In contrast, trees acclimated to prolonged irrigation by increasing their crown density and hence the canopy water demand. This study demonstrates that despite substantial canopy adjustments, P. sylvestris may be vulnerable to extreme droughts because of limited adjustment potential in its hydraulic system. While sparser canopies reduce water demand, such shifts take decades to occur under chronic water deficits and might not mitigate short-term extreme drought events.

A conceptual understanding on how the vegetation's carbon (C) balance is determined by source activity and sink demand is important to predict its C uptake and sequestration potential now and in the future. We have gathered trajectories of photosynthesis and growth as a function of environmental conditions described in the literature and compared them with current concepts of source and sink control. There is no clear evidence for pure source or sink control of the C balance, which contradicts recent hypotheses. Using model scenarios, we show how legacy effects via structural and functional traits and antecedent environmental conditions can alter the plant's carbon balance. We, thus, combined the concept of short-term source–sink coordination with long-term environmentally driven legacy effects that dynamically acclimate structural and functional traits over time. These acclimated traits feedback on the sensitivity of source and sink activity and thus change the plant physiological responses to environmental conditions. We postulate a whole plant C-coordination system that is primarily driven by stomatal optimization of growth to avoid a C source–sink mismatch. Therefore, we anticipate that C sequestration of forest ecosystems under future climate conditions will largely follow optimality principles that balance water and carbon resources to maximize growth in the long term.

Qua o là sta amo ün tal eroe plain fideltà sün seis post, cun büst sfrattamà, e stenda il bratsch our da spelm e grava, sco üna sgür pronta pel cumbat, o ch'el sta ingramüschà sco ün vegliet cha la burasca da la vita ha fat sgobar sün sias crotschas.

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.

Scots pine (Pinus sylvestris L.) is a common European tree species, and understanding its acclimation to the rapidly changing climate through physiological, biochemical or structural adjustments is vital for predicting future growth. We investigated a long-term irrigation experiment at a naturally dry forest in Switzerland, comparing Scots pine trees that have been continuously irrigated for 17 years (irrigated) with those for which irrigation was interrupted after 10 years (stop) and non-irrigated trees (control), using tree growth, xylogenesis, wood anatomy, and carbon, oxygen and hydrogen stable isotope measurements in the water, sugars and cellulose of plant tissues. The dendrochronological analyses highlighted three distinct acclimation phases to the treatments: irrigated trees experienced (i) a significant growth increase in the first 4 years of treatment, (ii) high growth rates but with a declining trend in the following 8 years and finally (iii) a regression to pre-irrigation growth rates, suggesting the development of a new growth limitation (i.e. acclimation). The introduction of the stop treatment resulted in further growth reductions to below-control levels during the third phase. Irrigated trees showed longer growth periods and lower tree-ring δ¹³C values, reflecting lower stomatal restrictions than control trees. Their strong tree-ring δ¹⁸O and δ²H (O–H) relationship reflected the hydrological signature similarly to the control. On the contrary, the stop trees had lower growth rates, conservative wood anatomical traits, and a weak O–H relationship, indicating a physiological imbalance. Tree vitality (identified by crown transparency) significantly modulated growth, wood anatomical traits and tree-ring δ¹³C, with low-vitality trees of all treatments performing similarly regardless of water availability. We thus provide quantitative indicators for assessing physiological imbalance and tree acclimation after environmental stresses. We also show that tree vitality is crucial in shaping such responses. These findings are fundamental for the early assessment of ecosystem imbalances and decline under climate change.

In an increasingly dry environment, it is crucial to understand how tree species use soil water and cope with drought. However, there is still a knowledge gap regarding the relationships between species-specific stomatal behaviour, spatial root distribution, and root water uptake (RWU) dynamics.
Our study aimed to investigate above- and below-ground aspects of water use during soil drying periods in four temperate tree species that differ in stomatal behaviour: two isohydric tracheid-bearing conifers, Scots pine and Norway spruce, and two more anisohydric deciduous species, the diffuse-porous European beech, and the ring-porous Downy oak. From 2015 to 2020, soil-tree-atmosphere-continuum parameters were measured for each species in monospecific forests where trees had no access to groundwater. The hourly time series included data on air temperature, vapor pressure deficit, soil water potential, soil hydraulic conductivity, and RWU to a depth of 2 m. Analysis of drought responses included data on stem radius, leaf water potential, estimated osmotically active compounds, and drought damage.
Our study reveals an inherent coordination between stomatal regulation, fine root distribution and water uptake. Compared to conifers, the more anisohydric water use of oak and beech was associated with less strict stomatal closure, greater investment in deep roots, four times higher maximum RWU, a shift of RWU to deeper soil layers as the topsoil dried, and a more pronounced soil drying below 1 m depth. Soil hydraulic conductivity started to limit RWU when values fell below 10⁻³ to 10⁻⁵ cm/d, depending on the soil. As drought progressed, oak and beech may also have benefited from their leaf osmoregulatory capacity, but at the cost of xylem embolism with around 50 % loss of hydraulic conductivity when soil water potential dropped below −1.25 MPa.
Consideration of species-specific water use is crucial for forest management and vegetation modelling to improve forest resilience to drought.

