Ectomycorrhizal fungal communities in late-successional Swedish boreal forests, and their composition following wildfire

This study was conducted to evaluate the effects of wildfires on ectomycorrhizal (EM) fungal communities in Scots pine ( Pinus sylvestris ) stands. Below- and above-ground communities were analysed in terms of species richness and evenness by examining mycorrhizas and sporocarps in a chronosequence of burned stands in comparison with adjacent unburned late-successional stands. The internal transcribed spacer (ITS)-region (rDNA) of mycobionts from single mycorrhizas was digested with three restriction enzymes and compared with an ITS–restriction fragment length polymorphism (RFLP) reference database of EM sporocarps. Spatial variation seemed to be more prominent than the effects of fire on the EM fungal species composition. Most of the common species tended to be found in all sites, suggesting that EM fungal communities show a high degree of continuity following low-intensity wildfires. Species richness was not affected by fire, whereas the evenness of species distributions of mycorrhizas was lower in the burned stands. The diversity of EM fungi was relatively high considering that there were only three EM tree species present in the stands. In total, 135 EM taxa were identified on the basis of their RFLP patterns; 66 species were recorded as sporocarps, but only 11 of these were also recorded as mycorrhizas. The species composition of the below-ground community of EM fungi did not reflect that of the sporocarps produced. EM fungal species present in our ITS–RFLP reference database accounted for 54–99% of the total sporocarp production in the stands, but only 0–32% of the mycorrhizal abundance.     

Genotype identity determines productivity and CO2 efflux across a genotype-species gradient of ectomycorrhizal fungi

Ectomycorrhizal (EM) fungal communities are taxonomically diverse, and independent manipulation of both intra- and interspecific diversity has previously been shown to positively influence the productivity and activity of EM fungi. Here, we combine manipulations of intra- and interspecific richness and test the effects of a genotype-species gradient on the biomass production and respiration of EM fungi in vitro. Genotype identity had the most pronounced effect on fungal productivity, and in some cases variation within species was greater than between species. We found small negative effects of both species and genotype richness on biomass production, CO₂ efflux and the final nitrogen (N) content of the fungal communities corresponding to mixed negative selection and complementarity effects. Our study highlights the degree of variability between individual EM fungi at the genotype level, and consequently emphasises the importance of individual genotypes for playing key roles in shaping belowground community functioning.     

Fire and local factors shape ectomycorrhizal fungal communities associated with Pinus ponderosa in mountains of the Madrean Sky Island Archipelago

With a warming and drying climate, coniferous forests worldwide are increasingly threatened by wildfires. We examined how fire impacts ectomycorrhizal (EM) fungi associated with Pinus ponderosa, an important tree species in western North America. In the biodiverse Madrean Sky Islands, P. ponderosa forests exist on insular mountains separated by arid lands. How do EM fungi in these isolated ranges respond to fire, and can data from individual ranges predict community shifts after fire at a regional scale? By comparing areas in two ranges that experienced moderate fires 12–16 y earlier, and proximate areas in each range without recent fire, we reveal pervasive effects on diversity and composition of EM communities more than a decade after moderate fires occurred. Post-fire differences in EM communities in different ranges highlight the challenge of predicting fungal community shifts in these isolated forests, despite similarities of climate, plant communities, and fire severity.     

Plant identity strongly structures the root-associated fungal community in a diverse subtropical forest

Revealing the relationship between plants and root-associated fungi is very important in understanding diversity maintenance and community assembly in ecosystems. However, the community assembly of root-associated fungi of focal plant species along a subtropical plant species diversity gradient is less documented. Here, we examined root-associated fungal communities associated with five ectomycorrhizal (EM) plant species (Betula luminifera, Castanea henryi, Castanopsis fargesii, C. sclerophylla, and Quercus serrate) in a Chinese subtropical woody plant species diversity (1, 2, 4, 8, 16 and 24 species) experiment, using paired-end Illumina MiSeq sequencing of the ITS2 region. In total, we detected 1933 root-associated fungal operational taxonomic units (OTUs) at a 97% sequence similarity level. Plant identity had a significant effect on total and saprotrophic fungal OTU richness, but plant species diversity level had a significant effect on saprotrophic and pathogenic fungal OTU richness. The community composition of total, saprotrophic and EM fungi was structured by plant identity and plant species diversity level. However, the community composition of pathogenic fungi was only shaped by plant identity. This study highlights that plant identity has a stronger effect on the root-associated fungal community than plant species diversity level in a diverse subtropical forest ecosystem.     

