Species-specific effects of mycorrhizal symbiosis on Populus trichocarpa after a lethal dose of copper.

Poplars have been identified as heavy metals hyperaccumulators and can be used for phytoremediation. We have previously established that their symbiosis with arbuscular mycorrhizal fungi (AMF) may alter their uptake, tolerance and distribution to excess concentrations of heavy metals in soils. In this study we hypothesised that mycorrhizal symbiosis improves the tolerance of poplars to lethal copper (Cu) concentrations, but this influence may vary among different AMF species. We conducted an experiment in a growth chamber with three Cu application levels of control (0 mg kg-1), threshold-lethal (729 mg kg-1) and supra-lethal (6561 mg kg-1), and three mycorrhizal treatments (non-mycorrhizal, Rhizophagus irregularis, and Paraglomus laccatum) in a completely randomized design with six replications. The poplars did not grow after application of 729 mg Cu kg-1 substrate, and mycorrhizal symbiosis did not help plants to tolerate this level of Cu. This can be explained by the toxicity suffered by mycorrhizal fungi. Translocation of Cu from roots to shoots increased when plants were colonised with R. irregularis and P. laccatum under threshold-lethal and supra-lethal applications of Cu, respectively. This result shows that mycorrhizal mediation of Cu partitioning in poplars depends on the fungal species and substrate Cu concentration. Multi-model inference analysis within each mycorrhizal treatment showed that in plants colonised with R. irregularis, a higher level of mycorrhizal colonisation may prevent Cu transfer to the shoots. We did not observe this effect in P. laccatum plants probably due to the relatively low colonisation rate (14%). Nutrient concentrations in roots and shoots were impacted by applied substrate Cu levels, but not by mycorrhizas. Magnesium (Mg), potassium (K), and manganese (Mn) concentrations in roots reduced with enhancing applied substrate Cu due to their similar ionic radii with Cu and having common transport mechanism. Synergistic effect on shoot concentration between applied substrate Cu levels and Mg, K, calcium, iron (Fe), and zinc was observed. Root Cu concentration was inversely related with root K and Mn concentrations, and shoot Cu concentration had a positive correlation with shoot Fe and K concentrations. Overall, mycorrhizal symbiosis has the potential to enhance plant health and their resilience to Cu toxicity in contamination events. However, it is important to note that the effectiveness of this symbiotic relationship varies among different mycorrhizal species and is influenced by the level of contamination.

A critical review of Pongamia pinnata multiple applications: From land remediation and carbon sequestration to socioeconomic benefits.

Pongamia pinnata (L.) Pierre (Pongamia) is a tree native to Southeast Asia. Recently, interest in Pongamia focused on its potential as a biofuel source as its seeds contain around 40% oil. However, Pongamia has multiple applications beyond biofuel production. It is a legume, can form symbiotic associations with mycorrhizal fungi, has been shown to be tolerant to drought, salinity, and heavy metals in soil, and has potential to mitigate climate change. Additionally, Pongamia oil has medicinal properties, can be used as biopesticide, insect repellent, to produce soap, and as a source of edible grade vegetable oil. The seed cake can be used as a source of bioenergy, food and feed protein, and organic fertiliser, and the flowers are a good source of pollen and nectar. Pongamia can also bring socio-economic benefits as its ability to restore degraded and contaminated land provides opportunities for local communities through novel valorisation pathways. These multiple applications have potential to form part of a circular bioeconomy in line with sustainable development goals. Although research on the multiple applications of Pongamia has grown considerably, knowledge gaps remain and these need to be addressed so that the full potential of Pongamia can be achieved. Further understanding of the mechanisms underlying its resilience to abiotic stresses, phytoremediation potential and biotic interactions should be a priority, and co-ordinated breeding efforts will be key. Here, we critically review the available literature on Pongamia and highlight gaps in knowledge in which future research should focus on to ensure that the full potential of this versatile tree can be achieved. We conclude that Pongamia can potentially form part of a circular bioeconomy and that harnessing the multiple applications of Pongamia in a holistic manner, with collaboration among key stakeholders, is crucial for the successful application of its benefits far beyond biofuel production.

