Dr. Stefan Klesse

It is generally recognized that time series of tree-ring widths must be detrended, i.e. the age- or size-related trend must be removed before any climate-growth analysis can be performed. However, in contrast to ring-width (RW) analyses, there are no best practice rules on how to use basal area increment (BAI). BAI as an important variable in forestry has become also an increasingly used variable in dendrochronology. BAI is considered a better representative of the overall tree growth than RW and it is supposedly less affected by age trends than RW.
In this paper, we provide guidance by investigating the common rhetoric surrounding the use of BAI in the literature, and by highlighting possible problems and consequences from apparent misconceptions. We performed a literature review for the period 2018-2023, whether age trends in BAI time series are considered in the analysis. We also conducted additional analyses with a published tree-ring dataset of Klesse et al. (2018) to investigate whether BAI or RW time series exhibit stronger trends. Both the literature review and the data analysis provide no support for the frequently used claim that BAI time series have smaller age trends which would preclude detrending. In fact, in 58% of the investigated time series, age trends in BAI are stronger than in RW when the trees are younger than 50 years old, and even between the ages of 51 and 100 years, we found stronger trends in BAI in 44% of the cases. We also provide an overview of three general methods for calculating the BAI together with their advantages and disadvantages. We discuss whether the inside-out method, the outside-in method, or the Bakker (2005) method are useful to convert RW to BAI when the pith offset estimate (POE) and the diameter at breast height (DBH) are available or missing, respectively. We conclude with a general recommendation to calculate BAI based on both POE and DBH information using the Bakker method. Further, analyses with BAI should always consider the size or age of each tree, especially when trends or growth rates are compared for trees of different age classes. For climate-growth analyses, the detrending of BAI is always necessary. By addressing these key issues, we aim to improve the reliability and consistency of BAI-based studies in dendrochronology and climate-growth research.

Introduction: Quantitative wood anatomy is critical for establishing climate reconstruction proxies, understanding tree hydraulics, and quantifying carbon allocation. Its accuracy depends upon the image acquisition methods, which allows for the identification of the number and dimensions of vessels, fibres, and tracheids within a tree ring. Angiosperm wood is analysed with a variety of different image acquisition methods, including surface pictures, wood anatomical micro-sections, or X-ray computed micro-tomography. Despite known advantages and disadvantages, the quantitative impact of method selection on wood anatomical parameters is not well understood.

Methods: In this study, we present a systematic uncertainty analysis of the impact of the image acquisition method on commonly used anatomical parameters. We analysed four wood samples, representing a range of wood porosity, using surface pictures, micro-CT scans, and wood anatomical micro-sections. Inter-annual patterns were analysed and compared between methods from the five most frequently used parameters, namely mean lumen area (MLA), vessel density (VD), number of vessels (VN), mean hydraulic diameter (D_h), and relative conductive area (RCA). A novel sectorial approach was applied on the wood samples to obtain intra-annual profiles of the lumen area (A_l), specific theoretical hydraulic conductivity (K_s), and wood density (ρ).

Results: Our quantitative vessel mapping revealed that values obtained for hydraulic wood anatomical parameters are comparable across different methods, supporting the use of easily applicable surface picture methods for ring-porous and specific diffuse-porous tree species. While intra-annual variability is well captured by the different methods across species, wood density (ρ) is overestimated due to the lack of fibre lumen area detection.

Discussion: Our study highlights the potential and limitations of different image acquisition methods for extracting wood anatomical parameters. Moreover, we present a standardized workflow for assessing radial tree ring profiles. These findings encourage the compilation of all studies using wood anatomical parameters and further research to refine these methods, ultimately enhancing the accuracy, replication, and spatial representation of wood anatomical studies.

