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.

Incorporating high-resolution climate, remote sensing and topographic data to map annual forest growth in central and eastern Europe.

To enhance our understanding of forest carbon sequestration, climate change mitigation and drought impact on forest ecosystems, the availability of high-resolution annual forest growth maps based on tree-ring width (TRW) would provide a significant advancement to the field. Site-specific characteristics, which can be approximated by high-resolution Earth observation by satellites (EOS), emerge as crucial drivers of forest growth, influencing how climate translates into tree growth. EOS provides information on surface reflectance related to forest characteristics and thus can potentially improve the accuracy of forest growth models based on TRW. Through the modelling of TRW using EOS, climate and topography data, we showed that species-specific models can explain up to 52 % of model variance (Quercus petraea), while combining different species results in relatively poor model performance (R2 = 13 %). The integration of EOS into models based solely on climate and elevation data improved the explained variance by 6 % on average. Leveraging these insights, we successfully generated a map of annual TRW for the year 2021. We employed the area of applicability (AOA) approach to delineate the range in which our models are deemed valid. The calculated AOA for the established forest-type models was 73 % of the study region, indicating robust spatial applicability. Notably, unreliable predictions predominantly occurred in the climate margins of our dataset. In conclusion, our large-scale assessment underscores the efficacy of combining climate, EOS and topographic data to develop robust models for mapping annual TRW. This research not only fills a critical void in the current understanding of forest growth dynamics but also highlights the potential of integrated data sources for comprehensive ecosystem assessments.

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.

Growth and resilience responses of Scots pine to extreme droughts across Europe depend on predrought growth conditions.

Global climate change is expected to further raise the frequency and severity of extreme events, such as droughts. The effects of extreme droughts on trees are difficult to disentangle given the inherent complexity of drought events (frequency, severity, duration, and timing during the growing season). Besides, drought effects might be modulated by trees' phenotypic variability, which is, in turn, affected by long-term local selective pressures and management legacies. Here we investigated the magnitude and the temporal changes of tree-level resilience (i.e., resistance, recovery, and resilience) to extreme droughts. Moreover, we assessed the tree-, site-, and drought-related factors and their interactions driving the tree-level resilience to extreme droughts. We used a tree-ring network of the widely distributed Scots pine (Pinus sylvestris) along a 2,800 km latitudinal gradient from southern Spain to northern Germany. We found that the resilience to extreme drought decreased in mid-elevation and low productivity sites from 1980-1999 to 2000-2011 likely due to more frequent and severe droughts in the later period. Our study showed that the impact of drought on tree-level resilience was not dependent on its latitudinal location, but rather on the type of sites trees were growing at and on their growth performances (i.e., magnitude and variability of growth) during the predrought period. We found significant interactive effects between drought duration and tree growth prior to drought, suggesting that Scots pine trees with higher magnitude and variability of growth in the long term are more vulnerable to long and severe droughts. Moreover, our results indicate that Scots pine trees that experienced more frequent droughts over the long-term were less resistant to extreme droughts. We, therefore, conclude that the physiological resilience to extreme droughts might be constrained by their growth prior to drought, and that more frequent and longer drought periods may overstrain their potential for acclimation.

Comparing soil inventory with modelling: Carbon balance in central European forest soils varies among forest types.

Forest soils represent a large carbon pool and already small changes in this pool may have an important effect on the global carbon cycle. To predict the future development of the soil organic carbon (SOC) pool, well-validated models are needed. We applied the litter and soil carbon model Yasso15 to 1838 plots of the German national forest soil inventory (NFSI) for the period between 1985 and 2014 to enables a direct comparison to the NFSI measurements. In addition, to provide data for the German Greenhouse Gas Inventory, we simulated the development of SOC with Yasso15 applying a climate projection based on the RCP8.5 scenario. The initial model-calculated SOC stocks were adjusted to the measured ones in the NFSI. On average, there were no significant differences between the simulated SOC changes (0.25 ±â€¯0.10 Mg C ha-1 a-1) and the NFSI data (0.39 ±â€¯0.11 Mg C ha-1 a-1). Comparing regional soil-unit-specific aggregates of the SOC changes, the correlation between both methods was significant (r2 = 0.49) although the NFSI values had a wider range and more negative values. In the majority of forest types, representing 75% of plots, both methods produced similar estimates of the SOC balance. Opposite trends were found in mountainous coniferous forests on acidic soils. These soils had lost carbon according to the NFSI (-0.89 ±â€¯0.30 Mg C ha-1 a-1) whereas they had gained it according to Yasso15 (0.21 ±â€¯0.10 Mg C ha-1 a-1). In oligotrophic pine forests, the NFSI indicated high SOC gains (1.36 ±â€¯0.17 Mg C ha-1 a-1) and Yasso15 much smaller (0.29 ±â€¯0.10 Mg C ha-1 a-1). According to our results, German forest soils are a large carbon sink. The application of the Yasso15 model supports the results of the NFSI. The sink strength differs between forest types possibly because of differences in organic matter stabilisation.

