No Future Growth Enhancement Expected at the Northern Edge for European Beech due to Continued Water Limitation.

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.

Non-linear response of Norway spruce to climate variation along elevational and age gradients in the Carpathians.

Climate change, namely increased warming coupled with a rise in extreme events (e.g., droughts, storms, heatwaves), is negatively affecting forest ecosystems worldwide. In these ecosystems, growth dynamics and biomass accumulation are driven mainly by environmental constraints, inter-tree competition, and disturbance regimes. Usually, climate-growth relationships are assessed by linear correlation due to the simplicity and straightforwardness of modeling. However, applying this method may bias results, since the ecological and physiological responses of trees to environmental factors are non-linear, and usually bell-shaped. In the Eastern Carpathian, Norway spruce is at the southeasternmost edge of its natural occurrence; this region is thus potentially vulnerable to climate change. A non-linear assessment of climate-growth relationships using machine-learning techniques for Norway spruce in this area had not been conducted prior to this study. To address this knowledge gap, we analyzed a large tree-ring network from 158 stands, with over 3000 trees of varying age distributed along an elevational gradient. Our results showed that non-linearity in the growth-climate response of spruce was season-specific: temperatures from the previous autumn and current growing season, along with water availability during winter, induced a bell-shaped response. Moreover, we found that at low elevations, spruce growth was mainly limited by water availability in the growing season, while winter temperatures are likely to have had a slight influence along the entire elevational gradient. Furthermore, at elevations lower than 1400 m, spruce trees were also found to be sensitive to previous autumn water availability. Overall, our results shed new light on the response of Norway spruce to climate in the Carpathians, which may aid in management decisions.

Oak tree-ring carbon stable isotopes from eastern Europe reveal significant drought signals along elevational gradients.

The importance of stable carbon and oxygen isotopes in tree-ring-based climate reconstructions is increasingly recognized, especially in regions where traditional dendrochronological parameters, such as tree-ring width, usually fail. However, the effects of elevation and other site conditions on climate signals in tree-ring stable isotope (TRSI) chronologies remain unclear. Here, we assess the sensitivities to precipitation and drought of tree-ring width (TRW) and δ13C and δ18O TRSI chronologies of 136 living oaks (Quercus spp.) from five different elevational zones between 130 and 630 m asl in eastern Slovakia. Our results show that while the mean δ13C values were stable across the elevational gradient, TRW gradually decreased with increasing elevation, and the mean δ18O values significantly differed between the lower and higher sites. Despite these variations, we observed strong coherency among all the elevation-specific TRW and TRSI chronologies. We also found that mainly mid-May to July precipitation and mid-May to August drought controlled TRW and the δ13C values, whereas the δ18O reflected an overall lower climate signal. Our results show a relatively stable drought signal across the elevational range, with shorter seasonal response windows at higher elevations. Furthermore, our study indicates that carbon TRSI and TRW oak chronologies capture distinct summer drought signals independently of elevation and therefore have a strong paleoclimatic potential across eastern Europe.

Impacts of elevated CO2 levels and temperature on photosynthesis and stomatal closure along an altitudinal gradient are counteracted by the rising atmospheric vapor pressure deficit.

The efficiency of water use in plants, a critical ecophysiological parameter closely related to water and carbon cycles, is essential for understanding the interactions between plants and their environment. This study investigates the effects of ongoing climate change and increasing atmospheric CO2 concentration on intrinsic (stomata-based; iWUE) and evaporative (transpiration-based; eWUE) water use efficiency in oak trees along a naturally small altitudinal gradient (130-630 m a.s.l.) of Vihorlat Mountains (eastern Slovakia, Central Europe). To assess changes in iWUE and eWUE values over the past 60 years (1961-2020), stable carbon isotope ratios in latewood cellulose (δ13Ccell) of annually resolved tree rings were analyzed. Such an approach was sensitive enough to distinguish tree responses to growth environments at different altitudes. Our findings revealed a rising trend in iWUE, particularly in oak trees at low and middle altitudes. However, this increase was negligible at high altitudes. Warmer and drier conditions at lower altitudes likely led to significant stomatal closure and enhanced efficiency in photosynthetic CO2 uptake due to rising CO2 concentration. Conversely, the increasing intracellular-to-ambient CO2 ratio (Ci/Ca) at higher altitudes indicated lower efficiency in photosynthetic CO2 uptake. In contrast to iWUE, eWUE showed no increasing trends over the last 60 years. This suggests that the positive impacts of elevated CO2 concentrations and temperature on photosynthesis and stomatal closure are counteracted by the rising atmospheric vapor pressure deficit (VPD). These differences underscore the importance of the correct interpretation of stomata-based and transpiration-based WUEs and highlight the necessity of atmospheric VPD correction when applying tree-ring δ13C-derived WUE at ecosystem and global levels.

Harmonised statistics and maps of forest biomass and increment in Europe.

