Soil carbon dioxide efflux in pure and mixed stands of oak and beech

Total Soil Respiration (TSR) was measured in pure and mixed stands of oak and beech and was partitioned into two contributions using the forest floor removal technique: Mineral Soil Respiration (MSR) and Forest Floor Respiration (FFR). In addition, laboratory incubations of the forest floor and the Ah horizon were performed to evaluate the heterotrophic respiration and the DOC production of these horizons. The relationships between soil temperature and the various soil respiration contributions in the three stands were compared using Q ₁₀ functions. In situ, significant differences (α = 0,05) between stands were observed for the R ₁₀ parameter (respiration rate at 10 °C) of the TSR, MSR and FFR contributions, while only the temperature sensitivity (Q ₁₀) of TSR was significantly affected by stand composition. The effect of soil water content was only significant on MSR and followed different patterns according to stand composition. Under controlled conditions, the R ₁₀ of the forest floor and of the Ah horizon varied with stand composition and the Q ₁₀ of the forest floor decreased in the order: oak (2.27) > mixture (2.01) > beech (1.71).     

Tree effects on the chemical topsoil features of oak, beech and pine forests

We aimed to study tree effects on the chemical properties of forest soils. We compared soil features of three types of forest ecosystems, each with four stands (replicates): beech forests (Fagus sylvatica), oak forests (dominated by Quercus pyrenaica) and pine plantations (Pinus sylvestris). Five samples from the top 10 cm of soil were taken per stand, from which pH, organic matter content (O.M.), total nitrogen (N) and available calcium (Ca²⁺), magnesium (Mg²⁺), potassium (K⁺) and sodium (Na⁺) were determined. Litter layer depth was measured at each soil sampling point. We also measured tree density and crown diameters at each stand. Our results indicated that soil samples from the four pine plantation stands were more similar while oak and beech stands were characterised by great variability in terms of soil properties and leaf litter depth. Although the identity of the dominant tree species significantly influenced several topsoil chemical properties (increase in pH and available cations in oak forests and higher organic matter and total nitrogen in beech and pine ecosystems), there were other important factors affecting soil features that may be taken under consideration. Differences between soil properties of the three types of forest ecosystems were mainly related to the characteristics of the litter layer and less related to the tree layer structure. Finally, the establishment of pine plantations in naturally deciduous tree areas made the topsoil features more homogeneous.     

Canopy composition as a measure to identify patterns of nutrient input in a mixed European beech and Norway spruce forest in central Europe

The influence of canopy composition on litterfall and throughfall was investigated in a mixed spruce beech forest in central Germany. We hypothesised that different parts of the mixed canopy created distinct patterns of element inputs via litterfall and throughfall. The investigation was carried out in two plots, representing the most contrasting cases of mixed forests: a stand greatly dominated by spruce (SDP) and a stand greatly dominated by beech (BDP). The canopies of the two plots were classified in four categories: pure beech, pure spruce, mixed canopy and gap. Amounts of throughfall water were lower and major element fluxes were higher under spruce than under beech in both plots, indicating that the nutrient inputs under the canopies of individual trees are driven by species-specific properties of the canopies and are quite independent of the degree of admixture. With the exception of K⁺, mixed canopies showed intermediate element inputs via throughfall, compared with pure canopy classes. The K⁺ input was significantly greater under mixed canopies, and these differences were more pronounced in the SDP than in the BDP. Results suggest that individual spruce trees in the BDP induce greater spatial heterogeneity of throughfall input than individual beech trees in the SDP. Nutrient inputs via foliar litterfall were similar among the different canopy classes, but the Mg input was lower under spruce canopy. This effect was balanced by higher Mg input via spruce throughfall. In our study, throughfall was the main source of heterogeneity in nutrient inputs, while foliar litterfall had a homogenising effect.     

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.     

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.     

