Modelling above and below ground carbon dynamics in a mixed beech and spruce stand influenced by climate

Tree growth and carbon dynamics are important issues especially in the context of climate change. However, we essentially lack knowledge about the effects on carbon dynamics especially in mixed stands. Thus, the objective of this study was to test the effects of climatic changes on the above and below ground carbon dynamics of a mixed stand of Norway spruce (Picea abies [L.] Karst.) and European beech (Fagus sylvatica L.) by means of scenario simulations. To account for the typical tree interactions in a mixed-species stand a spatial explicit tree growth model based on eco-physiological processes was applied. Three different climate scenarios considering altered precipitation, temperature, and radiation were calculated for an unthinned and a thinned stand. The results showed significant changes of above and belowground biomass over time, especially when temperature and radiation were increased additionally to decreased precipitation. The reduction in biomass increments of Norway spruce were more attenuated above than below ground. In contrast, the results for beech were the opposite: The belowground increments were reduced more. These results suggest a shift in the species contribution to above and belowground biomass under dryer and warmer conditions. Distinct effects were also found when thinned and unthinned stands were compared. A reduced stand density changed the proportions of above and below ground carbon allocation. As a main reason for the changed growth reactions the water balance of trees was identified which lead to changed biomass allocation pattern. This article belongs to the special issue “Growth and defence of Norway spruce and European beech in pure and mixed stands”.     

Stand density and growth of Norway spruce (<Emphasis Type="Italic">Picea abies</Emphasis> (L.) Karst.) and European beech (<Emphasis Type="Italic">Fagus sylvatica</Emphasis> L.): evidence from long-term experimental plots

On the basis of nine Norway spruce (Picea abies (L.) Karst.) and ten European beech (Fagus sylvatica L.) thinning experiments in Germany, for which both residual and removed stock had been registered first during 1870, I scrutinize how moderate and heavy thinning from below (B-, C-grade) affects the production of merchantable volume compared with light thinning (A-grade). In relation to A-grade, cumulative merchantable volume (CV) of B- and C-grade amounts in average to 103–107% in juvenile and to 97–102% in mature Norway spruce stands. The corresponding findings for European beech are 101–106% and 94–102%. CV of individual stands varies between 89% and 130% for Norway spruce and 73% and 155% for European beech (CV of A-grade = 100%). These findings are substantiated by the relation between stand density (SDI) and periodic annual increment (PAI). On the B- and C-grade plots of spruce and beech, respectively, SDI was reduced down to 41–91% and 31–83% of the A-grade. When SDI is reduced in young stands, PAI follows a unimodal curve. Norway spruce’s PAI culminates in 109% if SDI is reduced to 59%; European beech’s PAI culminates in 123% when density is reduced to 50%. Whereas Norway spruce’s growth reacts most positively on thinning under poor site conditions and with increment reduction on favourable sites, European beech behaves oppositely. With stand development the culmination point of the unimodal relation moves towards maximum density, so that in older stands PAI follows the increasing pattern, which is the left portion of a unimodal curve. A model is presented which apparently unifies contradictory patterns of stand density–growth reactions by integrating relative stand density, average tree size and site fertility effects, and makes the findings operable for forest management.     

The nutrient status of Norway spruce in pure and in mixed-species stands

Atmospheric deposition of N and S appears to have caused nutrient imbalance in Norway spruce stands in southern Sweden. This calls for a change of forest management to procedures that promote nutrient balance. Studies have shown lower soil acidity in Norway spruce/deciduous mixed stands than in spruce monocultures, but the tree nutrient status in such mixtures has not been much investigated so far.

The nutrient status of Norway spruce foliage and top mineral soil chemistry in monocultures and in stands mixed with beech, birch, or oak was investigated through paired comparisons on 30 sites in southern Sweden (27 sites) and eastern Denmark (three sites). In total, 45 mixed stands and 34 pure stands were included in the study.

Spruce needles from mixed stands had higher concentrations and ratios to N of K, P, and Zn than needles from pure spruce stands. Among the mixed stands, the K status appeared to be positively correlated with the percentage of deciduous tree basal area. Soil samples from mixed stands had a higher Mg concentration, base saturation, and BC/Al ratio than soil samples from pure stands. The spruce needle nutrient status was comparable in pure stands on fertile sites and in mixed stands on poor sites. We did not detect any differences in spruce tree growth between pure and mixed stands.