Given the importance of climate in shaping species' geographic distributions, climate change poses an existential threat to biodiversity. Climate envelope modeling, the predominant approach used to quantify this threat, presumes that individuals in populations respond to climate variability and change according to species-level responses inferred from spatial occurrence data—such that individuals at the cool edge of a species' distribution should benefit from warming (the "leading edge"), whereas individuals at the warm edge should suffer (the "trailing edge"). Using 1,558 tree-ring time series of an aridland pine (Pinus edulis) collected at 977 locations across the species’ distribution, we found that trees everywhere grow less in warmer-than-average and drier-than-average years. Ubiquitous negative temperature sensitivity indicates that individuals across the entire distribution should suffer with warming—the entire distribution is a trailing edge. Species-level responses to spatial climate variation are opposite in sign to individual-scale responses to time-varying climate for approximately half the species’ distribution with respect to temperature and the majority of the species’ distribution with respect to precipitation. These findings, added to evidence from the literature for scale-dependent climate responses in hundreds of species, suggest that correlative, equilibrium-based range forecasts may fail to accurately represent how individuals in populations will be impacted by changing climate. A scale-dependent view of the impact of climate change on biodiversity highlights the transient risk of extinction hidden inside climate envelope forecasts and the importance of evolution in rescuing species from extinction whenever local climate variability and change exceeds individual-scale climate tolerances.

Laut einer Studie hat das Bohrkernentnehmen keine nachweisbaren Auswirkungen auf das Wachstum und die Sterblichkeit von Bäumen. Die Methode könnte also öfter eingesetzt werden, um die Reaktion der Wälder auf veränderte Umweltbedingungen besser zu verstehen.

With ongoing global warming, increasing water deficits promote physiological stress on forest ecosystems with negative impacts on tree growth, vitality, and survival. How individual tree species will react to increased drought stress is therefore a key research question to address for carbon accounting and the development of climate change mitigation strategies. Recent tree-ring studies have shown that trees at higher latitudes will benefit from warmer temperatures, yet this is likely highly species-dependent and less well-known for more temperate tree species. Using a unique pan-European tree-ring network of 26,430 European beech (Fagus sylvatica L.) trees from 2118 sites, we applied a linear mixed-effects modeling framework to (i) explain variation in climate-dependent growth and (ii) project growth for the near future (2021–2050) across the entire distribution of beech. We modeled the spatial pattern of radial growth responses to annually varying climate as a function of mean climate conditions (mean annual temperature, mean annual climatic water balance, and continentality). Over the calibration period (1952–2011), the model yielded high regional explanatory power (R² = 0.38–0.72). Considering a moderate climate change scenario (CMIP6 SSP2-4.5), beech growth is projected to decrease in the future across most of its distribution range. In particular, projected growth decreases by 12%–18% (interquartile range) in northwestern Central Europe and by 11%–21% in the Mediterranean region. In contrast, climate-driven growth increases are limited to around 13% of the current occurrence, where the historical mean annual temperature was below ~6°C. More specifically, the model predicts a 3%–24% growth increase in the high-elevation clusters of the Alps and Carpathian Arc. Notably, we find little potential for future growth increases (−10 to +2%) at the poleward leading edge in southern Scandinavia. Because in this region beech growth is found to be primarily water-limited, a northward shift in its distributional range will be constrained by water availability.

The future performance of the widely abundant European beech (Fagus sylvatica L.) across its ecological amplitude is uncertain. Although beech is considered drought-sensitive and thus negatively affected by drought events, scientific evidence indicating increasing drought vulnerability under climate change on a cross-regional scale remains elusive. While evaluating changes in climate sensitivity of secondary growth offers a promising avenue, studies from productive, closed-canopy forests suffer from knowledge gaps, especially regarding the natural variability of climate sensitivity and how it relates to radial growth as an indicator of tree vitality. Since beech is sensitive to drought, we in this study use a drought index as a climate variable to account for the combined effects of temperature and water availability and explore how the drought sensitivity of secondary growth varies temporally in dependence on growth variability, growth trends, and climatic water availability across the species' ecological amplitude.
Our results show that drought sensitivity is highly variable and non-stationary, though consistently higher at dry sites compared to moist sites. Increasing drought sensitivity can largely be explained by increasing climatic aridity, especially as it is exacerbated by climate change and trees' rank progression within forest communities, as (co-)dominant trees are more sensitive to extra-canopy climatic conditions than trees embedded in understories. However, during the driest periods of the 20th century, growth showed clear signs of being decoupled from climate. This may indicate fundamental changes in system behavior and be early-warning signals of decreasing drought tolerance. The multiple significant interaction terms in our model elucidate the complexity of European beech's drought sensitivity, which needs to be taken into consideration when assessing this species' response to climate change.

