Unmanaged Beech Forest Research Articles

Strict forest protection: A meaningful contribution to Climate-Smart Forestry? An evaluation of temporal trends in the carbon balance of unmanaged forests in Germany

The role of unmanaged forests is discussed controversially in the context of climate change. One of the key questions is, whether they can contribute to the mitigation of climate change as a carbon sink and storage. However, carbon dynamics of forests after the cessation of management are not well understood. We analyzed a set of 64 forest sites, covering wide gradients of time since abandonment (0–68 years) and stand age (65–261 years) in even-aged, unmanaged beech forests. Five sites that were unmanaged for >100 years complemented the main dataset. We compiled site-specific carbon balances, distinguishing six carbon-compartments: Carbon in aboveground living and dead biomass, carbon in belowground living and dead biomass, and carbon in the organic layer and the mineral topsoil (0–30 cm). We found positive effects of increasing TSA on the carbon stock in living biomass and aboveground dead biomass for up to 50 years after management ceased. The average increase of the total carbon stock over 50 years of TSA was ≈ 80 Mg C ha–1. The effect of stand age on aboveground living biomass showed a convex relation. Aboveground dead biomass increased logistically with TSA, while belowground dead biomass decreased. On average, the five sites unmanaged for >100 years held lower total carbon stocks compared to the observed biomass peak around 50 years of TSA. However, they contained considerably higher amounts of deadwood. Carbon in the mineral soil did neither change with TSA nor with stand age and was driven by pH. Carbon stocks in newly unmanaged forests increased almost linearly for approximately 50 years after cessation of management. Subsequently, a stabilization or medium-term decrease in carbon stock was observed, likely due to the initiating transition from even-aged to multi-aged structures. We conclude that, besides their value for biodiversity and ecosystem functions, the potential of naturally developing forests as a medium-term carbon sink and long-term stable carbon storage should be considered as a valuable contribution to Climate-Smart Forestry.

Development of a site form equation for predicting and mapping site quality. A case study of unmanaged beech forests in the Cantabrian range (NW Spain)

Site form (SF), expressed as the dominant height of a stand at a reference dominant diameter, is used less often than site index (SI) to estimate forest site quality. However, it has the advantage that it is age-independent and can therefore be applied in a wider set of situations in forestry practice. Like SI, elaboration of SF has traditionally required stem analysis or repeated measurements in permanent plots; however, Airborne Laser Scanning (ALS) can nowadays be used to generate site quality practical maps, thereby simplifying the method. The aims of this study were to fit a dynamic equation to stem analysis data in order estimate SF in natural beech forests in NW Spain, as well as to compare the performance of SF for site quality estimation and to analyze the possibility of using ALS data for site quality prediction in these forests. The Algebraic Difference Approach formulation of the Bertalanffy-Richards model provided the best results and defined four curves for dominant heights of 5, 10, 15 and 20 m at a reference dominant diameter of 20 cm. A significant relationship between SF and SI was observed, and we therefore believe that SF is a good approach for site quality estimation. These results can be used directly at inventory plot level for establishing site quality classes without the need to know the stand age, either in terms of SF or translated into SI. On the other hand, ALS data allowed estimation of both dominant height and dominant diameter, although better results were obtained with the former. Applying the SF dynamic model to both data sets enabled prediction of SF at 25x25 m/pixel (and SI using the SI-SF relationship). The overall accuracy of the relationship between the observed SI and that predicted from ALS metrics yielded a coefficient of determination of 0.456 without bias, heteroscedasticity or absence of normality. The results of this unbiased raster model were considered rather good, as predictions were obtained for a pixel size of 25x25 m (0.0625 ha). Prediction of mean SI value for one hectare would thus be necessary to average the values obtained in 16 pixels with an expected error compensation. Use of the raster model based on ALS metrics will enable site quality estimation for current beech stands at high spatial resolution without the need for fieldwork, providing very valuable information for forest managers and researchers.