Carbohydrate depletion in roots impedes phosphorus nutrition in young forest trees.

Nutrient imbalances cause the deterioration of tree health in European forests, but the underlying physiological mechanisms are unknown. Here, we investigated the consequences of decreasing root carbohydrate reserves for phosphorus (P) mobilisation and uptake by forest trees. In P-rich and P-poor beech (Fagus sylvatica) forests, naturally grown, young trees were girdled and used to determine root, ectomycorrhizal and microbial activities related to P mobilisation in the organic layer and mineral topsoil in comparison with those in nongirdled trees. After girdling, root carbohydrate reserves decreased. Root phosphoenolpyruvate carboxylase activities linking carbon and P metabolism increased. Root and ectomycorrhizal phosphatase activities and the abundances of bacterial genes catalysing major steps in P turnover increased, but soil enzymes involved in P mobilisation were unaffected. The physiological responses to girdling were stronger in P-poor than in P-rich forests. P uptake was decreased after girdling. The soluble and total P concentrations in roots were stable, but fine root biomass declined after girdling. Our results support that carbohydrate depletion results in reduced P uptake, enhanced internal P remobilisation and root biomass trade-off to compensate for the P shortage. As reductions in root biomass render trees more susceptible to drought, our results link tree deterioration with disturbances in the P supply as a consequence of decreased belowground carbohydrate allocation.

Soil and root nutrient chemistry structure root-associated fungal assemblages in temperate forests.

Root-associated fungi (RAF) link nutrient fluxes between soil and roots and thus play important roles in ecosystem functioning. To enhance our understanding of the factors that control RAF, we fitted statistical models to explain variation in RAF community structure using data from 150 temperate forest sites covering a broad range of environmental conditions and chemical root traits. We found that variation in RAF communities was related to both root traits (e.g., cations, carbohydrates, NO3 - ) and soil properties (pH, cations, moisture, C/N). The identified drivers were the combined result of distinct response patterns of fungal taxa (determined at the rank of orders) to biotic and abiotic factors. Our results support that RAF community variation is related to evolutionary adaptedness of fungal lineages and consequently, drivers of RAF communities are context-dependent.

Specialisation and diversity of multiple trophic groups are promoted by different forest features.

While forest management strongly influences biodiversity, it remains unclear how the structural and compositional changes caused by management affect different community dimensions (e.g. richness, specialisation, abundance or completeness) and how this differs between taxa. We assessed the effects of nine forest features (representing stand structure, heterogeneity and tree composition) on thirteen above- and belowground trophic groups of plants, animals, fungi and bacteria in 150 temperate forest plots differing in their management type. Canopy cover decreased light resources, which increased community specialisation but reduced overall diversity and abundance. Features increasing resource types and diversifying microhabitats (admixing of oaks and conifers) were important and mostly affected richness. Belowground groups responded differently to those aboveground and had weaker responses to most forest features. Our results show that we need to consider forest features rather than broad management types and highlight the importance of considering several groups and community dimensions to better inform conservation.

Assembly processes of trophic guilds in the root mycobiome of temperate forests.

