Spatial and seasonal changes in species diversity of epilithic fungi along environmental gradients of a river

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Wood decomposing abilities of diverse lignicolous fungi on nondecayed and decayed beech wood

We tested the decay abilities of 28 isolates from 28 lignicolous fungal species (Basidiomycota, Ascomycota and Zygomycota) with the pure culture test. We used beech wood powder in varying moisture conditions and decay stages (nondecayed, intermediately decayed and well decayed) as substrates. The weight loss in wood powder was -0.2-17.8%. Five isolates of Basidiomycota (Bjerkandera adusta, Mycena haematopus, Omphalotus guepiniformis, Trametes hirsuta, Trametes versicolor) caused high weight losses in nondecayed wood. We detected significant effects of decay stage on weight loss in wood in most isolates tested, whereas moisture content rarely had an effect on weight loss. Among Basidiomycota and Xylariaceae in Ascomycota weight loss was greater for nondecayed wood than for intermediately and well decayed wood. In contrast four isolates in Ascomycota (Scytalidium lignicola, Trichoderma hamatum, T. harzianum, T. koningii) caused substantial weight loss in intermediately and well decayed wood, although they rarely caused weight loss in nondecayed wood. Zygomycota caused low weight loss in wood. Wood decay stages also affected decomposition of wood chemical components. Acid-unhydrolyzable residue (AUR) decomposition was reduced, whereas holocellulose decomposition was stimulated by some strains of Basidiomycota and Ascomycota in well decayed wood. T. harzianum in particular caused significant weight loss of holocellulose in well decayed wood, although this fungus caused negligible weight loss of both AUR and holocellulose in nondecayed wood. We discuss these changes in the decay patterns of AUR and holocellulose with varying wood decay stages in relation to the role of fungal decomposition of woody debris in forests.     

DNA分子标记技术在真菌系统学研究中的应用及影响

DNA分子标记技术为真菌系统进化研究提供了许多新的方法,真菌分子系统学已成为一门成熟的学科。简述了真菌分子系统学的发展简史和代表性的研究方法以及对真菌系统学的主要贡献,包括将广义的真菌划分为3个类群,粘菌和卵菌不再属于真菌界成员。真菌生命之树项目的研究结果对真菌界高阶分类系统作出重大调整,将先前的4个门(壶菌门、接合菌门、子囊菌门和担子菌门)变为7个门(微孢子虫门、壶菌门、新丽鞭毛菌门、芽枝霉门、球囊菌门、子囊菌门和担子菌门)和4个亚门,并对真菌各类群概念作出修订。此外,DNA分子标记技术对真菌种概念的认识、有性型-无性型关联及分子生态学等研究领域产生了重要影响。     

Distinct seasonality of chytrid-dominated benthic fungal communities in the neritic oceans (Bohai Sea and North Yellow Sea)

Benthic fungal diversities in Bohai Sea and North Yellow Sea were investigated using pyrosequencing of 18S rDNA. Overall, Chytridiomycota dominated, followed by Basidiomycota, Ascomycota and Cryptomycota, in terms of alpha diversities and relative abundance. The beta diversity of benthic fungi showed a significant seasonality but no regional differences, accounted for by contrasting relative abundances of Chytridiomycota and Basidiomycota. Significantly seasonal changes in Chytridiomycota and Basidiomycota assemblage structure were also observed, but not for Ascomycota and Cryptomycota. Environmental filtering was more important than water depth and geographic distance in shaping the distribution of benthic fungi in the neritic oceans. The overall fungal beta diversity co-varied with concentration of chlorophyll a, pH, and salinity, distance from land, and water depth. The assemblage structure of benthic Chytridiomycota, Basidiomycota, Ascomycota and Cryptomycota co-varied with different sets of environmental parameters, suggesting their niche differentiations in the coastal sediments.     