Wildfire severity reduces richness and alters composition of soil fungal communities in boreal forests of western Canada

Wildfire is the dominant disturbance in boreal forests and fire activity is increasing in these regions. Soil fungal communities are important for plant growth and nutrient cycling postfire but there is little understanding of how fires impact fungal communities across landscapes, fire severity gradients, and stand types in boreal forests. Understanding relationships between fungal community composition, particularly mycorrhizas, and understory plant composition is therefore important in predicting how future fire regimes may affect vegetation. We used an extreme wildfire event in boreal forests of Canada's Northwest Territories to test drivers of fungal communities and assess relationships with plant communities. We sampled soils from 39 plots 1 year after fire and 8 unburned plots. High‐throughput sequencing (MiSeq, ITS) revealed 2,034 fungal operational taxonomic units. We found soil pH and fire severity (proportion soil organic layer combusted), and interactions between these drivers were important for fungal community structure (composition, richness, diversity, functional groups). Where fire severity was low, samples with low pH had higher total fungal, mycorrhizal, and saprotroph richness compared to where severity was high. Increased fire severity caused declines in richness of total fungi, mycorrhizas, and saprotrophs, and declines in diversity of total fungi and mycorrhizas. The importance of stand age (a surrogate for fire return interval) for fungal composition suggests we could detect long‐term successional patterns even after fire. Mycorrhizal and plant community composition, richness, and diversity were weakly but significantly correlated. These weak relationships and the distribution of fungi across plots suggest that the underlying driver of fungal community structure is pH, which is modified by fire severity. This study shows the importance of edaphic factors in determining fungal community structure at large scales, but suggests these patterns are mediated by interactions between fire and forest stand composition.     

Comparison of root-associated communities of native and non-native ectomycorrhizal hosts in an urban landscape

Non-native tree species are often used as ornamentals in urban landscapes. However, their root-associated fungal communities remain yet to be examined in detail. Here, we compared richness, diversity and community composition of ectomycorrhizosphere fungi in general and ectomycorrhizal (EcM) fungi in particular between a non-native Pinus nigra and a native Quercus macrocarpa across a growing season in urban parks using 454-pyrosequencing. Our data show that, while the ectomycorrhizosphere community richness and diversity did not differ between the two host, the EcM communities associated with the native host were often more species rich and included more exclusive members than those of the non-native hosts. In contrast, the ectomycorrhizosphere communities of the two hosts were compositionally clearly distinct in nonmetric multidimensional ordination analyses, whereas the EcM communities were only marginally so. Taken together, our data suggest EcM communities with broad host compatibilities and with a limited numbers of taxa with preference to the non-native host. Furthermore, many common fungi in the non-native Pinus were not EcM taxa, suggesting that the fungal communities of the non-native host may be enriched in non-mycorrhizal fungi at the cost of the EcM taxa. Finally, while our colonization estimates did not suggest a shortage in EcM inoculum for either host in urban parks, the differences in the fungi associated with the two hosts emphasize the importance of using native hosts in urban environments as a tool to conserve endemic fungal diversity and richness in man-made systems.     

Fungal Community Shifts in Structure and Function across a Boreal Forest Fire Chronosequence

Forest fires are a common natural disturbance in forested ecosystems and have a large impact on the microbial communities in forest soils. The response of soil fungal communities to forest fire is poorly documented. Here, we investigated fungal community structure and function across a 152-year boreal forest fire chronosequence using high-throughput sequencing of the internal transcribed spacer 2 (ITS2) region and a functional gene array (GeoChip). Our results demonstrate that the boreal forest soil fungal community was most diverse soon after a fire disturbance and declined over time. The differences in the fungal communities were explained by changes in the abundance of basidiomycetes and ascomycetes. Ectomycorrhizal (ECM) fungi contributed to the increase in basidiomycete abundance over time, with the operational taxonomic units (OTUs) representing the genera Cortinarius and Piloderma dominating in abundance. Hierarchical cluster analysis by using gene signal intensity revealed that the sites with different fire histories formed separate clusters, suggesting differences in the potential to maintain essential biogeochemical soil processes. The site with the greatest biological diversity had also the most diverse genes. The genes involved in organic matter degradation in the mature forest, in which ECM fungi were the most abundant, were as common in the youngest site, in which saprotrophic fungi had a relatively higher abundance. This study provides insight into the impact of fire disturbance on soil fungal community dynamics.     

