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Author(s): Martin M. Gossner (corresponding author) [1, 2, 3]; Thomas M. Lewinsohn [1, 4]; Tiemo Kahl [5, 6]; Fabrice Grassein [7]; Steffen Boch [7]; Daniel Prati [6]; Klaus Birkhofer [8, [...]

Despite the important role of wood-inhabiting fungi (WIF) in deadwood decomposition, our knowledge of the factors shaping the dynamics of their species richness and community composition is scarce. This is due to limitations regarding the resolution of classical methods used for characterizing WIF communities and to a lack of well-replicated long-term experiments with sufficient numbers of tree species. Here, we used a large scale experiment with logs of 11 tree species at an early stage of decomposition, distributed across three regions of Germany, to identify the factors shaping WIF community composition and Operational Taxonomic Unit (OTU) richness using next generation sequencing. We found that tree species identity was the most significant factor, corresponding to (P < 0.001) and explaining 10% (representing 48% of the explainable variance) of the overall WIF community composition. The next important group of variables were wood-physicochemical properties, of which wood pH was the only factor that consistently corresponded to WIF community composition. For overall WIF richness patterns, we found that approximately 20% of the total variance was explained by wood N content, location, tree species identity and wood density. It is noteworthy that the importance of determinants of WIF community composition and richness appeared to depend greatly on tree species group (broadleaved vs. coniferous) and it differed between the fungal phyla Ascomycota and Basidiomycota. [ABSTRACT FROM AUTHOR]

Abstract: Knowledge on food web structure and function provides important information on trophic relationships, energy flux, and ecosystem response to environmental changes. The tracing of trophic marker fatty acids (FAs) to assign predator–prey interactions is well established in food web research. However, the application on the level of entire consumer communities has not been performed yet. Here, we demonstrate that lipid pattern analysis of entire consumer assemblages can provide a first‐line assessment for food web diagnostics. The trophic diverse group of nematodes was used as a model and community level lipid profiling (CLLP) was performed. As prerequisite a lipid library was constructed, assigning FAs to their predominant organismic origin to disentangle phylogenetic (i.e. consumer lipid metabolism) from diet type effects on CLLP. The suitability of CLLP analysis was tested using 150 forest sites differing in management type and intensity across three German regions. The nematode CLLP reflected ecosystem conditions and thereby separated regions as well as forest habitats, that is, conifer and deciduous stands. CLLP further enabled to address if environmental properties acted on the level of consumers or their resource use or both. However, forest management intensity was poorly assigned, likely due to a predominant species‐specific impact, not represented on community basis. We propose CLLP as a fast and robust biochemical method for food web diagnostics in cryptic habitats such as soil or benthos. Provided that organismic assemblages compared are substantially different in lipid metabolism, CLLP give insight into changes in the composition of consumer communities as well as their major diet under field conditions. [ABSTRACT FROM AUTHOR]

Summary: Nitrogen deposition can strongly affect biodiversity, but its specific effects on terrestrial microbial communities and their roles for ecosystem functions and processes are still unclear. Here, we investigated the impacts of N deposition on wood‐inhabiting fungi (WIF) and their related ecological functions and processes in a highly N‐limited deadwood habitat. Based on high‐throughput sequencing, enzymatic activity assay and measurements of wood decomposition rates, we show that N addition has no significant effect on the overall WIF community composition or on related ecosystem functions and processes in this habitat. Nevertheless, we detected several switches in presence/absence (gain/loss) of wood‐inhabiting fungal OTUs due to the effect of N addition. The responses of WIF differed from previous studies carried out with fungi living in soil and leaf‐litter, which represent less N‐limited fungal habitats. Our results suggest that adaptation at different levels of organization and functional redundancy may explain this buffered response and the resistant microbial‐mediated ecosystem function and processes against N deposition in highly N‐limited habitats. [ABSTRACT FROM AUTHOR]