Diversity and functional structure of soil animal communities suggest soil animal food webs to be buffered against changes in forest land use

Forest soil and litter is inhabited by a diverse community of animals, which directly and indirectly rely on dead organic matter as habitat and food resource. However, community composition may be driven by biotic or abiotic forces, and these vary with changes in habitat structure and resource supply associated with forest land use. To evaluate these changes, we compiled comprehensive data on the species composition of soil animal communities and environmental factors in forest types varying in land-use intensity in each of three regions in Germany, i.e., coniferous, young managed, old managed, and unmanaged beech forests. Coniferous forests featured high amounts of leaf litter and low microbial biomass concentrations contrasting in particular unmanaged beech forests. However, soil animal diversity and functional community composition differed little between forest types, indicating resilience against disturbance and forest land use. Structural equation modelling suggested that despite a significant influence of forest management on resource abundance and quality, the biomass of most soil fauna functional groups was not directly affected by forest management or resource abundance/quality, potentially because microorganisms hamper the propagation of nutrients to higher trophic levels. Instead, detritivore biomass depended heavily on soil pH. Macrofauna decomposers thrived at high pH, whereas mesofauna decomposers benefitted from low soil pH, but also from low biomass of macrofauna decomposers, potentially due to habitat modification by macrofauna decomposers. The strong influence of soil pH shows that decomposer communities are structured predominantly by regional abiotic factors exceeding the role of local biotic factors such as forest type.

Effects of forest structure on the interaction between avian hosts, dipteran vectors and haemosporidian parasites

BackgroundForest habitats are important biodiversity refuges for a wide variety of bird species. Parasitism may modulate host species presence and abundance, and parasite effects can change according to forest management practices. Such processes are not well studied in vector-borne avian haemosporidians. We analyzed the effects of forest management on bird-dipteran-haemosporidian interactions, using seven common bird species in managed and unmanaged beech forest habitats in northeastern Germany. We assumed that forest structural heterogeneity affects parasite population parameters in avian hosts (i.e., prevalence and parasitemia), through its effect on the condition of the avian host but also through varying vector abundances.ResultsParasite prevalence was high (about 80%) and homogeneous across different beech forest categories (i.e., young, old, unmanaged) and for all bird species, except Erithacus rubecula (35%). Parasitemia varied across bird species but not across forest categories within each avian species (lowest parasitemia were found in E. rubecula, Turdus merula, and Turdus philomelos). In our study system, we found that vector abundance was not the main driver of parasite dynamics. We found that forest structure affects parasite infection probability directly and potentially host condition via available resources that have to be used either to combat infections (i.e., high parasitemia) or to maintain a good body condition.ConclusionsThe effects of each of the predictors were bird species-specific, and we found that Diptera vectors were not the foremost influence in our host-vector-parasite system. Effects of forest habitat variables indicated that for most bird species in this study, habitat regulation of infection probability was more likely (i.e., E. rubecula, Fringilla coelebs, Sylvia atricapilla), whereas for Parus major habitat characteristics impacted first individuals' body condition and subsequently the probability of infection. Our findings emphasize the need of species-specific analyses and to use continuous forest structural parameters (e.g., the proportion of gap, south facing aspect) to better understand habitat and land use effects on host-vector-parasite dynamics.

Unimodal Relationships of Understory Alpha and Beta Diversity along Chronosequence in Coppiced and Unmanaged Beech Forests

Patterns of diversity across spatial scales in forest successions are being overlooked, despite their importance for developing sustainable management practices. Here, we tested the recently proposed U-shaped biodiversity model of forest succession. A chronosequence of 11 stands spanning from 5 to 400 years since the last disturbance was used. Understory species presence was recorded along 200 m long transects of 20 × 20 cm quadrates. Alpha diversity (species richness, Shannon and Simpson diversity indices) and three types of beta diversity indices were assessed at multiple scales. Beta diversity was expressed by a) spatial compositional variability (number and diversity of species combinations), b) pairwise spatial turnover (between plots Sorensen, Jaccard, and Bray–Curtis dissimilarity), and c) spatial variability coefficients (CV% of alpha diversity measures). Our results supported the U-shaped model for both alpha and beta diversity. The strongest differences appeared between active and abandoned coppices. The maximum beta diversity emerged at characteristic scales of 2 m in young coppices and 10 m in later successional stages. We conclude that traditional coppice management maintains high structural diversity and heterogeneity in the understory. The similarly high beta diversities in active coppices and old-growth forests suggest the presence of microhabitats for specialist species of high conservation value.

