Resistance of European tree species to drought stress in mixed versus pure forests: evidence of stress release by inter‐specific facilitation

While previous studies focused on tree growth in pure stands, we reveal that tree resistance and resilience to drought stress can be modified distinctly through species mixing. Our study is based on tree ring measurement on cores from increment boring of 559 trees of Norway spruce (Picea abies [L.] Karst.), European beech (Fagus sylvatica [L.]) and sessile oak (Quercus petraea (Matt.) Liebl.) in South Germany, with half sampled in pure, respectively, mixed stands. Indices for resistance, recovery and resilience were applied for quantifying the tree growth reaction on the episodic drought stress in 1976 and 2003. The following general reaction patterns were found. (i) In pure stands, spruce has the lowest resistance, but the quickest recovery; oak and beech were more resistant, but recover was much slower and they are less resilient. (ii) In mixture, spruce and oak perform as in pure stands, but beech was significantly more resistant and resilient than in monoculture. (iii) Especially when mixed with oak, beech is facilitated. We hypothesise that the revealed water stress release of beech emerges in mixture because of the asynchronous stress reaction pattern of beech and oak and a facilitation of beech by hydraulic lift of water by oak. This facilitation of beech in mixture with oak means a contribution to the frequently reported overyield of beech in mixed versus pure stands. We discuss the far‐reaching implications that these differences in stress response under intra‐ and inter‐specific environments have for forest ecosystem dynamics and management under climate change.     

Effects of changes in tree species composition on water flow dynamics – Model applications and their limitations

Water-plant relations play a key role in the water cycling in terrestrial ecosystems. Consequently, changes in tree species composition may have distinct effects on the water retention capacity as well as on the pattern of streamflow generation. Such changes may result from modified interception properties and transpiration related to differences in canopy properties and root distribution. In order to evaluate the potential hydrological effects of the current silvicultural conversion from monocultural conifer stands into mixed or pure deciduous stands the hydrological model BROOK90 was applied to two forested upland catchments in Germany. The Rotherdbach catchment (9.4 ha, 93 yr-old Norway spruce) is situated in the Eastern Ore Mountains. The Schluchsee catchment (11 ha, 55-yr-old Norway spruce) is located in the higher altitudes of the Black Forest. The calibrated model is capable to describe rather well the temporal variation of streamflow but also the portions of the individual flow components. Data for a beech scenario were adapted for each site using a standard parameter set for deciduous trees provided by BROOK90. The annual discharge in the fictional beech stand at Rotherdbach is 30 to 50% higher compared to spruce with an increase of soil moisture and especially the slow streamflow components. This mainly results from low interception rates during winter time. In contrast, the spruce stand has a permanently higher interception rate. Effects of tree species conversion are moderate at Schluchsee. The annual discharge of a fictional beech stand at Schluchsee is 7 to 14% higher compared to spruce. There in contrast to Rotherdbach, effects of tree species conversion on soil moisture dynamics are small since vertical percolation in the highly permeable soil dominates and precipitation is abundant. Practical forestry will favorably establish mixed beech–spruce rather than pure beech stands. However, it is critical to simulate mixed stands with BROOK90. Therefore, a simple summation of model results from spruce and beech according to their respective area in a fictional mixed stand can only be a first approximation. Advanced hydrological simulation of mixed stand conditions should regard interactions of tree species and spatial parameter distribution. However, this is not yet feasible due to a distinct lack of information. As a consequence, there is a strong need to collect relevant hydrological and ecophysiological data in mixed stands in the future.     

Relationships between tree dimension and coarse root biomass in mixed stands of European beech (Fagus sylvatica L.) and Norway spruce (Picea abies[L.] Karst.)