贝加尔针茅草原土壤真菌群落结构对氮素和水分添加的响应

研究氮沉降和降雨变化对土壤真菌群落结构的互作效应,对未来预测多个气候变化因子对草地生态系统的交互作用具有重要意义。以施氮和灌溉模拟氮沉降和降雨增加,采用裂区设计,应用高通量测序技术,研究8个氮添加水平(0、15、30、50、100、150、200、300 kg N hm~(-2)a~(-1))和2个水分添加水平(不灌溉、模拟夏季增雨100 mm灌溉)对土壤真菌群落结构的影响。结果表明,氮素和水分添加后,土壤真菌群落中占优势的门类分别为接合菌门Zygomycota(22.0%—48.9%)、担子菌门Basidiomycota(7.8%—18.5%)、子囊菌门Ascomycota(9.4%—20.1%)、球囊菌门Glomeromycota(0.7%—3.1%)、壶菌门Chytridiomycota(0.1%—1.3%)。常规降雨条件下,随着氮添加水平升高,接合菌门相对丰度呈现出先升高后降低的趋势,N50处理最高;子囊菌门相对丰度在高氮添加时(N100—N300)呈升高趋势。而在氮素和水分同时添加条件下,随着氮添加水平升高,接合菌门相对丰度呈降低趋势,子囊菌门相对丰度变化则不明显。在相同的氮添加水平下,水分添加使接合菌门相对丰度增加,而担子菌门、子囊菌门、球囊菌门和壶菌门的相对丰度降低。在不同氮素和水分添加条件下,有5个土壤真菌门类11个真菌纲相对丰度变化显著。接合菌门的Mortierella属,担子菌门的Entolomataceae科和Geastrum属相对丰度变化极显著,可作为土壤真菌群落结构变化的指示种。PCo A分析结果也表明氮素和水分添加改变了土壤真菌群落结构。植物-土壤-微生物系统的结构方程模型结果表明,植物群落组成及植物物种丰富度的变化是土壤真菌群落结构发生变化的主要影响因素,土壤无机氮及p H的变化主要通过影响植物群落间接影响真菌群落,其对真菌群落的直接影响则较小。综上,氮素和水分添加改变了土壤真菌群落结构,且两者存在明显的互作效应,水分添加可改变氮添加对土壤真菌群落的影响。     

西藏真菌资源调查概况

西藏是我国生物多样性最丰富的地区之一,地形复杂,生境多样,孕育了极为丰富的真菌资源。本文归纳统计出西藏真菌总计185科551属2 599种,其中包括真菌高阶分类单元壶菌门12种、接合菌门20种、球囊菌门70种、子囊菌门764种和担子菌门1 733种,并对各门进行了物种数量分析,列出了优势科。西藏大型真菌资源丰富,共统计出食用菌468种、药用菌243种和毒菌135种,对这些代表类群进行了概述。在此基础上展望西藏真菌资源调查以及生物多样性研究的发展趋势,为相关工作提供参考。     

Potential bias of fungal 18S rDNA and internal transcribed spacer polymerase chain reaction primers for estimating fungal biodiversity in soil

Four fungal 18S rDNA and internal transcribed spacer (ITS) polymerase chain reaction (PCR) primer pairs were tested for their specificity towards target fungal DNA in soil DNA extracts, and their ability to assess the diversity of fungal communities in a natural grassland soil was compared. Amplified PCR products were cloned, and approximately 50 clones from each library were sequenced. Phylogenetic analysis and database searches indicated that each of the sequenced cloned DNA fragments was of fungal origin for each primer pair, with the exception of the sequences generated using the 18S rDNA primers nu-SSU-0817 and nu-SSU-1196, where 35 of the 50 sequenced clones represented soil invertebrates. Although some of the primers have previously been suggested to be biased towards certain fungal taxonomic groups, the ratio of sequences representing each of the four main fungal phyla, Ascomycota, Basidiomycota, Chytridiomycota and Zygomycota, was similar for each of the primer pairs, suggesting that primer bias may be less significant than previously thought. Collector's curves were plotted to estimate the coverage obtained for each of the clone libraries after clustering the sequences into operational taxonomic units at a level of 99% sequence similarity. The curves indicated that good coverage of diversity was achieved, with the exception of the clone library constructed using primers nu-SSU-0817 and nu-SSU-1196, on account of the high number of non-fungal sequences obtained. The work demonstrates the usefulness of 18S rDNA and ITS PCR primers for assessing fungal diversity in environmental samples, and it also highlights some potential limitations of the approach with respect to PCR primer specificity and bias.     