Stand characteristics of mixed oriental beech (<Emphasis Type="Italic">Fagus orientalis</Emphasis> Lipsky) stands in the stem exclusion phase,northern Iran

The structure of forest stands changes through developmental phases. This study is carried out in the unmanaged, oriental beech (Fagus orientalis Lipsky) stands in the north of Iran. The aim of this research was to quantify structural characteristics of stands in the stem exclusion phase using common structural indices, which include mingling, tree–tree distance, stem diameter, and tree height differentiation. According to our measurements from three stands, naturally regenerated stands tend to be mixed in species composition have slightly heterogeneous diameter distributions and uniform tree height. The average distance between trees was 3.3 m. Stocking volume of the stands had an average of 540 m³ ha^?1 and 412 stem ha^?1. Dead wood volume was 24 m³ ha^?1, and as a standing volume, the most frequent species in dead wood pool was oriental beech (F. orientalis) (48 %). The common form of dead trees was snag (41 %). The mean value of mingling and tree-to-tree interval indices revealed that beech was mixed intensively with hornbeam and appears to be a more successful competitor for space and light compared with hornbeam; moreover, we found relatively high evidence of inter-species competition in this phase. A better understanding of stand characteristics in the stem exclusion phase as a critical part of the natural dynamics of forest ecosystems could facilitate predictions about the future changes within the stand.     

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.     

Biogeochemical cycling and chemical fluxes in a managed northern forested wetland, Michigan, USA

Forest harvesting and subsequent regeneration treatments may cause changes in soil and solution chemistry that adversely affect forest productivity and environmental quality. The objective of this study was to assess soil carbon (C), nitrogen (N), and base cation pools and fluxes, and to construct a hydrogen ion (H⁺) mass balance to identify major processes controlling acidity production and consumption 14 years following whole-tree harvesting and regeneration in a northern forested wetland with underlying mineral soils derived from calcareous glacial drift. Results for soil elemental and nutrient pools in the harvested/regenerated stand were compared to an adjacent non-harvested stand and a riparian zone. The riparian zone had the highest soil total C, total N, and exchangeable calcium (Ca) and magnesium (Mg) pools; however, no difference in exchangeable potassium (K) was evident among stand types. Moreover, no differences between the harvested/regenerated and uncut stands were evident in any of the soil chemical pools.Net export of base cations was minimal and the H⁺ mass balance indicated that net cation exchange was not a significant process in H⁺ production or consumption in either the uncut or harvested/regenerated stands. The most striking differences in the H⁺ mass balance were (1) eight times the H⁺ consumption from sulfate (SO₄²⁻) reduction in the harvested/regenerated stand compared to that in the uncut condition and (2) nearly twice the H⁺ production due to N immobilization in the harvested/regenerated stand. However, both stand types were net H⁺ sinks and increases in H⁺ export due to whole-tree harvesting were not evident.The riparian zone was a net exporter of base cations. This finding was attributed to a combination of base cation exchange and carbonate mineral weathering; data suggested the importance of the latter. More research, however, is required to isolate the contributions of cation exchange and carbonate weathering on base cation export from the riparian zone. Stream chemistry was consistent with that of the riparian zone, indicating a strong linkage between the riparian zone stream chemistry, and whole-tree harvesting had no intermediate term (i.e., 14 years) effects on stream acidification in this managed northern wetland ecosystem.     

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.     

Development of bioavailable pools of base cations and P after afforestation of volcanic soils in Iceland

Few long-term studies have been conducted on changes in soil nutrients after afforestation in Iceland, a country with a young history of forest management. While fertilization was shown to improve survival of seedlings in the first years after planting on the nutrient limited soils, knowledge about the nutrients status of the soils that develop under maturing forest stands is still scarce. In a chronosequence study, the development of base cations and Olsen-phosphorus (Olsen-P) in the mineral soil was followed in six forest stands of two different tree species of increasing age (14–97 years): native birch (Betula pubescens) and introduced Siberian larch (Larix sibirica). A treeless heathland was included to present soil conditions prior to forest establishment. The sites are part of the largest forest area in Iceland, located in the east of the country. Results revealed an effect of stand age on all soil nutrients investigated except for potassium (K). Olsen-P increased in 0–10 cm depth of the mineral soil, indicating a better availability and thus improved P supply in maturing forest stands. Calcium (Ca) and magnesium (Mg) concentrations decreased with stand age in 0–10 and 10–20 cm soil depth, while sodium (Na) decreased only in the upper soil layer. Only Olsen-P and K concentrations were higher in the upper soil layer as compared to 10–20 cm depth. This indicates a higher biotic control as opposed to the geochemical control of the other base cations.     

