Growth Of Beech Research Articles

Protection of understory vegetation by deer exclosure fences prevent the reduction of beech growth due to soil erosion

Overbrowsing by ungulates has degraded understory vegetation in worldwide forests. Understory degradation causes soil erosion and environmental degradation, causing mature tree decline. Although ungulate exclosure fences are installed to protect vegetation, their effectiveness in preventing tree decline associated with soil degradation remains unclear. We aimed to fill this gap on slopes of Mt. Shiraga, Japan, where sika deer have overbrowsed for >20 years. Tree-ring records in 1960–2022 for eight beeches per each of inside and outside fences were analyzed to compare stem basal area increment (BAI, cm−2 yr−1) and trend change timings for BAI (i.e., break-point). Soil properties, exposed root height (ERH) as cumulative soil erosion index, and intrinsic water use efficiency (iWUE) as water limitation proxy were also compared. Compared with areas inside fences, areas outside fences had no understory vegetation, higher ERH, and higher soil hardness and moisture. BAI outside fences turned their growth trend from positive to negative since 2004, but not inside fences. Break-point of BAI outside fences matched the understory degradation period in the field. The long-term BAI trend (i.e., regression slope of BAI) after the break-point was negatively related to ERH but not to soil properties (e.g., soil hardness and moisture). Furthermore, BAI outside fences was negatively related to iWUE after the break-point, whereas BAI inside fences was not. These results indicate that the maintenance of understory by fencing prevents overbrowsing-associated mature tree decline through mitigating root exposure and related water limitation, despite the potential of water competition with understory.

Climatic Effects on the Growth of Fagus sylvatica L. in Mixed Stands with Pinus sylvestris L. in Lithuania

Climate change and warming will potentially profoundly affect forest growth and yield. However, climate change is not only a threat but also an opportunity for tree species to be more productive or colonize new territories where previously they were fragmented or absent. European beech (Fagus sylvatica L.) is a dominant and significant tree species in European forests with a pronounced ability to grow on a great variety of sites and to form mixtures with other tree species due to its shade tolerance. So far, only a few studies have analysed its growth and reaction to changing climatic conditions outside its natural gradient in the north. The aim of this study was therefore to clarify the main growth patterns and tree ring formation characteristics of beech (outside their natural distribution) in mixed stands with Scots pine (Pinus sylvestris L.) The analyses were carried out in two-layered stands in the western part of Lithuania with tree-ring data measured on stem disks collected at 1 m height that were available from shelter-cut thinning activities in a number of forest stands in the region. Our results show that higher temperatures during the summer months of July and August generally inhibited the radial growth of beech under given site conditions. In contrast, warmer late-autumn and cooler early-winter periods together with warmer springs positively affected the radial increments. Above-average precipitation during the vegetation period contributed to the beech producing additional amounts of radial increments. The increasing temperature trends of the summer months and decreasing monthly amounts of precipitation during the vegetation period in recent decades tend to have adverse effects on the ecological conditions for beech growth and vitality in Lithuania. Despite these results, the European beech successfully grows in the sample region and has produced 200 m3 ha−1 in the analysed stands.

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.

Site conditions rather than provenance drive tree growth, climate sensitivity and drought responses in European beech in Germany

Ongoing climate change and associated extreme events strongly impact the growth and vitality of forest ecosystems in Europe. Because of its’ high drought sensitivity, European beech, which is considered as climax tree species in large parts of Central Europe, may specifically suffer. Hence, recent studies increasingly focus on the resistance and resilience of beech growth to climate change. Intra-specific variations in growth responses by comparing different beech provenances, however, received less attention, as did the question whether provenance selection can be used to mitigate potential future negative impacts of climate change. Therefore, we here investigated 24 provenances belonging to the International Beech Provenance Trial growing at three sites in Germany along a latitudinal gradient (study sites are referred to as ’North’, ‘Center’, ‘South’). Specifically, we compared tree-ring width (TRW), diameter breast height (DBH), climate-growth relationships, as well as drought resistance and resilience in the extreme years 2003 and 2018. Large differences in growth performance were observed between the three study sites. At site North, beech trees showed the highest DBH and TRW. Tree growth was predominantly driven by previous-year October and current-year winter temperature, whereas growth at sites Center and South was significantly impacted by summer SPEI and constrained by precipitation in late winter and early June, respectively. Overall, drought responses in 2003 were less variable than in 2018. We found increasing resistance and decreasing resilience from the wetter North to the drier South, but with minimal differences between the Center and the South. Whereas differences between study sites were large, provenance differentiation within sites was comparably low, substantiating that beech is a highly plastic tree species. Even though some provenances were found to perform slightly better or worse, differences were not statistically significant and show unclear patterns. Hence, we conclude that climate change will affect beech forests in Europe mainly depending upon site conditions, and that provenance selection may not ensure superior growth performance.

