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Mycena s.s. is a ubiquitous mushroom genus whose members degrade multiple dead plant substrates and opportunistically invade living plant roots. Having sequenced the nuclear genomes of 24 Mycena species, we find them to defy the expected patterns for fungi based on both their traditionally perceived saprotrophic ecology and substrate specializations. Mycena displayed massive genome expansions overall affecting all gene families, driven by novel gene family emergence, gene duplications, enlarged secretomes encoding polysaccharide degradation enzymes, transposable element (TE) proliferation, and horizontal gene transfers. Mainly due to TE proliferation, Arctic Mycena species display genomes of up to 502 Mbp (2-8× the temperate Mycena), the largest among mushroom-forming Agaricomycetes, indicating a possible evolutionary convergence to genomic expansions sometimes seen in Arctic plants. Overall, Mycena show highly unusual, varied mosaic-like genomic structures adaptable to multiple lifestyles, providing genomic illustration for the growing realization that fungal niche adaptations can be far more fluid than traditionally believed.

Despite host-fungal symbiotic interactions being ubiquitous in all ecosystems, understanding how symbiosis has shaped the ecology and evolution of fungal spores that are involved in dispersal and colonization of their hosts has been ignored in life-history studies. We assembled a spore morphology database covering over 26,000 species of free-living to symbiotic fungi of plants, insects and humans and found more than eight orders of variation in spore size. Evolutionary transitions in symbiotic status correlated with shifts in spore size, but the strength of this effect varied widely among phyla. Symbiotic status explained more variation than climatic variables in the current distribution of spore sizes of plant-associated fungi at a global scale while the dispersal potential of their spores is more restricted compared to free-living fungi. Our work advances life-history theory by highlighting how the interaction between symbiosis and offspring morphology shapes the reproductive and dispersal strategies among living forms.

Traditional strict separation of fungi into ecological niches as mutualist, parasite or saprotroph is increasingly called into question. Sequences of assumed saprotrophs have been amplified from plant root interiors, and several saprotrophic genera can invade and interact with host plants in laboratory growth experiments. However, it is uncertain if root invasion by saprotrophic fungi is a widespread phenomenon and if laboratory interactions mirror field conditions. Here, we focused on the widespread and speciose saprotrophic genus Mycena and performed (1) a systematic survey of their occurrences (in ITS1/ITS2 datasets) in mycorrhizal roots of 10 plant species, and (2) an analysis of natural abundances of 13C/15N stable isotope signatures of Mycena basidiocarps from five field locations to examine their trophic status. We found that Mycena was the only saprotrophic genus consistently found in 9 out of 10 plant host roots, with no indication that the host roots were senescent or otherwise vulnerable. Furthermore, Mycena basidiocarps displayed isotopic signatures consistent with published 13C/15N profiles of both saprotrophic and mutualistic lifestyles, supporting earlier laboratory-based studies. We argue that Mycena are widespread latent invaders of healthy plant roots and that Mycena species may form a spectrum of interactions besides saprotrophy also in the field.

Despite host-fungal symbiotic interactions being ubiquitous in all ecosystems, understanding how symbiosis has shaped the ecology and evolution of fungal spores that are involved in dispersal and colonization of their hosts has been ignored in life-history studies. We assembled a spore morphology database covering over 26,000 species of free-living to symbiotic fungi of plants, insects and humans and found more than eight orders of variation in spore size. Evolutionary transitions in symbiotic status correlated with shifts in spore size, but the strength of this effect varied widely among phyla. Symbiotic status explained more variation than climatic variables in the current distribution of spore sizes of plant-associated fungi at a global scale while the dispersal potential of their spores is more restricted compared to free-living fungi. Our work advances life-history theory by highlighting how the interaction between symbiosis and offspring morphology shapes the reproductive and dispersal strategies among living forms.

Traditional strict separation of fungi into ecological niches as mutualist, parasite or saprotroph is increasingly called into question. Sequences of assumed saprotrophs have been amplified from plant root interiors, and several saprotrophic genera can invade and interact with host plants in laboratory growth experiments. However, it is uncertain if root invasion by saprotrophic fungi is a widespread phenomenon and if laboratory interactions mirror field conditions. Here, we focused on the widespread and speciose saprotrophic genus Mycena and performed (1) a systematic survey of their occurrences (in ITS1/ITS2 datasets) in mycorrhizal roots of 10 plant species, and (2) an analysis of natural abundances of 13C/15N stable isotope signatures of Mycena basidiocarps from five field locations to examine their trophic status. We found that Mycena was the only saprotrophic genus consistently found in 9 out of 10 plant host roots, with no indication that the host roots were senescent or otherwise vulnerable. Furthermore, Mycena basidiocarps displayed isotopic signatures consistent with published 13C/15N profiles of both saprotrophic and mutualistic lifestyles, supporting earlier laboratory-based studies. We argue that Mycena are widespread latent invaders of healthy plant roots and that Mycena species may form a spectrum of interactions besides saprotrophy also in the field.

Despite host-fungal symbiotic interactions being ubiquitous in all ecosystems, understanding how symbiosis has shaped the ecology and evolution of fungal spores that are involved in dispersal and colonization of their hosts has been ignored in life-history studies. We assembled a spore morphology database covering over 26,000 species of free-living to symbiotic fungi of plants, insects and humans and found more than eight orders of variation in spore size. Evolutionary transitions in symbiotic status correlated with shifts in spore size, but the strength of this effect varied widely among phyla. Symbiotic status explained more variation than climatic variables in the current distribution of spore sizes of plant-associated fungi at a global scale while the dispersal potential of their spores is more restricted compared to free-living fungi. Our work advances life-history theory by highlighting how the interaction between symbiosis and offspring morphology shapes the reproductive and dispersal strategies among living forms.