Relationships between tree parameters above ground and the biomass of the coarse root system were examined in six mixed spruce-beech stands in the Solling Mountain region in northwest Germany. The selected stands were located on comparable sites and covered an age range of 44 to 114 years. Coarse roots (d?\ge?2 mm) of 42 spruce and 27 beech trees were sampled by excavating the entire root system. A linear model with logarithmic transformation of the variables was developed to describe the relationship between the coarse root biomass (CRB, dry weight) and the corresponding tree diameter at breast height (DBH). The coefficients of determination (R ²) attained values between 0.92 for spruce and 0.94 for beech; the logarithmic standard deviation values were between 0.29 and 0.43. A significantly different effect of tree species on the model estimates could not be detected by an analysis of co-variance (ANCOVA). For spruce, the derived relationships were similar to those reported in previous studies, but not for beech. Biomass partitioning in the tree compartments above and below ground differs significantly between spruce (coarse root/shoot ratio 0.16±0.06) and beech (coarse root/shoot ratio 0.10±0.03) in the mixed stands. These results are similar to those given in other studies involving pure spruce and beech stands on comparable sites in the region, although the ratios of pure stands in other regions growing under different site conditions are somewhat higher. Comparing trees of the same DBH classes, root/shoot ratios of spruce are 1.2 to 3 times higher than those of beech. Dominant spruce trees (DBH>60 cm) attained the highest ratios, suppressed beech trees (DBH<10 cm) the lowest. Site conditions of varying climate and soils and interspecific tree competition are likely to affect root/shoot ratio and DBH-coarse root biomass relationships. The greater variability in beech compared with spruce indicates a high 'plasticity' and adaptability of beech carbon allocation. Thus, the derived equations are useful for biomass estimates of coarse roots involving trees of different ages in mixed stands of spruce and beech in the Solling Mountains. However, application of these relationships to stands in other regions would need further testing.     

Size‐dependence of tree growth response to drought for Norway spruce and European beech individuals in monospecific and mixed‐species stands

• Climate anomalies have resulted in changing forest productivity, increasing tree mortality in Central and Southern Europe. This has resulted in more severe and frequent ecological disturbances to forest stands. This study analysed the size‐dependence of growth response to drought years based on 384 tree individuals of Norway spruce [Picea abies (L.) Karst.] and European beech [Fagus sylvatica ([L.)] in Bavaria, Germany.
• Samples were collected in both monospecific and mixed‐species stands. To quantify the growth response to drought stress, indices for basal area increment, resistance, recovery and resilience were calculated from tree ring measurements of increment cores. Linear mixed models were developed to estimate the influence of drought periods.
• The results show that ageing‐related growth decline is significant in drought years. Drought resilience and resistance decrease significantly with growth size among Norway spruce individuals. Evidence is also provided for robustness in the resilience capacity of European beech during drought stress. Spruce benefits from species mixing with deciduous beech, with over‐yielding spruce in pure stands.
• The importance of the influence of size‐dependence within tree growth studies during disturbances is highlighted and should be considered in future studies of disturbances, including drought.

     

Mixed Norway spruce (Picea abies [L.] Karst) and European beech (Fagus sylvatica [L.]) stands under drought: from reaction pattern to mechanism

Key message

We review causes of synergies in mixed-species stands, develop guiding hypotheses for revealing their mechanisms and present a rainfall exclusion experiment along with a transect (KROOF) for exploring drought effects.

Abstract

While monocultures have dominated forest research and practice in the past, in face of growing resource scarcity and climate change, mixed-species stands are on the advance. Long-term observations show that mixed-species stands frequently over-yield monocultures, and they further suggest that the over-yielding is often higher on poor than on fertile sites and in low-growth than in high-growth years. However, the underlying causes have not yet been clarified. We start with a review of, among others, hydraulic redistribution, complementary eco-physiological traits, and ectomycorrhizal networks as possible causes behind the observed productivity gains in mixed-species stands. Then, we develop guiding hypotheses for further exploration of synergies in mixed-species stands. Finally, we introduce into the concept and model approach of the rainfall exclusion experiment for examining the role of water supply in mixed versus pure forest stands of spruce and beech. At the Kranzberg site, six plots are equipped with water retention roofs, which only close during rain events. The remaining six plots serve as non-roofed control. Together with the rainfall exclusion experiment, an ecological gradient with five sites extending through SE-Germany forms the “Kranzberg Roof Experiment” (in short KROOF). Kranzberg Forest is a part of this gradient from moist to dry conditions, with each site providing mixed and pure associations each of spruce and beech. The ecophysiological approach will be complemented by tree ring analysis and modelling of productivity of the tree associations under intense drought.     