Seagrasses,seaweeds and plant debris: An extraordinary reservoir of fungal diversity in the Mediterranean Sea

This review gathers data derived from many research efforts on marine fungi associated with plant-origin substrates in the Mediterranean Sea. Overall, the review draws up a list of 378 taxa associated with seagrasses, seaweeds and wood.For each of the three substrates, on average, 92.7% of the taxa belonged to the phylum Ascomycota. Basidiomycota were better represented in seagrasses (9.8%) than in seaweeds (4.9%) and wood (1.2%). Mucoromycota and Mortirellomycota were scarce, while Chytridiomycota was detected only in association with phanerogames (3.7%). Dothideomycetes and Sordariomycetes were the dominant classes, while the orders Pleosporales and Hypocreales were significantly represented in the three sububstrates (42, 37, 10 taxa-30, 40, 16 taxa). Seagrasses with 210 associated taxa were the substrates with the richest fungal communities, immediately followed by seaweeds (180 taxa) and finally by wood (78 taxa). Out of the total, only 12 taxa were shared by the three varieties of substrates, including species that were widespread in marine environments. However, many algal species and seagrasses inhabiting the Mediterranean Sea remain unexplored. This gap indicates the need to apply more extensive surveys to explain the huge fungal biodiversity herein hosted, and increase the chances of describing novel fungal lineages.     

Loss of the flagellum happened only once in the fungal lineage: phylogenetic structure of Kingdom Fungi inferred from RNA polymerase II subunit genes

Background  

At present, there is not a widely accepted consensus view regarding the phylogenetic structure of kingdom Fungi although two major phyla, Ascomycota and Basidiomycota, are clearly delineated. Regarding the lower fungi, Zygomycota and Chytridiomycota, a variety of proposals have been advanced. Microsporidia may or may not be fungi; the Glomales (vesicular-arbuscular mycorrhizal fungi) may or may not constitute a fifth fungal phylum, and the loss of the flagellum may have occurred either once or multiple times during fungal evolution. All of these issues are capable of being resolved by a molecular phylogenetic analysis which achieves strong statistical support for major branches. To date, no fungal phylogeny based upon molecular characters has satisfied this criterion.     

Increasing ecological inference from high throughput sequencing of fungi in the environment through a tagging approach

High throughput sequencing methods are widely used in analyses of microbial diversity, but are generally applied to small numbers of samples, which precludes characterization of patterns of microbial diversity across space and time. We have designed a primer-tagging approach that allows pooling and subsequent sorting of numerous samples, which is directed to amplification of a region spanning the nuclear ribosomal internal transcribed spacers and partial large subunit from fungi in environmental samples. To test the method for phylogenetic biases, we constructed a controlled mixture of four taxa representing the Chytridiomycota, Zygomycota, Ascomycota and Basidiomycota. Following cloning and colony restriction fragment length polymorphism analysis, we found no significant difference in representation in 19 of the 23 tested primers. We also generated a clone library from two soil DNA extracts using two primers for each extract and compared 456 clone sequences. Community diversity statistics and contingency table tests applied to counts of operational taxonomic units revealed that the two DNA extracts differed significantly, while the pairs of tagged primers from each extract were indistinguishable. Similar results were obtained using UniFrac phylogenetic comparisons. Together, these results suggest that the pig-tagged primers can be used to increase ecological inference in high throughput sequencing projects on fungi.     