Despite host-fungal symbiotic interactions being ubiquitous in all ecosystems, understanding how symbiosis has shaped the ecology and evolution of fungal spores that are involved in dispersal and colonization of their hosts has been ignored in life-history studies. We assembled a spore morphology database covering over 26,000 species of free-living to symbiotic fungi of plants, insects and humans and found more than eight orders of variation in spore size. Evolutionary transitions in symbiotic status correlated with shifts in spore size, but the strength of this effect varied widely among phyla. Symbiotic status explained more variation than climatic variables in the current distribution of spore sizes of plant-associated fungi at a global scale while the dispersal potential of their spores is more restricted compared to free-living fungi. Our work advances life-history theory by highlighting how the interaction between symbiosis and offspring morphology shapes the reproductive and dispersal strategies among living forms.

Traditional strict separation of fungi into ecological niches as mutualist, parasite or saprotroph is increasingly called into question. Sequences of assumed saprotrophs have been amplified from plant root interiors, and several saprotrophic genera can invade and interact with host plants in laboratory growth experiments. However, it is uncertain if root invasion by saprotrophic fungi is a widespread phenomenon and if laboratory interactions mirror field conditions. Here, we focused on the widespread and speciose saprotrophic genus Mycena and performed (1) a systematic survey of their occurrences (in ITS1/ITS2 datasets) in mycorrhizal roots of 10 plant species, and (2) an analysis of natural abundances of 13C/15N stable isotope signatures of Mycena basidiocarps from five field locations to examine their trophic status. We found that Mycena was the only saprotrophic genus consistently found in 9 out of 10 plant host roots, with no indication that the host roots were senescent or otherwise vulnerable. Furthermore, Mycena basidiocarps displayed isotopic signatures consistent with published 13C/15N profiles of both saprotrophic and mutualistic lifestyles, supporting earlier laboratory-based studies. We argue that Mycena are widespread latent invaders of healthy plant roots and that Mycena species may form a spectrum of interactions besides saprotrophy also in the field.

Many children spend considerable time in daycare centers and may be influenced by the indoor microorganisms there, including fungi. In this study, we investigate the indoor mycobiomes of 125 daycare centers distributed along strong environmental gradients throughout Norway. Dust samples were collected from doorframes outside and inside buildings using a community science sampling approach. Fungal communities in the dust samples were analyzed using DNA metabarcoding of the internal transcribed spacer 2 (ITS2) region. We observed a marked difference between the outdoor and indoor mycobiomes. The indoor mycobiomes included considerably more yeasts and molds than the outdoor samples, with Saccharomyces, Mucor, Malassezia, and Penicillium being among the most dominant fungal genera. Changes in the indoor fungal richness and composition correlated with numerous variables related to both outdoor and indoor conditions; there was a clear geographic structure in the indoor mycobiome composition that mirrored the outdoor climate, ranging from humid areas in western Norway to drier and colder areas in eastern Norway. Moreover, the number of children in the daycare centers, as well as various building features, influenced the indoor mycobiome composition. We conclude that the indoor mycobiomes in Norwegian daycare centers are structured by multiple factors and are dominated by yeasts and molds. This study exemplifies how community science sampling enables DNA-based analyses of a high number of samples covering wide geographic areas.

Molds thrive in indoor environments, challenging the stability of building materials and occupants’ health. Diverse sampling and analytical techniques can be applied in the microbiology of buildings, with specific benefits and drawbacks. We evaluated the use of two methods, the microscopy of visible mold growth (hereinafter “mold” samples) (tape lifts) and the DNA metabarcoding of mold and dust samples (swabs), for mapping mold-damage indicator fungi in residential buildings in Oslo. Overall, both methods provided consistent results for the mold samples, where nearly 80% of the microscopy-identified taxa were confirmed by DNA analyses. Aspergillus was the most abundant genus colonizing all materials, while some taxa were associated with certain substrates: Acremonium with gypsum board, Chaetomium with chipboard, Stachybotrys with gypsum board and wood, and Trichoderma with wood. Based on the DNA data, the community composition was clearly different between the mold and the dust, with a much higher alpha diversity in the dust. Most genera identified in the mold were also detected with a low abundance in the dust from the same apartments. Their spatial distribution indicated some local spread from the mold growth to other areas, but there was no clear correlation between the relative abundances and the distance to the damages. To study mold damages, different microbiological analyses (microscopy, cultivation, DNA, and chemistry) should be combined with a thorough inspection of buildings. The interpretation of such datasets requires the collaboration of skilled mycologists and building consultants.

Recent global warming affects species compositions at an unprecedented rate. To predict climate-induced changes in species assemblages, a better understanding of the link between species occurrence and climate is needed. Macrofungal fruit body assemblages are correlated with the thermal environment at the European scale. However, it is still unknown whether macrofungal communities are also phylogenetically structured by thermal environments. Thermal environments are characterized by annual temperature means but also by intra-annual temperature variability (hereafter termed temperature seasonality), which are both considered in this study. Here, we used distribution data of 2882 species based on fruit body records across Europe to address two main questions: 1) are mushroom assemblages at the extremes of the mean (warm and cold) and seasonal (high intra-annual variability, i.e. continental) climate gradient phylogenetically more similar than expected (phylogenetic alpha diversity); 2) are mushroom assemblages, that are subject to different mean and seasonal temperature conditions, composed of different lineages (phylogenetic beta diversity). Our phylogenetic alpha diversity analysis shows that mushroom assemblages are phylogenetically structured by warm and cold environments, indicating that phylogenetically related species with similar traits thrive under more extreme conditions. In contrast, assemblages are phylogenetically more dissimilar (overdispersed) in temperature seasonal environments, indicating limiting similarity. Phylogenetic beta diversity was significantly correlated with mean and seasonal temperature differences, a response mainly driven by a few genera. Our results show that macrofungal assemblages are phylogenetically structured by temperature across Europe, suggesting phylogenetically constrained specialization towards temperature extremes. Predicted anthropogenic warming is likely to affect species composition and phylogenetic diversity with ad

