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Phylogenetic relationships among studied species of Chalara based on the EF-1α region using NJ, ML and BA analyses. Ex-type sequences are in boldface type, ex-epitype sequence is marked with an asterisk, strains whose sequences are not in GenBank are in smaller size font. Bootstrap values of NJ and ML greater than 50% are indicated along nodes. Thick lines show PP higher than 0.95. The tree was rooted with Tryblidiopsis pinastri DQ471106.1
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Chalara is a genus of anamorphic fungi with typical phialidic conidiogenous cells. Species of Chalara live mostly as litter saprotrophs, many of them on coniferous litter. In this study, the morphology and DNA sequences (ITS
rDNA, 28S rDNA and EF-1α) of strains of various species of Chalara isolated from coniferous litter needles were compared with...
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... Within Leotiomycetes, the chalara-like fungi are hyphomycetous anamorphs, and characterized by sporodochial or synnematous conidiomata, or solitary conidiophores, sessile or stalked phialidic conidiogenous cell with a basal venter and a cylindrical collarette bearing a deep-seated conidiogenous locus, and hyaline, cylindrical or obclavate or oblong, aseptate or septate conidia extruded in short or long chain (Nag Raj and Hughes 1974;Nag Raj and Kendrick 1975;Minter et al. 1982Minter et al. , 1983Seifert et al. 2011). In nature they are commonly found as saprobes on decaying plant material such as leaves, branches, stems, barks, woods, and fruits from terrestrial and submerged environments (Willoughby and Archer 1973;Nag Raj and Kendrick 1975;Tsui et al. 2001;McKenzie et al. 2002;Baschien et al. 2008;Koukol 2011;Friggens et al. 2017). However, they were reported as plant pathogens of trees and ferns, fungicolous or lichenicolous, endophytic fungi of plants and lichens, associated with roots of plants and eggs of nematodes, or from soil (Nag Raj and Kendrick 1975;Gams and Holubová-Jechová 1976;Morgan-Jones et al. 1984;Samuels and Rogerson 1990;Christiansen 1993;Etayo and Diererich 1996;Liu and Zhang 1998;Kowalski 2006;Etayo and Sancho 2008;Husson et al. 2011;Koukol et al. 2012;Gross and Han 2015;Guatimosim et al. 2016Friggens et al. 2017;Crous et al. 2020;Etayo and Silanes 2020;Newsham et al. 2020). ...
... The two genera were monographed by Nag Raj and Kendrick (1975) who accepted 66 species, including some species correctly placed under Thielaviopsis. Since then another 58 new species have been formally described under the two genera (Gams and Holubová-Jechová 1976;Morgan-Jones and Ingram 1976;Arambarri et al. 1981Arambarri et al. , 2007Holubová-Jechová 1982McKenzie 1982McKenzie , 1993McKenzie , 1997Kiffer and Delon 1983;Carris and Glawe 1984;Kirk 1984Kirk , 1985Kirk , 1986Morgan-Jones et al. 1984;Etayo 1996;Subramanian and Sudha 1986;Kile and Walker 1987;Carris 1988;Cabello 1989;Morgan-Jones et al. 1992;Christiansen 1993;Matsushima 1993;Sutton 1993;Kowalski and Halmschlager 1996;Kile et al. 1996;Gadhil and Dick 1999;Taylor et al. 2001;McKenzie et al. 2002;Wu 2004;Kowalski 2006;Shabunin 2007;Koukol 2011;Pratibha et al. 2005;Silva et al. 2015;Crous et al. 2016aCrous et al. , b, 2019Crous et al. , 2021Etayo and Silanes 2020). Species concepts in the two genera are mainly based on limited morphological characters of setae, conidiophores (absent or present, shape, size, color, septation, smooth or verrucose), conidiogenous cells (shape, size, color, venter/collarette ratio), and conidia (shape, size, color, septation, basal frill). ...
... Phylogenetic analyses showed that Chaetochalara and Chalara s. lat. were polyphyletic, while other recently established genera such as Lareunionomyces, Neochalara, Neolauriomyces and Zymochalara were monophyletic (Paulin and Harrington 2000;Cai et al. 2009;Koukol 2011Koukol , 2012Réblová et al. 2011;Guatimosim et al. 2016;Suija and Motiejünaité 2016;Ekanayaka et al. 2019;Crous et al. 2020Crous et al. , 2021Hosoya 2021;Mitchell et al. 2022). In their phylogenetic analyses of chalara-like fungi, Paulin and Harrington (2000) concluded that the 16 Chalara species without known teleomorphs had leotialian affinities, but the relationship among these Chalara specie and among the Leotiales were not resolved. ...