Root-associated mycobiomes (RAMs) link plant and soil ecological processes, thereby supporting ecosystem functions. Understanding the forces that govern the assembly of RAMs is key to sustainable ecosystem management. Here, we dissected RAMs according to functional guilds and combined phylogenetic and multivariate analyses to distinguish and quantify the forces driving RAM assembly processes. Across large biogeographic scales (>1,000 km) in temperate forests (>100 plots), RAMs were taxonomically highly distinct but composed of a stable trophic structure encompassing symbiotrophic, ectomycorrhizal (55%), saprotrophic (7%), endotrophic (3%) and pathotrophic fungi (<1%). Taxonomic community composition of RAMs is explained by abiotic factors, forest management intensity, dominant tree family (Fagaceae, Pinaceae) and root resource traits. Local RAM assemblies are phylogenetically clustered, indicating stronger habitat filtering on roots in dry, acid soils and in conifer stands than in other forest types. The local assembly of ectomycorrhizal communities is driven by forest management intensity. At larger scales, root resource traits and soil pH shift the assembly process of ectomycorrhizal fungi from deterministic to neutral. Neutral or weak deterministic assembly processes are prevalent in saprotrophic and endophytic guilds. The remarkable consistency of the trophic composition of the RAMs suggests that temperate forests attract fungal assemblages that afford functional resilience under the current range of climatic and edaphic conditions. At local scales, the filtering processes that structure symbiotrophic assemblies can be influenced by forest management and tree selection, but at larger scales, environmental cues and host resource traits are the most prevalent forces.

Leaf litter species identity influences biochemical composition of ectomycorrhizal fungi.

In forest ecosystems, ectomycorrhizal (ECM) fungi are important for plant growth and soil biogeochemical processes. The biochemical composition of ECM mycelium is an important fungal effect trait with consequences for its decomposition rate, and consequently on soil carbon pools and plant nutrition. Although the link between ECM fungi and leaf litter-released nutrients is well known, the response of ECM fungal biochemical composition to different leaf litter species remains poorly understood. To determine how leaf litter quality influences ECM fungi's biochemical profiles, we planted young beech trees in an oak forest and replaced the natural leaf litter with that of European beech (Fagus sylvatica), ash (Fraxinus excelsior), maple (Acer pseudoplatanus), or lime (Tilia cordata). We assessed the biochemical profiles of ECM root tips colonized by common fungal taxa in temperate forests (i.e., Cenococcum geophilum, Inocybe sp., and Lactarius subdulcis), using attenuated total reflectance Fourier transform infrared spectroscopy (ATR-FTIR). ECM fungal biochemical composition changed with leaf litter species. Changes were apparent in the infrared absorption bands assigned to functional groups of lipids, amides, and carbohydrates. C. geophilum and L. subdulcis exhibited large spectral differences corresponding to the initial pattern of leaf litter chemical composition between samples collected in the beech and ash leaf litter treatments. In contrast, Inocybe sp. was influenced by lime, but with no differences between samples from ash or beech leaf litter treatments. Although the spectral bands affected by leaf litter type differed among ECM fungi, they were mainly related to amides, indicating a dynamic response of the fungal proteome to soil nutritional changes. Overall, the results indicate that the biochemical response of ECM fungi to leaf litter species varies among ECM fungal species and suggests that the biochemical composition of ECM mycelium is a fungal response trait, sensitive to environmental changes such as shifts in leaf litter species.

Ectomycorrhizal fungal diversity increases phosphorus uptake efficiency of European beech.

Increases in summer droughts and nitrogen (N) deposition have raised concerns of widespread biodiversity loss and nutrient imbalances, but our understanding of the ecological role of ectomycorrhizal fungal (ECMF) diversity in mediating root functions remains a major knowledge gap. We used different global change scenarios to experimentally alter the composition of ECMF communities colonizing European beech saplings and examined the consequences for phosphorus (P) uptake (H333 PO4 feeding experiment) and use efficiencies of trees. Specifically, we simulated increases in temperature and N deposition and decreases in soil moisture and P availability in a factorial experiment. Here, we show that ECMF α diversity is a major factor contributing to root functioning under global change. P uptake efficiency of beech significantly increased with increasing ECMF species richness and diversity, as well as with decreasing P availability. As a consequence of decreases in ECMF diversity, P uptake efficiency decreased when soil moisture was limiting. By contrast, P use efficiencies were a direct (negative) function of P availability and not of ECMF diversity. We conclude that increasing summer droughts may reduce ECMF diversity and the complementarity of P uptake by ECMF species, which will add to negative growth effects expected from nutrient imbalances under global change.