Korištenje morfoloških značajki sadnica obične bukve (Fagus sylvatica L.) za selekciju reprodukcijskog materijala iz provenijencija vjerojatno zahvaćenih klimatskim promjenama

Global climate change causes the reduction of tree growth and spread of European beech (Fagus sylvatica L.) forests, thus indicating the need to preserve tree abundance due to the changes that have occurred. The comparison of beech seedlings from various provenances in southeast and central Europe was carried out based on the dimensions of root collar diameter and height to select the proper starting material that would reduce the negative consequences of the climate. There have been statistically significant differences in root collar diameter and height between various seedlings tested in European provenances. Clear differentiation was recorded between European countries in the growth performances of one-year-old and two-year-old seedlings of F. sylvatica based on the nursery test. The study's results indicate the potential for producing adaptable beech reproductive material from various European provenances. This finding lays the groundwork for future precise analyses that aim to select provenances tolerant to changing climate conditions.

Drought effects on growth and density of temperate tree regeneration under different levels of nitrogen deposition

Temperate forests in Central Europe suffer from climate change-induced productivity and vitality reductions and increased tree mortality. Most field work assessing climate change effects in forests refers to mature trees and does not cover interaction between climate change and nitrogen deposition. Here we show in a study of 54 forest sites representing different combinations of climatic conditions and atmospheric nitrogen deposition across Germany that nitrogen significantly affects the drought tolerance of tree regeneration in the field. We compared shoot length increment and regeneration density of Central Europe’s naturally most dominant tree species, European beech (Fagus sylvatica) with three species, which are discussed as potentially more drought-tolerant replacement tree species for climate change adaptation of forestry. Growth of beech was reduced with increasing nitrogen deposition identifying high reactive nitrogen loads as an additional threat for this species, but between-site variation of beech growth was not dependent on climatic parameters despite growth reductions after the drought summer 2018 across all study sites. Remarkably, the only species in the between-site comparisons where low growth was attributable to dry climate was Douglas fir (Pseudotsuga menziesii), as shoot length increment was reduced at sites with dry spring climate. Silver fir (Abies alba) showed increasing growth with increasing nitrogen deposition in combination with increasing drought, probably due to reduced ammonium uptake. Shoot length increment in sessile oak (Quercus petraea) was not affected by the between-site variation of either drought or nitrogen. Our results (combined with results of regeneration density) indicate that high atmospheric nitrogen deposition, which is primarily found in regions with intensive livestock farming in Central Europe, modifies the climate change response of temperate tree species. Interaction of drought and nitrogen deposition should thus be addressed in the debate on climate change adaptation of forestry.

The Dynamics of Stand Structure Development and Natural Regeneration of Common Beech (Fagus sylvatica L.) in Plitvice Lakes National Park

The authors investigate the structural characteristics, regeneration processes, growth, development, and survival of a young generation of common beech (Fagus sylvatica L.) based on three periodic measurements (1998, 2009, and 2018). The studied forest community (Lamio orvale-Fagetum sylvaticae (Ht. 1938) Bohridi 1963) is situated within a forest reserve in Plitvice Lakes National Park, Croatia. Monitoring was carried out according to UN/ECE (2000) for systematic and repeated research. The basic structural indicators, structural canopy elements, and appearance of the young generation were measured as the basic conditions of natural restoration in repeated phytocenological surveys (1980, 1988, 2004). The distribution of the number of trees of the first two measurements (1998–2009) indicates a distribution with pronounced right asymmetry. The structural dynamics observed during the surveys and alongside vegetation surveys indicate the dynamics of the growth and development of beech. The results show that the main drivers of development dynamics in protected forest ecosystems are structural breaks (gaps) that occur in stands due to the action of abiotic and biotic factors. The passive management model in effect in the national park should be replaced with a more active approach to facilitate natural processes with the aim of preserving and renewing the forest ecosystem.