Effects of phenolic acids from ginseng rhizosphere on soil fungi structure,richness and diversity in consecutive monoculturing of ginseng

Ginseng yield and quality are seriously compromised by consecutive monoculturing in northeastern China. The imbalance of soil fungi communities and autotoxicity of ginseng are the major factors in consecutive monoculturing ginseng crops. Soil fungal communities were identified using Illumina MiSeq sequencing, applied to soils that consecutively cultured ginseng (CCG) for six years and new forest soil (NFS), or receiving application of phenolic acids (PAs). The CCG field received five treatments with five different phenolic acids, including gallic acid (GA), salicylic acid (SA), 3-phenylpropionic acid (3-PA), benzoic acid (BA) and cinnamic acid (CA), which were detected from ginseng rhizosphere in consecutive cropping soil. Fungal richness, fungi diversity, community composition, relative taxon abundances, root rot disease, and growth rate were compared among the different treatments. 579 fungal operational taxonomic units at 97% ITS sequence identity were found among 201,617 sequence reads derived from 18 separate soil samples. Members of the phylum Ascomycota dominated the soil fungal communities, and putative pathogens, such as Fusarium, Gibberella and Nectriaceae_unclassified which may include the abundant sexual morph of Cylindrocarpon destructans, showed higher relative abundances in the CCG fields. Compared to the CCG and NFS fields, PAs (except CA) enhanced the fungi richness and decreased fungi diversity. Cluster analysis indicated that the PAs (except CA) changed the fungi structure in a uniform way. PAs stimulate root rot disease and enhance disease severity, restricting plant growth. The results suggest that the PAs (except CA) may enhance the fungi richness, decrease the fungi diversity and changed the fungi structure to increase fungal pathogen loads, which could explain the declined yield and quality of ginseng in consecutively monocultured ginseng crops.     

林火对植物根围丛枝菌根真菌多样性的影响

林火是森林生态系统的一种主要干扰因子,以青岛市三标山林火迹地为研究对象,采集荆条(Vitex negundo)、胡枝子(Lespedeza bicolor)、花木蓝(Indigofera kirilowii)、青花椒(Zanthoxylum schinifolium)和野青茅(Deyeuxia arundinacea)5种优势植物根围土壤,研究不同林火强度对丛枝菌根(AM)真菌多样性的影响。结果表明,AM真菌侵染率和孢子密度随火灾强度的加强而降低;非过火区植物根围土壤中,分离鉴定出AM真菌3属11种,轻度过火区分离鉴定出AM真菌3属10种,中度过火区分离鉴定出AM真菌3属9种,重度过火区分离鉴定出AM真菌3属8种。过火区AM真菌种丰度低于非过火区。过火区和非过火区AM真菌的重要值和优势种不同,非过火区植物根围的优势种是地球囊霉(Glomus geosporum)、台湾球囊霉(G.taiwanensis)、分支巨孢囊霉(Gigaspora ramisporophora)、极大巨孢囊霉(Gi.gigantean)、福摩萨球囊霉(G.formosanum)、悬钩子球囊霉(G.rubiforme)、柯氏无梗囊霉(Acaulospora koskei)和松蜜无梗囊霉(A.thomii);轻度过火区植物根围的优势种是地球囊霉和台湾球囊霉;中度过火区的是台湾球囊霉和地球囊霉(野青茅除外);重度过火区植物根围的优势种是地球囊霉。不同强度的过火区对AM真菌群落组成有不同程度的影响。认为林火降低植物根围土壤中AM真菌多样性。     

Co-invading ectomycorrhizal fungal succession in pine-invaded mountain grasslands