The chalara-like anamorphs of Leotiomycetes are phialidic hyphomycetes with cylindrical collarettes and deeply seated sporulating loci, and hyaline, aseptate or septate, cylindrical conidia. They are commonly found on plant litters in both terrestrial and submerged environments, and with broad geographical distribution. This paper reports our research result of diversity, taxonomy and phylogeny of these fungi in China, which is based on a systematic study by using an integrated approach of literature study, morphological observation and phylogenetic analyses of 153 chalara-like fungal species with diversified morphology in conidiomata, setae, conidiophores, phialides and conidia. The phylogenetic analyses employing different datasets of SSU, LSU and ITS sequences of 116 species showed that these chalara-like fungi were paraphyletic and scattered in 20 accepted genera belonging to five families of Leotiomycetes: Arachnopezizaceae, Hamatocanthoscyphaceae, Helotiaceae, Neolauriomycetaceae and Pezizellaceae. Additional six genera, Ascoconidium , Bioscypha , Chalarodendron , Didonia , Phaeoscypha and Tapesina , all reported with chalara-like anamorphs in literatures, are also accepted as members of Pezizellaceae or Leotiomycetes genera incertae sedis. Among of these 26 accepted genera of chalara-like fungi in Leotiomycetes, 17 genera are asexually typified genera ( Ascoconidium , Bloxamia , Chalara , Chalarodendron , Constrictochalara , Cylindrochalara , Cylindrocephalum , Leochalara , Lareunionomyces , Minichalara , Neochalara , Neolauriomyces , Nagrajchalara , Parachalara , Stipitochalara , Xenochalara and Zymochalara ), and 9 are sexually typified genera ( Bioscypha , Bloxamiella , Calycellina , Calycina , Didonia , Hymenoscyphus , Mollisina , Phaeoscypha and Tapesina ). The phylogenetic significance of conidial septation in generic delimitation was further confirmed; while other morphologies such as conidiomata, setae, conidiophores, phialides, conidial length, and conidial ornamentation have little phylogenetic significance, but could be used for species delimitation. The polyphyletic genus Chalara s. lat. is revised with monophyletic generic concepts by redelimitation of Chalara s. str. in a narrow concept, adaption of the emended Calycina to also include asexually typified chalara-like fungi, reinstatement of Cylindrocephalum , and introduction of six new genera: Constrictochalara W.P. Wu & Y.Z. Diao, Leochalara W.P. Wu & Y.Z. Diao, Minichalara W.P. Wu & Y.Z. Diao, Nagrajchalara W.P. Wu & Y.Z. Diao, Parachalara W.P. Wu & Y.Z. Diao and Stipitochalara W.P. Wu & Y.Z. Diao. Chaetochalara becomes a synonym of Chalara s. str., and the known species are disassembled into Chalara s. str. and Nagrajchalara . The polyphyletic genus Bloxamia is also redefined by introducing the new genus Bloxamiella W.P. Wu & Y.Z. Diao for B . cyatheicola . Five existing species of Chalara s. lat. were excluded from Leotiomycetes and reclassified: Chalara breviclavata as Chalarosphaeria breviclavata W.P. Wu & Y.Z. Diao gen. et sp. nov. in Chaetosphaeriaceae, C . vaccinii as Sordariochalara vaccinii W.P. Wu & Y.Z. Diao gen. et sp. nov. in Lasiosphaeriaceae, and three other Chalara species with hyaline phialides, C . hyalina , C . schoenoplecti and C . siamense as combinations of Pyxidiophora in Pyxidiophoraceae. For biodiversity of these fungi in China, a total of 80 species in 12 genera, including 60 new species, 17 new records and 1 new name, were discovered and documented in this paper. In addition, five species including three new species are reported from Japan. In connection to this revision, a total of 44 new combinations are made. The identification keys are provided for most of these genera. Future research area of these fungi should be the phylogenetic relationship of several sexually typified genera such as Bioscypha , Calycellina , Calycina , Didonia , Phaeoscypha , Rodwayella and Tapesina , and systematic revision of existing names under the genera Bloxamia , Chaetochalara and Chalara .