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

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

Plant Biodiversity under Impact of Slope Position in Managed and Unmanaged Beech Forest of Asalem-Gilan

Plant Biodiversity under Impact of Slope Position in Managed and Unmanaged Beech Forest of Asalem-Gilan

Dissolved organic matter characteristics of deciduous and coniferous forests with variable management: different at the source, aligned in the soil

Abstract. Dissolved organic matter (DOM) is part of the biogeochemical cycles of carbon and nutrients, carries pollutants and drives soil formation. The DOM concentration and properties along the water flow path through forest ecosystems depend on its sampling location and transformation processes. To improve our understanding of the effects of forest management, especially tree species selection and management intensity, on DOM concentrations and properties of samples from different ecosystem fluxes, we studied throughfall, stemflow, litter leachate and mineral soil solution at 26 forest sites in the three regions of the German Biodiversity Exploratories. We covered forest stands with three management categories (coniferous, deciduous age class and unmanaged beech forests). In water samples from these forests, we monitored DOC concentrations over 4 years and characterized the quality of DOM with UV-vis absorption, fluorescence spectroscopy combined with parallel factor analysis (PARAFAC) and Fourier transform ion cyclotron resonance mass spectrometry (FT-ICR-MS). Additionally, we performed incubation-based biodegradation assays. Multivariate statistics revealed strong significant effects of ecosystem fluxes and smaller effects of main tree species on DOM quality. Coniferous forests differed from deciduous forests by showing larger DOC concentrations, more lignin- and protein-like molecules, and fewer tannin-like molecules in throughfall, stemflow, and litter leachate. Cluster analysis of FT-ICR-MS data indicated that DOM compositions, which varied in aboveground samples depending on tree species, become aligned in mineral soil. This alignment of DOM composition along the water flow path in mineral soil is likely caused by microbial production and consumption of DOM in combination with its interaction with the solid phase, producing a characteristic pattern of organic compounds in forest mineral soils. We found similarly pronounced effects of ecosystem fluxes on the biodegradability of DOM, but surprisingly no differences between deciduous and coniferous forests. Forest management intensity, mainly determined by biomass extraction, contribution of species, which are not site-adapted, and deadwood mass, did not influence DOC concentrations, DOM composition and properties significantly.

Effects of management on ambrosia beetles and their antagonists in European beech forests

Land-use intensification has been shown to negatively affect biodiversity and ecosystem functioning. Because higher trophic levels are expected to suffer most strongly from intensification, negative consequences for ecosystem services, such as biological pest control by natural enemies, are likely. In European beech forests ambrosia beetles are common secondary pests of freshly cut wood and stored timber, but they may also infest living trees that are highly stressed.We examined whether forest management intensity, ranging from unmanaged beech forests to non-natural conifer plantations, affects the abundance, attack rate, and breeding success of ambrosia beetles across three regions of Germany. We applied pheromone trapping to estimate the abundance of ambrosia beetles and exposed experimental beech logs to measure infestation rates and breeding success.In general, we found decreasing abundance and attack probability by ambrosia beetles with increasing management intensity, which is most likely related to the availability of suitable breeding substrate. However, the abundance of the invasive species Xylosandrus germanus increased with increasing management intensity in one region, where high management intensity was represented by pine forests; the drier and warmer conditions in this forest type most likely increased population densities. Breeding success of Trypodendron domesticum increased with increasing management intensity, suggesting less effective pest control by natural antagonists, in particular by specialised parasitoids, at high management intensities.We conclude that the availability of breeding substrate, in combination with microclimate and antagonist effectiveness, shape ambrosia beetle populations in Central European forests. Forest management strategies should thus reduce the availability of breeding substrate in production forests, in particular where microclimatic conditions are favourable for beetle reproduction (i.e. warm, moist). Moreover, unmanaged forest patches should be maintained to promote parasitoid abundance and thus to mitigate negative effects of climate change, such as increasing drought stress of trees.