Phylogenetic and functional traits of ectomycorrhizal assemblages in top soil from different biogeographic regions and forest types.

Ectomycorrhizal (EM) fungal taxonomic, phylogenetic, and trait diversity (exploration types) were analyzed in beech and conifer forests along a north-to-south gradient in three biogeographic regions in Germany. The taxonomic community structures of the ectomycorrhizal assemblages in top soil were influenced by stand density and forest type, by biogeographic environmental factors (soil physical properties, temperature, and precipitation), and by nitrogen forms (amino acids, ammonium, and nitrate). While α-diversity did not differ between forest types, ß-diversity increased, leading to higher γ-diversity on the landscape level when both forest types were present. The highest taxonomic diversity of EM was found in forests in cool, moist climate on clay and silty soils and the lowest in the forests in warm, dry climate on sandy soils. In the region with higher taxonomic diversity, phylogenetic clustering was found, but not trait clustering. In the warm region, trait clustering occurred despite neutral phylogenetic effects. These results suggest that different forest types and favorable environmental conditions in forests promote high EM species richness in top soil presumably with both high functional diversity and phylogenetic redundancy, while stressful environmental conditions lead to lower species richness and functional redundancy.

Interspecific temporal and spatial differences in the acquisition of litter-derived nitrogen by ectomycorrhizal fungal assemblages.

The spatiotemporal dynamics of, and interspecific differences in, the acquisition of litter-derived nitrogen (N) by natural assemblages of ectomycorrhizal root tips are poorly understood. Small cylindrical mesh bags containing (15)N-labelled beech (Fagus sylvatica) leaf litter that permit hyphal but not root ingrowth were inserted vertically into the top soil layer of an old-growth beech forest. The lateral transfer of (15)N into the circumjacent soil, roots, microbes and ectomycorrhizas was measured during an 18-month exposure period. Ectomycorrhial fungi (EMF) showed large interspecific variation in the temporal pattern and extent of (15)N accumulation. Initially, when N was mainly available from the leachate, microbes were more efficient at N immobilization than the majority of EMF, but distinct fungal species also showed significant (15)N accumulation. During later phases, the enrichment of (15)N in Tomentella badia was higher than in microbes and other EMF species. Roots and soil accumulated (15)N with a large delay compared with microbes and EMF. Because approximately half of the studied fungal species had direct access to N from leaf litter and the remainder to N from leached compounds, we suggest that EMF diversity facilitates the N utilization of the host by capturing N originating from early-released solutes and late degradation products from a recalcitrant source.

An improved method for extraction of soil fungal mycelium.

Fungal mycelium is a major component of the soil microbiome. The soil hyphosphere represents a complex and dynamic niche for specific microorganisms, where multitrophic interactions occur, affecting ecosystem processes. However, extracting fungal mycelium from the soil to enable its taxonomical, chemical, and structural characterisation is challenging in the absence of a fast, efficient, and low-cost procedure. In this study, an old method (Bingle and Paul 1985), based on successive soil wet filtrations and density gradient centrifugation, was improved and tested in three different soil types (silty clay, silty clay loam, and loamy sand). The improved method reduced the number of filtrations by about five times and the centrifugation time from 40 min to 1 min. It avoided using any chemical substance which may impair further chemical analyses or DNA isolation and amplification. The method efficiency was about 50 % in the clay and 23 % in the sandy soils. However, a pre-step consisting of removing the fine-root fragments and other debris under the stereomicroscope may increase the method efficiency to more than 65 %, independent of the soil type.•A simple, efficient, and low-cost method suitable for extracting soil mycelium from a large number of samples.•The protocol includes successive soil wet filtrations and sucrose gradient centrifugation.•The method efficiency increases if the fine-root fragments and other debris are previously removed from the soil.