Ectomycorrhizal (EM) fungal communities that associate with invading pines (Pinus spp.) are expected to be poor in species diversity. However, long-term successional trajectories and the persistence of dispersal limitations of EM fungi in the exotic range are not well understood. We sampled the roots and surrounding soil of Pinus elliottii and P. taeda trees invading mountain grasslands of Argentina. We also sampled the EM fungal spore bank in grassland soil near (∼150 m) and far (∼850 m) from the original pine plantations. We found 86 different co-invasive EM fungal OTUs. Differential dispersal capacities among EM fungi were detected in the spore bank of grassland soil, but not under mature pines. After thirty years of invasion, the age, but not the degree of spatial isolation of pine individuals affected the EM fungal composition. We showed how EM fungal succession occurs during pine invasions, which may have clear consequences for ecosystem functioning of co-invaded sites.     

重度火烧迹地兴安落叶松(Larix gmelinii)根际土壤真菌群落研究

为探究内蒙古根河大兴安岭林区重度火烧迹地不同生长状态的兴安落叶松根际土壤真菌群落特征,选取火烧枯立木（BDW）、火烧存活木（BSW）、未火烧对照木（CK）,通过磷脂脂肪酸方法分析根际土壤真菌群落生物量变化,利用高通量测序技术对根际土壤真菌群落组成与功能进行分析,探讨影响根际土壤真菌群落的主要环境因子。结果表明：（1）相较于未火烧对照木,火烧枯立木与火烧存活木根际土壤真菌群落磷脂脂肪酸含量显著降低,真菌群落α多样性显著降低。β多样性分析与群落相似性分析结果显示,火烧枯立木、火烧存活木、未火烧对照木根际土壤真菌群落组成存在显著差异。（2）相较于未火烧对照木,火烧枯立木与火烧存活木根际土壤中担子菌门（Basidiomycota）真菌相对丰度显著下降,子囊菌门（Ascomycota）真菌相对丰度显著升高。子囊菌门（Ascomycota）内,火烧枯立木根际土壤中内生真菌相对丰度显著高于火烧存活木与未火烧对照木,而火烧存活木根际土壤中腐生-外生菌根复合型真菌相对丰度显著高于火烧枯立木与未火烧对照木。（3）根际土壤总磷、总钾含量是影响重度火烧迹地兴安落叶松根际土壤真菌群落优势菌门由担子菌门（Basidiomycota）转变为子囊菌门（Ascomycota）的主要环境因子,而根际土壤总碳、总氮、总磷含量则是造成火烧枯立木与火烧存活木根际土壤真菌群落组成与功能差异的主要环境因子。本研究有助于了解大兴安岭林区土壤真菌群落结构,对真菌群落多样性的维持与管理具有参考价值。     

Diversity and community structure of ectomycorrhizal fungi in a wooded meadow

Wooded meadows are seminatural plant communities that support high diversity of various taxa. Due to changes in land use, wooded meadows have severely declined during the last century. The dominant trees in wooded meadows acquire mineral nutrients via ectomycorrhizal fungi. Using anatomotyping and sequencing of root tips, interpolation and extrapolation methods, we studied the diversity and community structure of ectomycorrhizal fungi in two soil horizons of both managed and forested parts of a wooded meadow in Estonia. Species of Thelephoraceae, Sebacinaceae and the genus Inocybe dominated the whole ectomycorrhizal fungal community of 172 observed species. Forested and managed parts of the wooded meadow harboured different communities of ectomycorrhizal fungi, whereas soil horizon had a negligible effect on the fungal community composition. Diverse soil conditions and host trees likely support the high richness of ectomycorrhizal fungi in the wooded meadow ecosystem. Direct sequencing integrated with interpolation and extrapolation methods are promising to identify the fungi at the species level and to compare species richness between communities of ectomycorrhizal fungi.     