This paper discusses possible reasons for a positive effect on the tree nutrient status in mixed-species stands and the possibility of using mixed-species stands as a forest management procedure to avoid nutrient imbalance.     

Productivity of mixed versus pure stands of oak (Quercus petraea (Matt.) Liebl. and Quercus robur L.) and European beech (Fagus sylvatica L.) along an ecological gradient

The mixture of beech (Fagus sylvatica L.) and oak (sessile oak, Quercus petraea (Matt.) Liebl., and pedunculate oak, Q. robur L.) is of considerable importance in Europe and will probably become even more important under climate change. Therefore, the performance of oak and beech in mixture was compared with the species’ growth in pure stands. Data from 37 long-term mixing experiments in Poland, Germany and Switzerland were pooled for analysis of mixing effects on stand productivity and possible interrelationships with mixing portions or site conditions. We found that on average, mixed stands of oak and beech exceeded biomass productivity in pure stands by 30 % or 1.7 t ha^?1 year^?1, as the growth of both species was benefitted by the mixture. However, that the interaction actually ranged from facilitation and overyielding on poor sites to underyielding on fertile sites triggered by competition. An empirically derived interaction model showed volume and dry mass growth changing in mixed stands from gains of 50 % to losses of 10 % depending on site conditions. It is concluded that the analysed mixture grows in accordance with the stress-gradient hypothesis and that our results suggest a site-specific relationship between species mixture and biomass productivity. As a consequence, an adequate species mix should result in increased productivity under steady state as well as climate change.     

Effects of root trenching of overstorey Norway spruce (Piceaabies) on growth and biomass of underplanted beech (Fagussylvatica) and Douglas fir (Pseudotsugamenziesii) saplings

In Central Europe, the conversion of pure Norway spruce stands (Picea abies [L.] Karst.) into mixed stands with beech (Fagus silvatica L.) and other species like e.g. Douglas fir (Pseudotsuga menziesii [Mirb.] Franco) is accomplished mainly by underplanting of seedlings beneath the canopy of overstorey spruce trees after partial cutting treatments what means exposure to shade and below-ground root competition by the overstorey to the seedlings. Particularly about the second factor, our knowledge is limited. Therefore, we carried out a below-ground competition exclusion experiment by root trenching and investigated the effects on soil resources, growth, and biomass partitioning of underplanted beech and Douglas fir saplings under target diameter and strip cutting treatments. The exclusion of overstorey root competition by trenching increased the soil water potential in the second year that had a fairly dry growing season and led to significantly higher foliar concentrations of most nutrients, particularly in Douglas fir, indicating an amended nutrient supply. Both improvements were accompanied by an increase in length and diameter increment of the underplanted saplings, appearing in both species only after having surpassed a species-specific threshold light value (Douglas fir 16% of above canopy radiation, beech 22%). We also found significant interactions between trenching and light for specific fine root length and further biomass and morphological parameters. Judged by the much steeper increase in height and diameter growth with increasing light after release from below-ground competition, Douglas fir saplings appeared to be more sensitive to root competition than beech saplings what conforms to older findings for beech. According to our results, a strip cutting seems to be more appropriate than a target diameter cutting treatment to replace a pure spruce stand by a mixed stand with beech and Douglas fir.     

Climate change impacts on stand structure and competitive interactions in a southern Swedish spruce–beech forest