Recent global warming affects species compositions at an unprecedented rate. To predict climate-induced changes in species assemblages, a better understanding of the link between species occurrence and climate is needed. Macrofungal fruit body assemblages are correlated with the thermal environment at the European scale. However, it is still unknown whether macrofungal communities are also phylogenetically structured by thermal environments. Thermal environments are characterized by annual temperature means but also by intra-annual temperature variability (hereafter termed temperature seasonality), which are both considered in this study. Here, we used distribution data of 2882 species based on fruit body records across Europe to address two main questions: 1) are mushroom assemblages at the extremes of the mean (warm and cold) and seasonal (high intra-annual variability, i.e. continental) climate gradient phylogenetically more similar than expected (phylogenetic alpha diversity); 2) are mushroom assemblages, that are subject to different mean and seasonal temperature conditions, composed of different lineages (phylogenetic beta diversity). Our phylogenetic alpha diversity analysis shows that mushroom assemblages are phylogenetically structured by warm and cold environments, indicating that phylogenetically related species with similar traits thrive under more extreme conditions. In contrast, assemblages are phylogenetically more dissimilar (overdispersed) in temperature seasonal environments, indicating limiting similarity. Phylogenetic beta diversity was significantly correlated with mean and seasonal temperature differences, a response mainly driven by a few genera. Our results show that macrofungal assemblages are phylogenetically structured by temperature across Europe, suggesting phylogenetically constrained specialization towards temperature extremes. Predicted anthropogenic warming is likely to affect species composition and phylogenetic diversity with addit

Recent global warming affects species compositions at an unprecedented rate. To predict climate-induced changes in species assemblages, a better understanding of the link between species occurrence and climate is needed. Macrofungal fruit body assemblages are correlated with the thermal environment at the European scale. However, it is still unknown whether macrofungal communities are also phylogenetically structured by thermal environments. Thermal environments are characterized by annual temperature means but also by intra-annual temperature variability (hereafter termed temperature seasonality), which are both considered in this study. Here, we used distribution data of 2,882 species based on fruit body records across Europe to address two main questions: (i) Are mushroom assemblages at the extremes of the mean (warm and cold) and seasonal (high intra-annual variability, i.e. continental) climate gradient phylogenetically more similar than expected (phylogenetic alpha diversity); (ii) are mushroom assemblages, that are subject to different mean and seasonal temperature conditions, composed of different lineages (phylogenetic beta diversity). Our phylogenetic alpha diversity analysis shows that mushroom assemblages are phylogenetically structured by warm and cold environments, indicating that phylogenetically related species with similar traits thrive under more extreme conditions. In contrast, assemblages are phylogenetically more dissimilar (overdispersed) in temperature seasonal environments, indicating limiting similarity. Phylogenetic beta diversity was significantly correlated with mean and seasonal temperature differences, a response mainly driven by a few genera. Our results show that macrofungal assemblages are phylogenetically structured by temperature across Europe, suggesting phylogenetically constrained specialization towards temperature extremes. Predicted anthropogenic warming is likely to affect species composition and phylogenetic diversity with a, Recent global warming affects species compositions at an unprecedented rate. To predict climate-induced changes in species assemblages, a better understanding of the link between species occurrence and climate is needed. Macrofungal fruit body assemblages are correlated with the thermal environment at the European scale. However, it is still unknown whether macrofungal communities are also phylogenetically structured by thermal environments. Thermal environments are characterized by annual temperature means but also by intra-annual temperature variability (hereafter termed temperature seasonality), which are both considered in this study. Here, we used distribution data of 2,882 species based on fruit body records across Europe to address two main questions: (i) Are mushroom assemblages at the extremes of the mean (warm and cold) and seasonal (high intra-annual variability, i.e. continental) climate gradient phylogenetically more similar than expected (phylogenetic alpha diversity); (ii) are mushroom assemblages, that are subject to different mean and seasonal temperature conditions, composed of different lineages (phylogenetic beta diversity). Our phylogenetic alpha diversity analysis shows that mushroom assemblages are phylogenetically structured by warm and cold environments, indicating that phylogenetically related species with similar traits thrive under more extreme conditions. In contrast, assemblages are phylogenetically more dissimilar (overdispersed) in temperature seasonal environments, indicating limiting similarity. Phylogenetic beta diversity was significantly correlated with mean and seasonal temperature differences, a response mainly driven by a few genera. Our results show that macrofungal assemblages are phylogenetically structured by temperature across Europe, suggesting phylogenetically constrained specialization towards temperature extremes. Predicted anthropogenic warming is likely to affect species composition and phylogenetic diversity with a

Mycorrhiza is a beneficial association between plant roots and fungi. This mutualistic symbiosis is essential for plant growth in most natural terrestrial ecosystems and in agriculture. Commercial mycorrhizal products containing fungi and bacteria may promote plant growth, especially on sites without a natural microbial community. Due to the risk of unintended negative effects, introduction of new species or genetically different isolates of native species should always be considered carefully. This report assesses the risk of establishment and spread of six fungal species and six bacterial species included in different commercial mycorrhizal products, as well as the species’ potential impact on Norwegian biodiversity. Most of the evaluated fungi and bacteria are probably present in Norway, even though presence at present data only exist for two of the six fungal species. Establishment of the assessed fungi on the plants and sites where they are applied is considered moderately likely, with medium uncertainty, while establishment of the bacterial species is considered to range from very unlikely to very likely depending on the bacterial group, with low uncertainty. The probability of spread to the wider environment ranges from unlikely (four fungal species), to moderately likely (two fungal species), to very likely (five of the six bacterial species). However, for all species it is considered unlikely that establishment and spread would have negative effects on other native species, habitats and ecosystems in Norway.