Relative Importance of Deterministic and Stochastic Processes in Driving Arbuscular Mycorrhizal Fungal Assemblage during the Spreading of a Toxic Plant

Both deterministic and stochastic processes are expected to drive the assemblages of arbuscular mycorrhizal (AM) fungi, but little is known about the relative importance of these processes during the spreading of toxic plants. Here, the species composition and phylogenetic structure of AM fungal communities colonizing the roots of a toxic plant, Ligularia virgaurea, and its neighborhood plants, were analyzed in patches with different individual densities of L. virgaurea (represents the spreading degree). Community compositions of AM fungi in both root systems were changed significantly by the L. virgaurea spreading, and also these communities fitted the neutral model very well. AM fungal communities in patches with absence and presence of L. virgaurea were phylogenetically random and clustered, respectively, suggesting that the principal ecological process determining AM fungal assemblage shifted from stochastic process to environmental filtering when this toxic plant was present. Our results indicate that deterministic and stochastic processes together determine the assemblage of AM fungi, but the dominant process would be changed by the spreading of toxic plants, and suggest that the spreading of toxic plants in alpine meadow ecosystems might be involving the mycorrhizal symbionts.     

Changes of butterfly communities after forest fire

Fires change the diversity and composition of insects in forest ecosystems. In the present study, we examined the change of butterfly communities after a fire including the increase of butterfly richness, grassland species, and generalist species, and more changed communities. Butterflies were surveyed for 5 years after the big Uljin fire in 2007. During each year, butterflies were counted monthly by the line transect method from April to October at two sites (burned vs. unburned, ~ 1.5 km routes). Specialist grassland species decreased in the year of the fire but generalist species did not increase significantly. Butterfly richness did not change but butterfly diversity decreased due to a sudden increase of a species, Polygonia c-aureum. The butterfly community in the year of the fire was different from those in later years, showing temporary change of community in the year of the fire. Species composition was significantly different between burned and unburned sites, but this phenomenon cannot be interpreted as an influence of fire due to highly variable species composition of local butterfly assemblages and the non-repetitive sampling site of the present study.     

Fungal communities associated with roots of two closely related Juglandaceae species with a disjunct distribution in the tropics

We studied the biogeography and community structure of root-associated and ectomycorrhizal fungal communities in two related species of tropical Juglandaceae that have disjunct distributions in Asia and Mesoamerica. We tested the effects of environmental and dispersal factors in structuring root-associated fungi at a regional scale. We used Illumina sequencing to document fungi on the roots of Oreomunnea mexicana in Panama and Mexico and Alfaropsis roxburghiana in China. Ectomycorrhizal fungi dominated the communities with both hosts but we detected a more diverse root-associated fungal community in Alfaropsis but higher ectomycorrhizal fungi richness in Oreomunnea. Geographic distance was the best predictor of variation in fungal species composition, when including both hosts and when analyzing each host independently. However, our results showed a high correlation between geographic distance and abiotic variables, and therefore we were not able to determine if the observed changes in fungal community composition were explained also by spatially structured environmental or phylogenetic factors.     

The annual invasive plant Impatiens glandulifera reduces hyphal biomass of soil fungi in deciduous forests

Soil fungi play a crucial role in ecosystem functioning and there is increasing evidence that exotic plants invading forests can affect soil fungal communities. We examined potential effects of the invasive plant Impatiens glandulifera on hyphal biomass of ectomycorrhizal fungi, their genetic diversity and the diversity of other soil fungi in deciduous forests in Switzerland. We compared invaded patches with patches where I. glandulifera had been removed, by establishing pairs of 3-m long transect lines at the edge of seven areas of either type. Along the transects we assessed the length of ectomycorrhizal fungal hyphae using the ‘ingrowth mesh bag method’, and used terminal restriction fragment length polymorphism (T-RFLP) analysis to examine fungal genetic diversity. The invasive plant reduced fungal hyphal biomass by 30–80%: the reduction was largest in the centre of the patch. I. glandulifera did not alter fungal richness, but affected the composition of fungal communities. This is probably the result of a decrease of mycorrhizal fungi, coupled with an increase of saprotrophic fungi. Our findings demonstrate the adverse impacts of an annual invasive plant species on both fungal hyphal biomass and the composition of soil fungal communities. This may negatively affect forest nutrient and carbon cycling, soil stability and the functionality of the fungal community, with major consequences for forest ecosystem functioning.     