It is believed that European beech (Fagus sylvatica L.) will increase its competitive ability at its northern range margin in Scandinavia due to climate change. In mixed old-growth forests of beech and Norway spruce (Picea abies (L.) Karst.) at Siggaboda nature reserve (southern Sweden), stand structure characteristics were sequentially recorded in the years 2004, 2005 and 2007 as well as growth in stem diameter using tree-coring analyses. Using these measurements, we studied the effects on stand dynamics of an extreme storm event (2005 “Gudrun” hurricane), drought and heat (mid-summer 2006, spring 2007) and subsequent bark beetle attacks on spruce (growing season 2007), overlaid with warming tendencies. The storm, which caused disastrous damage in many stands nearby, had comparatively little impact on the structure of the spruce–beech stand. All together, only 32 trees (19 spruces, 10 beeches, 3 other species) per hectare were thrown or broken mainly in the leeward direction (NE) or impacted by secondary damage by uprooted neighbour trees; this represents 7% of the total tree number and 11% of the growing stock. Diameter and height structure did not change significantly. However, the 2006 drought and the 2007 attack of biotic agents changed the stand structure and composition strongly due to the death of about 19% of the dominating older spruce trees that accounted for 35% of total stand volume. This resulted in a considerable increase in beech’s contribution to stem number (4% increase) and wood volume of the living stand (7% increase). A comparison of diameter growth of beech and spruce during the periods 1894–1949 and 1950–2005 showed a distinct decrease in growth superiority of spruce during the last 50 years. These results support the idea of a northward migration of European beech as a nemoral tree species in Sweden, due to a higher tolerance to the abiotic and biotic threats accompanying climate change and an increased competitive ability compared to boreal tree species Norway spruce.     

Tree species composition influences differences in water use efficiency of upland forested microwatersheds

Water use efficiency (WUE) was compared in three upland South Moravian forested microwatersheds in the light of effects of global climate change on forest ecosystems (GCC). The experimental catchments were characterized as upland headwater forested microwatersheds of similar size and morphology and silvicultural system, but each with different dominant tree species in the stands (over 50% of forest stand composition in living stock): Norway spruce, European beech and mixed forest. WUE was evaluated according to mean daily streamflow reduction, measured at the discharge points of the recipients of the individual catchments in precipitation-free periods lasting more than 5 days. During these times, streamflow dynamics are mainly influenced by evapotranspiration processes occurring in the forest stands. Four precipitation-free periods were observed, two in the middle of the growing season and two at its end. Two of these periods were long (15 days or more), and two were shorter (6 days). The results indicated that WUE of upland forested catchments can be very different, depending upon the dominant tree species and the seasonal phase. Highest WUE at the catchment scale (never decreasing below 80%) was exhibited by beech predominating site. WUE of mixed forest was high as well, never decreasing below 69%. The lowest WUE was exhibited by spruce predominating site, especially during a long precipitation-free period in the summer where it decreased down to 39%. In the context of the landscape, upland microwatersheds with pure spruce stands could cause its accelerated dry out in the summer and pose a significant threat to sustainable water and forest management of these areas. In comparison, mixed forests stands where spruce is not the dominant species or beech stands should still be a viable option even under the effects of GCC.     

Intra-annual growth response of adult Norway spruce (<Emphasis Type="Italic">Picea abies</Emphasis> [L.] KARST.) and European beech (<Emphasis Type="Italic">Fagus sylvatica</Emphasis> L.) to an experimentally enhanced,free-air ozone regime

Several findings indicate an impact of ozone on stem diameter growth leaving the question unanswered, if and how the intra-annual growth pattern is changed. In this study the hypotheses are tested, that (1) ozone will alter the absolute growth and (2) alter and shift the period of growth activity within a year. Our data originates from the free air ozone fumigation experiment ‘Kranzberger Forst’ in a mixed stand of Norway spruce and common beech near Freising/Germany. Annual and intra-annual growth reactions of a sample of five adult beech and five spruce trees, exposed to double ambient ozone were examined and compared to the same number of untreated reference trees. Diameter increments were measured with plastic diameter girth bands and high-resolution, automatically logging micro-dendrometers, mounted at breast height (1.3 m). We used the increment data from the growth periods 2000 to 2005. The high-resolution micro-dendrometer data were examined by fitting a Weibull function to the standardized annual growth profiles to obtain curve parameters for statistical tests. We estimated the parameters ‘T’ which represents the point of time, when 63% of the annual diameter increment is performed and the parameter ‘m’, the Weibull module, which was used as an indicator for the span of time needed to complete the annual growth. The statistical significance of these curve parameters, together with the absolute diameter increment, was tested by use of mixed regression models. The analysis of the growth curve parameters revealed a significantly altered intra-annual growth pattern of both species induced by ozone. Spruce under ozone showed reduced absolute annual diameter increment and a preponed growth activity compared to untreated trees. Beech’s absolute diameter increment was not affected under ozone, but its growth activity was delayed. For both species, ozone fumigation did not alter the individual length of the annual growing season. These results are discussed with respect to drought, tree ring anatomy and tree allometry. The study shows that ozone is able to change growth behaviour of trees even if increment losses are not obvious. This article belongs to the special issue "Growth and defence of Norway spruce and European beech in pure and mixed stands".     