Mycorrhiza is a beneficial association between plant roots and fungi. This mutualistic symbiosis is essential for plant growth in most natural terrestrial ecosystems and in agriculture. Commercial mycorrhizal products containing fungi and bacteria may promote plant growth, especially on sites without a natural microbial community. Due to the risk of unintended negative effects, introduction of new species or genetically different isolates of native species should always be considered carefully. This report assesses the risk of establishment and spread of six fungal species and six bacterial species included in different commercial mycorrhizal products, as well as the species’ potential impact on Norwegian biodiversity. Most of the evaluated fungi and bacteria are probably present in Norway, even though presence at present data only exist for two of the six fungal species. Establishment of the assessed fungi on the plants and sites where they are applied is considered moderately likely, with medium uncertainty, while establishment of the bacterial species is considered to range from very unlikely to very likely depending on the bacterial group, with low uncertainty. The probability of spread to the wider environment ranges from unlikely (four fungal species), to moderately likely (two fungal species), to very likely (five of the six bacterial species). However, for all species it is considered unlikely that establishment and spread would have negative effects on other native species, habitats and ecosystems in Norway.

Mycorrhiza is a beneficial association between plant roots and fungi. This mutualistic symbiosis is essential for plant growth in most natural terrestrial ecosystems and in agriculture. Commercial mycorrhizal products containing fungi and bacteria may promote plant growth, especially on sites without a natural microbial community. Due to the risk of unintended negative effects, introduction of new species or genetically different isolates of native species should always be considered carefully. This report assesses the risk of establishment and spread of six fungal species and six bacterial species included in different commercial mycorrhizal products, as well as the species’ potential impact on Norwegian biodiversity. Most of the evaluated fungi and bacteria are probably present in Norway, even though presence at present data only exist for two of the six fungal species. Establishment of the assessed fungi on the plants and sites where they are applied is considered moderately likely, with medium uncertainty, while establishment of the bacterial species is considered to range from very unlikely to very likely depending on the bacterial group, with low uncertainty. The probability of spread to the wider environment ranges from unlikely (four fungal species), to moderately likely (two fungal species), to very likely (five of the six bacterial species). However, for all species it is considered unlikely that establishment and spread would have negative effects on other native species, habitats and ecosystems in Norway.

Thermal melanism theory states that dark-colored ectotherm organisms are at an advantage at low temperature due to increased warming. This theory is generally supported for ectotherm animals, however, the function of colors in the fungal kingdom is largely unknown. Here, we test whether the color lightness of mushroom assemblages is related to climate using a dataset of 3.2 million observations of 3,054 species across Europe. Consistent with the thermal melanism theory, mushroom assemblages are significantly darker in areas with cold climates. We further show differences in color phenotype between fungal lifestyles and a lifestyle differentiated response to seasonality. These results indicate a more complex ecological role of mushroom colors and suggest functions beyond thermal adaption. Because fungi play a crucial role in terrestrial carbon and nutrient cycles, understanding the links between the thermal environment, functional coloration and species’ geographical distributions will be critical in predicting ecosystem responses to global warming.

Premise of the Study: Fungal diversity (richness) trends at large scales are in urgent need of investigation, especially through novel situations that combine long-term observational with environmental and remotely sensed open-source data. Methods: We modeled fungal richness, with collections-based records of saprotrophic (decaying) and ectomycorrhizal (plant mutualistic) fungi, using an array of environmental variables across geographical gradients from northern to central Europe. Temporal differences in covariables granted insight into the impacts of the shorter- versus longer-term environment on fungal richness. Results: Fungal richness varied significantly across different land-use types, with highest richness in forests and lowest in urban areas. Latitudinal trends supported a unimodal pattern in diversity across Europe. Temperature, both annual mean and range, was positively correlated with richness, indicating the importance of seasonality in increasing richness amounts. Precipitation seasonality notably affected saprotrophic fungal diversity (a unimodal relationship), as did daily precipitation of the collection day (negatively correlated). Ectomycorrhizal fungal richness differed from that of saprotrophs by being positively associated with tree species richness. Discussion: Our results demonstrate that fungal richness is strongly correlated with land use and climate conditions, especially concerning seasonality, and that ongoing global change processes will affect fungal richness patterns at large scales.

The home-field advantage hypothesis (HFA) predicts that plant litter decomposes faster than expected underneath the plant from which it originates. We tested this hypothesis in a decomposition experiment where litters were incubated reciprocally in neighbouring European beech and Norway spruce forests. We analysed fungal communities in the litter through DNA metabarcoding and evaluated the effect of mesofauna (mites and springtails) on litter mass loss by using different litter-bag mesh sizes. Accounting for general differences in decomposition between litter and forest types, we found a significant home field advantage of 24%. Litter decomposed faster in the beech forest but spruce litter decomposed faster than beech litter. Fungal communities showed a clear dependency on both forest and litter type. Mesofauna did not affect litter mass loss rates or microbial species composition.

Insufficient reference database coverage is a widely recognized limitation of molecular ecology approaches which are reliant on database matches for assignment of function or identity. Here, we use data from 65 amplicon high-throughput sequencing (HIS) datasets targeting the internal transcribed spacer (ITS) region of fungal rDNA to identify substrates and geographic areas whose underrepresentation in the available reference databases could have meaningful impact on our ability to draw ecological conclusions. A total of 14 different substrates were investigated. Database representation was particularly poor for the fungal communities found in aquatic (freshwater and marine) and soil ecosystems. Aquatic ecosystems are identified as priority targets for the recovery of novel fungal lineages. A subset of the data representing soil samples with global distribution were used to identify geographic locations and terrestrial biomes with poor database representation. Database coverage was especially poor in tropical, subtropical, and Antarctic latitudes, and the Amazon, Southeast Asia, Australasia, and the Indian subcontinent are identified as priority areas for improving database coverage in fungi. (C) 2018 Elsevier Ltd and British Mycological Society.

