Changes in Root-Shoot Allometric Relations in Alpine Norway Spruce Trees After Strip Cutting.

Silvicultural interventions such as strip cuttings can change the resource availability of the edge trees. This may alter tree allometry, as light regime, water, and nutrient availability can change at the forest edge. Increased root growth may optimize resource uptake and/or enhance tree anchorage to withstand the altered wind regime. However, little is known about the patterns of the root-shoot allometric responses to strip cuttings. In three alpine stands differing in climate, site productivity, and stand characteristics, we selected 71 Norway spruce trees and took increment cores from stems, root collars, and main roots. This enabled us to study changes in the long-term root-stem allometry for 46 years and short-term allometric responses to intervention. The effects of cutting were compared between edge trees and trees from the stand interior in 10 years before and after the intervention. The long-term allocation to roots increased with stem diameter, with the strongest effects on the regularly managed stand with the tallest and largest trees. These results support the allometric biomass partitioning theory, which postulates resource allocation patterns between different plant organs to depend on plant size. Strip cutting on north-facing slopes boosted edge-tree growth in all plant compartments and enhanced allocation to roots. This change in allometry started 2 years after cutting but disappeared 7-8 years later. In the post-cutting period, the highest root-shoot increase was observed in the small trees independent of the site. This indicates the change in growing conditions to have the strongest effects in formerly suppressed trees. Thus, the effect of such acclimation on the wind firmness of subdominant spruce trees is a question with high importance for optimizing cutting layouts in lowering post-cutting vulnerability to disturbance. The results from this case study contribute to a better understanding of the structural acclimation of spruce trees from high-elevation forests to new forest edges. However, for a more mechanistic understanding of environmental drivers, further analyses of tree-ring stable isotopes are recommended.

Grow slowly, persist, dominate-Explaining beech dominance in a primeval forest.

Being able to persist in deep shade is an important characteristic of juvenile trees, often leading to a strong dominance of shade-tolerant species in forests with low canopy turnover and a low disturbance rate. While leaf, growth, and storage traits are known to be key components of shade tolerance, their interplay during regeneration development and their influence on juveniles' survival time remains unclear. We assessed the ontogenetic effects of these three traits on the survival time of beech (Fagus sylvatica), and Norway and sycamore maples (Acer pseudoplatanus, Acer platanoides) in a primeval beech forest. Biomass allocation, age, and content of nonstructural carbohydrates (NSC) were measured in the stems and roots of 289 seedlings and saplings in high- and low-vitality classes. Saplings experienced a trade-off between absolute growth rate (AGR) and storage (NSC) as the leaf area ratio (LAR) decreases with biomass development. High LAR but low AGR and low NSC corresponded to beech with a marked ability to persist in deep shade while awaiting canopy release. In turn, a comparably small LAR in combination with a high AGR and higher storage (NSC), as observed in Norway maple and sycamore maple, reduced sapling survival time, thus offering an explanation for beech dominance and maple disappearance in the undergrowth of old-growth beech forests.

Stand inventory data from the 10-ha forest research plot in Uholka: 15 yr of primeval beech forest development.

Forests dominated by European beech (Fagus sylvatica L.) are among both the most widespread in Europe and the most intensely exploited globally. One of the largest remnants of unmanaged beech forests in Europe is the Uholka-Shyrokyi Luh forest in Transcarpathia, Ukraine, covering 8,800 ha of primeval forest. In 2000, a permanent forest plot of 10 ha has been established in the Uholka massif. All living and dead trees with a diameter at breast height (DBH) ≥ 60 mm were identified to species, DBH measured, stems tagged and mapped. Since then, the plot has been remeasured in 2005, 2010, and 2015. In total, 4,820 individual trees were measured with 14,116 individual measurements throughout all four inventories. In spring 2018, an Airborne Laser Scan was carried out, covering the Uholka-Shyrokyi Luh forest. This data set allows us to derive a high-resolution digital elevation model (DEM) of the plot area. European beech covers a share of ≈ 95% of the basal area (BA) of all living stems. While BA was relatively stable throughout all inventories (≈ 38 m2 /ha), the number of stems increased considerably between 2010 and 2015 from 290 to 430 stems/ha. Additionally, the proportion of beech stems decreased from 95% in 2010 to 86% in 2015. The continuity of the share of beech on BA and the decrease in number of stems can be traced back to disturbance events, which led to an increase of more light demanding species in the recruitment but did not alter the distribution of BA as these small trees contribute very little to BA. The data set allows for important insights into the development and the spatial and temporal dynamics of primeval beech forests. It can be used to quantify the demographic processes growth, mortality, and recruitment, and to study inter- and intraspecific effects on demographic rates, stand structure, and species composition. No copyright or proprietary restrictions are associated with the use of this data set other than citation of this Data Paper.

How to kill a tree: empirical mortality models for 18 species and their performance in a dynamic forest model.