Forest biomass is an essential resource in relation to the green transition and its assessment is key for the sustainable management of forest resources. Here, we present a forest biomass dataset for Europe based on the best available inventory and satellite data, with a higher level of harmonisation and spatial resolution than other existing data. This database provides statistics and maps of the forest area, biomass stock and their share available for wood supply in the year 2020, and statistics on gross and net volume increment in 2010-2020, for 38 European countries. The statistics of most countries are available at a sub-national scale and are derived from National Forest Inventory data, harmonised using common reference definitions and estimation methodology, and updated to a common year using a modelling approach. For those counties without harmonised statistics, data were derived from the State of Europe's Forest 2020 Report at the national scale. The maps are coherent with the statistics and depict the spatial distribution of the forest variables at 100 m resolution.

Empirical and process-based models predict enhanced beech growth in European mountains under climate change scenarios: A multimodel approach.

Process-based models and empirical modelling techniques are frequently used to (i) explore the sensitivity of tree growth to environmental variables, and (ii) predict the future growth of trees and forest stands under climate change scenarios. However, modelling approaches substantially influence predictions of the sensitivity of trees to environmental factors. Here, we used tree-ring width (TRW) data from 1630 beech trees from a network of 70 plots established across European mountains to build empirical predictive growth models using various modelling approaches. In addition, we used 3-PG and Biome-BGCMuSo process-based models to compare growth predictions with derived empirical models. Results revealed similar prediction errors (RMSE) across models ranging between 3.71 and 7.54 cm2 of basal area increment (BAI). The models explained most of the variability in BAI ranging from 54 % to 87 %. Selected explanatory variables (despite being statistically highly significant) and the pattern of the growth sensitivity differed between models substantially. We identified only five factors with the same effect and the same sensitivity pattern in all empirical models: tree DBH, competition index, elevation, Gini index of DBH, and soil silt content. However, the sensitivity to most of the climate variables was low and inconsistent among the empirical models. Both empirical and process-based models suggest that beech in European mountains will, on average, likely experience better growth conditions under both 4.5 and 8.5 RCP scenarios. The process-based models indicated that beech may grow better across European mountains by 1.05 to 1.4 times in warmer conditions. The empirical models identified several drivers of tree growth that are not included in the current process-based models (e.g., different nutrients) but may have a substantial effect on final results, particularly if they are limiting factors. Hence, future development of process-based models may build upon our findings to increase their ability to correctly capture ecosystem dynamics.

Plot size matters: Toward comparable species richness estimates across plot-based inventories.

To understand the state and trends in biodiversity beyond the scope of monitoring programs, biodiversity indicators must be comparable across inventories. Species richness (SR) is one of the most widely used biodiversity indicators. However, as SR increases with the size of the area sampled, inventories using different plot sizes are hardly comparable. This study aims at producing a methodological framework that enables SR comparisons across plot-based inventories with differing plot sizes. We used National Forest Inventory (NFI) data from Norway, Slovakia, Spain, and Switzerland to build sample-based rarefaction curves by randomly incrementally aggregating plots, representing the relationship between SR and sampled area. As aggregated plots can be far apart and subject to different environmental conditions, we estimated the amount of environmental heterogeneity (EH) introduced in the aggregation process. By correcting for this EH, we produced adjusted rarefaction curves mimicking the sampling of environmentally homogeneous forest stands, thus reducing the effect of plot size and enabling reliable SR comparisons between inventories. Models were built using the Conway-Maxell-Poisson distribution to account for the underdispersed SR data. Our method successfully corrected for the EH introduced during the aggregation process in all countries, with better performances in Norway and Switzerland. We further found that SR comparisons across countries based on the country-specific NFI plot sizes are misleading, and that our approach offers an opportunity to harmonize pan-European SR monitoring. Our method provides reliable and comparable SR estimates for inventories that use different plot sizes. Our approach can be applied to any plot-based inventory and count data other than SR, thus allowing a more comprehensive assessment of biodiversity across various scales and ecosystems.

Competition for soil resources forces a trade-off between enhancing tree productivity and understorey species richness in managed beech forests.

Traditionally focussed on maximising productivity, forest management increasingly has to consider other functions performed by the forest stands, such as biodiversity conservation. Terrestrial plant communities typically possess a hump-back relationship between biomass productivity and plant species richness. However, there is evidence of a reverse relationship in forests dominated by beech, one of the most competitive and widespread tree species in temperate Europe. To fully explore the tree productivity-species richness relationship, we investigated above- and below-ground drivers of understorey plant species richness. We focussed on managed beech forests growing along an elevation gradient in Central Europe. We found that the lowest understorey plant diversity was under conditions optimal for beech. Tree fine root mass, canopy openness, soil C/N ratio, the interaction between tree fine root mass and stoniness, and stand structural diversity explain the variation of understorey species richness. We show that the competition for soil resources is the main driver of plant species diversity in managed forests; maximising beech growth in optimal conditions may thus come at the expense of understorey plant richness.

Climate warming induced synchronous growth decline in Norway spruce populations across biogeographical gradients since 2000.