Structural crown properties of Norway spruce (Picea abies [L.] Karst.) and European beech (Fagus sylvatica [L.]) in mixed versus pure stands revealed by terrestrial laser scanning

How tree morphology develops in mixed-species stands is essential for understanding and modelling mixed-stand dynamics. However, research so far focused on the morphological variation between tree species and neglected the variation within a species depending on intra- and interspecific competition. Our study, in contrast, addresses crown properties of nine mature Norway spruces (Picea abies [L.] Karst.) of a pure stand and compares them with ten spruces growing in mixture with European beech (Fagus sylvatica [L.]). The same was done with 11 pure stand beeches and 12 beeches growing in mixture with spruce. Through application of a terrestrial laser scanner and a new skeletonization approach, we deal with both species’-specific morphological traits such as branch angle, branch length, branch bending, crown volume and space occupation of branches within the crown, some of which were hardly accessible so far. Special attention is paid to distinct differences between trees growing in mixed and pure stands: for spruce, our study reveals significantly longer branches and greater crown volumes in the mixed stand when compared to the pure stand. In case of European beech, individuals growing in mixture show flatter branch angles, more distinct ramification, greater crown volumes and a lower share of a single branch’s space occupation in the total crown volume. The results show that the presented methods yield detailed information on the morphological traits analyzed in this study and that interspecific competition on its own may have a significant impact on crown structures. Implications for production ecology and stand dynamics of mixed-species forests are discussed.     

Greater accumulation of litter in spruce (Picea abies) compared to beech (Fagus sylvatica) stands is not a consequence of the inherent recalcitrance of needles

Background and aims

Replacement of beech by spruce is associated with changes in soil acidity, soil structure and humus form, which are commonly ascribed to the recalcitrance of spruce needles. It is of practical relevance to know how much beech must be admixed to pure spruce stands in order to increase litter decomposition and associated nutrient cycling. We addressed the impact of tree species mixture within forest stands and within litter on mass loss and nutritional release from litter.

Methods

Litter decomposition was measured in three adjacent stands of pure spruce (Picea abies), mixed beech-spruce and pure beech (Fagus sylvatica) on three nutrient-rich sites and three nutrient-poor sites over a three-year period using the litterbag method (single species and mixed species bags).

Results

Mass loss of beech litter was not higher than mass loss of spruce litter. Mass loss and nutrient release were not affected by litter mixing. Litter decay indicated non-additive patterns, since similar remaining masses under pure beech (47%) and mixed beech-spruce (48%) were significantly lower than under pure spruce stands (67%). Release of the main components of the organic substance (C_org, N_tot, P, S, lignin) and associated K were related to mass loss, while release of other nutrients was not related to mass loss.

Conclusions

In contradiction to the widely held assumption of slow decomposition of spruce needles, we conclude that accumulation of litter in spruce stands is not caused by recalcitrance of spruce needles to decay; rather adverse environmental conditions in spruce stands retard decomposition. Mixed beech-spruce stands appear to be as effective as pure beech stands in counteracting these adverse conditions.     

Crown allometry and growing space efficiency of Norway spruce (Picea abies [L.] Karst.) and European beech (Fagus sylvatica L.) in pure and mixed stands