Impacts of changing climate on the productivity of Norway spruce dominant stands with a mixture of Scots pine and birch in relation to water availability in southern and northern Finland

A process-based ecosystem model was used to assess the impacts of changing climate on net photosynthesis and total stem wood growth in relation to water availability in two unmanaged Norway spruce (Picea abies) dominant stands with a mixture of Scots pine (Pinus sylvestris) and birch (Betula sp.). The mixed stands were grown over a 100-year rotation (2000-99) in southern and northern Finland with initial species shares of 50, 25 and 25% for Norway spruce, Scots pine and birch, respectively. In addition, pure Norway spruce, Scots pine and birch stands were used as a comparison to identify whether species' response is different in mixed and pure stands. Soil type and moisture conditions (moderate drought) were expected to be the same at the beginning of the simulations irrespective of site location. Regardless of tree species, both annual net canopy photosynthesis (P(nc)) and total stem wood growth (V(s)) were, on average, lower on the southern site under the changing climate compared with the current climate (difference increasing toward the end of the rotation); the opposite was the case for the northern site. Regarding the stand water budget, evapotranspiration (E(T)) was higher under the changing climate regardless of site location. Transpiration and evaporation from the canopy affected water depletion the most. Norway spruce and birch accounted for most of the water depletion in mixed stands on both sites regardless of climatic condition. The annual soil water deficit (W(d)) was higher on the southern site under the changing climate. On the northern site, the situation was the opposite. According to our results, the growth of pure Norway spruce stands in southern Finland could be even lower than the growth of Norway spruce in mixed stands under the changing climate. The opposite was found for pure Scots pine and birch stands due to lower water depletion. This indicates that in the future the management should be properly adapted to climate change in order to sustain the productivity of mixed stands dominated by Norway spruce.     

Interspecific competition impacts on the morphology and distribution of fine roots in European beech (Fagus sylvatica L.) and Norway spruce (Picea abies (L.) Karst.)

Morphology and vertical distribution patterns of spruce and beech live fine roots (diameter ≤2 mm) were studied using a soil core method in three comparable mature stands in the Solling: (1) pure beech, (2) pure spruce and (3) mixed spruce–beech. This study was aimed at determining the effects of interspecific competition on fine root structure and spatial fine root distribution of both species. A vertical stratification of beech and spruce fine root systems was found in the mixed stand due to a shift in beech fine roots from upper to lower soil layers. Moreover, compared to pure beech, a significantly higher specific root length (SRL, P<0.05) and specific surface area (SSA, P<0.05) were found for beech admixed with spruce (pure beech/mixed beech SRL 16.1–23.4 m g⁻¹, SSA 286–367 cm² g⁻¹). Both indicate a flexible ‘foraging’ strategy of beech tending to increase soil exploitation and space sequestration efficiency in soil layers less occupied by competitors. Spruce, in contrast, followed a more conservative strategy keeping the shallow vertical rooting and the root morphology quite constant in both pure and mixed stands (pure spruce/mixed spruce SRL 9.6/7.7 m g⁻¹, P>0.10; SSA 225/212 cm² g⁻¹, P>0.10). Symmetric competition belowground between mixed beech and spruce was observed since live fine roots of both species were under-represented compared to pure stand. However, the higher space sequestration efficiency suggests a higher competitive ability of beech belowground.     