Insufficient reference database coverage is a widely recognized limitation of molecular ecology approaches which are reliant on database matches for assignment of function or identity. Here, we use data from 65 amplicon high-throughput sequencing (HIS) datasets targeting the internal transcribed spacer (ITS) region of fungal rDNA to identify substrates and geographic areas whose underrepresentation in the available reference databases could have meaningful impact on our ability to draw ecological conclusions. A total of 14 different substrates were investigated. Database representation was particularly poor for the fungal communities found in aquatic (freshwater and marine) and soil ecosystems. Aquatic ecosystems are identified as priority targets for the recovery of novel fungal lineages. A subset of the data representing soil samples with global distribution were used to identify geographic locations and terrestrial biomes with poor database representation. Database coverage was especially poor in tropical, subtropical, and Antarctic latitudes, and the Amazon, Southeast Asia, Australasia, and the Indian subcontinent are identified as priority areas for improving database coverage in fungi. (C) 2018 Elsevier Ltd and British Mycological Society.

The home-field advantage hypothesis (HFA) predicts that plant litter decomposes faster than expected underneath the plant from which it originates. We tested this hypothesis in a decomposition experiment where litters were incubated reciprocally in neighbouring European beech and Norway spruce forests. We analysed fungal communities in the litter through DNA metabarcoding and evaluated the effect of mesofauna (mites and springtails) on litter mass loss by using different litter-bag mesh sizes. Accounting for general differences in decomposition between litter and forest types, we found a significant home field advantage of 24%. Litter decomposed faster in the beech forest but spruce litter decomposed faster than beech litter. Fungal communities showed a clear dependency on both forest and litter type. Mesofauna did not affect litter mass loss rates or microbial species composition.

Aim: Macroecological scales of species compositional trends are well documented for a variety of plant and animal groups, but remain sparse for fungi, despite their ecological importance in carbon and nutrient cycling. It is, thus, essential to understand the composition of fungal assemblages across broad geographical scales and the underlying drivers. Our overall aim was to describe these patterns for fungi across two nutritional modes (saprotrophic and ectomycorrhizal). Furthermore, we aimed to elucidate the temporal component of fruiting patterns and to relate these to soil carbon and nitrogen deposition. Location: Central and Northern Europe. Methods: A total of 4.9 million fungal fruit body observations throughout Europe, collected between 1970 and 2010, were analysed to determine the two main environmental and geographical gradients structuring fungal assemblages for two main nutritional modes, saprotrophic and ectomycorrhizal fungi. Results: Two main gradients explaining the geography of compositional patterns were identified, for each nutritional mode. Mean annual temperature (and related collinear, seasonal measures) correlated most strongly with the first gradient for both nutritional modes. Soil organic carbon was the highest correlate of the second compositional gradient for ectomycorrhizal fungi, suspected as an indicator of vegetation- and pH-related covariates. In contrast, nitrogen deposition constituted a second gradient for saprotrophic fungi, likely a proxy for anthropogenic pollution. Compositional gradients and environmental conditions correlated similarly when the data were divided into two time intervals of 1970–1990 and 1991–2010. Evidence of compositional temporal change was highest with increasing elevation and latitude. Main conclusions: Fungal assemblage patterns demonstrate clear biogeographical patterns that relate the nutritional modes to their main environmental correlates of temperature, soil organic carbon and nitrogen deposition. W

Here we assess the impact of geographically dependent (latitude, longitude and altitude) changes in bioclimatic (temperature, precipitation and primary productivity) variability on fungal fruiting phenology across Europe. Two main nutritional guilds of fungi, saprotrophic and ectomycorrhizal, were further separated into spring and autumn fruiters. We used a path‐analysis to investigate how biogeographic patterns in fungal fruiting phenology coincided with seasonal changes in climate and primary production. Across central to northern Europe, mean fruiting varied by approximately 25 days, primarily with latitude. Altitude affected fruiting by up to 30 days, with spring delays and autumnal accelerations. Fruiting was as much explained by the effects of bioclimatic variability as by their large‐scale spatial patterns. Temperature drove fruiting of autumnal ectomycorrhizal and saprotrophic, as well as spring saprotrophic groups, while primary production and precipitation were major drivers for spring‐fruiting ectomycorrhizal fungi. Species‐specific phenology predictors were not stable, instead deviating from the overall mean. There is significant likelihood that further climatic change, especially in temperature, will impact fungal phenology patterns at large spatial scales. The ecological implications are diverse, potentially affecting food webs (asynchrony), nutrient cycling and the timing of nutrient availability in ecosystems.

Despite their obvious importance, our knowledge about the eukaryotic microbial diversity of inland waters is still limited and poorly documented. We applied 18S rDNA amplicon sequencing to provide a comprehensive analysis of eukaryotic diversity in 74 low-productivity lakes along a 750 km longitudinal transect (5.40-18.52°E) across southern Scandinavia. We detected a wide diversity of pelagic microbial eukaryotes, classified into 1882 operational taxonomic units (OTUs). The highest OTU richness was found in traditional phytoplankton groups like Dinoflagellata, Chrysophyceae, Chlorophyta and Cryptophyta. A total of 53.6% OTUs were primarily autotrophic, while 19.4% of the heterotrophic OTUs belonged to putative parasitic taxa. Except for a longitudinal trend in the relative influence of mixotrophs, there were no significant associations between major functional groups (autotrophs, heterotrophs and parasites) and spatial and environmental variables. Community dissimilarity increased significantly with increasing geographical distance between lakes. In accordance with earlier, microscopy-based surveys in this region, we demonstrate distinct gradients in protistan diversity and community composition, which are better explained by spatial structure than local environment. The strong association between longitude and protistan diversity is probably better explained by differences in regional species pools due to differences in landscape productivity than by dispersal limitation or climatic constraints

Changing climate is expected to alter precipitation patterns in the Arctic, with consequences for subsurface temperature and moisture conditions, community structure, and nutrient mobilization through microbial belowground processes. Here, we address the effect of increased snow depth on the variation in species richness and community structure of ectomycorrhizal (ECM) and saprotrophic fungi. Soil samples were collected weekly from mid-July to mid-September in both control and deep snow plots. Richness of ECM fungi was higher, while saprotrophic fungi was lower in increased snow depth plots relative to controls. ECM fungal richness was related to soil NO3-N, NH4-N, and K; and saprotrophic fungi to NO3-N and pH. Small but significant changes in the composition of saprotrophic fungi could be attributed to snow treatment and sampling time, but not so for the ECM fungi. Delayed snow melt did not influence the temporal variation in fungal communities between the treatments. Results suggest that some fungal species are favored, while others are disfavored resulting in their local extinction due to long-term changes in snow amount. Shifts in species composition of fungal functional groups are likely to affect nutrient cycling, ecosystem respiration, and stored permafrost carbon.