The effects of species mixture on the growth and yield of mid-rotation mixed stands of Scots pine and silver birch

The effect of tree species mixture on stand volume yield and on tree-species-specific diameter and height growth rates were analysed in managed mixed stands of Scots pine (Pinus sylvestris L.) and silver birch (Betula pendula Ehrn.).Data were obtained from 14 repeatedly measured stands located in Southern Finland on mineral soil sites with varying admixture of Scots pine and silver birch. Statistical analysis was carried out for studying the effect of species mixture on the development of stand characteristics. For the analysis, the plots were categorised into three groups (plot types) according to the species dominance. In order to analyse species-specific growth rates, individual-tree mixed linear growth models for tree diameter and height growth were developed for both tree species.The results clearly show that the yield of the managed mid-rotation, mixed stands was greater for stands dominated by Scots pine than for stands dominated by birch, and the stand volume increment decreased with an increasing proportion of silver birch. Analysis of diameter and height growth by tree species revealed that the main reason for this pattern is the negative impact of birch competition on the growth of pine trees. The increase in diameter of pine was clearly hampered if the proportion of birch was high. An abundance of birch also slightly decreased the growth in height of Scots pine, although the effect was less than on diameter growth. Species mixture did not affect the diameter growth of birch but did have a significant effect on height development. Height growth of birch was considerably greater in pine-dominated stands than in birch-dominated stands. In pine-dominated mixed stands, the height growth of birch was quite close to that of dominant pine trees, and birches can endure in competition with pines for light.The results apply for even-aged and single-storey managed stands, where stocking density and structure are controlled with pre-commercial and commercial thinnings. The results are not applicable to unmanaged mixed stands undergoing self-thinning. This study provides new information on mixed stands from a silvicultural perspective, which can be applied in decisions involving the management of mixed stands.     

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.     

Habitat factors for land snails in European beech forests with a special focus on coarse woody debris

Analyses of land snails and habitat factors in acid beech forests were conducted in southern Germany (northern Bavaria). The objectives were to study the effects of habitat characteristics on snail density and species richness. Habitat structures were determined for 37 plots in one big forest. We found a significant relationship between the number of snail species and individuals and the following set of habitat factors coverage of herbaceous layer, growing stock, mean diameter at breast height of the three largest trees (DBHmax), stand age, total dead wood volume per ha, and advanced decomposed dead wood volume per ha. We use maximally selected rank statistics to estimate cutpoints separating stands with low densities, from stands with high snail densities. Here, we define cutpoints for a significant higher snail density at a stand age of 187 years, 57 m³/ha dead wood, 40 m³/ha advanced decomposed dead wood, 63 cm DBHmax and more than 1% herbaceous layer. For species richness, cutpoints are estimated at 338 m³/ha stand volume, 170 years stand age, 50 m³/ha total dead wood amount, 15 m³/ha advanced decomposed dead wood and 56 cm DBHmax. The microhabitat analysis shows a higher pH value and a higher Calcium content at the bottom of large snags and under large lying dead wood pieces in comparison to litter, upper mineral soil and at the bottom of vital living trees. Snail species and individual density are significantly linked to these patterns of chemical parameters. The identified cutpoints are a good base for ecological management decisions in forest management.     

Influence of stand,site and meteorological variables on the maximum leaf area index of beech,oak and Scots pine

Different multiple linear regression models of maximum leaf area index (LAI_max) based on stand characteristics, site quality, meteorological variables and their combinations were constructed and cross-validated for three economically important tree species in Flanders, Belgium: European beech (Fagus sylvatica L.), Pedunculate oak (Quercus robur L.) and Scots pine (Pinus sylvestris L.). The models were successfully tested on similar datasets of experimental sites across Europe. For each species, ten homogeneous and mature stands were selected, covering the species’ entire stand productivity range based on an a priori site index classification. LAI_max was derived from measurements of leaf area index (LAI) made by means of hemispherical digital photography over the whole growing season (mid-April till end October 2008). Species-specific models of LAI_max for beech and oak were mostly driven by management practice affecting stand characteristics and tree growth. Tree density and dominant height were main predictors for beech, while stand age and tree-ring growth were important in the oak models. Scots pine models were more affected by site quality and meteorological variables. The beech meteorological model showed very good agreement with LAI at several European sites. Scots pine’s stand model predicted well LAI across Europe. Since the species-specific models did not share common predictors, generic models of LAI_max were developed for the 30 studied sites. Dominant height was found to be the best predictor in those generic models. As expected, they showed a lower predictive performance than species-specific ones.     