Radial stem growth is a key ecosystem process resulting in long-term carbon sequestration. Despite recognition of its importance to global carbon cycling, high uncertainties remain regarding how radial growth phenology (e.g., the onset, mid, and cessation of radial growth) will be affected by climate change. In this study, we evaluated to what extent high spatially (3 × 3 m) and temporally (up to daily) resolved satellite imagery from PlanetScope can be used to monitor stem growth phenology. For this, we made use of detailed stem growth phenological observations of six common European tree species measured by automated point dendrometers at 14 distinct sites across Switzerland between 2017 and 2021. These growth phenological observations were then linked through multiple regression modeling with metrics extracted from spectral index time series. Our results show that the remote sensing-based models enable monitoring the onset (root mean squared deviation (RMSD) ranges from 5.96 to 27.04 days) and mid-stages of stem growth (RMSD ranges from 10.20 to 36.34 days) with reasonable accuracy as opposed to the cessation of stem growth that showed low accuracy (RMSD ranges from 16.02 to 153.63 days). The accuracy of the remote sensing-based prediction models and their optimal suite of predictors varied across species. The latter has important implications for the remote sensing of stem growth phenology in mixed forests, suggesting that it is important for satellite sensors to resolve individual tree crowns. Overall, our results suggest the need for novel spectral indices that capture the spectral components of mechanistic linkages between stem growth and canopy properties that go beyond the mere detection of leaf phenology. When employing such spectral indices, remote sensing could make it possible to detect not only shifts in leaf phenology caused by climate change but also those in stem growth on a broad spatial scale.

Trees remain sufficiently hydrated during drought by closing stomata and reducing canopy conductance (G_c) in response to variations in atmospheric water demand and soil water availability. Thresholds that control the reduction of G_c are proposed to optimize hydraulic safety against carbon assimilation efficiency. However, the link between G_c and the ability of stem tissues to rehydrate at night remains unclear.
We investigated whether species-specific G_c responses aim to prevent branch embolisms, or enable night-time stem rehydration, which is critical for turgor-dependent growth. For this, we used a unique combination of concurrent dendrometer, sap flow and leaf water potential measurements and collected branch-vulnerability curves of six common European tree species.
Species-specific G_c reduction was weakly related to the water potentials at which 50% of branch xylem conductivity is lost (P₅₀). Instead, we found a stronger relationship with stem rehydration. Species with a stronger G_c control were less effective at refilling stem-water storage as the soil dries, which appeared related to their xylem architecture.
Our findings highlight the importance of stem rehydration for water-use regulation in mature trees, which likely relates to the maintenance of adequate stem turgor. We thus conclude that stem rehydration must complement the widely accepted safety–efficiency stomatal control paradigm.

Wood is a sustainable natural resource and an important global commodity. According to the 'moon wood theory', the properties of wood, including its growth and water content, are believed to oscillate with the lunar cycle. Despite contradicting our current understanding of plant functioning, this theory is commonly exploited for marketing wooden products. To examine the moon wood theory, we applied a wavelet power transformation to series of 2,000,000 hourly stem radius records from dendrometers. We separated the influence of 74 consecutive lunar cycles and meteorological conditions on the stem variation of 62 trees and six species. We show that the dynamics of stem radius consist of overlapping oscillations with periods of 1 day, 6 months, and 1 year. These oscillations in stem dimensions were tightly coupled to oscillations in the series of air temperature and vapour pressure deficit. By contrast, we revealed no imprint of the lunar cycle on the stem radius variation of any species. We call for scepticism towards the moon wood theory, at least as far as the stem water content and radial growth are concerned. We foresee that similar studies employing robust scientific approaches will be increasingly needed in the future to cope with misleading concepts.

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.