The record-breaking drought in 2018 caused premature leaf discoloration and shedding (early browning) in many beech (Fagus sylvatica L.) dominated forests in Central Europe. However, a high degree of variability in drought response among individual beech trees was observed. While some trees were severely impacted by the prolonged water deficits and high temperatures, others remained vital with no or only minor signs of crown vitality loss. Why some beech trees were more susceptible to drought-induced crown damage than others and whether growth recovery is possible are poorly understood. Here, we aimed to identify growth characteristics associated with the variability in drought response between individual beech trees based on a sample of 470 trees in northern Switzerland. By combining tree growth measurements and crown condition assessments, we also investigated the possible link between crown dieback and growth recovery after drought. Beech trees with early browning exhibited an overall lower growth vigor before the 2018 drought than co-occurring vital beech trees. This lower vigor is mainly indicated by lower overall growth rates, stronger growth declines in the past decades, and higher growth-climate sensitivity. Particularly, warm previous year summer conditions negatively affected current growth of the early-browning trees. These findings suggest that the affected trees had less access to critical resources and were physiologically limited in their growth predisposing them to early browning. Following the 2018 drought, observed growth recovery potential corresponded to the amount of crown dieback and the local climatic water balance. Overall, our findings emphasize that beech-dominated forests in Central Europe are under increasing pressure from severe droughts, ultimately reducing the competitive ability of this species, especially on lowland sites with shallow soils and low water holding capacity.

Selon une étude, le prélèvement de carottes n'a pas d'effet détectable sur la croissance et la mortalité des arbres. Cette méthode pourrait donc être utilisée plus fréquemment pour aider à mieux comprendre la réaction des forêts à l'évolution des conditions environnementales.

Tree cores are a highly valuable scientific resource. Annual growth ring data can, for example, improve our understanding of climate change impacts and effects of environmental pollution, allow for better annual estimations of tree growth patterns and carbon storage, and help quantify dynamics and changes in forest ecosystems. The value of coring trees for research has been weighted against concerns around the potential harm coring might cause to trees. To date, there is indeed limited research accurately quantifying the potential effects of coring on tree growth and only a handful of studies assessing its influence on mortality. Consequently, many European long-term forest inventorying and monitoring programs are concerned that tree coring might bias the repeated tree measurements in permanent plots, which they rely on for assessments of states and changes of forests. In this study, we assessed the effects of tree coring on the growth and mortality of three widespread European tree species approximately 10 years after they were cored. We used repeated tree measurements from permanent research sites in Switzerland and Ukraine. In Switzerland, we assessed 35 cored and 159 uncored Norway spruce (Picea abies) trees as well as 147 cored and 332 uncored silver fir (Abies alba) trees. In Ukraine, we assessed 348 cored and 6'611 uncored European beech (Fagus sylvatica) trees. We found no statistical evidence that coring negatively affected the growth or mortality of the three tree species assessed. Although we cannot rule out subtle effects on tree health and wood quality, our findings do not provide any evidence that coring affects or biases repeated measurements (such as DBH measures and recording of mortality) performed on the investigated tree species. Tree coring could therefore be considered more often for routine incorporation, particularly in long-term forest inventorying and monitoring programs and initiatives.

Der Schweizer Wald beherbergt zahlreiche Baumarten mit unterschiedlichen ökologischen Ansprüchen an Licht, Nährstoffen und Klima. In der empirischen Waldmodellierung werden ähnliche Baumarten oft gruppiert, weil zu wenige Beobachtungen vorliegen, um artspezifische Modelle für Einwuchs, Wachstum und Mortalität zu entwickeln. Wir präsentieren eine neue Baumartengruppierung, die ähnliche ökologische Ansprüche und deren Einfluss auf das Wachsen und Absterben berücksichtigt.