天山一号冰川前沿生态系统真菌群落结构演替及分布格局

【目的】分析真菌群落结构和多样性随着一号冰川退缩前沿年代序列的变化,揭示真菌群落的演替轨迹及环境因子对群落组成的影响。【方法】采用宏基因组学研究方法,结合生物信息学和统计学分析技术,对取自一号冰川末端表面冰尘,底部和前沿14个样品进行总DNA的提取,ITS基因的扩增并使用Illumina Miseq平台测序,通过相关生物地理化学特性综合分析在不同年代序列下真菌群落结构及其演替规律。【结果】经测序,筛选和质控分析获得185103条rawreads,占78.3%的非单序列在97%的相似度聚类分析共得到300个操作分类单元(OTU),共划分为6个门:子囊菌门(Ascomycota,52.7%)、担子菌门(Basidiomycota,16.9%)、壶菌门(Chytridiomycota,15.1%)、接合菌门(Zygomycota,2.4%)和球囊菌门(Glomeromycota,1.2%)。从演替初期到后期阶段虽然子囊菌的序列数逐渐下降而担子菌出现缓慢上升趋势,但子囊菌随着土壤年代序列的增加始终为优势类群,壶菌在冰川底部和前沿基层普遍存在且丰度仅次于子囊菌和担子菌。我们在缺乏植被的最新退缩基层发现依靠自养型宿主存活的活体营养菌,如Taphrinomycetes、Urediniomycetes和Ustilaginomycetes。从冰川底部和前沿基层检测到丰度较高的酵母菌,而粪生真菌(coprophilous fungi)仅仅出现在冰川前沿基层,共23个操作分类单元。球囊菌仅在前沿部分样品中存在,有着十分狭小的生态位分布。【结论】一号冰川前沿随着年代序列的增加真菌群落存在明显的演替轨迹和多样性的显著变化,不同生态位真菌类群组成的相似性较低且都存在明显的指示性真菌类群。     

《中国真菌志》50年编研成果及展望

历经几代真菌人的艰苦奋斗,《中国真菌志》目前已出版65卷,其中子囊菌类有35卷、担子菌类26卷、接合菌类1卷、卵菌1卷、黏菌2卷;上述卷册记录923属9 228种及种下分类群。编研涉及大型和小型的类群、腐生菌、食药用菌、菌根真菌、作物和林木病原菌、捕食性真菌、虫生菌、菌生真菌以及毒菌等。编研过程中发现并发表了大量新分类群,丰富了对菌物物种多样性的认知。归纳已有成果,已立项卷册应加快编研和出版进度;资源调查与分类学研究需进一步加强,按分类群(专科、专属)继续开展编研工作,注重拟参编类群的前期研究积累;以下分类群值得在后续工作中予以关注：水霉目等部分卵菌、壶菌类、座囊菌纲部分类群、盘菌纲和锤舌菌纲中尚未参编的主要类群、伞菌纲中具有小型子实体的类群等。由于分类系统在不断更新和完善,已出版卷册中使用的部分名称和分类地位的处理需依据被大多数学者所公认的分类系统给予适时的订正。期待建设一支继往开来的菌物分类学研究队伍,不断取得令世人瞩目的编研成果,为我国菌物资源利用和物种保护提供科学依据。     

Determining diversity of freshwater fungi on decaying leaves: comparison of traditional and molecular approaches

Traditional microscope-based estimates of species richness of aquatic hyphomycetes depend upon the ability of the species in the community to sporulate. Molecular techniques which detect DNA from all stages of the life cycle could potentially circumvent the problems associated with traditional methods. Leaf disks from red maple, alder, linden, beech, and oak as well as birch wood sticks were submerged in a stream in southeastern Canada for 7, 14, and 28 days. Fungal biomass, estimated by the amount of ergosterol present, increased with time on all substrates. Alder, linden, and maple leaves were colonized earlier and accumulated the highest fungal biomass. Counts and identifications of released conidia suggested that fungal species richness increased, while community evenness decreased, with time (up to 11 species on day 28). Conidia of Articulospora tetracladia dominated. Modifications of two molecular methods-denaturing gradient gel electrophoresis (DGGE) and terminal restriction fragment length polymorphism (T-RFLP) analysis-suggested that both species richness and community evenness decreased with time. The dominant ribotype matched that of A. tetracladia. Species richness estimates based on DGGE were consistently higher than those based on T-RFLP analysis and exceeded those based on spore identification on days 7 and 14. Since traditional and molecular techniques assess different aspects of the fungal organism, both are essential for a balanced view of fungal succession on leaves decaying in streams.     