Effects of forest management on bryophyte species richness in Central European forests

We studied the effect of three major forest management types (unmanaged beech, selection beech, and age class forests) and stand variables (SMId, soil pH, proportion of conifers, litter cover, deadwood cover, rock cover and cumulative cover of woody trees and shrubs) on bryophyte species richness in 1050 forest plots in three regions in Germany. In addition, we analysed the species richness of four ecological guilds of bryophytes according to their colonized substrates (deadwood, rock, soil, bark) and the number of woodland indicator bryophyte species. Beech selection forests turned out to be the most species rich management type, whereas unmanaged beech forests revealed even lower species numbers than age-class forests. Increasing conifer proportion increased bryophyte species richness but not the number of woodland indicator bryophyte species. The richness of the four ecological guilds mainly responded to the abundance of their respective substrate. We conclude that the permanent availability of suitable substrates is most important for bryophyte species richness in forests, which is not stringently linked to management type. Therefore, managed age-class forests and selection forests may even exceed unmanaged forests in bryophyte species richness due to higher substrate supply and therefore represent important habitats for bryophytes. Typical woodland indicator bryophytes and their species richness were negatively affected by SMId (management intensity) and therefore better indicate forest integrity than the species richness of all bryophytes. Nature conservation efforts should focus on the reduction of management intensity. Moreover, maintaining and increasing a variability of substrates and habitats, such as coarse woody debris, increasing structural heterogeneity by retaining patches with groups of old, mature to over-mature trees in managed forests, maintaining forest climate conditions by silvicultural methods that assure stand continuity, e.g. by selection cutting rather than clear cutting and shelterwood logging might promote bryophyte diversity and in particular the one of woodland indicator bryophytes.

Plastic tree crowns contribute to small-scale heterogeneity in virgin beech forests—An individual-based modeling approach

Old-growth beech (Fagus sylvatica L.) forests in Europe show a structural heterogeneity, which distinguish from managed ones. Recent investigations revealed that facilitative belowground interactions might counteract aboveground competition for light. However, the exact interaction of these mechanisms is unclear. We developed the BEEchPlasticity (BEEP) model to investigate whether the mere focus on the aboveground competition for canopy space with subsequent modeling of the light transmittance through the forest canopy suffices to reproduce the observed structural attributes. The BEEP model is individual-based, and explicitly describes the plasticity of tree crowns in a three-dimensional space through a geometric approach. This conceptual design allows tracing neighborhood effects to the forest stand level.We ran 10 simulation experiments with 1000 time steps on a simulation area of 0.5öha. We analyzed the emerging spatial point patterns, gap dynamics as well as distributions of tree age, tree height, tree crown projection area, and tree diameter. By applying various indices that revealed the emergent horizontal and vertical forest structure, we were able to show a strong mutual link between beech crown plasticity and forest heterogeneity. The crown plasticity enabled the trees to close small gaps, which resulted in a highly dynamic tree regeneration, which in turn led to a small-scale heterogeneous forest structure. We found that, over long periods, crown centroids of canopy trees taller than 20öm were more regularly distributed than stem foot points. Absolute crown displacements were greater than reported from beech stands in North-Germany and the Sudetes in the Czech Republic. This indicates a need to restrain the model crown growth in order to recognize tree architecture and stability. We could simulate an increasing forest structural heterogeneity. Starting with a structural homogeneous layer of tree saplings, we showed that forest structures similar to those in unmanaged beech forests took several tree generations or approximately 700 time steps to emerge. We recommend enhancing this model approach with spatial explicit modeling of the leaf-area distribution, which has already been done for single beech trees.