Mycorrhizal C/N ratio determines plant-derived carbon and nitrogen allocation to symbiosis.

Carbon allocation of trees to ectomycorrhizas is thought to shape forest nutrient cycling, but the sink activities of different fungal taxa for host resources are unknown. Here, we investigate fungal taxon-specific differences in naturally composed ectomycorrhizal (EM) communities for plant-derived carbon and nitrogen. After aboveground dual labeling of young beech with 15N and 13C, ectomycorrhizas formed with different fungal taxa exhibit strong differences in label enrichment. Secondary Ion Mass Spectrometry (SIMS) imaging of nitrogen in cross sections of ectomycorrhizas demonstrates plant-derived 15N in both root and fungal structures. Isotope enrichment in ectomycorrhizas correlates with that in the corresponding ectomycorrhiza-attached lateral root, supporting fungal taxon-specific N and C fluxes in ectomycorrhizas. The enrichments with 13C and 15N in the symbiosis decrease with increasing C/N ratio of ectomycorrhizas, converging to zero at high C/N. The relative abundances of EM fungal species on roots are positively correlated with 13C enrichment, demonstrating higher fitness of stronger than of less C-demanding symbioses. Overall, our results support that differences among the C/N ratios in ectomycorrhizas formed with different fungal species regulate the supply of the symbioses with host-derived carbon and provide insights on functional traits of ectomycorrhizas, which are important for major ecosystem processes.

Contrasting distribution of enzyme activities in the rhizosphere of European beech and Norway spruce.

Recent policies and silvicultural management call for forest regeneration that involve the selection of tree species able to cope with low soil nutrient availability in forest ecosystems. Understanding the impact of different tree species on the rhizosphere processes (e.g., enzyme activities) involved in nutrient mobilisation is critical in selecting suitable species to adapt forests to environmental change. Here, we visualised and investigated the rhizosphere distribution of enzyme activities (cellobiohydrolase, leucine-aminopeptidase, and acid phosphomonoesterase) using zymography. We related the distribution of enzyme activities to the seedling root morphological traits of European beech (Fagus sylvatica) and Norway spruce (Picea abies), the two most cultivated temperate tree species that employ contrasting strategies in soil nutrient acquisition. We found that spruce showed a higher morphological heterogeneity along the roots than beech, resulting in a more robust relationship between rhizoplane-associated enzyme activities and the longitudinal distance from the root apex. The rhizoplane enzyme activities decreased in spruce and increased in beech with the distance from the root apex over a power-law equation. Spruce revealed broader rhizosphere extents of all three enzymes, but only acid phosphomonoesterase activity was higher compared with beech. This latter result was determined by a larger root system found in beech compared with spruce that enhanced cellobiohydrolase and leucine-aminopeptidase activities. The root hair zone and hair lengths were significant variables determining the distribution of enzyme activities in the rhizosphere. Our findings indicate that spruce has a more substantial influence on rhizosphere enzyme production and diffusion than beech, enabling spruce to better mobilise nutrients from organic sources in heterogeneous forest soils.

Long-Term Nitrogen Deposition Alters Ectomycorrhizal Community Composition and Function in a Poplar Plantation.

The continuous upsurge in soil nitrogen (N) enrichment has had strong impacts on the structure and function of ecosystems. Elucidating how plant ectomycorrhizal fungi (EMF) mutualists respond to this additional N will facilitate the rapid development and implementation of more broadly applicable management and remediation strategies. For this study, we investigated the responses of EMF communities to increased N, and how other abiotic environmental factors impacted them. Consequently, we conducted an eight-year N addition experiment in a poplar plantation in coastal eastern China that included five N addition levels: 0 (N0), 50 (N1), 100 (N2), 150 (N3), and 300 (N4) kg N ha-1 yr-1. We observed that excessive N inputs reduced the colonization rate and species richness of EMF, and altered its community structure and functional traits. The total carbon content of the humus layer and available phosphorus in the mineral soil were important drivers of EMF abundance, while the content of ammonium in the humus layer and mineral soil determined the variations in the EMF community structure and mycelium foraging type. Our findings indicated that long-term N addition induced soil nutrient imbalances that resulted in a severe decline in EMF abundance and loss of functional diversity in poplar plantations.