Microorganisms in forest soils provide essential ecosystem services, such as decomposition of organic matter and nutrient mineralization. However, microbial community structure and function can be affected by environmental conditions, such as regional climate and soil properties and, moreover, by human activity through forest management. We examined the biomass and composition of microbial assemblages in 150 forest stands in the organic layer (Oi, Oe, Oa) and upper mineral soil (0–10 cm) in three regions across Germany (Schwäbische Alb, Hainich-Dün, Schorfheide-Chorin) by phospholipid fatty acid (PLFA) analysis. Different explanatory environmental variables (total C, N, S, P, organic C, inorganic C, pH, water content) were identified. The intensity of land use was characterized with the Forest Management Intensity Index (ForMI). The total amount of PLFAs, as measure for microbial biomass, was different among the three regions both in the organic layer and mineral soil. In the organic layer, total PLFAs decreased from Schwäbische Alb over Hainich-Dün to Schorfheide-Chorin, with the latter comprising a fourfold and twofold lower amount in fungal and bacterial PLFAs, respectively. In contrast, in the mineral soil the forests in the Hainich-Dün showed the highest microbial biomass. Discriminant function analysis of PLFA pattern indicated that Gram-positive bacteria and fungi accounted mainly for the regional differences in the organic layer, whereas in the mineral soil additionally Gram-negative and actinobacteria were important. Redundancy analysis showed that PLFA profiles were predominantly affected by sampling site and environmental variables, with the water content in the organic layer and the soil texture in the mineral soil explaining most of the variability in microbial communities between the three regions. Additionally, forest stands were classified into four management groups (conifer; deciduous with low, medium, and high intensity) based on the ForMI. In the mineral soil, forest management accounted for a small proportion of the observed regional differences. Within regions, fungal biomass in the organic layer decreased with management intensity at the Schwäbische Alb and increased in the mineral soils of Hainich-Dün region. Microbial community structure discriminated coniferous and deciduous forests in all three regions, and moreover showed a separation based on forest management intensity in the Schorfheide-Chorin. In conclusion, microbial biomass and community composition in forest organic layer and mineral soil were more influenced by regional conditions, including environmental properties such as moisture, soil texture, C/N ratio and pH, than by forest management intensity. However, within given environments, microbial assemblages can be influenced by forest management, in particular through changes in the tree species composition. [ABSTRACT FROM AUTHOR]

Deadwood decay is an important ecosystem process in forest ecosystems, but the relative contribution of specific wood properties of tree species, activities of wood-degrading enzymes, and decomposer communities such as fungi and insects is unclear. We ask whether wood properties, in particular differences between angiosperms and gymnosperms, and organismic diversity of colonizers contribute to wood decomposition. To test this, we exposed deadwood logs of 13 tree species, covering four gymnosperms and nine angiosperm species, in 30 plots under different forest management in three regions in Germany. After a decomposition time of 6.5 years Carpinus betulus and Fagus sylvatica showed the highest decay rates. We found a positive correlation of decay rate with enzyme activities, chemical wood properties (S, K concentration) and organismic diversity, while, heartwood character, lignin content, extractive concentration and phenol content were negatively correlated with decay rate across all 13 tree species. By applying a multi-model inference approach we found that the activity of the wood-degrading enzymes laccase and endocellulase, beetle diversity, heartwood presence, wood ray height and fungal diversity were the most important predictor variables for wood decay. Although we were not able to identify direct cause and effect relations by our approach, we conclude that enzyme activity and organismic diversity are the main drivers of wood decay rate, which greatly differed among tree species. Maintaining high tree species diversity will therefore result in high structural deadwood diversity in terms of decay rate and decay stage. [ABSTRACT FROM AUTHOR]

In forest ecosystems, deadwood is an important component that provides habitat and contributes to nutrient cycles, as well as to carbon and water storage. The change of wood constituents, nutrients and microbial species richness in the field over the whole time of decomposition has only rarely been studied, in particular not in relation to oxidative enzyme activities (mediating lignin degradation) and different forest management regimes. To describe wood decomposition, we selected coarse woody debris (CWD) in form of 197 logs of Fagus sylvatica , Picea abies and Pinus sylvestris in forests with different management regimes across three regions in Germany. They were sampled and analyzed for wood density, water content, wood constituents (Klason and acid-soluble lignin, organic extractives, water-soluble lignin fragments), carbon, nitrogen and metals (Al, Ca, Cu, K, Mg, Mn and Zn). Furthermore, the activities of oxidative enzymes like laccase, manganese peroxidase, and general peroxidase were measured. Since filamentous fungi (Basidiomycota, Ascomycota) are the major biological agents of wood decomposition, fungal species richness based on sporocarps and molecular fingerprints was recorded. Higher forest management intensity had a negative effect on deadwood volume and in consequence on fungal species richness (sporocarps), but hardly to other analyzed variables. Furthermore, there were significant differences between the tree species for the concentrations of wood constituents and most nutrients as well as the activities of oxidative enzymes, although their course during decomposition was mostly similar among the tree species. We found that molecular species richness increased with the period of decomposition in contrast to the number of fruiting species, which was highest in the intermediate stage of decomposition. Both types of species richness increased with increasing volume of the CWD logs. Regarding the entire period of decomposition, white-rot fungi (WRF), based on identification of sporocarps, were the most abundant group of wood-decaying fungi in all three tree species. This corresponds well with the overall presence of laccase and peroxidases and the concomitant substantial loss of lignin, which points to the importance of these enzymes in deadwood decomposition. We found a continuous decomposition and decline of volume-related concentrations in wood constituents and nutrients with time of decomposition. Contrary to volume-related concentrations, the concentrations related to dry mass frequently increased. [ABSTRACT FROM AUTHOR]