Changing climate is expected to alter precipitation patterns in the Arctic, with consequences for subsurface temperature and moisture conditions, community structure, and nutrient mobilization through microbial belowground processes. Here, we address the effect of increased snow depth on the variation in species richness and community structure of ectomycorrhizal (ECM) and saprotrophic fungi. Soil samples were collected weekly from mid-July to mid-September in both control and deep snow plots. Richness of ECM fungi was higher, while saprotrophic fungi was lower in increased snow depth plots relative to controls. ECM fungal richness was related to soil NO3-N, NH4-N, and K; and saprotrophic fungi to NO3-N and pH. Small but significant changes in the composition of saprotrophic fungi could be attributed to snow treatment and sampling time, but not so for the ECM fungi. Delayed snow melt did not influence the temporal variation in fungal communities between the treatments. Results suggest that some fungal species are favored, while others are disfavored resulting in their local extinction due to long-term changes in snow amount. Shifts in species composition of fungal functional groups are likely to affect nutrient cycling, ecosystem respiration, and stored permafrost carbon.

Changing climate is expected to alter precipitation patterns in the Arctic, with consequences for subsurface temperature and moisture conditions, community structure, and nutrient mobilization through microbial belowground processes. Here, we address the effect of increased snow depth on the variation in species richness and community structure of ectomycorrhizal (ECM) and saprotrophic fungi. Soil samples were collected weekly from mid-July to mid-September in both control and deep snow plots. Richness of ECM fungi was higher, while saprotrophic fungi was lower in increased snow depth plots relative to controls. ECM fungal richness was related to soil NO3-N, NH4-N, and K; and saprotrophic fungi to NO3-N and pH. Small but significant changes in the composition of saprotrophic fungi could be attributed to snow treatment and sampling time, but not so for the ECM fungi. Delayed snow melt did not influence the temporal variation in fungal communities between the treatments. Results suggest that some fungal species are favored, while others are disfavored resulting in their local extinction due to long-term changes in snow amount. Shifts in species composition of fungal functional groups are likely to affect nutrient cycling, ecosystem respiration, and stored permafrost carbon.

Changing climate is expected to alter precipitation patterns in the Arctic, with consequences for subsurface temperature and moisture conditions, community structure, and nutrient mobilization through microbial belowground processes. Here, we address the effect of increased snow depth on the variation in species richness and community structure of ectomycorrhizal (ECM) and saprotrophic fungi. Soil samples were collected weekly from mid-July to mid-September in both control and deep snow plots. Richness of ECM fungi was lower, while saprotrophic fungi was higher in increased snow depth plots relative to controls. [Correction added on 23 September 2016 after first online publication: In the preceding sentence, the richness of ECM and saprotrophic fungi were wrongly interchanged and have been fixed in this current version.] ECM fungal richness was related to soil NO3-N, NH4-N, and K; and saprotrophic fungi to NO3-N and pH. Small but significant changes in the composition of saprotrophic fungi could be attributed to snow treatment and sampling time, but not so for the ECM fungi. Delayed snow melt did not influence the temporal variation in fungal communities between the treatments. Results suggest that some fungal species are favored, while others are disfavored resulting in their local extinction due to long-term changes in snow amount. Shifts in species composition of fungal functional groups are likely to affect nutrient cycling, ecosystem respiration, and stored permafrost carbon.

Changing climate is expected to alter precipitation patterns in the Arctic, with consequences for subsurface temperature and moisture conditions, community structure, and nutrient mobilization through microbial belowground processes. Here, we address the effect of increased snow depth on the variation in species richness and community structure of ectomycorrhizal (ECM) and saprotrophic fungi. Soil samples were collected weekly from mid-July to mid-September in both control and deep snow plots. Richness of ECM fungi was higher, while saprotrophic fungi was lower in increased snow depth plots relative to controls. ECM fungal richness was related to soil NO3-N, NH4-N, and K; and saprotrophic fungi to NO3-N and pH. Small but significant changes in the composition of saprotrophic fungi could be attributed to snow treatment and sampling time, but not so for the ECM fungi. Delayed snow melt did not influence the temporal variation in fungal communities between the treatments. Results suggest that some fungal species are favored, while others are disfavored resulting in their local extinction due to long-term changes in snow amount. Shifts in species composition of fungal functional groups are likely to affect nutrient cycling, ecosystem respiration, and stored permafrost carbon.

Changing climate is expected to alter precipitation patterns in the Arctic, with consequences for subsurface temperature and moisture conditions, community structure, and nutrient mobilization through microbial belowground processes. Here, we address the effect of increased snow depth on the variation in species richness and community structure of ectomycorrhizal (ECM) and saprotrophic fungi. Soil samples were collected weekly from mid-July to mid-September in both control and deep snow plots. Richness of ECM fungi was higher, while saprotrophic fungi was lower in increased snow depth plots relative to controls. ECM fungal richness was related to soil NO3-N, NH4-N, and K; and saprotrophic fungi to NO3-N and pH. Small but significant changes in the composition of saprotrophic fungi could be attributed to snow treatment and sampling time, but not so for the ECM fungi. Delayed snow melt did not influence the temporal variation in fungal communities between the treatments. Results suggest that some fungal species are favored, while others are disfavored resulting in their local extinction due to long-term changes in snow amount. Shifts in species composition of fungal functional groups are likely to affect nutrient cycling, ecosystem respiration, and stored permafrost carbon.