Effects of Drought, Phosphorus Fertilization and Provenance on the Growth of Common Beech and Sessile Oak

The negative impact of drought on plant growth may be modified by the different availability of mineral nutrients and by their adaptation to different local habitat conditions. In this study, we examine the impact of drought, fertilization with phosphorus and provenance, as well as their interactions, on the growth and allometric growth relationships between the belowground and aboveground organs of common beech (Fagus sylvatica L.) and sessile oak (Quercus petraea (Matt.) Liebl.). The research was conducted on saplings originating from two mature mixed stands (dry and wet provenances) dominated by these species. In the common garden experiment, saplings were exposed to regular watering and drought in interaction with moderate and high phosphorus concentrations in the growing substrate (achieved by phosphorus fertilization). The obtained results indicate the negative impact of drought and phosphorus fertilization on the growth of both species. In common beech, a negative impact of phosphorus fertilization on the adaptive capacity to drought was demonstrated by unfavorable ratios between fine root mass and the mass of other organs. The sessile oak provenances under the impact of drought showed a different root collar diameter/stem height increment ratio, which indicates their different phenotypic plasticity as a consequence of adaptation to different frequencies of dry periods in their natural habitats.

The effects of geomorphology, soil and climate on the trajectory of aboveground biomass accumulation of beech (Fagus sylvatica L.) at the southern range margin

Forest ecosystems are among the most important carbon sinks in the terrestrial biomes. Therefore, the estimation of aboveground biomass accumulation is of fundamental importance for understanding the contribution of forest stands to the global carbon budget. In this study, we proposed a modelling approach to estimate aboveground biomass of beech forests in the Central Apennine based on stand age, climatic variables, topographic features and soil parameters. Using forest inventory data from the local forest management plans, international databases, and mixed-effect linear model, we identified stand age as a major driver of beech forests aboveground biomass. Climatic variables had generally higher influence than soil and topographic features, probably as a consequence of the homogeneous calcareous substrate which favoured the development of soils which are highly suitable for the growth of beech. Temperature range and seasonality were the most important climatic variables. Interestingly, we found that the aboveground biomass in Apennine beech forests is strongly site-specific. Different management approaches during the past centuries, i.e. presence or absence of conversion interventions of the analysed stands, are probably responsible for the growing site's significant influence. By highlighting the ideal locations for allocating the functions of forest production or evaluating the value of ecosystem services that regulate the climate, this study will help to improve both the precision of carbon budget modelling and decision-making in nature conservation and forest management.

Root overlap and allocation of above- and belowground growth of European beech in pure and mixed stands of Douglas fir and Norway spruce

Site conditions and species identity have a combined effect on fine root growth of trees in pure and mixed stands. However, mechanisms that may contribute to this effect are rarely studied, even though they are essential to assess the potential of species to cope with climate change. This study examined fine root overlap and the linkage between fine root and stem growth of European beech (Fagus sylvatica) growing in pure and mixed stands with Douglas fir (Pseudotsuga menziesii) or Norway spruce (Picea abies) at two different study sites in northwestern Germany.The study sites represented substantially different soil and climate conditions. At each site, three stands, and at each stand, three pairs of trees were studied. In the pure beech stand, the pairs consisted of two beech trees, while in the mixed stands each pair was composed of a beech tree and a conifer. Between each pair, three evenly spaced soil cores were taken monthly throughout the growing season. In the pure beech stands, microsatellite markers were used to assign the fine roots to individual trees. Changes in stem diameter of beech were quantified and then upscaled to aboveground wood productivity with automatic high-resolution circumference dendrometers.We found that fine root overlap between neighboring trees varied independently of the distance between the paired trees or the stand types (pure versus mixed stands), indicating that there was no territorial competition. Aboveground wood productivity (wood NPP) and fine root productivity (root NPP) showed similar unimodal seasonal patterns, peaking in June. However, this pattern was more distinct for root NPP, and root NPP started earlier and lasted longer than wood NPP. The influence of site conditions on the variation in wood and root NPP of beech was stronger than that of stand type. Wood NPP was, as expected, higher at the richer site than at the poorer site. In contrast, root NPP was higher at the poorer than at the richer site.We concluded that beech can respond to limited resources not only above- but also belowground and that the negative relationship between above- and belowground growth across the study sites suggests an ‘optimal partitioning’ of growth under stress.