The vapor pressure deficit reflects the difference between how much moisture the atmosphere could and actually does hold, a factor that fundamentally affects evapotranspiration, ecosystem functioning, and vegetation carbon uptake. Its spatial variability and long-term trends under natural versus human-influenced climate are poorly known despite being essential for predicting future effects on natural ecosystems and human societies such as crop yield, wildfires, and health. Here we combine regionally distinct reconstructions of pre-industrial summer vapor pressure deficit variability from Europe’s largest oxygen-isotope network of tree-ring cellulose with observational records and Earth system model simulations with and without human forcing included. We demonstrate that an intensification of atmospheric drying during the recent decades across different European target regions is unprecedented in a pre-industrial context and that it is attributed to human influence with more than 98% probability. The magnitude of this trend is largest in Western and Central Europe, the Alps and Pyrenees region, and the smallest in southern Fennoscandia. In view of the extreme drought and compound events of the recent years, further atmospheric drying poses an enhanced risk to vegetation, specifically in the densely populated areas of the European temperate lowlands.

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.

1. Drought legacies in radial tree growth are an important feature of variability in biomass accumulation and are widely used to characterize forest resilience to climate change. Defined as a deviation from normal growth, the statistical significance of legacy effects depends on the definition of "normal" - expected growth under average conditions - which has not received sufficient scrutiny.
2 We re-examined legacy effect analyses using the International Tree-Ring Data Bank (ITRDB) and then produced synthetic tree-ring data to disentangle four key variables influencing the magnitude of legacy effects. We hypothesized that legacy effects (i) are mainly influenced by the auto-correlation of the radial growth time series (phi), (ii) depend on climate-growth cross-correlation (rho), (iii) are directly proportional to the inherent variability of the growth time series (standard deviation, SD), and (iv) scale with the chosen extreme event threshold.
3. Using a data simulation approach, we were able to reproduce observed lag patterns, demonstrating that legacy effects are a direct outcome of ubiquitous biological memory. We found that stronger legacy effects for conifers compared to angiosperms is a consequence of their higher auto-correlation, and that the detectability of legacy effects following rare drought events at individual sites is compromised by strong background stochasticity.
4. Synthesis. We propose two pathways forward to improve the assessment and interpretation of legacy effects: First, we highlight the need to account for auto-correlated residuals of climate-growth regression models a posteriori, thereby retrospectively adjusting expectations for "normal" growth variability. Alternatively, we recommend including lagged climate variables in regression models a priori. By doing so, the magnitude of detected legacy effects is greatly reduced and biological memory is directly attributed to antecedent climatic drivers. We argue that future analyses should focus on understanding the functional reasons for how and why key statistical parameters describing this biological memory differ across species and sites. These two pathways should also stimulate improved process-based representation of vegetation carbon dynamics in mechanistic models.

Tree growth varies closely with high–frequency climate variability. Since the 1930s detrending climate data prior to comparing them with tree growth data has been shown to better capture tree growth sensitivity to climate. However, in a context of increasingly pronounced trends in climate, this practice remains surprisingly rare in dendroecology. In a review of Dendrochronologia over the 2018-2021 period, we found that less than 20 % of dendroecological studies detrended climate data prior to climate-growth analyses. With an illustrative study, we want to remind the dendroecology community that such a procedure is still, if not more than ever, rational and relevant. We investigated the effects of detrending climate data on climate-growth relationships across North America over the 1951–2000 period. We used a network of 2536 tree individual ring-width series from the Canadian and Western US forest inventories. We compared correlations between tree growth and seasonal climate data (Tmin, Tmax, Prec) both raw and detrended. Detrending approaches included a linear regression, 30-yr and 100-yr cubic smoothing splines. Our results indicate that on average the detrending of climate data increased climate–growth correlations. In addition, we observed that strong trends in climate data translated to higher variability in inferred correlations based on raw vs. detrended climate data. We provide further evidence that our results hold true for the entire spectrum of dendroecological studies using either mean site chronologies and correlations coefficients, or individual tree time series within a mixed-effects model framework where regression coefficients are used more commonly. We show that even without a change in correlation, regression coefficients can change a lot and we tend to underestimate the true climate impact on growth in case of climate variables containing trends. This study demonstrates that treating climate and tree-ring time series “like-for-like” is a necessary procedure to reduce false negatives and positives in dendroecological studies. Concluding, we recommend using the same detrending for climate and tree growth data when tree-ring time series are detrended with splines or similar frequency-based filters.