Organic carbon stocks and sequestration rates of forest soils in Germany.

The National Forest Soil Inventory (NFSI) provides the Greenhouse Gas Reporting in Germany with a quantitative assessment of organic carbon (C) stocks and changes in forest soils. Carbon stocks of the organic layer and the mineral topsoil (30 cm) were estimated on the basis of ca. 1.800 plots sampled from 1987 to 1992 and resampled from 2006 to 2008 on a nationwide grid of 8 × 8 km. Organic layer C stock estimates were attributed to surveyed forest stands and CORINE land cover data. Mineral soil C stock estimates were linked with the distribution of dominant soil types according to the Soil Map of Germany (1 : 1 000 000) and subsequently related to the forest area. It appears that the C pool of the organic layer was largely depending on tree species and parent material, whereas the C pool of the mineral soil varied among soil groups. We identified the organic layer C pool as stable although C was significantly sequestered under coniferous forest at lowland sites. The mineral soils, however, sequestered 0.41 Mg C ha(-1) yr(-1) . Carbon pool changes were supposed to depend on stand age and forest transformation as well as an enhanced biomass input. Carbon stock changes were clearly attributed to parent material and soil groups as sandy soils sequestered higher amounts of C, whereas clayey and calcareous soils showed small gains and in some cases even losses of soil C. We further showed that the largest part of the overall sample variance was not explained by fine-earth stock variances, rather by the C concentrations variance. The applied uncertainty analyses in this study link the variability of strata with measurement errors. In accordance to other studies for Central Europe, the results showed that the applied method enabled a reliable nationwide quantification of the soil C pool development for a certain period.

Temperature responses of growth and wood anatomy in European beech saplings grown in different carbon dioxide concentrations.

Effects of temperature on growth and wood anatomy were studied in young European beech (Fagus sylvatica L.) grown in 7-l pots for 2.5 years in field-phytotron chambers supplied with an ambient (approximately 400 micromol mol-1) or elevated (approximately 700 micromol mol-1) carbon dioxide concentration ([CO2]). Temperatures in the chambers ranged in increments of 2 degrees C from -4 degrees C to +4 degrees C relative to the long-term mean monthly (day and night) air temperature in Berlin-Dahlem. Soil was not fertilized and soil water and air humidity were kept constant. Data were evaluated by regression analysis. At final harvest, stem diameter was significantly greater at increased temperature (Delta1 degrees C: 2.4%), stems were taller (Delta1 degrees C: 8.5%) and stem mass tree-1 (Delta1 degrees C: 10.9%) and leaf area tree-1(Delta1 degrees C: 6.5%) were greater. Allocation pattern was slightly influenced by temperature: leaf mass ratio and leaf area ratio decreased with increasing temperature (Delta1 degrees C: 2.3% and 2.2% respectively), whereas stem mass/total mass increased (Delta1 degrees C: 2.1%). Elevated [CO2] enhanced height growth by 8.8% and decreased coarse root mass/total mass by 10.3% and root/shoot ratio by 11.7%. Additional carbon was mainly invested in aboveground growth. At final harvest a synergistic interaction between elevated [CO2] and temperature yielded trees that were 3.2% taller at -4 degrees C and 12.7% taller at +4 degrees C than trees in ambient [CO2]. After 2.5 seasons, cross-sectional area of the oldest stem part was approximately 32% greater in the +4 degrees C treatment than in the -4 degrees C treatment, and in the last year approximately 67% more leaf area/unit tree ring area was produced in the highest temperature regime compared with the lowest. Elevated [CO2] decreased mean vessel area of the 120 largest vessels per mm2 by 5.8%, causing a decrease in water conducting capacity. There was a positive interaction between temperature and elevated [CO2] for relative vessel area, which was approximately 38% higher at +4 degrees C than at -4 degrees C in elevated [CO2] compared with ambient [CO2]. Overall, temperature had a greater effect on growth than [CO2], but elevated [CO2] caused quantitative changes in wood anatomy.