Spatial patterns of tree species in Suserup Skov – a semi-natural forest in Denmark

Abstract In Europe, concerns regarding the provision of ecosystem services from forests, such as biodiversity, has led to a widespread conversion from even-aged to close-to-nature forest management. These concerns have also led to a demand for quantifying stand structure, dynamics, and species diversity in order to evaluate ecosystem status and devise management action. In this study, species distribution, species association, and size diversity in a semi-natural, unmanaged beech forest in Denmark were analyzed using spatial indices including uniform angle index, mingling index, and differentiation index and spatial functions including pair correlation function, bivariate pair correlation function, mark variogram, and the Wiegand scheme. To analyze the spatial patterns of tree species, two plots, one in a less disturbed part of the forest and the other in a more recently disturbed part, was selected. Based on the spatial functions, all species showed a clustered pattern as a result of forest dynamics. Fagus sylvatica L. was segregated from other species and was more likely to form pure groups. In contrast, Ulmus glabra L.-Fraxinus excelsior L. and Ulmus glabra L.-Acer psedoplatanous L. showed positive association. All species in the less disturbed part of the forest showed positive spatial correlation of tree diameters, probably as a result of gap-phase dynamics. However, a lack of spatial correlation of tree diameters was observed for beech and elm trees in the more recently disturbed part of the forest. Our results indicated that spatial functions using stem-mapped data provide more reliable results and additional insights into the stand structure than neighborhood based indices. The study findings can be used to align forest management practices with goals of protecting biodiversity in managed forest.

Driving factors and temporal fluctuation of Collembola communities and reproductive mode across forest types and regions.

Despite the major role of Collembola in forest soil animal food webs, ecological and evolutionary determinants of their community composition are not well understood. We investigated abundance, community structure, life forms, and reproductive mode of Collembola in four different forest types (coniferous, young managed beech, old managed beech, and unmanaged beech forests) representing different management intensities. Forest types were replicated within three regions across Germany: the Schorfheide‐Chorin, the Hainich, and the Swabian Alb, differing in geology, altitude, and climate. To account for temporal variation, samples were taken twice with an interval of 3 years. To identify driving factors of Collembola community structure, we applied structural equation modeling, including an index of forest management intensity, abiotic and biotic factors such as pH, C‐to‐N ratio of leaf litter, microbial biomass, and fungal‐to‐bacterial ratio. Collembola abundance, biomass, and community composition differed markedly between years, with most pronounced differences in the Schorfheide, the region with the harshest climatic conditions. There, temporal fluctuations of parthenogenetic Collembola were significantly higher than in the other regions. In the year with the more favorable conditions, parthenogenetic species flourished, with their abundance depending mainly on abiotic, density‐independent factors. This is in line with the “Structured Resource Theory of Sexual Reproduction,” stating that parthenogenetic species are favored if density‐independent factors, such as desiccation, frost or flooding, prevail. In contrast, sexual species in the same year were mainly influenced by resource quality‐related factors such as the fungal‐to‐bacterial ratio and the C‐to‐N ratio of leaf litter. The influence of forest management intensity on abundances was low, indicating that disturbance through forest management plays a minor role. Accordingly, differences in community composition were more pronounced between regions than between different forest types, pointing to the importance of regional factors.

Deadwood enrichment in European forests – Which tree species should be used to promote saproxylic beetle diversity?

Modification of natural ecosystems has threatened biodiversity worldwide, with forests suffering especially. Strategies aimed at mitigating such loss in forests often include enrichment of deadwood, a critical resource for many decomposer species. However, it remains unclear how deadwood can best be enriched to most effectively promote the diversity of saproxylic species. In this study, we investigated saproxylic beetle diversity in experimentally exposed deadwood logs of 13 different tree species across 30 forests in three regions of Germany. We tested whether gamma-diversity differs between tree species and whether the alpha-diversity within an individual log depended on whether logs were placed in unmanaged beech forests, managed beech forests, or managed conifer forests. We found significant differences in gamma- and alpha-diversity of saproxylic beetles among tree species, but the ranking of tree species differed between regions, suggesting differences in regional beetle species pools. Randomization tests aiming to identify how many and which deadwood logs would need to be exposed to best conserve saproxylic beetle diversity, showed that the overall diversity of beetles increased with the number of tree species exposed, due to turnover of beetle species between tree species. However, some species (e.g. Carpinus) and species combinations (e.g. Carpinus-Picea) reached exceptionally high beetle diversity. Alpha-diversity was higher in conifer than in beech forests, but did not differ between managed and unmanaged beech forests. Canopy cover above logs and average stand temperature strongly influenced alpha-diversity, suggesting that environmental conditions that may be affected by management act as habitat filters for species assemblages. We conclude that deadwood enrichment strategies would be most effective when combining particular tree species that support highest diversity.