Dynamic Vegetation Models (DVMs) are designed to be suitable for simulating forest succession and species range dynamics under current and future conditions based on mathematical representations of the three key processes regeneration, growth, and mortality. However, mortality formulations in DVMs are typically coarse and often lack an empirical basis, which increases the uncertainty of projections of future forest dynamics and hinders their use for developing adaptation strategies to climate change. Thus, sound tree mortality models are highly needed. We developed parsimonious, species-specific mortality models for 18 European tree species using >90,000 records from inventories in Swiss and German strict forest reserves along a considerable environmental gradient. We comprehensively evaluated model performance and incorporated the new mortality functions in the dynamic forest model ForClim. Tree mortality was successfully predicted by tree size and growth. Only a few species required additional covariates in their final model to consider aspects of stand structure or climate. The relationships between mortality and its predictors reflect the indirect influences of resource availability and tree vitality, which are further shaped by species-specific attributes such as maximum longevity and shade tolerance. Considering that the behavior of the models was biologically meaningful, and that their performance was reasonably high and not impacted by changes in the sampling design, we suggest that the mortality algorithms developed here are suitable for implementation and evaluation in DVMs. In the DVM ForClim, the new mortality functions resulted in simulations of stand basal area and species composition that were generally close to historical observations. However, ForClim performance was poorer than when using the original, coarse mortality formulation. The difficulties of simulating stand structure and species composition, which were most evident for Fagus sylvatica L. and in long-term simulations, resulted from feedbacks between simulated growth and mortality as well as from extrapolation to very small and very large trees. Growth and mortality processes and their species-specific differences should thus be revisited jointly, with a particular focus on small and very large trees in relation to their shade tolerance.

Risk of genetic maladaptation due to climate change in three major European tree species.

Tree populations usually show adaptations to their local environments as a result of natural selection. As climates change, populations can become locally maladapted and decline in fitness. Evaluating the expected degree of genetic maladaptation due to climate change will allow forest managers to assess forest vulnerability, and develop strategies to preserve forest health and productivity. We studied potential genetic maladaptation to future climates in three major European tree species, Norway spruce (Picea abies), silver fir (Abies alba), and European beech (Fagus sylvatica). A common garden experiment was conducted to evaluate the quantitative genetic variation in growth and phenology of seedlings from 77 to 92 native populations of each species from across Switzerland. We used multivariate genecological models to associate population variation with past seed source climates, and to estimate relative risk of maladaptation to current and future climates based on key phenotypic traits and three regional climate projections within the A1B scenario. Current risks from climate change were similar to average risks from current seed transfer practices. For all three climate models, future risks increased in spruce and beech until the end of the century, but remained low in fir. Largest average risks associated with climate projections for the period 2061-2090 were found for spruce seedling height (0.64), and for beech bud break and leaf senescence (0.52 and 0.46). Future risks for spruce were high across Switzerland. However, areas of high risk were also found in drought-prone regions for beech and in the southern Alps for fir. Genetic maladaptation to future climates is likely to become a problem for spruce and beech by the end of this century, but probably not for fir. Consequently, forest management strategies should be adjusted in the study area for spruce and beech to maintain productive and healthy forests in the future.

Distinct genecological patterns in seedlings of Norway spruce and silver fir from a mountainous landscape.

Understanding the genecology of forest trees is critical for gene conservation, for predicting the effects of climate change and climate change adaptation, and for successful reforestation. Although common genecological patterns have emerged, species-specific details are also important. Which species are most vulnerable to climate change? Which are the most important adaptive traits and environmental drivers of natural selection? Even though species have been classified as adaptive specialists vs. adaptive generalists, large-scale studies comparing different species in the same experiment are rare. We studied the genecology of Norway spruce (Picea abies) and silver fir (Abies alba), two co-occurring but ecologically distinct European conifers in Central Europe. For each species, we collected seed from more than 90 populations across Switzerland, established a seedling common-garden test, and developed genecological models that associate population variation in seedling growth and phenology to climate, soil properties, and site water balance. Population differentiation and associations between seedling traits and environmental variables were much stronger for Norway spruce than for silver fir, and stronger for seedling height growth than for bud phenology. In Norway spruce, height growth and second flushing were strongly associated with temperature and elevation, with seedlings from the lowlands being taller and more prone to second flush than seedlings from the Alps. In silver fir, height growth was more weakly associated with temperature and elevation, but also associated with water availability. Soil characteristics explained little population variation in both species. We conclude that Norway spruce has become an adaptive specialist because trade-offs between rapid juvenile growth and frost avoidance have subjected it to strong diversifying natural selection based on temperature. In contrast, because silver fir has a more conservative growth habit, it has evolved to become an adaptive generalist. This study demonstrates that co-occurring tree species can develop very different adaptive strategies under identical environmental conditions, and suggests that Norway spruce might be more vulnerable to future maladaptation due to rapid climate change than silver fir.