Evaluation of mature conifer plantations as secondary habitat for epigeic forest arthropods (Coleoptera: Carabidae; Araneae)

Spiders (Araneae) and ground beetles (Coleoptera, Carabidae) were studied in a woodland of the Northwest German lowland. An ancient oak–beech stand (170 years old) growing on mineral soil as well as a 110-year-old Scots pine (Pinus sylvestris) and a 55-year-old spruce (Picea spp.) forest growing on peat were investigated by pitfall trapping. A total of 155 species (39 carabids, 116 spiders) and 16,887 individuals (5269 carabids, 11,618 spiders) was recorded. Beetle diversity was high in the oak–beech stand and the spruce forest, but significantly lower in the pine forest. In both conifer plantations the activity density of carabids was considerably lower. Spider diversity was significantly higher in the spruce forest when compared to the beech and pine stand, respectively. Analyses of assemblage similarity distinguished clearly the fauna of all three stands. For each forest type, indicator species were detected. Although both conifer plantations were planted on former bogs, spider species typical of bogs were present only in the pine stand, not in the spruce stand. In both animal taxa, species typical of deciduous forests were more numerous and abundant in the oak–beech stand when compared to the conifer plantations. Although they were in direct contact, the conifer stands on peat only to a very low extent serve as secondary habitats for the epigeic fauna of the autochthonous deciduous woodland. During the 1990s, various agricultural programmes in Central Europe promoted such conifer plantations – in contrast, such afforestation measures on extensively used or fallow land of former bogs are not supported by the results of this study.     

Spatiotemporal dynamic of European beech (Fagus sylvatica L.) in Slovenia, 1970–2005

Based on data acquired from the spatial information system Silva-SI, the majority of the entire forest area in Slovenia (22,220 forest compartments with a total area of 7446 km², classified into eight forest categories) was analysed for changes in the distribution of European beech (Fagus sylvatica L.) in the period 1975–2005 using a binary logistic regression model in terms of selected site, stand and management variables. Additionally, changes in the abundance of beech in forest stands in which beech was present at the beginning and the end of the analysed period were analysed. Beech expanded its area by more than 1200 ha per year on average, i.e. the annual expansion rate amounted to 0.24%. Among the 18 studied variables, three site, four stand and no management variables were included in the binary logistic regression model of beech expansion. Beech expansion was more pronounced at lower altitudes, on sites with steep topography, and on sites with a higher proportion of beech in potential natural vegetation. The probability of beech expansion reduced by a factor of 0.54 when the distance to the nearest compartment with beech increased by 1 km. Among other stand variables, the proportion of early successional phases and the proportion of silver fir (Abies alba Mill.) and Norway spruce (Picea abies (L.) Karst.) also influenced the expansion of beech. During the observed period the growing stock of beech almost doubled, its proportion in total growing stock increasing from 27% to 32%. Significant differences were found in changes of beech proportion in the total growing stock among different forest categories; a decrease in the beech proportion was registered in alpine coniferous forests and thermophilous deciduous forests, while in other forest categories the proportion of beech increased. The recent development of forest stands and their current structure indicate a further expansion of beech and an increase in the proportion of beech in forest stands.     