Determining Diversity of Freshwater Fungi on Decaying Leaves: Comparison of Traditional and Molecular Approaches

Traditional microscope-based estimates of species richness of aquatic hyphomycetes depend upon the ability of the species in the community to sporulate. Molecular techniques which detect DNA from all stages of the life cycle could potentially circumvent the problems associated with traditional methods. Leaf disks from red maple, alder, linden, beech, and oak as well as birch wood sticks were submerged in a stream in southeastern Canada for 7, 14, and 28 days. Fungal biomass, estimated by the amount of ergosterol present, increased with time on all substrates. Alder, linden, and maple leaves were colonized earlier and accumulated the highest fungal biomass. Counts and identifications of released conidia suggested that fungal species richness increased, while community evenness decreased, with time (up to 11 species on day 28). Conidia of Articulospora tetracladia dominated. Modifications of two molecular methods—denaturing gradient gel electrophoresis (DGGE) and terminal restriction fragment length polymorphism (T-RFLP) analysis—suggested that both species richness and community evenness decreased with time. The dominant ribotype matched that of A. tetracladia. Species richness estimates based on DGGE were consistently higher than those based on T-RFLP analysis and exceeded those based on spore identification on days 7 and 14. Since traditional and molecular techniques assess different aspects of the fungal organism, both are essential for a balanced view of fungal succession on leaves decaying in streams.     

Diversity and antimicrobial activities of the fungal endophyte community associated with the traditional Brazilian medicinal plant Solanum cernuum Vell. (Solanaceae)

The diversity and antimicrobial activity of endophytic fungi associated with the Brazilian medicinal plant Solanum cernuum Vell. were studied during summer and winter seasons. A total of 246 fungal isolates were obtained, including 225 filamentous fungi and 21 yeasts. They were identified by morphological, physiological, and molecular methods. Fifty-five different taxa represented by the phyla Ascomycota (33 taxa), Basidiomycota (21 taxa), and Zygomycota (one taxon) were identified. The most abundant taxa were closely related to Arthrobotrys foliicola , Colletotrichum gloeosporioides , Coprinellus radians , Glomerella acutata , Diatrypella frostii , Phoma glomerata , Mucor sp., Phlebia subserialis , Phoma moricola , Phanerochaete sordida , and Colletotrichum sp. A total of 265 fungal extracts were screened and 64 (26.01%) displayed antimicrobial activities. Among these extracts, 18 (28.12%) presented antibacterial and antifungal activities, 42 (65.62%) displayed selective antibacterial activity, and four (6.25%) exhibited only antifungal activity. The best values of minimum inhibitory concentration were obtained from extracts of Cryptococcus rajasthanensis , Glomerella acutata, Leptosphaeria sp., and Phoma glomerata ranging from 7.8 to 15.62?μg/mL. This study is the first survey of the endophytic fungi community associated with S. cernuum, and our results show that they can represent a promising source of bioactive compounds.     

Manganese superoxide dismutase in pathogenic fungi: an issue with pathophysiological and phylogenetic involvements

Manganese-containing superoxide dismutases (MnSODs) are ubiquitous metalloenzymes involved in cell defence against endogenous and exogenous reactive oxygen species. In fungi, using this essential enzyme for phylogenetic analysis of Pneumocystis and Ganoderma genera, and of species selected among Ascomycota, Basidiomycota and Zygomycota, provided interesting results in taxonomy and evolution. The role of mitochondrial and cytosolic MnSODs was explored in some pathogenic Basidiomycota yeasts (Cryptococcus neoformans var. grubii, Cryptococcus neoformans var. gattii, Malassezia sympodialis), Ascomycota filamentous fungi (Aspergillus fumigatus), and Ascomycota yeasts (Candida albicans). MnSOD-based phylogenetic and pathogenic data are confronted in order to evaluate the roles of fungal MnSODs in pathophysiological mechanisms.     