Differences in carbon uptake and water use between a managed and an unmanaged beech forest in central Germany

Net atmospheric carbon dioxide exchange, total evapotranspiration and net primary production of two neighbouring beech (Fagus sylvatica L.) forests in central Germany differing in site management were measured using the eddy covariance technique and biometric methods. The unmanaged site was an old-growth forest with admixtures of ash (Fraxinus excelsior L.) and sycamore (Acer pseudoplatanus L.) trees whereas the managed site was a regularly thinned, even-aged (about 130years old), pure beech stand. Average carbon fluxes measured over seven years did not differ significantly between the two forests. Evapotranspiration was slightly higher (and consequently water use efficiency lower) at the unmanaged site. The maximum rates as well as the interannual variability of both net ecosystem exchange and net primary production were considerably larger at the managed, even-aged stand. The lowest annual carbon sequestration rates were observed in years with high fruit production in beech combined with cold and dry spells during leaf development which affected the carbon balance of the managed forest more than that of the unmanaged forest. In contrast, an extraordinarily dry period in late summer 2003 caused a stronger reduction in net carbon uptake in the old-growth forest which can probably be attributed to the contribution of ash to the ecosystem fluxes. Ash has a different phenology which made it more susceptible to the late summer drought. The relative importance of tree and structural (age, size) diversity, leaf area index and regenerative growth as well as the temporal frame and extent of single weather extremes for the forest–atmosphere exchange is discussed and it is concluded that site management and forest structure needs to be included in soil–vegetation–atmosphere-transfer models as it can often override effects of land use or plant functional type.

Association of tree and plot characteristics with microhabitat formation in European beech and Douglas-fir forests

Process-orientated, unmanaged forest remnants are not sufficient for halting the loss of forest biodiversity. Thus, integrated biodiversity-promoting management for forest inhabitants is needed. Microhabitats, such as tree cavities or bark pockets, are essential for the preservation of saproxylic species and of critical importance for endangered ones. This study investigates (1) which factors trigger the formation of microhabitats at both the individual tree and aggregated plot level, and (2) whether the co-occurrence of microhabitats differs between managed (=logged) and unmanaged forests. Relationships between the occurrence of 17 microhabitat types and individual tree features (e.g. light availability, and tree vitality) and plot characteristics (e.g. stand density index and stand age) in 398 plots dominated by Fagus sylvatica or Pseudotsuga menziesii in Germany and the USA were studied using random-effects logistic and normal regression modelling. Separate analyses were performed for German beech forests, German Douglas-fir forests, and the US Douglas-fir forests. Our results show that (1) tree diameter in breast height (DBH), tree vitality and branchiness or epicormic branches are highly related with the occurrence of one or more microhabitats on individual trees in managed and unmanaged beech and US Douglas-fir forests. In managed German Douglas-fir forests, vitality is not a predictor for the occurrence of microhabitats on a tree, but tree density and the maximum age of trees in a stand in addition to DBH and branchiness have an effect. Time since last management is not a statistically significant predictor for the presence of microhabitats at the tree level, but it is for German beech at the plot level. In Douglas-fir-dominated forests both in Germany and in the USA, the stand density index was the only common predictor at the plot level. (2) Unmanaged German beech and Douglas-fir forests exhibit more statistically significant and positive correlations with microhabitat groups than managed stands, implying that the presence of one microhabitat group on a tree is associated with the presence of other microhabitat groups. We finally conclude that measures for supporting microhabitat inhabitants in managed forests are scale and species dependent (tree versus plot level; beech versus Douglas-fir-dominated forests). Trees that carry microhabitats seem to have similar features independently of forest management. At the plot level, density management may trigger the accumulation of microhabitats. Our results indicate that in forest management, it is possible to consider the factors influencing the formation of microhabitats and implement adequate forest practices to advance their formation.