... Furthermore, OTUs assigned to the genera Mycena and Chalara correlated directly with the activity of litter-degrading CAZymes (Fig. 6). Chalara species live mostly as litter saprotrophs (Koukol 2011), while several Mycena species are reported to switch between saprotrophic and biotrophic lifestyles (Thoen et al. 2020). These results further highlight the robustness of our approach to link fungal identity with ecological processes. ...
In forest ecosystems, decomposition is essential for carbon and nutrient cycling and therefore a key process for ecosystem functioning. During the decomposition process, litter chemistry, involved decomposer organisms, and enzymatic activity change interdependently. Chemical composition of the litter is the most complex and dynamic component in the decomposition process and therefore challenging to assess holistically. In this study, we aimed to characterize chemical shifts during decomposition and link them to changes in decomposer fungal activity. We characterized the chemical composition of freshly fallen autumn leaves of European beech (Fagus sylvatica) and the corresponding leaf litter after 1 year of decomposition by proton nuclear magnetic resonance spectroscopy. We further tested the applicability of spiking experiments for qualitative and quantitative characterization of leaves and litter chemistry. The composition and transcriptional activity of fungal communities was assessed by high-throughput Illumina sequencing in the same litter samples. We were able to distinguish freshly fallen leaves from 1-year-old litter based on their chemical composition. Chemical composition of leaves converged among regions with progressing decomposition. Fungal litter communities differed in composition among regions, but they were functionally redundant according to the expression of genes encoding litter degrading enzymes (CAZymes). Fungi of the saprotrophic genera Mycena and Chalara correlated with transcription of litter-degrading CAZymes in 1-year-old litter. Forestry measures influenced the diversity and transcription rate of the detected CAZymes transcripts in litter. Their expression was primarily predicted by composition of the soluble chemical fraction of the litter. Environmental NMR fingerprints thus proved valuable for inferring ecological contexts. We propose and discuss a holistic framework to link fungal activity, enzyme expression, and chemical composition.
... (DNASTAR, Madison, WI), and then aligned in BioEdit v 7.0.9.0 (Hall 1999). The newly generated sequences were confirmed their identity via NCBI Blastn while reference sequences from related studies (Cantrell & Hanlin 1997, Fukami et al. 2010, Koukol 2011, Han et al. 2014, Baral & Rämä 2015, Guatimosim et al. 2016, Hosoya & Zhao 2016, Suija & MotieJűnaitë 2017, Zhuang et al. 2017, Vu et al. 2019, Mitchell 2021, Senn-Irlett et al. 2021 were retrieved from the GenBank (Table 1). Each data set was aligned using MAFFT v. 7, edited manually using BioEdit v. 7.2 (Hall 1999) Gutiérrez et al. 2009). ...
Calycina-like specimens were collected in Doi Inthanon National Park, Thailand. The combined phylogeny of ITS and LSU based on maximum likelihood and bayesian inference analyses confirmed the taxonomic placement of our specimen in Pezizellaceae. Based on morphology and multi-gene phylogeny, the new discomycete specimen was identified as Calycina montana comb. nov., and was documented for the first time in Thailand. Calycina montana is characterized by the amber color, stipitate apothecia, gelatinized ectal excipulum, cylindrical asci and ellipsoidal ascospores. The morphology was observed together with the type strain, Bisporella montana. Our phylogenetic analyses and observation of the morphology of the type strain also confirmed the new combination of Bisporella shangrilana as Calycina shangrilana comb. nov., which resolves the confusion of Bisporella placement in Pezizellaceae.
... A total of four fungal isolates with sequenced genomes were selected to represent various fungal classes and life strategies: Trichoderma harzianum Rifai-an opportunistic mycoparasite (Weindling 1932) in the class Sordariomycetes, Chalara longipes (Preuss) Cooke-an ubiquitous colonizer of needle litter (Koukol 2011) in the class Leotiomycetes, Lacaria bicolor (Maire) P. D. Orton-an ectomycorrhizal fungus (Martin et al. 2008) in the order Agaricales (Agaricomycetes) and Serpula lacrymans (Wulfen)-a brown-rot wood decomposer (Kauserud et al. 2007) in the order Boletales (Agaricomycetes). Fungal stock cultures were kept on Modified Melin-Norkrans (MMN) agar (Marx 1969) at room temperature in darkness. ...