Covering large parts of Europe, Norway spruce (Picea abies L Karst.) plays an important role in the adaptation strategy of forest services to future climate change. Although dendroecology can provide valuable information on the past relationships between tree growth and climate, most previous studies were biased towards species-specific distribution limits, where old individuals grow slowly under extreme conditions. In the present study, we investigated the growth variability and climate sensitivity of 2851 Norway spruce trees along longitudinal (E 12-26°), latitudinal (N 45-51°), and elevation (118-1591 m a.s.l.) gradients in central-eastern Europe. We reveal that summer weather significantly affects the radial growth of spruce trees, but the effects strongly vary along biogeographical gradients. Extreme summer heatwaves in 2000 and 2003 reduced the growth rates by 10-35%, most pronounced in the southern Carpathians. In contrast to the population in the Czech Republic, climate warming induced a synchronous decline in the growth rates across biogeographical gradients in the Carpathian arc. By demonstrating the increased vulnerability of Norway spruce under warmer climate conditions, we recommended that the forest services and conservation managers replace or admix monocultures of this species with more drought-resilient mixtures including fir, beech and other broadleaved species.

Non-destructive monitoring of annual trunk increments by terrestrial structure from motion photogrammetry.

Annual trunk increments are essential for short-term analyses of the response of trees to various factors. For instance, based on annual trunk increments, it is possible to develop and calibrate forest growth models. We investigated the possibility of estimating annual trunk increments from the terrestrial structure from motion (SfM) photogrammetry. Obtaining the annual trunk increments of mature trees is challenging due to the relatively small growth of trunks within one year. In our experiment, annual trunk increments were obtained by two conventional methods: measuring tape (perimeter increment) at heights of 0.8, 1.3, and 1.8 m on the trunk and increment borer (diameter increment) at a height of 1.3 m on the trunk. The following tree species were investigated: Fagus sylvatica L. (beech), Quercus petraea (Matt.) Liebl. (oak), Picea abies (L.) H. Karst (spruce), and Abies alba Mill (fir). The annual trunk increments ranged from 0.9 cm to 2.4 cm (tape/perimeter) and from 0.7 mm to 3.1 mm (borer/diameter). The data were collected before- and after-vegetation season, besides the data collection increment borer. When the estimated perimeters from the terrestrial SfM photogrammetry were compared to those obtained using the measuring tape, the root mean square error (RMSE) was 0.25-1.33 cm. The relative RMSE did not exceed 1% for all tree species. No statistically significant differences were found between the annual trunk increments obtained using the measuring tape and terrestrial SfM photogrammetry for beech, spruce, and fir. Only in the case of oak, the difference was statistically significant. Furthermore, the correlation coefficient between the annual trunk increments collected using the increment borer and those derived from terrestrial SfM photogrammetry was positive and equal to 0.6501. Terrestrial SfM photogrammetry is a hardware low-demanding technique that provides accurate three-dimensional data that can, based on our results, even detect small temporal tree trunk changes.

Contrasting effects of environmental change on the radial growth of co-occurring beech and fir trees across Europe.

Under predicted climate change, native silver fir (Abies alba) and European beech (Fagus sylvatica) are the most likely replacement species for the Norway spruce (Picea abies) monocultures planted across large parts of continental Europe. Our current understanding of the adaptation potential of fir-beech mixed forests to climate change is limited because long-term responses of the two species to environmental changes have not yet been comprehensively quantified. We compiled and analysed tree-ring width (TRW) series from 2855 dominant, co-dominant, sub-dominant and suppressed fir and beech trees sampled in 17 managed and unmanaged mixed beech-fir forest sites across Continental Europe, including Bosnia and Herzegovina, Germany, Italy, Romania and Slovakia. Dendroecological techniques that combine various detrending methods were used to investigate variation in radial growth of co-occurring fir and beech trees. Coincidental with peak SO2 emissions, the growth of silver fir declined between 1950 and 1980 at most sites, whereas beech growth increased during this period. Correspondent to a significant warming trend from 1990-2010, average beech growth declined, but silver fir growth increased. Long-term growth patterns and growth-climate sensitivity of fir and beech trees did not significantly differ between managed and unmanaged forests. Multi-decadal changes in the growth rate of all vertical tree classes were similar. In contrast to previous indications of limited drought susceptibility of beech mixed stands, this study suggests that the mixture of tree species in forest stands does not necessarily prevent growth depressions induced by long-term environmental change. Our results further imply that forest management does not necessarily alter their sensitivity to environmental changes.

Possible causes of the recent rapid increase in the radial increment of silver fir in the Western Carpathians.

Silver fir is one of the most productive and ecologically valuable native European tree species, however, it has been experiencing decline which has periodically occurred over its natural range. This paper aims to investigate the recent climate-growth relationships of silver fir (Abies alba Mill.) and its temporal change along the course of its life. Long-term tree-ring databases, as well as records on climate, atmospheric SO2, NO3 and acid concentrations from four different regions in the Western Carpathians were used. The results provide clear evidence of significant increase of silver fir's radial increment over the entire Western Carpathian area since 1970-1980. The results indicated that the most probable factors behind the rapid recovery of tree radial increment were reductions in emissions of NO3 and SO2, alongside a significant increase in mean June, July and April temperatures.