Does one model fit all? Patterns of beech mortality in natural forests of three European regions.

Large uncertainties characterize forest development under global climate change. Although recent studies have found widespread increased tree mortality, the patterns and processes associated with tree death remain poorly understood, thus restricting accurate mortality predictions. Yet, projections of future forest dynamics depend critically on robust mortality models, preferably based on empirical data rather than theoretical, not well-constrained assumptions. We developed parsimonious mortality models for individual beech (Fagus sylvatica L.) trees and evaluated their potential for incorporation in dynamic vegetation models (DVMs). We used inventory data from nearly 19,000 trees from unmanaged forests in Switzerland, Germany, and Ukraine, representing the largest dataset used to date for calibrating such models. Tree death was modelled as a function of size and growth, i.e., stem diameter (dbh) and relative basal area increment (relBAI), using generalized logistic regression accounting for unequal re-measurement intervals. To explain the spatial and temporal variability in mortality patterns, we considered a large set of environmental and stand characteristics. Validation with independent datasets was performed to assess model generality. Our results demonstrate strong variability in beech mortality that was independent of environmental or stand characteristics. Mortality patterns in Swiss and German strict forest reserves were dominated by competition processes as indicated by J-shaped mortality over tree size and growth. The Ukrainian primeval beech forest was additionally characterized by windthrow and a U-shaped size-mortality function. Unlike the mortality model based on Ukrainian data, the Swiss and German models achieved good discrimination and acceptable transferability when validated against each other. We thus recommend these two models to be incorporated and examined in DVMs. Their mortality predictions respond to climate change via tree growth, which is sufficient to capture the adverse effects of water availability and competition on the mortality probability of beech under current conditions.

Drought response of five conifer species under contrasting water availability suggests high vulnerability of Norway spruce and European larch.

The ability of tree species to cope with anticipated decrease in water availability is still poorly understood. We evaluated the potential of Norway spruce, Scots pine, European larch, black pine, and Douglas-fir to withstand drought in a drier future climate by analyzing their past growth and physiological responses at a xeric and a mesic site in Central Europe using dendroecological methods. Earlywood, latewood, and total ring width, as well as the δ(13) C and δ(18) O in early- and latewood were measured and statistically related to a multiscalar soil water deficit index from 1961 to 2009. At the xeric site, δ(13) C values of all species were strongly linked to water deficits that lasted longer than 11 months, indicating a long-term cumulative effect on the carbon pool. Trees at the xeric site were particularly sensitive to soil water recharge in the preceding autumn and early spring. The native species European larch and Norway spruce, growing close to their dry distribution limit at the xeric site, were found to be the most vulnerable species to soil water deficits. At the mesic site, summer water availability was critical for all species, whereas water availability prior to the growing season was less important. Trees at the mesic were more vulnerable to water deficits of shorter duration than the xeric site. We conclude that if summers become drier, trees growing on mesic sites will undergo significant growth reductions, whereas at their dry distribution limit in the Alps, tree growth of the highly sensitive spruce and larch may collapse, likely inducing dieback and compromising the provision of ecosystem services. However, the magnitude of these changes will be mediated strongly by soil water recharge in winter and thus water availability at the beginning of the growing season.

Ammonia monitoring in Switzerland with passive samplers: patterns, determinants and comparison with modelled concentrations.

Gaseous ammonia (NH3) is an important form of N deposition to ecosystems, but it is not being routinely monitored in Switzerland. Therefore, a study was conducted to estimate annual means and seasonal patterns of NH3 concentrations for different site types in Switzerland, and to compare annual measured and modelled NH3 concentrations. NH3 concentrations were measured using the 'Zürcher' passive sampler, a Palmes type sampler with an acidic solution as absorbent. Twenty-four sampling sites were run for one year, and 17 for two years. The samplers were changed fortnightly or monthly. Spatial emission patterns were mapped by combining information on (1) the location of emission sources, (2) national statistics on NH3-emitting activities and (3) activity-specific emission factors. The spatial resolution was one hectare. The mean annual NH3 concentration in the ambient air of the 41 sites was 2.5+/-0.3 microg m(-3) (mean+/-standard error). It ranged from 0.4 to 7.5 microg m(-3). The site type and the season were the most important factors explaining the variation in the seasonal mean concentration. NH3 concentrations were highest in intensively used agricultural areas and in cities, and lowest in Alpine sites remote from emission sources. At 39 out of 41 sites, the NH3 concentrations were higher in summer (3.1+/-0.3 microg m(-3)) than in winter (2.0+/-0.3 microg m(-3)). Modelled NH3 concentrations did not systematically deviate from measured concentrations (r2 = 0.69). With the combined monitoring and modelling approach, it is now possible to obtain a reasonable and consolidated picture of the overall NH3 situation in Switzerland.