Changing climate is expected to alter precipitation patterns in the Arctic, with consequences for subsurface temperature and moisture conditions, community structure, and nutrient mobilization through microbial belowground processes. Here, we address the effect of increased snow depth on the variation in species richness and community structure of ectomycorrhizal (ECM) and saprotrophic fungi. Soil samples were collected weekly from mid-July to mid-September in both control and deep snow plots. Richness of ECM fungi was lower, while saprotrophic fungi was higher in increased snow depth plots relative to controls. [Correction added on 23 September 2016 after first online publication: In the preceding sentence, the richness of ECM and saprotrophic fungi were wrongly interchanged and have been fixed in this current version.] ECM fungal richness was related to soil NO3-N, NH4-N, and K; and saprotrophic fungi to NO3-N and pH. Small but significant changes in the composition of saprotrophic fungi could be attributed to snow treatment and sampling time, but not so for the ECM fungi. Delayed snow melt did not influence the temporal variation in fungal communities between the treatments. Results suggest that some fungal species are favored, while others are disfavored resulting in their local extinction due to long-term changes in snow amount. Shifts in species composition of fungal functional groups are likely to affect nutrient cycling, ecosystem respiration, and stored permafrost carbon.

Plant pathogenic fungi are a large and diverse assemblage of eukaryotes with substantial impacts on natural ecosystems and human endeavours. These taxa often have complex and poorly understood life cycles, lack observable, discriminatory morphological characters, and may not be amenable to in vitro culturing. As a result, species identification is frequently difficult. Molecular (DNA sequence) data have emerged as crucial information for the taxonomic identification of plant pathogenic fungi, with the nuclear ribosomal internal transcribed spacer (ITS) region being the most popular marker. However, international nucleotide sequence databases are accumulating numerous sequences of compromised or low-resolution taxonomic annotations and substandard technical quality, making their use in the molecular identification of plant pathogenic fungi problematic. Here we report on a concerted effort to identify high-quality reference sequences for various plant pathogenic fungi and to re-annotate incorrectly or insufficiently annotated public ITS sequences from these fungal lineages. A third objective was to enrich the sequences with geographical and ecological metadata. The results – a total of 31,954 changes – are incorporated in and made available through the UNITE database for molecular identification of fungi (http://unite.ut.ee), including standalone FASTA files of sequence data for local BLAST searches, use in the next-generation sequencing analysis platforms QIIME and mothur, and related applications. The present initiative is just a beginning to cover the wide spectrum of plant pathogenic fungi, and we invite all researchers with pertinent expertise to join the annotation effort.

Plant pathogenic fungi are a large and diverse assemblage of eukaryotes with substantial impacts on natural ecosystems and human endeavours. These taxa often have complex and poorly understood life cycles, lack observable, discriminatory morphological characters, and may not be amenable to in vitro culturing. As a result, species identification is frequently difficult. Molecular (DNA sequence) data have emerged as crucial information for the taxonomic identification of plant pathogenic fungi, with the nuclear ribosomal internal transcribed spacer (ITS) region being the most popular marker. However, international nucleotide sequence databases are accumulating numerous sequences of compromised or low-resolution taxonomic annotations and substandard technical quality, making their use in the molecular identification of plant pathogenic fungi problematic. Here we report on a concerted effort to identify high-quality reference sequences for various plant pathogenic fungi and to re-annotate incorrectly or insufficiently annotated public ITS sequences from these fungal lineages. A third objective was to enrich the sequences with geographical and ecological metadata. The results – a total of 31,954 changes – are incorporated in and made available through the UNITE database for molecular identification of fungi (http://unite.ut.ee), including standalone FASTA files of sequence data for local BLAST searches, use in the next-generation sequencing analysis platforms QIIME and mothur, and related applications. The present initiative is just a beginning to cover the wide spectrum of plant pathogenic fungi, and we invite all researchers with pertinent expertise to join the annotation effort.

The taxonomic and ecological diversity of ancient fungal communities was assessed by combining next generation sequencing and metabarcoding of DNA preserved in permafrost. Twenty-six sediment samples dated 16000-32000 radiocarbon years old from two localities in Siberia were analysed for fungal ITS. We detected 75 fungal OTUs from 21 orders representing three phyla, although rarefaction analyses suggested that the full diversity was not recovered despite generating an average of 6677±3811 (mean±SD) sequences per sample and that preservation bias likely has considerable effect on the recovered DNA. Most OTUs (75.4%) represented ascomycetes. Due to insufficient sequencing depth, DNA degradation and putative preservation biases in our samples, the recovered taxa probably do not represent the complete historic fungal community, and it is difficult to determine whether the fungal communities varied geographically or experienced a composition shift within the period of 16000-32000 bp. However, annotation of OTUs to functional ecological groups provided a wealth of information on the historic communities. About one-third of the OTUs are presumed plant-associates (pathogens, saprotrophs and endophytes) typical of graminoid- and forb-rich habitats. We also detected putative insect pathogens, coprophiles and keratinophiles likely associated with ancient insect and herbivore faunas. The detection of putative insect pathogens, mycoparasites, aquatic fungi and endophytes broadens our previous knowledge of the diversity of fungi present in Beringian palaeoecosystems. A large group of putatively psychrophilic/psychrotolerant fungi was also detected, most likely representing a modern, metabolically active fungal community.