Fungal metabolic carbon acquisition and its subsequent partitioning between biomass production and respiration, i.e. the carbon-use efficiency (CUE), are central parameters in biogeochemical modelling. However, current available techniques for estimating these parameters are all associated with practical and theoretical shortcomings, making assessments unreliable. Gene expression analyses hold the prospect of phenotype prediction by indirect means, providing new opportunities to obtain information about metabolic priorities. We cultured four different fungal isolates (Chalara longipes, Laccaria bicolor, Serpula lacrymans, and Trichoderma harzianum) in liquid media with contrasting nitrogen availability and measured growth rates and respiration to calculate CUE. By relating gene expression markers to measured carbon fluxes, we identified genes coding for 1,3-β-glucan synthase and 2-oxoglutarate dehydrogenase as suitable markers for growth and respiration, respectively, capturing both intraspecific variation as well as within-strain variation dependent on growth medium. A transcript index based on these markers correlated significantly with differences in CUE between the fungal isolates. Our study paves the way for the use of these markers to assess differences in growth, respiration, and CUE in natural fungal communities, using metatranscriptomic or the RT-qPCR approach.
... A fungus identified as Chalara fungorum was amplified by the Ascomycete-specific primers with high levels of occurrence across many farms; it was significantly associated with high yield sites in farms 6 and 10 and found in high relative abundance in farm 3. The anamorphic genus Chalara represents a polyphyletic assemblage of species with similar morphologies (Koukol, 2011), and the true identity of this OTU remains to be determined. The genus contains some aggressive primary plant pathogens responsible for oak-wilt (conidial stage Chalara quercina) (Raj and Kendrick, 2006), the causal agent of narrowleaved ash dieback (C. ...
... fraxinea) (Kowalski and Holdenrieder, 2009), myrtle beech dieback in Australia (C. australis) (Kile and Walker, 1987), and on Pseudocyphellaria "speckle belly" lichens (C. pseudocyphellariae) (Etayo and Sancho, 2008;Koukol, 2011). However, most C. fungorum observations are as saprobic dematiaceous anamorphs on plant litter-coniferous needles, cones, wood or bark, in temperate and tropical regions, with several recurring on particular substrates (Koukol, 2011). ...
... pseudocyphellariae) (Etayo and Sancho, 2008;Koukol, 2011). However, most C. fungorum observations are as saprobic dematiaceous anamorphs on plant litter-coniferous needles, cones, wood or bark, in temperate and tropical regions, with several recurring on particular substrates (Koukol, 2011). It may also be fungicolous as it was originally described on an old fruiting body of a Hydnellum (Saccardo, 1886). ...
Soil bacteria and fungi are integral parts of healthy ecosystem functioning in production agriculture. The effects of fungal abundance and diversity on crop productivity is poorly understood. We sampled 10 corn farms at the V10 growth stage across southwestern Ontario, Canada, using aerial infrared imaging to identify zones of low and high productive corn plants. Roots and soils were sampled from low and high yield zones and soil physical and chemical properties were measured in conjunction with assessment of the root mycobiome communities using Illumina MiSeq sequencing of 4 rRNA amplicons. Higher crop yields were associated with sites having greater fungal phylogenetic diversity and Fisher's α diversity. Indicator species associated with high and low yield sites within a farm could be identified but there were no shared fungal indicators of productivity differences across farms. Communities largely varied across locations despite crop genetics, demonstrating a major influence of soil texture and chemistry in shaping the mycobiome in a site-specific manner. Across all 4 primers, roots from high-yielding sites shared 35 major OTUs including Penicillium spp., Trichoderma, Chalara fungorum , and Gibellulopsis . Low-yielding sites shared 31 OTUs including Fusarium spp., Pythium, Setophoma terrestris , and Neonectria . Soil physical and chemical parameters that contributed to broad scale differences in yield and mycobiome diversity included: %clay, %sand, %phosphorus saturation, cation exchange capacity, aluminum, pH, iron, potassium, %moisture, organic matter, and chlorine. The results show the importance of physicochemical balance in shaping the relationship between root mycobiome and plant productivity.