Girdling affects ectomycorrhizal fungal (EMF) diversity and reveals functional differences in EMF community composition in a beech forest.

The relationships between plant carbon resources, soil carbon and nitrogen content, and ectomycorrhizal fungal (EMF) diversity in a monospecific, old-growth beech (Fagus sylvatica) forest were investigated by manipulating carbon flux by girdling. We hypothesized that disruption of the carbon supply would not affect diversity and EMF species numbers if EM fungi can be supplied by plant internal carbohydrate resources or would result in selective disappearance of EMF taxa because of differences in carbon demand of different fungi. Tree carbohydrate status, root demography, EMF colonization, and EMF taxon abundance were measured repeatedly during 1 year after girdling. Girdling did not affect root colonization but decreased EMF species richness of an estimated 79 to 90 taxa to about 40 taxa. Cenococcum geophilum, Lactarius blennius, and Tomentella lapida were dominant, colonizing about 70% of the root tips, and remained unaffected by girdling. Mainly cryptic EMF species disappeared. Therefore, the Shannon-Wiener index (H') decreased but evenness was unaffected. H' was positively correlated with glucose, fructose, and starch concentrations of fine roots and also with the ratio of dissolved organic carbon to dissolved organic nitrogen (DOC/DON), suggesting that both H' and DOC/DON were governed by changes in belowground carbon allocation. Our results suggest that beech maintains numerous rare EMF species by recent photosynthate. These EM fungi may constitute biological insurance for adaptation to changing environmental conditions. The preservation of taxa previously not known to colonize beech may, thus, form an important reservoir for future forest development.

Differences in Root Nitrogen Uptake Between Tropical Lowland Rainforests and Oil Palm Plantations.

Conversion of lowland tropical rainforests to intensely fertilized agricultural land-use systems such as oil palm (Elaeis guineensis) plantations leads to changes in nitrogen (N) cycling. Although soil microbial-driven N dynamics has been largely studied, the role of the plant as a major component in N uptake has rarely been considered. We address this gap by comparing the root N contents and uptake in lowland rainforests with that in oil palm plantations on Sumatra, Indonesia. To this aim, we applied 15N-labeled ammonium to intact soil, measured the 15N recovery in soil and roots, and calculated the root relative N uptake efficiency for 10 days after label application. We found that root N contents were by one third higher in the rainforest than oil palm plantations. However, 15N uptake efficiency was similar in the two systems. This finding suggests that lower N contents in oil palm roots were likely caused by plant internal utilization of the absorbed N (e.g., N export to fruit bunches) than by lower ability to take up N from the soil. 15N recovery in roots was primarily driven by the amount of root biomass, which was higher in oil palm plantation than rainforest. The oil palms unveiled a high capacity to acquire N, offering the possibility of enhancing sustainable plantation management by reducing N fertilizer application.

Early stage root-Associated fungi show a high temporal turnover, but Are independent of beech progeny.

The relationship between trees and root-associated fungal communities is complex. By specific root deposits and other signal cues, different tree species are able to attract divergent sets of fungal species. Plant intraspecific differences can lead to variable fungal patterns in the root's proximity. Therefore, within the Beech Transplant Experiment, we analyzed the impact of three different European beech ecotypes on the fungal communities in roots and the surrounding rhizosphere soil at two time points. Beech nuts were collected in three German sites in 2011. After one year, seedlings of the different progenies were out-planted on one site and eventually re-sampled in 2014 and 2017. We applied high-throughput sequencing of the fungal ITS2 to determine the correlation between tree progeny, a possible home-field advantage, plant development and root-associated fungal guilds under field conditions. Our result showed no effect of beech progeny on either fungal OTU richness or fungal community structure. However, over time the fungal OTU richness in roots increased and the fungal communities changed significantly, also in rhizosphere. In both plant compartments, the fungal communities displayed a high temporal turnover, indicating a permanent development and functional adaption of the root mycobiome of young beeches.