Effects of the extreme drought in 2003 on soil respiration in a mixed forest

We present a field study on the drought effects on total soil respiration (SR_t) and its components, i.e., “autotrophic” (SR_a: by roots/mycorrhizosphere) and “heterotrophic” respiration (SR_h: by microorganisms and soil fauna in bulk soil), in a mature European beech/Norway spruce forest. SR_a and SR_h were distinguished underneath groups of beech and spruce trees using the root exclusion method. Seasonal courses of SR_a and SR_h were studied from 2002 to 2004, with the summer of 2003 being extraordinarily warm and dry in Central Europe. We (1) analyzed the soil temperature (T _s) and moisture sensitivity of SR_a and SR_h underneath both tree species, and (2) examined whether drought caused differential decline of SR_a between spruce and beech. Throughout the study period, SR_a of beech accounted for 45–55% of SR_t, independent of the soil water regime; in contrast, SR_a was significantly reduced during drought in spruce, and amounted then to only 25% of SR_t. In parallel, fine-root production was decreased during 2003 by a factor of six in spruce (from 750 to 130 mg l⁻¹ a⁻¹), but remained at levels similar to those in 2002 in beech (about 470 mg l⁻¹ a⁻¹). This species-specific root response to drought was related to a stronger decline of SR_a in spruce (by about 70%) compared to beech (by about 50%). The sensitivity of SR_a and SR_h to changing T _s and available soil water was stronger in SR_a than SR_h in spruce, but not so in beech. It is concluded that SR_a determines the effect of prolonged drought on the C efflux from soil to a larger extent in spruce than beech, having potential implications for respective forest types. This article belongs to the special issue "Growth and defence of Norway spruce and European beech in pure and mixed stands."     

Changes in forest-floor chemistry caused by a birch admixture in Norway spruce stands

The aim of this study was to determine how soil chemistry and the distribution of fine roots (<1 mm) in the organic and upper mineral soil horizons were affected by an admixture of birch (Betula pendula Roth and B. pubescens Ehrh.) in Norway spruce (Picea abies (L.) Karst) stands. The surface organic horizons (LF and H) and mineral soil were characterized to a depth of 10 cm on three sites in southern and central Sweden. On these sites, replicated plots had been established that contained either ca. 30-year-old birch growing as a shelter over similar-aged spruce (mixed plots) or spruce only. The treatments had been created 8–11 years before this study was done. A fourth site, with plots containing ca. 90-year-old spruce or birch/spruce, and a fifth site, with 30-year-old spruce and a low admixture (12% by basal area) of birch, were also included in the study. Concentrations of Ca and Mg and pH in the LF layer were significantly higher in plots with a birch admixture. In the H-horizon, concentrations of K, Ca and Mg were significantly higher in mixed plots than in plots with pure spruce. Consequently, base saturation was higher in mixed plots than in pure spruce plots. A shelter of birch decreased the total amount of spruce fine roots (<1 mm), as revealed at one of the sites. Total fine root biomass (birch + spruce) in the organic and mineral soil horizons (to 10 cm) did not differ significantly between the pure spruce stands and the spruce stands with a birch shelter.     

Regeneration growth in different light environments of mixed species, multiaged, mountainous forests of Romania

Growth of regenerating trees in different light environments was studied for the mountainous, mixed-species forests in the Carpathian Mountains of Romania. The primary species in these mixtures were silver fir (Abies alba Mill.), European beech (Fagus sylvatica L.) and Norway spruce (Picea abies (L.) Karst). Seedlings/saplings of these species were selected and measured in different stands from two different geographical locations. Regenerating trees were measured for height and diameter growth during the summer of 2002. For each seedling/sapling, percentage of above canopy light (PACL) and stand basal area (BA) were used to assess available and occupied growing space respectively. Regeneration growth was compared against these two variables and regression relationships were developed. Using these models, we predicted the dynamics of regeneration as both growth and species composition. Our results showed that in low-light environments (PACL<20–35%; BA>30 m²/ha), shade tolerant fir and beech clearly outcompeted the spruce. Therefore, in dense stands, spruce could be eliminated by the shade tolerant species. For intermediate levels of cover (PACL=35–70%; BA=15–35 m²/ha) the spruce grew at comparable rates as the beech and fir. All three species showed similar growth rates in open conditions (PACL>80–90%; BA<15–20 m²/ha) with the spruce having a tendency to outgrow the others. However, in terms of establishment, such conditions favor spruce and inhibit fir and beech.     