... The monotypic genus Hyalodendriella forms pigmented micro-and macroconidiophores directly on hyphae as well as pigmented limoniform to ellipsoid and prominently apiculate conidia ), while Chalara is characterised by forming sessile or stalked phialides with basal ventres, long collarettes and deep-seated conidiogenous loci; conidia are cylindrical and often produced in basipetal chains (Holubová-Jechová 1984;Kowalski 2006). However, Chalara, is highly polyphyletic; species have been placed in different classes of Ascomycota (Paulin and Harrington 2000;Koukol 2011). As the type species belongs to the Sordariomycetes (Chalara fusidioides), strain CBS 880.73 probably does not even belong in Chalara s. str. ...
During a survey on fungi associated with wood necroses of Prunus trees in Germany, strains belonging to the Leotiomycetes and Eurotiomycetes were detected by preliminary analyses of ITS sequences. Multi-locus phylogenetic analyses (LSU, ITS, TUB, EF-1α, depending on genus) of 31 of the 45 strains from Prunus and reference strains revealed several new taxa, including Arboricolonus gen. nov., a new genus in the Helotiales (Leotiomycetes) with a collophorina-like asexual morph. Seven Cadophora species (Heloti-ales, Leotiomycetes) were treated. The 29 strains from Prunus belonged to five species, of which C. luteo-olivacea and C. novi-eboraci were dominating; C. africana sp. nov., C. prunicola sp. nov. and C. ramosa sp. nov. were revealed as new species. The genus Cadophora was reported from Prunus for the first time. Phialophora bubakii was combined in Cadophora and differentiated from C. obscura, which was resurrected. Asexual morphs of two Proliferodiscus species (Helotiales, Leotiomycetes) were described, including one new species, Pr. ingens sp. nov. Two Minutiella species (Phaeomoniellales, Eurotiomycetes) were detected, including the new species M. pruni-avium sp. nov. Prunus avium and P. domestica are reported as host plants of Minutiella.
... by FUNGuild. It is generally considered to be a litter saprotroph (Koukol, 2011), but certain Chalara species can become pathogenic, causing ash dieback (Pautasso, Aas, Queloz, & Holdenrieder, 2013). Some RAF in our study exhibited a similar phenomenon. ...
Root-associated fungi (RAF) are an important factor affecting the host's growth, and their contribution to Pinus sylvestris var. mongolica plantation decline is substantial. Therefore, we selected three age groups of P. sylvestris plantations (26, 33, and 43 years), in the Mu Us Desert, to characterize the community structure and functional groups of RAF, identified by Illumina high-throughput sequencing and FUNGuild platform, respectively. The effects of soil properties and enzyme activities on fungal diversity and functional groups were also examined. The results indicated that (a) 805 operational taxonomic units of RAF associated with P. sylvestris belonged to six phyla and 163 genera. Diversity and richness were not significantly different in the three age groups, but community composition showed significant differences. Ascomycota and Basidiomycota dominated the fungal community, while Rhizopogon dominated in each plot. (b) The proportion of pathotrophs decreased with increasing age, while that of symbiotrophs increased sharply, which were mainly represented by ectomycorrhizal fungi. (c) Stand age and soil enzyme activity had a greater influence on fungal community composition than did soil properties, whereas environmental variables were not significantly correlated with fungal diversity and richness. Dynamics of fungal community composition and functional groups with the aging plantations reflected the growth state of P. sylvestris and were related to plantation degradation.
... and Chalara longipes (Preuss) Cooke are both common colonizers of pine litter but with different ecological strategies and abilities regarding decomposition (Baskaran et al., 2019). G. androsaceus is an efficient decomposer with high ligninolytic capacity (Boberg et al., 2011), whereas C. longipes is characterized by stress tolerance, endophytic capacity (Koukol, 2011) and lack of ligninolytic ability (Boberg et al., 2011). Baskaran et al. (2019) found that in the presence of G. androsaceus, 40% of the needle litter mass was lost during 10 months of incubation, but only 10% in the presence of C. longipes. ...
... Generally, Leotiomycetes (to which C. longipes belongs) have a broader tolerance to constrained nutrient availability and low pH than Agaricomycetes (to which G. androsaceus belongs) (Sterkenburg et al., 2015). Furthermore, C. longipes may colonise living needles as an endophyte (Koukol, 2011), and this versatile ecology may Fig. 4. Needle litter microcosms. The pictures of the litter microcosms have been taken after 10 months in presence of Gymnopus androsaceus (left side, blue) and Chalara longipes (right side, red). ...