Commercial forestry increasingly aims at both optimizing timber production and maintaining species diversity. To maintain the diversity of the species-rich group of wood-inhabiting fungi, effective forest conservation concepts that include the enrichment of dead wood in commercial forests are required. However, which type of dead wood should be enriched in which type of forest stand (coniferous or broad-leaved) is still debated. Our study aimed at (1) disentangling the relative importance of forest-stand type, dead-wood origin (tree species) and time since death and (2) determining whether fungal species richness on logs of broad-leaved trees is higher in broad-leaved stands than in coniferous stands and whether fungal species richness on logs of coniferous trees is higher in coniferous stands than in broad-leaved stands (home-field advantage). We exposed logs of 9 broad-leaved and 4 coniferous tree species in 19 broad-leaved and 9 coniferous forest stands in 2009 and surveyed the logs in 2012 and 2014 for wood-inhabiting fungi. Across all logs, fungal species richness was mainly driven by the tree species of the dead wood and time since death, whereas fungal community composition was solely driven by the tree species of the dead wood. The fungal species richness and community composition of broad-leaved logs was significantly correlated to time since death but not to forest-stand type. The fungal species richness and community composition of coniferous logs was neither affected by forest-stand type nor time since death. When individual tree species were considered, forest-stand type did not affect fungal species richness or community compositions, but fungal species richness on logs of Acer , Fagus , Carpinus and Populus increased with time since death. To increase the species richness of wood-inhabiting fungi in commercial forests, we recommend that the tree species diversity of dead wood should be increased and should especially originate from different lineages (angiosperms and gymnosperms), and that a broad variety of successional stages of dead wood should be maintained. Our results suggest that such a strategy would be effective irrespective of the tree species composition of the forest stand, as we found no support for home-field advantages in the early stage of decomposition. [ABSTRACT FROM AUTHOR]

Wood-inhabiting fungi are major agents of wood decomposition. However, it is unclear which factors determine their distribution and enzyme production. Many studies that have addressed this issue suffer from a lack of geographic extent. Here, we investigate the fungal community structure of 117 Fagus sylvatica logs in relation to wood physico-chemical properties and secreted ligninolytic enzymes, across three distinct geographical regions of Germany. Our results revealed that fungal community structure was similar across different regions, but was nevertheless variable in all regions. The relationships between fungal community structure, wood physico-chemical properties and enzyme activities were not consistent across different regions. However, we identified that the wood physico-chemical properties (i.e. decay class, remaining mass, density, extractives, total lignin and pH) were the most important factors associated with the fungal community structure in all three regions. In contrast, the wood physico-chemical properties and the fungal community structure did not sufficiently explain variation in the detected enzymatic activities. Thus, we assume that interspecific interactions and recently described priority effects play more important roles in the production of lignin modifying enzymes. [ABSTRACT FROM AUTHOR]

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

Large dead wood is an important structural component of forest ecosystems and a main component of forest carbon cycles. CO2 emissions from dead wood can be used as a proxy for actual decomposition rates. The main drivers of CO2 emission rates for dead wood of temperate European tree species are largely unknown. We applied a novel, closed chamber measurement technique to 360 dead wood logs of 13 important tree species in three regions in Germany. We found that tree species identity was with 71% independent contribution to the model (R² = 0.62) the most important driver of volume-based CO2 emission rates, with angiosperms having on average higher rates than conifers. Wood temperature and fungal species richness had a positive effect on CO2 emission rates, whereas wood density had a negative effect. This is the first time that positive fungal species richness-wood decomposition relationship in temperate forests was shown. Certain fungal species were associated with high or low CO2 emission rates. In addition, as indicated by separate models for each tree species, forest management intensity, study region, and the water content as well as C and N concentration of dead wood influenced CO2 emission rates. [ABSTRACT FROM AUTHOR]