In pure and mixed stands of Norway spruce ( Picea abies [L.] Karst.) and European beech ( Fagus sylvatica L.) we have analyzed crown allometry and growing space efficiency at the tree level and have scaled this from tree level to stand level production. Allometry is quantified by the ratio A between the relative growth rates of laterally and vertically oriented tree dimensions. Efficiency parameters, EOC for efficiency in space occupation, EEX for efficiency in space exploitation, and EBI for efficiency in biomass investment, were evaluated, based on quantity and quality of growing space and were measured using crown size and competition index. The evaluation reveals why pure stands of spruce are preferred by foresters, even though the natural vegetation would be dominated by beech. Spruce occupies its share of resources intensively by means of tightly packed pillar-like crowns, whereas beech seizes resources extensively by means of a multi-layered, veil-like canopy. With a given relative biomass increment, beech achieves a 57 % higher increment in crown projection area and a 127 % higher increment in height due to its particular capacity of lateral and vertical expansion. Beech trees are approximately 60 % more efficient in space occupation than spruce trees, however, on average, they are about 70 % less efficient in space exploitation. As a vertical fast growing tree, spruce is efficient in space exploitation under constant conditions, but far more susceptible to disturbances and less well equipped to overcome them when compared with beech. Beech is weaker in terms of space exploitation, while being superior in space occupation, where it encircles competitors and fills gaps after disturbances, which is a successful long-term strategy. A mixture of the two species reduces stand level production by 24 % in comparison to a pure spruce stand, however, when considering enhanced stabilization of the whole stand and risk distribution in the long term, the mixed stand may exceed the production level of pure spruce stands. EEX reflects a strong ontogenetic drift and competition effect that should be considered when scaling from tree to stand level production.     

Silver fir stand productivity is enhanced when mixed with Norway spruce: evidence based on large‐scale inventory data and a generic modelling approach

Questions: How to evaluate the mixture effect on basal area increment in two‐species forest stands? Is a mixed Norway spruce–silver fir stand more productive than pure adjacent stands of either species? How to develop generic modelling approaches to assess mixture effects in forest stands? Location: In addition to a case study on Norway spruce–silver fir stands in French mountain forests, the generic approach used goes beyond local applications. Methods: We took advantage of National Forest Inventory data to develop a unique stand basal‐area‐increment model for pure and mixed stands of Norway spruce and silver fir that responds to ecological site conditions. The database was made up of 284 pure Norway spruce stands, 196 pure silver fir stands, and 323 mixed stands of these species. Results: Pure silver fir basal area increment is strongly influenced by spring climatic conditions, whereas pure Norway spruce is more influenced by soil conditions. The mixture of these species has a positive effect on silver fir, which decreases as the proportion of fir increases. In contrast, the mixture has no noticeable effect on Norway spruce. Conclusion: We developed a stand basal‐area‐increment model evidencing an advantage of the mixture on silver fir basal area increment, but not on Norway spruce. The mathematical formulation of the model developed is generic and can be used in all two‐species mixture situations. It also makes it possible to compare different mixture situations with each other.     

Plant-soil feedback in spruce (Picea abies) and mixed spruce-beech (Fagus sylvatica) stands as indicated by dendrochemistry

Ten pairs of secondary pure spruce (Picea abies) and adjacent mixed spruce-beech (Fagus sylvatica) stands on comparable sites were selected on two different bedrocks for soil formation (Flysch: nutrient rich and high soil pH; Molasse: poor nutrient supply and acidic) to study how an admixture of beech to spruce stands affects nutrient cycling and consequently soil chemistry. Soil analyses indicated accumulation of Ca under the mixed stands while the top soil under pure spruce was acidified. It was hypothesized that changes of soil chemical properties due to species composition over the last six decades are reflected in the stem wood of spruce. Three healthy dominant spruce trees per plot were selected for coring. Cores were crossdated and half-decadal samples were analyzed for Ca, Mg, Mn and Al. Calcium and Mg concentrations in stem wood of spruce were significantly higher for the pure spruce than for the mixed stands in spite of lower Ca and Mg stores in the soil. We assume that acidification caused by pure spruce mobilized these cations temporarily, increasing soil solution contents and consequently stem wood concentrations. It was possible to reconstruct soil pH from the element ratios Ca/Al (pure stands) and Ca/Mg (mixed stands), since these ratios in the stem wood of the last half-decade did correlate with soil pH for selected soil depths. Reconstructed soil pH showed a decline over the last 60 years under both species compositions due to accumulation of base cations in the increasing biomass. Comparisons of reconstructed soil pH in 0–5 and 10–20 cm soil depth indicated more pronounced top soil acidification (lower soil pH in 0–5 cm) by spruce on the nutrient rich soil (Flysch) than on the acidic soil (Molasse). However, admixture of beech caused higher pH values in 0–5 cm than in 10–20 cm soil depth on Flysch due to the observed Ca-pump effect of beech (uptake of Ca from deeper soil horizons).     