Microbial communities interplay with their environment through their functional traits that can be as a response or an effect to the environment. Here we explore how a functional trait – the decomposition of organic matter, can be addressed based on genetic markers and how the expression of these markers reflect ecological strategies of two fungal litter decomposer Gymnopus androsaceus and Chalara longipes. We sequenced the genomes of these two fungi, as well as their transcriptomes at different steps of Pinus sylvestris needles decomposition in microcosms. Our results highlighted that if the gene content of the two species could indicate similar potential decomposition abilities, the expression levels of specific gene families belonging to the glycoside hydrolase (GH) category reflected contrasting ecological strategies. Actually C. longipes, the weaker decomposer in this experiment, turned‐out to have a high content of genes involved in cell wall polysaccharides decomposition but low expression levels, reflecting a versatile ecology compare to the more competitive G. androsaceus with high expression levels of keystone functional genes. Thus we established that sequential expression of genes coding for different components of the decomposer machinery indicated adaptation to chemical changes in the substrate as decomposition progressed. This article is protected by copyright. All rights reserved.
... Foliophoma sp. has already been reported as the most abundant genera in trees infected with V. oleaginea [10]. Like Chalara sp., species within the Foliophoma genus have been described as saprophytes but also as pathogens causing dieback and shoot necrosis in olive [36][37][38]. No such symptoms were observed in the olive trees used in this study, however these two highly detected genera in symptomatic plants, can be a secondary effect of the disease, behaving as opportunistic pathogens and increasing the severity of the infection. ...
Venturia oleaginea and Pseudocercospora cladosporioides are two of the most important olive fungal pathogens causing leaf spots: peacock spot, and cercosporiosis, respectively. In the present study, fungal communities associated with the presence of these pathogens were investigated. Overall, 300 symptomatic and asymptomatic trees from different cultivars were sampled from Alentejo, Portugal. A total of 788 fungal isolates were obtained and classified into 21 OTUs; Ascomycota was clearly the predominant phylum (96.6%). Trees from cultivar ‘Galega vulgar’ showed a significant higher fungal richness when compared to ‘Cobrançosa’, which in turn showed significant higher values than ‘Picual’. Concerning plant health status, symptomatic plants showed significant higher fungal richness, mainly due to the high number of isolates of the pathogens V. oleaginea and P. cladosporioides. In terms of fungal diversity, there were two major groups: ca. 90% of the isolates found in symptomatic plants belonged to V. oleaginea, P. cladosporioides, Chalara sp., and Foliophoma sp. while ca. 90% of the isolates found in asymptomatic plants, belonged to Alternaria sp. and Epicoccum sp. This study highlights the existence of different fungal communities in olive trees, including potential antagonistic organisms that can have a significant impact on diseases and consequently on olive production.
... In that study, we showed that litter-bags dominated by Mycena had higher decomposition rates, which in turn could reduce carbon sequestration (Asplund et al. 2018). Lophodermium and Chalara include several species that are commonly found on coniferous litter or as needle endophytes, and they are thus expected to be more abundant below spruce (Korkama-Rajala, Müller and Pennanen 2007;Koukol 2011). In line with our findings, litter samples have been found to have a higher percentage of tree species-specific OTUs than soil samples (Urbanová,Šnajdr and Baldrian 2015;Bahnmann et al. 2018). ...
Beech forests reaches its native distribution limit in SE Norway, but is expected to expand substantially northwards due to climate warming. This may potentially result in a fundamental transformation of contemporary Northern European forests, with tentative effects on the associated belowground fungi. Fungal communities mediate vital ecosystem processes such as ecosystem productivity and carbon sequestration in boreal forests. To investigate how soil fungi is affected by the vegetation transition from spruce to beech forest, we sampled litter, humus and mineral soil in a forest landscape dominated by beech, spruce or a mixture of these. The fungal communities in the soil samples were analyzed by DNA metabarcoding of the rDNA ITS2 region. Although soil layers were the most important structuring gradient, we found clear differences in fungal species composition between spruce and beech plots. The differences in fungal community composition were most evident in the litter and least in the mineral soil. Decomposers, most notably Mycena, dominated the litter layer while various mycorrhizal fungi dominated the humus and mineral layers. Some ectomycorrhizal taxa, such as Cenoccocum and Russula, were more abundant in spruce forests. Differences in fungal community composition between forest types can potentially have large impacts on carbon sequestration rates.