Strong coupling of plant and fungal community structure across western Amazonian rainforests

The Amazon basin harbors a diverse ecological community that has a critical role in the maintenance of the biosphere. Although plant and animal communities have received much attention, basic information is lacking for fungal or prokaryotic communities. This is despite the fact that recent ecological studies have suggested a prominent role for interactions with soil fungi in structuring the diversity and abundance of tropical rainforest trees. In this study, we characterize soil fungal communities across three major tropical forest types in the western Amazon basin (terra firme, seasonally flooded and white sand) using 454 pyrosequencing. Using these data, we examine the relationship between fungal diversity and tree species richness, and between fungal community composition and tree species composition, soil environment and spatial proximity. We find that the fungal community in these ecosystems is diverse, with high degrees of spatial variability related to forest type. We also find strong correlations between α- and β-diversity of soil fungi and trees. Both fungal and plant community β-diversity were also correlated with differences in environmental conditions. The correlation between plant and fungal richness was stronger in fungal lineages known for biotrophic strategies (for example, pathogens, mycorrhizas) compared with a lineage known primarily for saprotrophy (yeasts), suggesting that this coupling is, at least in part, due to direct plant–fungal interactions. These data provide a much-needed look at an understudied dimension of the biota in an important ecosystem and supports the hypothesis that fungal communities are involved in the regulation of tropical tree diversity.     

Patterns of fungal diversity and composition along a salinity gradient

Estuarine salinity gradients are known to influence plant, bacterial and archaeal community structure. We sequenced 18S rRNA genes to investigate patterns in sediment fungal diversity (richness and evenness of taxa) and composition (taxonomic and phylogenetic) along an estuarine salinity gradient. We sampled three marshes—a salt, brackish and freshwater marsh—in Rhode Island. To compare the relative effect of the salinity gradient with that of plants, we sampled fungi in plots with Spartina patens and in plots from which plants were removed 2 years prior to sampling. The fungal sediment community was unique compared with previously sampled fungal communities; we detected more Ascomycota (78%), fewer Basidiomycota (6%) and more fungi from basal lineages (16%) (Chytridiomycota, Glomeromycota and four additional groups) than typically found in soil. Across marshes, fungal composition changed substantially, whereas fungal diversity differed only at the finest level of genetic resolution, and was highest in the intermediate, brackish marsh. In contrast, the presence of plants had a highly significant effect on fungal diversity at all levels of genetic resolution, but less of an effect on fungal composition. These results suggest that salinity (or other covarying parameters) selects for a distinctive fungal composition, and plants provide additional niches upon which taxa within these communities can specialize and coexist. Given the number of sequences from basal fungal lineages, the study also suggests that further sampling of estuarine sediments may help in understanding early fungal evolution.     

High richness of ectomycorrhizal fungi and low host specificity in a coastal sand dune ecosystem revealed by network analysis

Ectomycorrhizal (EM) fungi are ubiquitous in temperate and boreal forests, comprising over 20,000 species forming root symbiotic associations with Pinaceae and woody angiosperms. As much as 100 different EM fungal species can coexist and interact with the same tree species, forming complex multispecies networks in soils. The degree of host specificity and structural properties of these interaction networks (e.g., nestedness and modularity) may influence plant and fungal community assembly and species coexistence, yet their structure has been little studied in northern coniferous forests, where trees depend on EM fungi for nutrient acquisition. We used high‐throughput sequencing to characterize the composition and diversity of bulk soil and root‐associated fungal communities in four co‐occurring Pinaceae in a relic foredune plain located at Îles de la Madeleine, Québec, Canada. We found high EM fungal richness across the four hosts, with a total of 200 EM operational taxonomic units (OTUs), mainly belonging to the Agaricomycetes. Network analysis revealed an antinested pattern in both bulk soil and roots EM fungal communities. However, there was no detectable modularity (i.e., subgroups of interacting species) in the interaction networks, indicating a low level of specificity in these EM associations. In addition, there were no differences in EM fungal OTU richness or community structure among the four tree species. Limited shared resources and competitive exclusion typically restrict the number of taxa coexisting within the same niche. As such, our finding of high EM fungal richness and low host specificity highlights the need for further studies to determine the mechanisms enabling such a large number of EM fungal species to coexist locally on the same hosts.