Several eukaryotic symbioses have shown to host a rich diversity of prokaryotes that interact with their hosts. Here, we study bacterial communities associated with ectomycorrhizal root systems of Bistorta vivipara compared to bacterial communities in bulk soil using pyrosequencing of 16S rRNA amplicons. A high richness of Operational Taxonomic Units (OTUs) was found in plant roots (3,571 OTUs) and surrounding soil (3,476 OTUs). The community composition differed markedly between these two environments. Actinobacteria, Armatimonadetes, Chloroflexi and OTUs unclassified at phylum level were significantly more abundant in plant roots than in soil. A large proportion of the OTUs, especially those in plant roots, presented low similarity to Sanger 16S rRNA reference sequences, suggesting novel bacterial diversity in ectomycorrhizae. Furthermore, the bacterial communities of the plant roots were spatially structured up to a distance of 60 cm, which may be explained by bacteria using fungal hyphae as a transport vector. The analyzed ectomycorrhizae presents a distinct microbiome, which likely influence the functioning of the plant-fungus symbiosis

The taxonomic and ecological diversity of ancient fungal communities was assessed by combining next generation sequencing and metabarcoding of DNA preserved in permafrost. Twenty-six sediment samples dated 16000-32000 radiocarbon years old from two localities in Siberia were analysed for fungal ITS. We detected 75 fungal OTUs from 21 orders representing three phyla, although rarefaction analyses suggested that the full diversity was not recovered despite generating an average of 6677±3811 (mean±SD) sequences per sample and that preservation bias likely has considerable effect on the recovered DNA. Most OTUs (75.4%) represented ascomycetes. Due to insufficient sequencing depth, DNA degradation and putative preservation biases in our samples, the recovered taxa probably do not represent the complete historic fungal community, and it is difficult to determine whether the fungal communities varied geographically or experienced a composition shift within the period of 16000-32000 bp. However, annotation of OTUs to functional ecological groups provided a wealth of information on the historic communities. About one-third of the OTUs are presumed plant-associates (pathogens, saprotrophs and endophytes) typical of graminoid- and forb-rich habitats. We also detected putative insect pathogens, coprophiles and keratinophiles likely associated with ancient insect and herbivore faunas. The detection of putative insect pathogens, mycoparasites, aquatic fungi and endophytes broadens our previous knowledge of the diversity of fungi present in Beringian palaeoecosystems. A large group of putatively psychrophilic/psychrotolerant fungi was also detected, most likely representing a modern, metabolically active fungal community.

Metabarcoding approaches use total and typically degraded DNA from environmental samples to analyse biotic assemblages and can potentially be carried out for any kinds of organisms in an ecosystem. These analyses rely on specific markers, here called metabarcodes, which should be optimized for taxonomic resolution, minimal bias in amplification of the target organism group and short sequence length. Using bioinformatic tools, we developed metabarcodes for several groups of organisms: fungi, bryophytes, enchytraeids, beetles and birds. The ability of these metabarcodes to amplify the target groups was systematically evaluated by (i) in silico PCRs using all standard sequences in the EMBL public database as templates, (ii) in vitro PCRs of DNA extracts from surface soil samples from a site in Varanger, northern Norway and (iii) in vitro PCRs of DNA extracts from permanently frozen sediment samples of late-Pleistocene age (similar to 16 00050 000 years bp) from two Siberian sites, Duvanny Yar and Main River. Comparison of the results from the in silico PCR with those obtained in vitro showed that the in silico approach offered a reliable estimate of the suitability of a marker. All target groups were detected in the environmental DNA, but we found large variation in the level of detection among the groups and between modern and ancient samples. Success rates for the Pleistocene samples were highest for fungal DNA, whereas bryophyte, beetle and bird sequences could also be retrieved, but to a much lesser degree. The metabarcoding approach has considerable potential for biodiversity screening of modern samples and also as a palaeoecological tool.

The incursion of High-Throughput Sequencing (HTS) in environmental microbiology brings unique opportunities and challenges. HTS now allows a high-resolution exploration of the vast taxonomic and metabolic diversity present in the microbial world, which can provide an exceptional insight on global ecosystem functioning, ecological processes and evolution. This exploration has also economic potential, as we will have access to the evolutionary innovation present in microbial metabolisms, which could be used for biotechnological development. HTS is also challenging the research community, and the current bottleneck is present in the data analysis side. At the moment, researchers are in a sequence data deluge, with sequencing throughput advancing faster than the computer power needed for data analysis. However, new tools and approaches are being developed constantly and the whole process could be depicted as a fast co-evolution between sequencing technology, informatics and microbiologists. In this work, we examine the most popular and recently commercialized HTS platforms as well as bioinformatics methods for data handling and analysis used in microbial metagenomics. This non-exhaustive review is intended to serve as a broad state-of-the-art guide to researchers expanding into this rapidly evolving field. © 2012 Elsevier B.V.

Metabarcoding approaches use total and typically degraded DNA from environmental samples to analyse biotic assemblages and can potentially be carried out for any kinds of organisms in an ecosystem. These analyses rely on specific markers, here called metabarcodes, which should be optimized for taxonomic resolution, minimal bias in amplification of the target organism group and short sequence length. Using bioinformatic tools, we developed metabarcodes for several groups of organisms: fungi, bryophytes, enchytraeids, beetles and birds. The ability of these metabarcodes to amplify the target groups was systematically evaluated by (i) in silico PCRs using all standard sequences in the EMBL public database as templates, (ii) in vitro PCRs of DNA extracts from surface soil samples from a site in Varanger, northern Norway and (iii) in vitro PCRs of DNA extracts from permanently frozen sediment samples of late-Pleistocene age (similar to 16 00050 000 years bp) from two Siberian sites, Duvanny Yar and Main River. Comparison of the results from the in silico PCR with those obtained in vitro showed that the in silico approach offered a reliable estimate of the suitability of a marker. All target groups were detected in the environmental DNA, but we found large variation in the level of detection among the groups and between modern and ancient samples. Success rates for the Pleistocene samples were highest for fungal DNA, whereas bryophyte, beetle and bird sequences could also be retrieved, but to a much lesser degree. The metabarcoding approach has considerable potential for biodiversity screening of modern samples and also as a palaeoecological tool.