Low but significant evolutionary potential for growth, phenology and reproduction traits in European beech

AbstractLocal survival of forest tree populations under climate change depends on existing genetic variation and their adaptability to changing environments. Responses to selection were studied in European beech (Fagus sylvatica) under field conditions. A total of 1087 adult trees, seeds, 1‐year‐old seedlings and established multiyear saplings were genotyped with 16 nuSSRs. Adult trees were assessed for phenotypic traits related to growth, phenology and reproduction. Parentage and paternity analyses were used to estimate effective female and male fecundity as a proxy of fitness and showed that few parents contributed to successful regeneration. Selection gradients were estimated from the relationship between traits and fecundity, while heritability and evolvability were estimated using mixed models and the breeder's equation. Larger trees bearing more fruit and early male flowering had higher total fecundity, while trees with longer growth season had lower total fecundity (directional selection). Stabilizing selection on spring phenology was found for female fecundity, highlighting the role of late frosts as a selection driver. Selection gradients for other traits varied between measurement years and the offspring cohort used to estimate parental fecundity. Compared to other studies in natural populations, we found low to moderate heritability and evolvability for most traits. Response to selection was higher for growth than for budburst, leaf senescence or reproduction traits, reflecting more consistent selection gradients across years and sex functions, and higher phenotypic variability in the population. Our study provides empirical evidence suggesting that populations of long‐lived organisms such as forest trees can adapt locally, even at short‐time scales.

Chronic warming and dry soils limit carbon uptake and growth despite a longer growing season in beech and oak.

Progressively warmer and drier climatic conditions impact tree phenology and carbon cycling with large consequences for forest carbon balance. However, it remains unclear how individual impacts of warming and drier soils differ from their combined effects and how species interactions modulate tree responses. Using mesocosms, we assessed the multiyear impact of continuous air warming and lower soil moisture alone or in combination on phenology, leaf-level photosynthesis, nonstructural carbohydrate concentrations, and aboveground growth of young European beech (Fagus sylvatica L.) and Downy oak (Quercus pubescens Willd.) trees. We further tested how species interactions (in monocultures and in mixtures) modulated these effects. Warming prolonged the growing season of both species but reduced growth in oak. In contrast, lower moisture did not impact phenology but reduced carbon assimilation and growth in both species. Combined impacts of warming and drier soils did not differ from their single effects. Under warmer and drier conditions, performances of both species were enhanced in mixtures compared to monocultures. Our work revealed that higher temperature and lower soil moisture have contrasting impacts on phenology vs. leaf-level assimilation and growth, with the former being driven by temperature and the latter by moisture. Furthermore, we showed a compensation in the negative impacts of chronic heat and drought by tree species interactions.

Ecophysiological Behavior of Fagus sylvatica L. Growing at Its Southern Distribution Limit: Insights for Understanding the Fate of the European Beech under Warmer and Dryer Growth Conditions

In the last 20 years, a significant mortality of Fagus sylvatica L. (European beech) has been documented in central and northern European forests. Surprisingly, no beech die off occurred at the southern limit. This fact leads us to hypothesize that European beech populations growing at the southern limit of the distribution might have a significant phenotypic plasticity to better cope with low water availability and rising temperatures. To check this hypothesis, we evaluated the ecophysiological behavior of F. sylvatica growing along an altitudinal transect in Calabria (Italy). We selected three study sites (750 m a.s.l., 976 m a.sl. m a.s.l., 1450 m a.s.l.) showing narrow ranges of temperature, rainfall and air humidity. Trees growing at 976 m a.s.l. showed the highest stomatal conductance values during the entire experimental period. The lowest gas exchange and highest leaf mass area were recorded in plants growing at 750 m a.s.l. In the European beech growing at 1450 m a.s.l., higher vessel density, lower mean vessel diameter and higher vessel grouping index values were recorded. Overall, our results highlighted that the measured populations show a considerable phenotypic plasticity leading them to adjust anatomical and physiological traits in response to narrow ranges of environmental parameters. Despite that, the distribution of F. sylvatica seems to be limited to areas with a growing season rainfall of at least 400 mm and vapor pressure deficit (VPD) values < 3 kPa, which may represent the main environmental thresholds which strongly limit the beech growth and, therefore, influence the ability of this species to cope with future environmental conditions.