Temporal variation of competition and facilitation in mixed species forests in Central Europe

Facilitation, reduced competition or increased competition can arise in mixed stands and become essential to the performance of these stands when compared to pure stands. Facilitation and over‐yielding are widely held to prevail on poor sites, whereas neutral interactions or competition, leading to under‐yielding of mixed versus pure stands, can occur on fertile sites. While previous studies have focused on the spatial variation of mixing effects, we examine the temporal variation of facilitation and competition and its effect on growth. The study is based on tree ring measurement on cores from increment borings from 559 trees of Norway spruce (Picea abies [L.] Karst.), European beech (Fagus sylvatica [L.]) and sessile oak (Quercus petraea (Matt.) Liebl.) in southern Germany, half of which were in pure stands and half in adjacent mixed stands. Mean basal area growth indices were calculated from tree ring measurements for pure and mixed stands for every species and site. The temporal variation, with positive correlations between species‐specific growth indices during periods of low growth and neutral or negative correlations during periods of high growth, is more distinct in mixed than in neighbouring pure stands. We provide evidence that years with low growth trigger over‐yielding of trees in mixed as opposed to pure stands, while years with high growth lead to under‐yielding. We discuss the relevance of the results in terms of advancing our understanding and modelling of mixed stands, extension of the stress gradient hypothesis, and the performance of mixed versus pure stands in the face of climate change.     

Survival of Norway spruce remains higher in mixed stands under a dryer and warmer climate

Shifts in tree species distributions caused by climatic change are expected to cause severe losses in the economic value of European forestland. However, this projection disregards potential adaptation options such as tree species conversion, shorter production periods, or establishment of mixed species forests. The effect of tree species mixture has, as yet, not been quantitatively investigated for its potential to mitigate future increases in production risks. For the first time, we use survival time analysis to assess the effects of climate, species mixture and soil condition on survival probabilities for Norway spruce and European beech. Accelerated Failure Time (AFT) models based on an extensive dataset of almost 65 000 trees from the European Forest Damage Survey (FDS) – part of the European‐wide Level I monitoring network – predicted a 24% decrease in survival probability for Norway spruce in pure stands at age 120 when unfavorable changes in climate conditions were assumed. Increasing species admixture greatly reduced the negative effects of unfavorable climate conditions, resulting in a decline in survival probabilities of only 7%. We conclude that future studies of forest management under climate change as well as forest policy measures need to take this, as yet unconsidered, strongly advantageous effect of tree species mixture into account.     

Variation in fine root biomass of three European tree species: Beech (Fagus sylvatica L.), Norway spruce (Picea abies L. Karst.), and Scots pine (Pinus sylvestris L.)

Abstract

Fine roots (<2 mm) are very dynamic and play a key role in forest ecosystem carbon and nutrient cycling and accumulation. We reviewed root biomass data of three main European tree species European beech, (Fagus sylvatica L.), Norway spruce (Picea abies L. Karst.) and Scots pine (Pinus sylvestris L.), in order to identify the differences between species, and within and between vegetation zones, and to show the relationships between root biomass and the climatic, site and stand factors. The collected literature consisted of data from 36 beech, 71 spruce and 43 pine stands. The mean fine root biomass of beech was 389 g m^?2, and that of spruce and pine 297 g m^?2 and 277 g m^?2, respectively. Data from pine stands supported the hypothesis that root biomass is higher in the temperate than in the boreal zone. The results indicated that the root biomass of deciduous trees is higher than that of conifers. The correlations between root biomass and site fertility characteristics seemed to be species specific. There was no correlation between soil acidity and root biomass. Beech fine root biomass decreased with stand age whereas pine root biomass increased with stand age. Fine root biomass at tree level correlated better than stand level root biomass with stand characteristics. The results showed that there exists a strong relationship between the fine root biomass and the above-ground biomass.     

