Smíšené lesy jako strategie adaptace na klimatickou změnu: budoucnost hospodaření v Krušných horách

The temperate, humid climate and nutrient-rich soils in the pre-alpine areas of southern Bavaria represent conditions where European beech and Norway spruce come out with rather equal above ground biomass production when cultivated in pure stands. In order to reveal the effect of mixture we established 37 experimental plots in even-aged pure and mixed stands of Norway spruce and European beech covering an age span of 37–155 years. The site conditions ranged from warm, dry and base-rich to cool, wet and acidic sites. The ratio of above ground biomass growth of Norway spruce in relation to European beech decreases from 1.14:1 in the monocultures to 1.04:1 in the mixed stands. The mixing of spruce and beech results in a mutual stimulation of biomass production and acceleration of size growth. Together both species produce up to 59% more above ground biomass than the neighboring pure stands. On average the overyielding amounts to 21% in the case of Norway spruce and 37% in the case of European beech. A total of 67% out of the plots indicate overyielding and 57% transgressive overyielding. In mixed stands both species’ tree sizes are significantly ahead of the corresponding pure stands. Facilitation of spruce and competitive reduction of beech yields mutualism with respect to growth on tree and stand level. Consequences for analyzing and modeling interspecific competition and for silvicultural prescriptions are discussed. Ecological implications of the mixing effect on the occurrence and stability of natural and man-made mixed stands of spruce and beech are considered.

The aim of this study was to compare increment in mixed stands to the respective pure stands adopting, as an example, mixed stands comprising Norway spruce (Picea abies [L.] Karst.) and European beech (Fagus sylvatica L.). Using data from two mixed stand age series, in the Solling and in Harz Mountains, first we investigated whether the growth reduction effect arising from competing trees is dependent on the species of the competitors. We than explored whether increment predictions from the forest growth simulation package TreeGrOSS revealed differences between pure and mixed stands and, if so, what factors play a role and what may cause these increment differences. The analysis was based on a simulated trial series in which the parameters such as percentage mixture, age, site quality class, and stocking level were varied. The results indicate that neighboring Norway spruce restrict the basal area increment of both European beech and Norway spruce more than European beech competitors with the same crown cross-sectional area. In the majority of cases, the simulation produced an increment gain in mixed stands compared with pure stands. The amount of increment gain in a mixed stand depended on the percentage mixture, age, site quality class, and stocking level and could be related to changes in stand structure and their effects on aboveground competition.

Aim of the study: This paper presents the most appropriate ways to estimate the species proportions by area in mixed stands of Norway spruce (Picea abies L. Karst.) and European beech (Fagus sylvatica L.) by comparing stand level and individual tree level approaches. It also investigates whether different ways of describing species proportions by area can result in different judgments on the over- or under-yielding of species in mixtures.Area of the study: Three triplets of pure and mixed stands of Norway spruce and European beech in three locations in the northeast of Austria are investigated. The three locations differ considerably in slope, bedrock and soil type as well as in site index.Material and Methods: In all 9 plots the coordinates of all trees, their dbh, height, height to the crown base and five year increment were measured. The potentially available areas of individual trees are calculated by Voronoi- diagrams and potential densities are estimated from the comparable pure stands, yield tables, and published equations for maximum basal area and Reineke’s maximum density line.Main results: The species proportions estimated by the individual tree approach with leaf area as growth characteristic gave the best fit with the stand approach with the most appropriate, regional maximum basal area equations. By using various definitions of species proportions, in the worst case the mixing effects on individual species can be seriously over- or underestimated while the mixing effects on the total increment is only negligibly affected.Research highlightsMeasures of species proportions by area are needed for comparing growth per hectare of a species in a mixed stand with that of the same species in a pure standSpecies proportions at the stand level are based on estimates of the species’ potential densities, either in terms of maximum basal area or of maximum stand density indexSpecies proportions at the tree level are derived from the area potentially available (APA) to the individual trees, based on the coordinates of trees in the stands, and on their growth characteristics, such as crown projection area or leaf areaFor the examples of Norway spruce - European beech stands, the species proportions derived according to the individual tree approach using leaf area as growth characteristics fits best with the stand approach using the most appropriate maximum basal area equationsKeywords: Picea abies; Fagus sylvatica; mixture proportion; growth efficiency; mixing effect.Abbreviations used: APA – area potentially available.