Soil erosion under forest hampers beech growth: Impacts of understory vegetation degradation by sika deer

The overpopulation of large herbivorous mammals degrades understory vegetation and hence promotes soil erosion in some northern hemisphere forests. Topsoil removal via erosion reduces soil water and nitrogen (N) content as well as exposes tree roots. However, whether topsoil removal inhibits tree growth through water and N uptake limitations has not been examined. This study aimed to measure the vertical length from exposed roots surface to the soil surface (exposed root height; ERH) under degraded forest understory vegetation and to evaluate its influence on tree growth. Specifically, we examined the relationships between ERH and beech growth and identified the timing of growth reduction based on annual tree-ring analysis at the individual scale. Furthermore, we discussed the effects of water and N limitations on beech growth reduction. We studied 12 Japanese beech trees on Mt. Sanpo, southern Kyushu Island, Japan, where understory vegetation was degraded and lost due to herbivory by Japanese sika deer from 1980 to 2003. Lower amounts of leaf production and relative growth rates of stem basal area increment (BAI) were observed in beech trees with higher ERH. Segmented regression analysis, which detects the changing time of a temporal trend (i.e., break-point), revealed that the BAI has resulted in a decreasing trend after 1997.6 ± 9.0 (yr), which coincided with the field observations on understory degradation in the study site. After the break-point of the BAI, the BAI of each study tree was negatively related to intrinsic water use efficiency, an integrated index of water limitation. These results suggest that soil erosion after the degradation of understory vegetation reduced beech growth, possibly because of the water limitations associated with root exposure. Reduced beech growth will reduce the supply of the litter layer that prevents topsoil erosion, which may enhance further soil erosion and tree growth reduction.

Growth of European beech recovered faster than that of Norway spruce after a five-year experimental drought in a mixed forest stand

Key MessageBeech growth acclimated better during severe drought and recovered faster than spruce after drought ended. This was associated with a shift in performance along relative tree size towards small trees.The effects of several consecutive drought years and the recovery reaction of mature trees in particular after a long-term drought have been poorly studied so far. In this study, we demonstrate the growth reactions of mature trees during and after a five-year treatment of extended summer droughts, followed by controlled irrigation in a very productive mixed forest stand. We exposed 70-year-old Norway spruce (Picea abies [L.] Karst) and 90-year-old European beech (Fagus sylvatica [L.]) trees to reduced precipitation using automatic throughfall exclusion (TE) roofs during the growing seasons from 2014 to 2018, irrigated the trees in early summer 2019 and removed the roofs thereafter. From 2009 to 2022, we monitored annual tree growth and precipitation on 6 plots with throughfall exclusion and on 6 plots with ambient Control conditions (CO) of the KROOF canopy experiment. Norway spruce lost significant growth during drought, with some trees dying and others remaining at a low growth level without significant recovery from the effects of drought stress. European beech also significantly reduced growth at the beginning of the drought but emerged stronger in growth from the drought than the Control group. Spruce and beech showed a non-significant trend of increased inter-specific growth compared to intra-specific growth during drought. We found that spruce benefitted more from mixture than beech in the recovery phase after drought than during the drought phase itself. Most importantly, we observed a shift in growth performance along the relative tree size towards smaller trees in the TE plots for both species. This change in the relationship between diameter increment and tree size during and after drought is a major finding of our study and suggests a possible response mechanism to prolonged drought. This key observation requires further investigation and should be considered in future forest management strategies under changing climatic conditions.

Spatial and temporal variation of Fagus sylvatica growth in marginal areas under progressive climate change

The escalating decline in growth trends of European beech (Fagus sylvatica) observed across its distribution area pose a major ecological and economic challenge for countries with a high proportion of beech, such as Slovenia. In this study, the effects of climate change were examined at a high-resolution scale, encompassing the large climatic, orographic, and ecological variability of beech forests in Slovenia. Using basal area increment data (BAI) from a tree-ring network (48 sites in Slovenia), modelled climate data, and generalized linear mixed models (GLMM), we found an average growth decline of 11% between the 1953–1985 and 1986–2018 subperiods, affecting 90.5% of the forest stands. Based on climate data, we defined two contrasted marginal areas of beech presence (warm and cold) and analysed the growth changes over time. The warm marginal areas, which predominate near the geographical margin of beech distribution with a sub-Mediterranean climatic regime, were most affected by growth decline, threatening the survival of beech populations in the area. In contrast, cold marginal areas, mainly at high elevations in the Alps, where beech growth had previously been limited by low temperatures, turned out to be the only ones where growth of beech increased under prevailing warming conditions. Consequently, high elevation regions harbour climatic potential for increased beech growth performance, and may represent areas of future expansion of the species.