Tree species differ in their physiological responses to drought, but the underlying causes are often unclear. Here we explored responses of radial growth to centennial drought events and sap flow (F_s) to seasonal drought in four mixed forests on either moist or drier sites in northwestern Switzerland. While the diffuse-porous species (Fagus sylvatica, Prunus avium, Tilia platyphyllos) showed marked growth reductions in 1976 and 2003, two known marker years for severe drought, growth of the two ring-porous species (Quercus petraea and Fraxinus excelsior) was less severely affected. During a dry early to midsummer, diffuse-porous species strongly reduced F_s at the two drier sites but not (or less so) at the two moister sites. Regardless of soil moisture availability, the deep-rooting, ring-porous trees invariably down-regulated F_s to 60–70% of their maxima in response to vapour pressure deficit (VPD) and maintained similar fluxes across sites, irrespective of upper soil moisture conditions. A generalised additive model of normalised F_s as a function of VPD and soil matric potential yielded a drought-sensitivity ranking of F_s led by the two insensitive ring-porous species followed by the diffuse-porous trees (ordered by increasing sensitivity: Fraxinus excelsior < Quercus petraea < Prunus avium < Acer pseudoplatanus < Fagus sylvatica < Tilia platyphyllos). In conclusion, ring-porous tree species exhibited stronger VPD-driven stomatal control over F_s, and tree-ring formation was less sensitive to severe drought than in their neighbouring diffuse-porous species. The F_s regulation explained the greater drought tolerance of the ring-porous trees.

Climate change exposes ecosystems to strong and rapid changes in their environmental boundary conditions mainly due to the altered temperature and precipitation patterns. It is still poorly understood how fast interlinked ecosystem processes respond to altered environmental conditions, if these responses occur gradually or suddenly when thresholds are exceeded, and if the patterns of the responses will reach a stable state. We conducted an irrigation experiment in the Pfynwald, Switzerland from 2003–2018. A naturally dry Scots pine (Pinus sylvestris L.) forest was irrigated with amounts that doubled natural precipitation, thus releasing the forest stand from water limitation. The aim of this study was to provide a quantitative understanding on how different traits and functions of individual trees and the whole ecosystem responded to increased water availability, and how the patterns and magnitudes of these responses developed over time. We found that the response magnitude, the temporal trajectory of responses, and the length of initial lag period prior to significant response largely varied across traits. We detected rapid and stronger responses from aboveground tree traits (e.g., tree-ring width, needle length, and crown transparency) compared to belowground tree traits (e.g., fine-root biomass). The altered aboveground traits during the initial years of irrigation increased the water demand and trees adjusted by increasing root biomass during the later years of irrigation, resulting in an increased survival rate of Scots pine trees in irrigated plots. The irrigation also stimulated ecosystem-level foliar decomposition rate, fungal fruit body biomass, and regeneration abundances of broadleaved tree species. However, irrigation did not promote the regeneration of Scots pine trees, which are reported to be vulnerable to extreme droughts. Our results provide extensive evidence that tree- and ecosystem-level responses were pervasive across a number of traits on long-term temporal scales. However, after reaching a peak, the magnitude of these responses either decreased or reached a new stable state, providing important insights into how resource alterations could change the system functioning and its boundary conditions.

Der Klimawandel hat erste Spuren im Wald hinterlassen. Trockene und teils auch heisse Sommer wie 2003, 2015 und 2018 häufen sich und führen verbreitet zum Tod von Bäumen; dazu tragen oft auch Borkenkäfer bei, die nicht nur geschwächte Fichten befallen, sondern auch Weisstannen. Neu war 2018, dass viele Buchen abstarben. Das hat Folgen für das Holzangebot.

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.

• The intensity and frequency of droughts events are projected to increase in future with expected adverse effects for forests. Thus, information on the dynamics of tree water uptake from different soil layers during and after drought is crucial.
• We applied an in situ water isotopologue monitoring system to determine the oxygen isotope composition in soil and xylem water of European beech with a 2-hour resolution together with measurements of soil water content, transpiration and tree water deficit. Using a Bayesian isotope mixing model, we inferred the relative and absolute contribution of water from four different soil layers to tree water use.
• Beech took up more than 50% of its water from the uppermost 5 cm soil layer at the beginning of the 2018 drought, but then reduced absolute water uptake from the drying topsoil by 84%. The trees were not able to quantitatively compensate for restricted topsoil water availability by additional uptake from deeper soil layers, which is related to the fine root depth distribution. Absolute water uptake from the topsoil was restored to pre-drought levels within 3 wk after rewetting.
• These uptake patterns help to explain both the drought sensitivity of beech and its high recovery potential after drought release.