Controls on soil carbon storage and turnover in German landscapes

Soil organic carbon (OC) storage across regions is influenced by climate and parent materials, which determine soil properties like clay content and mineralogy. Within homogeneous soil regions, land use and management practices are further important controls for soil OC contents and turnover. Here, we studied the impact of study region, land use (forest, grassland), forest management (spruce and beech forest under age-class management, unmanaged beech forest), and grassland management (meadow, mown pasture, unmown pasture) on stocks and turnover (based on Δ14C values) of soil OC in density frations of topsoil horizons. Samples were taken from 36 plots in the regions Hainich–Dun (HAI) and the Schwabische Alb (ALB) in Germany. They were separated into two light fractions (free light fraction (LF1), occluded light fraction (LF2)) and the mineral-associated organic matter (MOM) fraction using sodium polytungstate with a density of 1.6 g cm−3. Overall most soil OC was stored in the MOM fraction (73%). Soil OC concentrations and stocks in the MOM fraction differed between study regions, probably due to larger amounts of pedogenic Al- and Fe-oxides in the ALB than in the HAI region. Within each region, forest soils stored significantly higher proportions of total OC in the two LF (33±1.9 %) than grassland soils (20±2.3 %). Different management of forests and grasslands affected the C:N ratio of density fractions, but not OC storage. While modelled soil OC turnover in the MOM was longest of all fractions, all fractions had average Δ14C values above atmospheric values, suggesting a significant fast-cycling component in all of them. Different from stocks, turnover of OC in the MOM fraction were not affected by study region or contents of pedogenic oxides. Radiocarbon contents in the LF were higher for forest than for grassland sites, indicating faster turnover of OC at grassland sites. However, some of the observed difference could originate from different average lifetimes of roots in forests and grasslands. Applying different lag-times for OC input for forests and grasslands significantly reduced the differences in modelled turnover times. Lower Δ14C values of mown pastures than pasture soils in both regions suggest a management effect on soil C turnover in grasslands.We conclude that OC storage in the MOM of topsoil layers is more affected by regional differences in soil texture and mineralogy than by land use and management, while its turnover could not be explained with the studied soil properties. Soil OC storage and turnover in the two LFs is influenced by land use (forest or grassland) and management, but ecosystem specific lag-times have to be considered for modelling OC turnover in these fractions.

Trophic shift of soil animal species with forest type as indicated by stable isotope analysis

Anthropogenic land use shapes the dynamics and composition of central European forests and changes the quality and availability of resources of the decomposer system. These changes likely alter the structure and functioning of soil animal food webs. Using stable isotope analysis (13C, 15N) we investigated the trophic position and resource use of soil animal species in each of four forest types (coniferous, young managed beech, old managed beech and unmanaged beech forests) across three regions in Germany. Twenty‐eight species of soil invertebrates were analyzed covering three consumer levels and a representative spectrum of feeding types and morphologies. Data on stable isotope signatures of leaf litter, fine roots and soil were included to evaluate to which extent signatures of soil animals vary with those of local resources. Soil animal δ15N and δ13C signatures varied with the respective signatures of leaf litter and fine roots. After calibration to leaf litter signatures, soil animal stable isotope signatures of the different beech forests did not differ significantly. However, thick leaf litter layers, such as those in coniferous forests, were associated with low animal stable isotope signatures presumably due to reduced access of decomposer animals to root‐derived resources, suggesting that the decomposer food web is shifted towards leaf litter based energy pathways with the shift affecting all consumer levels. Variation in stable isotope signatures of soil animal species with litter quality parameters suggests that nutrition of third level but not first and second level consumers is related to litter quality, potentially due to microorganisms locking up litter resources thereby hampering their propagation to higher trophic levels.