Site‐adapted admixed tree species reduce drought susceptibility of mature European beech

Some forest‐related studies on possible effects of climate change conclude that growth potential of European beech (Fagus sylvatica L.) might be impaired by the predicted increase in future serious drought events during the growing season. Other recent research suggests that not only multiyear increment rates but also growth resistance and recovery of beech during, respectively, after dry years may differ between pure and mixed stands. Thus, we combined dendrochronological investigations and wood stable isotope measurements to further investigate the impact of neighborhood diversity on long‐term performance, short‐term drought response and soil water availability of European beech in three major geographic regions of Germany. During the last four decades, target trees whose competitive neighborhood consisted of co‐occurring species exhibited a superior growth performance compared to beeches in pure stands of the same investigation area. This general pattern was also found in exceptional dry years. Although the summer droughts of 1976 and 2003 predominantly caused stronger relative growth declines if target trees were exposed to interspecific competition, with few exceptions they still formed wider annual rings than beeches growing in close‐by monocultures. Within the same study region, recovery of standardized beech target tree radial growth was consistently slower in monospecific stands than in the neighborhood of other competitor species. These findings suggest an improved water availability of beech in mixtures what is in line with the results of the stable isotope analysis. Apparently, the magnitude of competitive complementarity determines the growth response of target beech trees in mixtures. Our investigation strongly suggest that the sensitivity of European beech to environmental constrains depends on neighborhood identity. Therefore, the systematic formation of mixed stands tends to be an appropriate silvicultural measure to mitigate the effects of global warming and droughts on growth patterns of Fagus sylvatica.     

Deposition and soil leaching in stands of Norway spruce and European Beech: Results from the Höglwald research in comparison with other European case studies

Stands of Norway spruce (Picea abies K.) and European beech (Fagus sylvatica L.) were investigated at the Höglwald research area, Southern Germany from 1985–1988 and from 1994–1997 in order to determine the effects of tree species on deposition and soil solution fluxes. The results were compared to 15 European case studies representing different deposition levels and site conditions. At the Höglwald site, which is characterised by a high nitrogen and a moderate sulphur load, throughfall deposition of nitrogen and sulphur compounds was about two-fold higher in spruce stands compared to beech stands. The differences in elemental input were clearly reflected in soil solution chemistry with a higher leaching of nitrate and sulphate in the spruce stands. The turnover of sulphur and nitrogen compounds induced a strong soil internal production of protons especially in the spruce stands. These results are in accordance with the other European case studies. Throughfall deposition and soil leaching of nitrogen and sulphur compounds was generally higher for spruce stands compared to beech stands. The species-related differences were mainly caused by dry deposition and were relatively small in remote areas. The consequences for the forest ecosystem itself and for the hydrosphere are discussed.     

Growth of adult Norway spruce (Picea abies [L.] Karst.) and European beech (Fagus sylvatica L.) under free-air ozone fumigation

This study attempted to detect the impact of ozone on adult trees of Norway spruce ( Picea abies [L.] Karst.) and European beech ( Fagus sylvatica L.) in an experimental mixed stand in Southern Bavaria, Germany. The aim was to examine whether there is a decrease in growth when trees are exposed to higher than atmospheric concentrations of ozone. This exposure was put into effect using a free-air fumigation system at tree crown level. Growth analysis was carried out on a group of 47 spruce and 36 beech trees, where radial stem increment at breast height - a sensitive index for stress - was measured. The ozone monitoring system allowed values to be obtained for the accumulated ozone exposure (SUM00) of each individual tree, so that their radial increment over three years could be correlated with the corresponding ozone exposure for the same time period. Correlation and regression analysis were then carried out to test the influence of ozone on diameter increment. In both spruce and beech, the initial stem diameter was the most influential factor on radial increment in the following year. A linear model was applied, including the diameter of the preceding year and the ozone exposure of the current year as predicting factors. For spruce trees, a significant negative influence of ozone exposure was found. In contrast, no significant ozone effect on diameter increment of beech was detected. The effect of ozone stress on a large spruce tree can lead to a decrease in potential radial increment of 22 %. The results are discussed in relation to other stress factors such as drought and lack of light.     