Long-term nutrient cycling patterns in Douglas-fir and red alder stands: a simulation study

Long-term patterns in nutrient cycling in regrowing Douglas-fir (Pseudosuga menziesii Mirb. Franco) and red alder (Alnus rubra Bong.) on native soils plus soils previously occupied by other species were simulated using the nutrient cycling model. Simulations of regrowing stands were also compared with observations of nutrient cycling in mature Douglas-fir and red alder. We hypothesized that (1) prolonged presence of red alder will cause a depletion in soil base cations due to increased nitrification and NO₃⁻ leaching; (2) lower base cation availability under red alder will ultimately cause biomass production to decline; (3) high N availability in red alder soils will favor regrowth of Douglas-fir; (4) higher base cation and P status of the Douglas-fir soils will favor growth of red alder both in the short- and long-term, since N is not limiting to red alder; and (5) in regrowing red alder, NO₃⁻ leaching will increase with time as a result of increased N fixation. All hypotheses were confirmed, but the effect of soil type on biomass production was minimal both for red alder and Douglas-fir. The higher soil organic matter content in the mature red alder stand most likely reflected previous occupation by old-growth Douglas-fir and also a large litter input from the understory vegetation. In general, the nutrient cycling model simulated differences in nutrient cycling patterns at least qualitatively between Douglas-fir and red alder and was helpful in identifying potential gaps in the understanding of biogeochemical cycling as well as uncertainties in the data. The nutrient cycling model did not fully elucidate differences in P cycling between Douglas-fir and red alder and overestimated weathering rates under Douglas-fir. Uncertainties in the data included: (1) temporal patterns in N fixation in the regrowing stands; (2) understory litterfall; and (3) site history and, consequently, presence of pre-existing differences in site conditions.     

Nutrient cycling and soil leaching in eighteen pure and mixed stands of beech (Fagus sylvatica) and spruce (Picea abies)

Studies on the combined effects of beech–spruce mixtures are very rare. Hence, forest nutrition (soil, foliage) and nutrient fluxes via throughfall and soil solution were measured in adjacent stands of pure spruce, mixed spruce–beech and pure beech on three nutrient rich sites (Flysch) and three nutrient poor sites (Molasse) over a 2-year period. At low deposition rates (highest throughfall fluxes: 17 kg N ha⁻¹ year⁻¹ and 5 kg S ha⁻¹ year⁻¹) there was hardly any linkage between nutrient inputs and outputs. Element outputs were rather driven by internal N (mineralization, nitrification) and S (net mineralization of organic S compounds, desorption of historically deposited S) sources. Nitrate and sulfate seepage losses of spruce–beech mixtures were higher than expected from the corresponding single-species stands due to an unfavorable combination of spruce-similar soil solution concentrations coupled with beech-similar water fluxes on Flysch, while most processes on Molasse showed linear responses. Our data show that nutrient leaching through the soil is not simply a “wash through” but is mediated by a complex set of reactions within the plant–soil system.     

Species proportions by area in mixtures of Scots pine (<Emphasis Type="Italic">Pinus sylvestris</Emphasis> L.) and European beech (<Emphasis Type="Italic">Fagus sylvatica</Emphasis> L.)

Scots pine (Pinus sylvestris L.) and European beech (Fagus sylvatica L.) dominate many of the European forest stands. Also, mixtures of European beech and Scots pine more or less occur over all European countries, but have been scarcely investigated. The area occupied by each species is of high relevance, especially for growth evaluation and comparison of different species in mixed and monospecific stands. Thus, we studied different methods to describe species proportions and their definition as proportion by area. 25 triplets consisting of mixed and monospecific stands were established across Europe ranging from Lithuania to Spain in northern to southern direction and from Bulgaria to Belgium in eastern to western direction. On stand level, the conclusive method for estimating the species proportion as a fraction of the stand area relates the observed density (tree number or basal area) to its potential. This stand-level estimation makes use of the potential from comparable neighboring monospecific stands or from maximum density lines derived from other data, e.g. forest inventories or permanent observations plots. At tree level, the fraction of the stand area occupied by a species can be derived from the proportions of their crown projection area or of their leaf area. The estimates of the potentials obtained from neighboring monospecific stands, especially in older stands, were poorer than those from the maximum density line depending on the Martonne aridity index. Therefore, the stand-level method in combination with the Martonne aridity index for potential densities can be highly recommended. The species’ proportions estimated with this method are best approximated by the proportions of the species’ leaf areas. In forest practice, the most commonly applied method is an ocular estimation of the proportions by crown projection area. Even though the proportions of pine were calculated here by measuring crown projection areas in the field, we found this method to underestimate the proportion by 25% compared to the stand-level approach.     

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.