Partitioned estimates of the boreal forest carbon (C) sink components are crucial for understanding processes and developing science-driven adaptation and mitigation strategies under climate change. Here, we provide a concise tree-centered overview of the boreal forest C balance and offer a circumpolar perspective on the contribution of trees to boreal forest C dynamics. We combine an ant’s-eye view, based on quantitative in situ observations of C balance, with a bird’s-eye perspective on C dynamics across the circumboreal region using large-scale data sets. We conclude with an outlook addressing the trajectories of the circumboreal C dynamics in response to projected environmental changes.

Tree-ring time series provide long-term, annually resolved information on the growth of trees. When sampled in a systematic context, tree-ring data can be scaled to estimate the forest carbon capture and storage of landscapes, biomes, and - ultimately - the globe. A systematic effort to sample tree rings in national forest inventories would yield unprecedented temporal and spatial resolution of forest carbon dynamics and help resolve key scientific uncertainties, which we highlight in terms of evidence for forest greening (enhanced growth) versus browning (reduced growth, increased mortality). We describe jump-starting a tree-ring collection across the continent of North America, given the commitments of Canada, the United States, and Mexico to visit forest inventory plots, along with existing legacy collections. Failing to do so would be a missed opportunity to help chart an evidence-based path toward meeting national commitments to reduce net greenhouse gas emissions, urgently needed for climate stabilization and repair.

Robust ecological forecasting of tree growth under future climate conditions is critical to anticipate future forest carbon storage and flux. Here, we apply three ingredients of ecological forecasting that are key to improving forecast skill: data fusion, confronting model predictions with new data, and partitioning forecast uncertainty. Specifically, we present the first fusion of tree-ring and forest inventory data within a Bayesian state-space model at a multi-site, regional scale, focusing on Pinus ponderosa var. brachyptera in the southwestern US. Leveraging the complementarity of these two data sources, we parsed the ecological complexity of tree growth into the effects of climate, tree size, stand density, site quality, and their interactions, and quantified uncertainties associated with these effects. New measurements of trees, an ongoing process in forest inventories, were used to confront forecasts of tree diameter with observations, and evaluate alternative tree growth models. We forecasted tree diameter and increment in response to an ensemble of climate change projections, and separated forecast uncertainty into four different causes: initial conditions, parameters, climate drivers, and process error. We found a strong negative effect of fall–spring maximum temperature, and a positive effect of water-year precipitation on tree growth. Furthermore, tree vulnerability to climate stress increases with greater competition, with tree size, and at poor sites. Under future climate scenarios, we forecast increment declines of 22%–117%, while the combined effect of climate and size-related trends results in a 56%–91% decline. Partitioning of forecast uncertainty showed that diameter forecast uncertainty is primarily caused by parameter and initial conditions uncertainty, but increment forecast uncertainty is mostly caused by process error and climate driver uncertainty. This fusion of tree-ring and forest inventory data lays the foundation for robust ecological forecasting of aboveground biomass and carbon accounting at tree, plot, and regional scales, including iterative improvement of model skill.