Thirteen-year monitoring of liming and PK fertilization effects on tree vitality in Norway spruce and European beech stands

The chemical fertility of the forest soils in the Belgian Ardenne is threatened by acidifying and eutrophying deposition and by the nutrient removal due to timber harvesting. Experiments were launched to evaluate the ability of liming and fertilization to improve foliar nutrition, maintain or restore crown condition and promote tree growth. In 1995, 10 liming and fertilization trials were installed in even-aged stands of European beech (5) and Norway spruce (5) distributed throughout the Ardenne. In each stand, two treatments were tested: liming with 3,000 kg ha⁻¹ of dolomitic limestone and liming plus fertilization with 0 to 800 kg ha⁻¹ of rock phosphate and 0 to 350 kg ha⁻¹ of K₂SO₄. Between 1995 and 2006, the foliar Ca and Mg status of spruce and beech trees improved in the limed stands, which limited significantly but did not prevent the decline in crown condition triggered by the summer drought in 2003. For spruce, liming also increased the increment in basal area. The additional fertilization increased the foliar nutrition in P but had no significant effects on soil chemistry and tree vitality.     

A comparison of litterfall and element fluxes in even aged Norway spruce,sitka spruce and beech stands in Denmark

Litterfall was investigated in three even-aged Norway spruce (Picea abies), sitka spruce (Picea sitchensis) and beech (Fagus sylvatica) stands on a nutrient poor-soil in Southern Denmark. Dry weights and N, P, K, S, Mg, Ca, Na, Al, and Fe concentrations and fluxes were examined in litterfall fractions. Foliage litter amounted to 90% of total litterfall. The tree stands showed a similar mean annual litterfall. In the spruce stands, annual litterfall was correlated negatively with the current year increment and positively with the previous year increment. Annual litterfall in beech was constant during the 6 study years whereas Norway spruce and sitka spruce showed large fluctuations between years caused by drought, spruce aphid infestations and probably sea salt stress. Norway spruce responded with a long lasting elevated needle loss. Sitka spruce responded to infestations with premature needle loss during short periods. The presence of a large syrphid (Coccinellidae) population was important in regulating aphid (Elatobium abietinum) population density. The between-year variation in element concentrations of litterfall was small whereas variations during the year were large. Interspecific levels were recognized: Norway spruce>beech>sitka spruce. High concentrations in Norway spruce were ascribed to a combination of drought, sea salt stress and elevated transpiration. In sitka spruce, aphid infestations reduced the litterfall N content. Sitka spruce showed the smallest amount of base cation fluxes with litterfall. In contrast, spruce and beech exhibited even litterfall element fluxes. Litterfall studies revealed reduced vitality in the non-native spruce stands and underlined the perception of a healthy stand of native beech.     

Nutritional effects triggered by the extreme summer 2003 in the free air ozone fumigation experiment at the Kranzberger Forst

Since the year 2000 mature beech and spruce trees were treated in a field experiment with double ambient ozone concentrations. Elevated ozone had no influence on average single leaf biomass and there were also no ozone effects on leaf nutrient concentrations in climatic normal years. However, the extraordinary dry summer 2003 triggered significant differences between the fumigated and control trees. For beech in the year after the drought event the control trees surprisingly had significantly lower foliar levels of K and P than in former years, whereas the ozone exposed trees showed no significant nutritional effects. There are indications, that the trees exposed to double ambient ozone were already adapted to higher ozone values, whereas the control trees experienced extraordinary high ambient ozone concentrations in the dry and sunny summer 2003. For spruce in autumn 2003 and 2004 ozone treated trees had significantly higher foliar levels of K in current year needles than control trees, an effect which cannot be thoroughly interpreted yet on the basis of the dataset available. This article belongs to the special issue „Growth and defence of Norway spruce and European beech in pure and mixed stands“.     

The impact of atmospheric deposition and climate on forest growth in European monitoring plots: An individual tree growth model