Soil water depletion and recharge patterns in mixed and pure forest stands of European beech and Norway spruce

Root distribution of Norway spruce in monospecific and mixed stands on different soils

Analysis of O F-layer humus mass variation in a mixed stand of European beech and Norway spruce: An application of structural equation modelling

Stands of Norway spruce (Picea abies K.) and European beech (Fagus sylvatica L.) were investigated at the Hoglwald 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 Hoglwald 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.

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 (R2) 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.

BackgroundClimate change triggered many studies showing that trends and events of environmental conditions can reduce but also accelerate growth at the stand and individual tree level. However, it is still rather unknown how climate change modifies the growth partitioning between the trees in forest stands.MethodsBased on long-term girth-tape measurements in mature monospecific and mixed-species stands of Norway spruce (Picea abies (L.) Karst.) and European beech (Fagus sylvatica L.) we traced the effect of the severe droughts in 2003 and 2015 from the stand down to the tree level.ResultsStand growth of Norway spruce decreased by about 30% in the once-in-a-century drought 2015, while European beech was much more drought resistant. Water availability generally amplified size-asymmetric growth partitioning. Especially in case of Norway spruce water availability primarily fostered the growth of predominant trees, whereas drought favoured the growth of small trees at the expense of the predominant ones. We could not detect significant differences between mixed and monospecific stands in this regard.ConclusionsThe drought-induced reallocation of growth in favour of small trees in case of spruce may result from its isohydric character. We hypothesize that as small trees are shaded, they can benefit from the reduced water consumption of their sun-exposed taller neighbours. In case of beech, as an anisohydric species, tall trees suffer less and smaller trees benefit less under drought. The discussion elaborates the consequences of the water dependent growth allocation for forest monitoring, growth modelling, and silviculture.

• Existing growth and yield plots of pure and mixed stands of Norway spruce (Picea abies (L.) H. Karst.) and European beech (Fagus sylvatica L.) were aggregated in order to unify the somewhat scattered sources of information currently available, as well as to develop a sound working hypothesis about mixing effects. The database contains information from 23 long-term plots, covering an ecological gradient from nutrient poor and dry to nutrient rich and moist sites throughout Central Europe. • An empirically formed interaction model showed, that depending on the site conditions, dry mass growth in mixed stands can range from −46% to +138 % of the growth yielded by a scaled combination of pure stands at equal mixing proportions. • Drawing from the interaction model, overyielding of the mixed stands appears to be triggered by two separate mechanisms. On poor sites, where significant overyielding is commonly found, facilitation by beech offsets nutrient-related growth limitations in spruce. In contrast, overyielding of mixed stands occurs less frequently on rich sites, and appears to be based on an admixture effect, with spruce reducing the severe intra-specific competition common in pure beech stands. • It was concluded that silviculture can accelerate growth of spruce by beech admixtures on poor sites, while growth of beech can be promoted by admixture of spruce, particularly on excellent sites.

We examined how soil organisms and C, N and P mineralisation are affected by admixing deciduous tree species, silver birch (Betula pendula) and woollen birch (B. pubescens), in managed Norway spruce (Picea abies) stands. Pure spruce and mixed spruce–birch stands were examined at four sites in southern and central Sweden. Soil macroarthropods and enchytraeids were sampled in litter and soil. In the uppermost 5 cm of soil humus we determined microbial biomass and microbial respiration; we estimated the rate of C, N and P mineralisation under laboratory conditions. The densities of Coleoptera, Diptera and Collembola were larger in mixed stands than in spruce stands. Soil fauna composition differed between mixed and spruce stands (as revealed by redundancy analysis). Staphyliniidae, Elateridae, Cecidiomyidae larvae and Onychiuridae were the families that increased most strongly in mixed stands. There were no differences in microbial biomass and microbial respiration, nor in the C, N and P mineralisation rates, between mixed and spruce stands. However, within mixed stands microbial biomass, microbial activity and C mineralisation were approximately 15% higher under birch trees than under spruce trees. We propose that the presence of birch leaf litter was likely to be the most important factor causing differences in soil fauna composition. Birch may also influence the quality and the decomposition rate of humus in mixed stands. However, when the proportion of birch trees is low, the short-term (decades) effect of this species on decomposition is likely to be small in mixed stands on acid forest soils.