Ongoing climate warming is increasing evapotranspiration, a process that reduces plant-available water and aggravates the impact of extreme droughts during the growing season. Such an exceptional hot drought occurred in Central Europe in 2018 and caused widespread defoliation in mid-summer in European beech (Fagus sylvatica L.) forests.
Here, we recorded crown damage in 2021 in nine mature even-aged beech-dominated stands in northwestern Switzerland along a crown damage severity gradient (low, medium, high) and analyzed tree-ring widths of 21 mature trees per stand. We aimed at identifying predisposing factors responsible for differences in crown damage across and within stands such as tree growth characteristics (average growth rates and year-to-year variability) and site-level variables (mean canopy height, soil properties).
We found that stand-level crown damage severity was strongly related to soil water availability, inferred from tree canopy height and plant available soil water storage capacity (AWC). Trees were shorter in drier stands, had higher year-to-year variability in radial growth, and showed higher growth sensitivity to moisture conditions of previous late summer than trees growing on soils with sufficient AWC, indicating that radial growth in these forests is principally limited by soil water availability. Within-stand variation of post-drought crown damage corresponded to growth rate and tree size (diameter at breast height, DBH), i.e., smaller and slower-growing trees that face more competition, were associated with increased crown damage after the 2018 drought. These findings point to tree vigor before the extreme 2018 drought (long-term relative growth rate) as an important driver of damage severity within and across stands. Our results suggest that European beech is less likely to be able to cope with future climate change-induced extreme droughts on shallow soils with limited water retention capacity.

The growth of past, present, and future forests was, is and will be affected by climate variability. This multifaceted relationship has been assessed in several regional studies, but spatially resolved, large-scale analyses are largely missing so far. Here we estimate recent changes in growth of 5800 beech trees (Fagus sylvatica L.) from 324 sites, representing the full geographic and climatic range of species. Future growth trends were predicted considering state-of-the-art climate scenarios. The validated models indicate growth declines across large region of the distribution in recent decades, and project severe future growth declines ranging from −20% to more than −50% by 2090, depending on the region and climate change scenario (i.e. CMIP6 SSP1-2.6 and SSP5-8.5). Forecasted forest productivity losses are most striking towards the southern distribution limit of Fagus sylvatica, in regions where persisting atmospheric high-pressure systems are expected to increase drought severity. The projected 21^st century growth changes across Europe indicate serious ecological and economic consequences that require immediate forest adaptation.

Sapwood characteristics, such as sapwood area as well as thermal and hydraulic conductivity, are linked to species-specific hydraulic function and resource allocation to water transport tissues (xylem). These characteristics are often unknown and thus a major source of uncertainty in sap flow data processing and transpiration estimates because bulk rather than species-specific values are usually applied. Here, we analyzed the sapwood characteristics of fifteen common tree species in eastern North America from different taxonomic (i.e., angiosperms and gymnosperms) and xylem porosity groups (i.e., tracheid-bearing, diffuse- or ring-porous species) and we assessed how uncertainties in sapwood characteristics involved in sap flow calculations are propagated in tree water use estimates. We quantified their sapwood area changes with stem diameter (allometric scaling) and thermal conductivity. We combined these measurements with species-specific values of wood density and hydraulic conductivity found in the literature and assessed the role of wood anatomy in orchestrating their covariation. Using an example sap flow dataset from tree species with different xylem porosity, we assessed the sensitivity of tree water use estimates to sapwood characteristics and their interactions. Angiosperms (ring- and diffuse-porous species), with specialized vessels for water transport, showed a steeper relationship (scaling) between tree stem diameter and sapwood area in comparison to gymnosperms (tracheid-bearing species). Gymnosperms (angiosperms) were characterized by lower (higher) wood density and higher (lower) sapwood moisture content, resulting in non-significant differences in sapwood thermal conductivity between taxonomic and xylem porosity groups. Clustering of species sapwood characteristics based on taxonomic or xylem porosity groups and constraining these parameters could facilitate more accurate sap flow calculations and tree water use estimates. When combined with an increasing number of sap flow observations, these findings should improve tree- and landscape-level transpiration estimates, leading to more robust partitioning of terrestrial water fluxes.