Many studies have examined whether communities are structured by random or deterministic processes, and both are likely to play a role, but relatively few studies have attempted to quantify the degree of randomness in species composition. We quantified, for the first time, the degree of randomness in forest bird communities based on an analysis of spatial autocorrelation in three regions of Germany. The compositional dissimilarity between pairs of forest patches was regressed against the distance between them. We then calculated the y-intercept of the curve, i.e. the ‘nugget’, which represents the compositional dissimilarity at zero spatial distance. We therefore assume, following similar work on plant communities, that this represents the degree of randomness in species composition. We then analysed how the degree of randomness in community composition varied over time and with forest management intensity, which we expected to reduce the importance of random processes by increasing the strength of environmental drivers. We found that a high portion of the bird community composition could be explained by chance (overall mean of 0.63), implying that most of the variation in local bird community composition is driven by stochastic processes. Forest management intensity did not consistently affect the mean degree of randomness in community composition, perhaps because the bird communities were relatively insensitive to management intensity. We found a high temporal variation in the degree of randomness, which may indicate temporal variation in assembly processes and in the importance of key environmental drivers. We conclude that the degree of randomness in community composition should be considered in bird community studies, and the high values we find may indicate that bird community composition is relatively hard to predict at the regional scale. [ABSTRACT FROM AUTHOR]

Future climates are likely to include extreme events, which in turn have great impacts on ecological systems. In this study, we investigated possible effects that could mitigate stem breakage caused by a rare and extreme ice storm in a Chinese subtropical forest across a gradient of forest diversity. We used Bayesian modeling to correct stem breakage for tree size and variance components analysis to quantify the influence of taxon, leaf and wood functional traits, and stand level properties on the probability of stem breakage. We show that the taxon explained four times more variance in individual stem breakage than did stand level properties; trees with higher specific leaf area (SLA) were less susceptible to breakage. However, a large part of the variation at the taxon scale remained unexplained, implying that unmeasured or undefined traits could be used to predict damage caused by ice storms. When aggregated at the plot level, functional diversity and wood density increased after the ice storm. We suggest that for the adaption of forest management to climate change, much can still be learned from looking at functional traits at the taxon level. [ABSTRACT FROM AUTHOR]

Nitrogen availability in dead wood is highly restricted and associations with N-fixing bacteria are thought to enable wood-decaying fungi to meet their nitrogen requirements for vegetative and generative growth. We assessed the diversity of nifH (dinitrogenase reductase) genes in dead wood of the common temperate tree species Fagus sylvatica and Picea abies from differently managed forest plots in Germany using molecular tools. By incorporating these genes into a large compilation of published nifH sequences and subsequent phylogenetic analyses of deduced proteins we verified the presence of diverse pools corresponding to functional nifH, almost all of which are new to science. The distribution of nifH genes strongly correlated with tree species and decay class, but not with forest management, while higher fungal fructification was correlated with decreasing nitrogen content of the dead wood and positively correlated with nifH diversity, especially during the intermediate stage of wood decay. Network analyses based on non-random species co-occurrence patterns revealed interactions among fungi and N-fixing bacteria in the dead wood and strongly indicate the occurrence of at least commensal relationships between these taxa. [ABSTRACT FROM AUTHOR]

Unicellular saprobic fungi (yeasts) inhabit soils worldwide. Although yeast species typically occupy defined areas on the biome scale, their distribution patterns within a single type of vegetation, such as forests, are more complex. In order to understand factors that shape soil yeast communities, soils collected underneath decaying wood logs and under forest litter were analyzed. We isolated and identified molecularly a total of 25 yeast species, including three new species. Occurrence and distribution of yeasts isolated from these soils provide new insights into ecology and niche specialization of several soil-borne species. Although abundance of typical soil yeast species varied among experimental plots, the analysis of species abundance and community composition revealed a strong influence of wood log deposition and leakage of organic carbon. Unlike soils underneath logs, yeast communities in adjacent areas harbored a considerable number of transient (phylloplane-related) yeasts reaching 30% of the total yeast quantity. We showed that distinguishing autochthonous community members and species transient in soils is essential to estimate appropriate effects of environmental factors on soil fungi. Furthermore, a better understanding of species niches is crucial for analyses of culture-independent data, and may hint to the discovery of unifying patterns of microbial species distribution. [ABSTRACT FROM AUTHOR]

Deadwood is an important biodiversity hotspot in forest ecosystems. While saproxylic insects and wood-inhabiting fungi have been studied extensively, little is known about deadwood-inhabiting bacteria. The study we present is among the first to compare bacterial diversity and community structure of deadwood under field conditions. We therefore compared deadwood logs of two temperate forest tree species Fagus sylvatica and Picea abies using 16S rDNA pyrosequencing to identify changes in bacterial diversity and community structure at different stages of decay in forest plots under different management regimes. Alphaproteobacteria, Acidobacteria and Actinobacteria were the dominant taxonomic groups in both tree species. There were no differences in bacterial OTU richness between deadwood of Fagus sylvatica and Picea abies. Bacteria from the order Rhizobiales became more abundant during the intermediate and advanced stages of decay, accounting for up to 25% of the entire bacterial community in such logs. The most dominant OTU was taxonomically assigned to the genus Methylovirgula, which was recently described in a woodblock experiment of Fagus sylvatica. Besides tree species we were able to demonstrate that deadwood physico-chemical properties, in particular remaining mass, relative wood moisture, pH, and C/N ratio serve as drivers of community composition of deadwood-inhabiting bacteria. [ABSTRACT FROM AUTHOR]