Effects of Douglas fir cultivation in German forests on soil seepage water quantity and quality

Key messageWe applied a modified forest gap model (ForClim) to depict changes in stand water transpiration via density reduction as a forest adaptation strategy. This approach is the key to analyzing the ecological resilience to drought, stress-induced mortality, and economic efficiency of managed mixed forest stands in Central Europe. The results show that specific geographic conditions and forest composition define the optimal stand density of drought-resilient forests.ContextReducing stand density has been recognized as a valid strategy to increase forest resilience to drought. Moreover, to develop adaptive management strategies (AMS) under climate change, it is crucial to consider not only drought resilience but also the economic efficiency of alternative AMS proposed to alleviate drought effects.AimsTo analyze how decreased inter-tree competition among overstorey trees affects stand vulnerability to drought and its expected yield.MethodsWe integrated experimental thinning data and historical responses to drought years in a climate-sensitive forest gap model, ForClim. We tested a business as usual (BAU) and three alternative AMS (“do-nothing,” low- and high-intensity overstorey removal) in mixed stands of Norway spruce (Picea abies), silver fir (Abies alba), and European beech (Fagus sylvatica) along an elevational gradient of 520–1020 m a.s.l. in Central Europe.ResultsHigh-intensity overstorey removal in mixed stands of all three species considerably increased forest volume growth resilience to drought and decreased stress-induced mortality by two-thirds vis à vis a “do-nothing” strategy. In sites including only conifer species, forest resilience was equally improved by high- and low-intensity overstorey removal compared to that in the BAU strategy. Regarding the timber economy, high-intensity overstorey removal resulted in a higher economic revenue of mixed stands (~ 22% higher net present value than other strategies) on the high-elevation sites (> 1000 m a.s.l.).ConclusionModifying forest density and structure by overstorey removal is principally suitable to increase forest resilience to drought and improve its economic efficiency. The magnitude of the effect however depends on the geographical setting and forest composition.

The Collembola community of a Central European forest: Influence of tree species composition

Understanding the interactions within the soil-plant-atmosphere-continuum becomes more important considering the eco- and hydrological impacts of climate change. Especially stand specific flow pathways and the characteristic timescales of water movement potentially provide important information on drought resilience of different forest ecosystems. This study analysed tree stand specific water uptake dynamics through water stable isotopy at high temporal resolution for two isotopic labelling events (7mm with &amp;#948;2H +1000 &amp;#8240; and 23mm with &amp;#948;2H +800&amp;#8240;) during the 2022 drought in south-west Germany. Measurements in pure and mixed tree stands of European beech (Fagus sylvatica, n=18) and Norway spruce (Picea abies, n=18) included sap flow, in-situ water isotopy of soil and xylem water, radial stem growth and microclimatic conditions. Our central hypothesis is that species identity and water competition between tree species are major drivers for ecohydrological flux dynamics. The results of the two labelling events showed differences in label water uptake of the two tree species and the transit times of the label in the system. The labelling events showed different transit times in the tree xylem depending on the label intensity. P. abies showed a slightly higher uptake of shallow soil water label in mixed stands than in pure patches. When labelled water infiltrated into deeper soil layers water was taken up faster by F. sylvatica in mixed forest patches than in pure forest patches and showed a generally slower uptake in P. abies than in F. sylvatica. The faster response time in water uptake during the second labelling was supported through an increase in measured sap flux and modelled branch water potential. Those dynamics of water isotopy measured in a high temporal resolution allow for a better understanding of root water uptake dynamics and water use strategies but also show species interaction effects on ecohydrological fluxes.