Deprivation of root-derived resources affects microbial biomass but not community structure in litter and soil.

The input of plant leaf litter has been assumed to be the most important resource for soil organisms of forest ecosystems, but there is increasing evidence that root-derived resources may be more important. By trenching roots of trees in deciduous and coniferous forests, we cut-off the input of root-derived resources and investigated the response of microorganisms using substrate-induced respiration and phospholipid fatty acid (PLFA) analysis. After one and three years, root trenching strongly decreased microbial biomass and concentrations of PLFAs by about 20%, but the microbial community structure was little affected and the effects were similar in deciduous and coniferous forests. However, the reduction in microbial biomass varied between regions and was more pronounced in forests on limestone soils (Hainich) than in those on sandy soils (Schorfheide). Trenching also reduced microbial biomass in the litter layer but only in the Hainich after one year, whereas fungal and bacterial marker PLFAs as well as the fungal-to-plant marker ratio in litter were reduced in the Schorfheide both after one and three years. The pronounced differences between forests of the two regions suggest that root-derived resources are more important in fueling soil microorganisms of base-rich forests characterized by mull humus than in forests poor in base cations characterized by moder soils. The reduction in microbial biomass and changes in microbial community characteristics in the litter layer suggests that litter microorganisms do not exclusively rely on resources from decomposing litter but also from roots, i.e. from resources based on labile recently fixed carbon. Our results suggest that both bacteria and fungi heavily depend on root-derived resources with both suffering to a similar extent to deprivation of these resources. Further, the results indicate that the community structure of microorganisms is remarkably resistant to changes in resource supply and adapts quickly to new conditions irrespective of tree species composition and forest management.

Multiple forest attributes underpin the supply of multiple ecosystem services.

Trade-offs and synergies in the supply of forest ecosystem services are common but the drivers of these relationships are poorly understood. To guide management that seeks to promote multiple services, we investigated the relationships between 12 stand-level forest attributes, including structure, composition, heterogeneity and plant diversity, plus 4 environmental factors, and proxies for 14 ecosystem services in 150 temperate forest plots. Our results show that forest attributes are the best predictors of most ecosystem services and are also good predictors of several synergies and trade-offs between services. Environmental factors also play an important role, mostly in combination with forest attributes. Our study suggests that managing forests to increase structural heterogeneity, maintain large trees, and canopy gaps would promote the supply of multiple ecosystem services. These results highlight the potential for forest management to encourage multifunctional forests and suggest that a coordinated landscape-scale strategy could help to mitigate trade-offs in human-dominated landscapes.

Divergent habitat filtering of root and soil fungal communities in temperate beech forests.

Distance decay, the general reduction in similarity of community composition with increasing geographical distance, is known as predictor of spatial variation and distribution patterns of organisms. However, changes in fungal communities along environmental gradients are little known. Here we show that distance decays of soil-inhabiting and root-associated fungal assemblages differ, and identify explanatory environmental variables. High-throughput sequencing analysis of fungal communities of beech-dominated forests at three study sites across Germany shows that root-associated fungi are recruited from the soil fungal community. However, distance decay is substantially weaker in the root-associated than in the soil community. Variance partitioning of factors contributing to the observed distance decay patterns support the hypothesis that host trees stabilize the composition of root-associated fungi communities, relative to soil communities. Thus, they not only have selective impacts on associated communities, but also buffer effects of changes in microclimatic and environmental variables that directly influence fungal community composition.