In the climate change discussion, the possibility of carbon sequestration of forests plays an important role. Therefore, research on the effects of environmental changes on net primary productivity is interesting. In this study we investigated the influence of changing temperature, precipitation and deposition of sulphur and nitrogen compounds on forest growth. The database consisted of 654 plots of the European intensive monitoring program (Level II plots) with 5-year growth data for the period 1994–1999. Among these 654 plots only 382 plots in 18 European countries met the requirements necessary to be used in our analysis. Our analysis was done for common beech (Fagus sylvatica), oak (Quercus petraea and Q. robur), Scots pine (Pinus sylvestris) and Norway spruce (Picea abies). We developed an individual tree growth model with measured basal area increment of each individual tree as responding growth factor and tree size (diameter at breast height), tree competition (basal area of larger trees and stand density index), site factors (soil C/N ratio, temperature), and environmental factors (temperature change compared to long-term average, nitrogen and sulphur deposition) as influencing parameters. Using a mixed model approach, all models for the tree species show a high goodness of fit with Pseudo-R² between 0.33 and 0.44. Diameter at breast height and basal area of larger trees were highly influential variables in all models. Increasing temperature shows a positive effect on growth for all species except Norway spruce. Nitrogen deposition shows a positive impact on growth for all four species. This influence was significant with p < 0.05 for all species except common beech. For beech the effect was nearly significant (p = 0.077). An increase of 1 kg N ha⁻¹ yr⁻¹ corresponds to an increase in basal area increment between 1.20% and 1.49% depending on species. Considering an average total carbon uptake for European forests near 1730 kg per hectare and year, this implies an estimated sequestration of approximately 21–26 kg carbon per kg nitrogen deposition.     

Bioeconomic modeling of mixed Norway spruce—European beech stands: economic consequences of considering ecological effects

The study provides an improved bioeconomic model of mixed-species stands in order to test the economic consequences of tree species mixtures composed of spruce (Picea abies [L.] Karst.) and beech (Fagus sylvatica L.), when the impact of mixing tree species on stand resistance against hazards is considered. We used survival probabilities of spruce in mixed- and mono-species stands derived by Griess et al. (For Ecol Manag 267:284–296, 2012) to expand an existing bioeconomic model by Knoke and Seifert (Ecol Model 210:487–498, 2008). Monte-Carlo simulations showed the highest average net present value (sum of all net revenues during one rotation period discounted by 2 %) for spruce stands with a small beech admixture of 7 % of total stems. The net present value of this mixed stand type was 8 % higher than that reached with a spruce monoculture, while risk—measured as standard deviation of net present value—was 18 % lower than that of a pure spruce stand. A mixed stand with 51 % beech led to a decrease of 23 % of the average net present value when compared to a spruce monoculture. However, the stabilizing effect of this high proportion of beech trees on spruce together with general economic diversification effects (as a result from low correlation of hazard and market risks of both tree species) reduced the standard deviation of the net present value by 55 %. Generally, an intensive species mixture in groups, with interactions between species throughout the stand, led to higher net present values and lower risks when compared to mixtures of identical proportions in large homogenous blocks without species interactions. For calculating survival probability, data from Rhineland-Palatinate were used, for modeling growth, data were based on information gained in Bavaria. It can be assumed that susceptibility toward windthrow was underestimated with respect to Bavarian conditions or growth was overestimated with respect to Rhineland-Palatinate conditions. This narrows the explanatory power of the study at hand and—once again—highlights the importance of a standardization of inventory processes all over Europe. Still the strong potential of mixing species is proven by the results. The hypothesis, derived from simpler, more traditional bioeconomic models that mixed-species stands are economically inferior to mono-species stands, could therefore be rejected. Of particular relevance to practicing foresters is the result that even mild admixtures may lead to substantial positive economic consequences. With relatively small initial investments then, a considerable increase in efficiency can be expected. This finding makes the admixture of at least small proportions interesting for the privately owned forest sector.     

Sustainability in multi-aged stands: an analysis of long-term plenter systems

Long-term research plots in multi-aged stands managed with theplenter system were assessed to evaluate sustainability of theplenter system in Central Europe. Plots primarily consistedof Norway spruce (Picea abies (L.) Karst., silver fir (Abiesalba Mill.) or European beech (Fagus sylvatica L.) and weremeasured for seven to 16 measurement intervals over 60–91years. Sustainability was assessed with four types of criteria:stand density, tree species diversity, basal area increment,and stand structure. Comparable even-aged stands were also analysedto compare and evaluate the performance of the measures of sustainability.Measures of species diversity, increment and stand structuraldiversity generally experienced increasing trends over timein these even-aged stands. Basal area generally increased andtrees ha^-1 decreased in multi-aged stands following similarpatterns as in even-aged stands. These results suggest thatthe plenter system is still evolving and is not the model ofsustainability often assumed. Many of the measures used havepotential as indicators of sustainability in multi-aged stands.