The dependency of the size-growth relationship of Norway spruce (Picea abies [L.] Karst.) and European beech (Fagus sylvatica [L.]) in forest stands on long-term site conditions, drought events, and ozone stress

Against a backdrop of increasing climate change, the effects of site conditions, drought events and ozone stress on the size-growth relationship in Norway spruce (Picea abies [L.] Karst.) and European beech (Fagus sylvatica [L.]) stands are analyzed. The size-growth relationship is represented by a straight line defined by intercept and slope of a simple linear equation with stem diameter at height 1.30 m as independent variable and annual stem diameter increment at height 1.30 as dependent variable. On the basis of 64 long-term experimental plots dating back to 1871 and representing an ecological gradient from fertile to poor sites, it is shown that poorer sites exhibit shallower slopes of the linear size-growth relationships than fertile sites. Annual measurements of the size-growth relationship, including the extremely dry years of 1976 and 2003, also showed that lower stand growth rates result in shallower size-growth relationship slopes. By comparing stands with and without experimental twice-ambient ozone exposure between 2000 and 2007, it was found that ozone stress can significantly reduce the slope of the size-growth relationship. This indicates that limiting site condition, whether acute or chronic in nature, distinctly reduces the superiority of tall trees, and that a lower degree of resource limitation increases the steepness of the size-growth relationship. The causes for this behavior and the consequences for stand dynamics, silvicultural treatment and prognostication by models are discussed.     

Effects of airborne sea salts on soil water acidification and leaching of aluminium in different forest ecosystems in Denmark

Studies on biogeochemical cycling of elements are performed in even aged forest ecosystems of Norway spruce, Sitka spruce and beech at several locations in Denmark. Episodes with high sea salt deposition and unusual low water surplus percolation caused extreme salt concentrations in the soil solutions. The changes were smallest beneath beech stands and most pronounced at the most salt-affected Sitka spruce stands. Contemporary, the pH drops and the Al³⁺ concentration increased above 20 mg L^-1 owing to cation exchange with Na⁺. The Cl^- and Al³⁺ concentrations reached levels reported as toxic for Norway spruce, but not for Sitka spruce and beech. The changes in the soil water chemistry must be considered as important factors for the reduced vitality in Norway spruce ecosystems in Denmark.     

Vapor–pressure deficit and extreme climatic variables limit tree growth

Assessing the effect of global warming on forest growth requires a better understanding of species‐specific responses to climate change conditions. Norway spruce and European beech are among the dominant tree species in Europe and are largely used by the timber industry. Their sensitivity to changes in climate and extreme climatic events, however, endangers their future sustainability. Identifying the key climatic factors limiting their growth and survival is therefore crucial for assessing the responses of these two species to ongoing climate change. We studied the vulnerability of beech and spruce to warmer and drier conditions by transplanting saplings from the top to the bottom of an elevational gradient in the Jura Mountains in Switzerland. We (1) demonstrated that a longer growing season due to warming could not fully account for the positive growth responses, and the positive effect on sapling productivity was species‐dependent, (2) demonstrated that the contrasting growth responses of beech and spruce were mainly due to different sensitivities to elevated vapor–pressure deficits (VPD), (3) determined the species‐specific limits to VPD above which growth rate began to decline, and (4) demonstrated that models incorporating extreme climatic events could account for the response of growth to warming better than models using only average values. These results support that the sustainability of forest trees in the coming decades will depend on how extreme climatic events will change, irrespective of the overall warming trend.