The Future of European Beech in Northern Germany—Climate Change Vulnerability and Adaptation Potential

Earth’s forests are increasingly exposed to climate risks through climate change-related drought and heat waves. Here, we review the climate vulnerability of European beech (Fagus sylvatica), the most abundant tree species of Central Europe’s temperate forests, and its adaptation potential to a hotter and drier climate in northern Germany, close to the center of the species’ distribution range. About two-thirds of the studied beech forests show persistent negative growth trends of their dominant trees since the onset of rapid warming in the early 1980s, driven primarily by long-term deterioration of the climatic water balance (CWB) in summer and, locally, long-term decrease in June precipitation. Lower water storage capacity of the soil increases the climate sensitivity of growth. Even though beech populations of drier sub-regions reveal some acclimation to drought (reduced stomatal conductance, turgor loss point reduction, higher C allocation to roots), they are more climate-vulnerable than populations at moister sites, visible in larger growth decreases and greater CWB sensitivity of growth. Even though beech mortality is still lower in the study region than in other parts of Central Europe, our results identify beech forest regions with <350 mm growing-season precipitation as vulnerable to recent climate conditions, which refers to half of the study region. With further climate warming and aridification, most of northern Germany likely will become unfavorable for beech growth within the next tree generation, which suggests choosing in many production forests alternative, more drought- and heat-resistant timber species.

Water use efficiency and climate legacies dominate beech growth at its rear edge

Abstract Rear‐edge tree populations are experiencing a combination of higher temperatures and more intense droughts that might push individuals beyond their tolerance limits. This trend towards rising atmospheric [CO2] is concurrent with an increase in intrinsic water use efficiency (iWUE), which theoretically enhances photosynthesis and decrease evapotranspiration rates, consequently improving tree resistance to drought. However, it remains unclear whether iWUE is favouring tree growth under current climate conditions, particularly when climate and iWUE legacy effects are simultaneously considered. We evaluated this question with an extensive sampling along Iberian rear‐edge (dry) populations comprising four mountain ranges and two distinct altitudes. We simultaneously examined the effects of climate and iWUE on secondary growth using annually resolved basal area increments (BAIs) for the period 1901–2017. We used linear mixed models including second‐order autocorrelation and 1‐year legacy effects of iWUE and summer drought. BAI and iWUE increased across the studied period. iWUE increase was driven by changes in atmospheric CO2 concentration and water availability during the growing season. Climate and iWUE exerted direct and lagged effects on beech growth. Water availability during growing season was the main driver of tree growth, combining direct and indirect effects through its impact on iWUE. Legacy effects of water availability and iWUE were more important than growing season conditions. The net effect of iWUE shifted when lagged effects were considered, resulting in a net negative impact on tree growth. Synthesis: Our results reveal that climate and iWUE legacy effects must be considered to assess the net iWUE effect on secondary growth. Considering lagged effects, the current increase in iWUE is constraining tree growth. Modelling efforts of tree growth response to climate warming should include climate and iWUE legacy effects to adequately assess terrestrial ecosystem carbon balance.

Growth Response of European Beech (Fagus sylvatica L.) and Silver Fir (Abies alba Mill.) to Climate Factors along the Carpathian Massive

European forests are becoming increasingly threatened by climate change and more frequent droughts. The likely responses of species to climate change will vary, affecting their competitiveness, their existence, and consequently, forest management decisions and measures. We determined the influence of climate on the radial growth of European beech and silver fir along the Carpathians to find similarities between the two species and the main differences. Along the Carpathian Mountains, seven sites with mature fir–beech stands above 800 m above sea level were selected and analyzed. Our study confirmed different responses depending on species and location. A more pronounced response of tree growth to climate was observed on the eastern side of the Carpathians, while it was less expressed or even absent on the southern sites. Both beech and fir show better radial growth with higher precipitation in July and slower growth with higher average and maximum temperatures in June of the current year. Fir demonstrates a positive correlation between radial growth and temperature in winter, while beech demonstrates a negative correlation between radial growth and temperature in summer. In the 1951–1960 decade, the average tree ring widths in fir and beech were largest at the southern sites compared to the other sites, but since 2011, the southern sites have had the lowest increase while northern sites have had the largest. Both species respond differently to climate and are likely to follow different competitive paths in the future.

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.