The Contribution of Managed and Unmanaged Forests to Climate Change Mitigation—A Model Approach at Stand Level for the Main Tree Species in Bavaria

Forestry-based carbon sequestration projects demand a comprehensive quantification of the different climate change mitigation effects. In our study, we modeled a life cycle of managed pure stands consisting of the four main tree species in Bavaria (spruce, pine, beech and oak). For spruce and beech, an unmanaged stand was additionally integrated in order to analyze the differences in climate change mitigation effects compared to the managed stands. We developed a climate change mitigation model, where stand development and silvicultural treatments including harvested timber volumes were conducted using the tree growth model Silva 2.3. The harvested wood products (HWP), including their substitution effects were calculated with a subsequent model. For unmanaged beech forests, we compiled measured data from the literature, and Bavarian strict forest reserves for validating our model results. The results for the managed stands reveal that spruce provides the highest total climate change mitigation effects. After a simulation period of 180 years, one hectare leads to a mean mitigation benefit of 13.5 Mg CO2 ha−1 year−1. In comparison, results for pine, beech and oak reveal lesser benefits with 10.1 Mg CO2 ha−1 year−1, 9.1 Mg CO2 ha−1 year−1 and 7.2 Mg CO2 ha−1 year−1, respectively. However, these results assume current growing conditions. Considering climate change, it is very likely that spruce will not be suitable in several regions of Bavaria in the future. Furthermore, excessive disturbances could affect spruce more drastically than the other tree species. In that case, the order could change and beech could exceed spruce. Thus the results cannot be seen as a general recommendation to establish spruce stands in order to achieve optimal climate change mitigation benefits. Nevertheless, results for spruce illustrate that high increment and especially wood use in long-lived products is crucial for high climate change mitigation effects. Mitigation effects in unmanaged spruce and beech stands do not differ in the first decades from their managed counterparts, but are below them in the long term with a total climate change mitigation benefit of 8.0 Mg CO2 ha−1 year−1 and 7.2 Mg CO2 ha−1 year−1, respectively. These differences are mainly caused by the missing substitution effects in the unmanaged stands. However, the precise dimensions of substitution effects still remain uncertain and the lack of data should be reduced via additional life cycle assessments for more products and product classes. However, neglecting substitution effects in climate change mitigation models leads to severe underestimations of the mitigation effects in managed forests.

Management tradeoff between endangered species and biodiversity conservation: The case of Carabus olympiae (Coleoptera: Carabidae) and carabid diversity in north-western Italian Alps

The worldwide loss of biodiversity is of great concern, and this has lead to strong policy frameworks to promote the maintenance of biodiversity through protecting species and habitats in many countries. Often however, rare species with legal protection provide a conservation focus, and therefore may dictate management policy. Co-occurring species, especially those without formal protected status, may contribute significantly to overall biodiversity, but may nonetheless receive limited attention. In such cases, conflicts may arise between the needs of individual species and those of wider biodiversity. We examine such a conflict in alpine ground beetles of the Sessera Valley (NW Italian Alps). This area is the type locality of the endangered species Carabus olympiae Sella, 1855, an endemic species known only from two restricted sites in the Alps. In order to assess priorities for C. olympiae conservation we used habitat suitability modeling, and identified potential indicator species that could serve as an indicator of the occurrence of the target species. In order to assess priorities for ground beetle diversity conservation we described the local ground beetle community by calculating abundance, species richness, average taxonomic distinctness (Δ+), taxonomic distinctness (Δ*) and functional diversity (FD). The best management option for the conservation of ground beetle diversity was found to be maintenance of unforested habitats, in particular pastures, which showed the highest values for all the parameters considered. Forested habitats (artificial plantation and beech forest) had the lowest ground beetle diversity. However, the preferred habitat for C. olympiae was beech forest, especially at medium altitudes (800–1400m) and with sloping terrain. Moreover, C. olympiae presence could be indirectly assessed by the co-occurrence of four indicator species: Tanythrix senilis, Carabus depressuss, Cychrus italicus and Pterostichus appenninus. Our main conclusion is that the most promising landscape-scale approach to preserve both the relict steno-endemic species C. olympiae and the high local syntopic ground beetle diversity is to maintain traditional pastoral activities along with the preservation of the unmanaged beech forests.