Abstract Functionally, ectomycorrhizal (ECM) and saprotrophic (SAP) fungi belong to different guilds, and they play contrasting roles in forest ecosystem C-cycling. SAP fungi acquire C by degrading the soil organic material, which precipitates massive CO 2 release, whereas, as plant symbionts, ECM fungi receive C from plants representing a channel of recently assimilated C to the soil. In this study, we aim to measure the amounts and identify the drivers of ECM and SAP fungal biomass in temperate forest topsoil. To this end, we measured ECM and SAP fungal biomass in mineral topsoils (0–12 cm depth) of different forest types (pure European beech, pure conifers, and mixed European beech with other broadleaf trees or conifers) in a range of about 800 km across Germany; moreover, we conducted multi-model inference analyses using variables for forest and vegetation, nutritive resources from soil and roots, and soil conditions as potential drivers of fungal biomass. Total fungal biomass ranged from 2.4 ± 0.3 mg g−1 (soil dry weight) in pure European beech to 5.2 ± 0.8 mg g−1 in pure conifer forests. Forest type, particularly the conifer presence, had a strong effect on SAP biomass, which ranged from a mean value of 1.5 ± 0.1 mg g−1 in broadleaf to 3.3 ± 0.6 mg g−1 in conifer forests. The European beech forests had the lowest ECM fungal biomass (1.1 ± 0.3 mg g−1), but in mixtures with other broadleaf species, ECM biomass had the highest value (2.3 ± 0.2 mg g−1) among other forest types. Resources from soil and roots such as N and C concentrations or C:N ratios were the most influential variables for both SAP and ECM biomass. Furthermore, SAP biomass were driven by factors related to forest structure and vegetation, whereas ECM biomass was mainly influenced by factors related to soil conditions, such as soil temperature, moisture, and pH. Our results show that we need to consider a complex of factors differentially affecting biomass of soil fungal functional groups and highlight the potential of forest management to control forest C-storage and the consequences of changes in soil fungal biomass. Highlights • Saprotrophic and ectomycorrhizal fungal biomass vary with forest tree composition. • Nutritive resources from soil and roots are the main drivers of fungal biomass. • Forest-related factors primarily influence the saprotrophic fungal biomass. • Soil conditions mainly influence ectomycorrhizal fungal biomass. [ABSTRACT FROM AUTHOR]

Forestry in Europe changed the tree species composition and reduced dead-wood amount and heterogeneity, and therefore negatively affected saproxylic diversity. Efficient conservation requires knowledge about the importance of the relevant diversity drivers across taxa. We examined the relative importance of space vs. host for saproxylic diversity at a spatial extend of 600 km in Germany. Further, we disentangled effects of among regions, forest stands, host clades, and tree species on saproxylic diversity. This allows inferences for spatial- and host tree-related conservation strategies. Beetle, fungal sporocarp, molecular-derived fungal, and bacterial communities were studied in a large nested dead-wood experiment comprising 11 tree species. We used multiplicative diversity partitioning to assess the diversity of rare, typical, and dominant species. The beta-diversity of beetles and fungal sporocarps was equally explained by space and host, but that of molecular fungi and bacteria mainly by the host. Across taxa, beta-diversity was higher among forest stands than among regions. However, for beetles and fungal sporocarps, differences among regions were also important. Host tree clade and host tree species were important for beetle and host clade for fungal sporocarp beta-diversity. Host tree species was more important than host clade for the beta-diversity of molecular fungi and bacteria. The divergent response of different taxa to space and host calls into question the use of a simple spatially-centered or host-centered strategy. Instead, a high dead-wood tree species diversity on a broad spatial coverage at the national scale in temperate European forests is necessary to maintain rare and abundant species. • Relative importance of space versus host across saproxylic taxa is unknown. • Beetle diversity is driven by space and host, microbial diversity mainly by the host. • Different fungal diversity measures yielded different results. • Conservation requires dead-wood enrichment with different clades and tree species. • Dead-wood enrichment must be established in different forest types across regions. [ABSTRACT FROM AUTHOR]