Evolutionary relationships within the fungi: analyses of nuclear small subunit rRNA sequences.

Nucleotide sequences of the small subunit ribosomal RNA (18S) gene were used to investigate evolutionary relationships within the Fungi. The inferred tree topologies are in general agreement with traditional classifications in the following ways: (1) the Chytridiomycota and Zygomycota appear to be basal groups within the Fungi. (2) The Ascomycota and Basidiomycota are a derived monophyletic group. (3) Relationships within the Ascomycota are concordant with traditional orders and divide the hemi- and euascomycetes into distinct lineages. (4) The Basidiomycota is divided between the holobasidiomycetes and phragmobasidiomycetes. Conflicts with traditional classification were limited to weakly supported branches of the tree. Strongly supported relationships were robust to minor changes in alignment, method of analysis, and various weighting schemes. Weighting, either of transversions or by site, did not convincingly improve the status of poorly supported portions of the tree. The rate of variation at particular sites does not appear to be independent of lineage, suggesting that covariation of sites may be an important phenomenon in these genes.     

Isolation of wood-inhabiting fungi from Canadian hardwood logs

Wood-inhabiting fungi include many molds, wood-staining fungi, and decay fungi. Most of these fungal species can result in economic losses to wood users. Studies on molds, staining fungi, and decay fungi are necessary to be able to control their growth on wood and wood products. In this study, wood-inhabiting fungi were isolated from logs of 3 major Canadian hardwood species: sugar maple, white birch, and yellow birch. Two media were used for isolation. From these 3 wood species, a total of 1198 fungal cultures were obtained from summer- and winter-harvested logs in dry storage and under water sprinkling. The results showed that most fungal species were not host specific and affected all of the wood species tested. Frequently isolated molds were Alternaria alternata, Trichoderma species, and Mucor/Rhizopus (Zygomycota) species, frequently isolated staining fungi were Ophiostoma piceae and Ophiostoma piliferum, a frequently isolated bark saprophyte was Nectria cinnabarina, and frequently isolated decay fungi were taxa of the phylum Basidiomycota. More fungal species were isolated from summer-harvested logs than from winter-harvested logs. Fewer fungal cultures, especially decay fungi, were isolated from logs in early storage than from logs in late storage.     

Assembling the fungal tree of life: progress, classification, and evolution of subcellular traits

Based on an overview of progress in molecular systematics of the true fungi (Fungi/Eumycota) since 1990, little overlap was found among single-locus data matrices, which explains why no large-scale multilocus phylogenetic analysis had been undertaken to reveal deep relationships among fungi. As part of the project "Assembling the Fungal Tree of Life" (AFTOL), results of four Bayesian analyses are reported with complementary bootstrap assessment of phylogenetic confidence based on (1) a combined two-locus data set (nucSSU and nucLSU rDNA) with 558 species representing all traditionally recognized fungal phyla (Ascomycota, Basidiomycota, Chytridiomycota, Zygomycota) and the Glomeromycota, (2) a combined three-locus data set (nucSSU, nucLSU, and mitSSU rDNA) with 236 species, (3) a combined three-locus data set (nucSSU, nucLSU rDNA, and RPB2) with 157 species, and (4) a combined four-locus data set (nucSSU, nucLSU, mitSSU rDNA, and RPB2) with 103 species. Because of the lack of complementarity among single-locus data sets, the last three analyses included only members of the Ascomycota and Basidiomycota. The four-locus analysis resolved multiple deep relationships within the Ascomycota and Basidiomycota that were not revealed previously or that received only weak support in previous studies. The impact of this newly discovered phylogenetic structure on supraordinal classifications is discussed. Based on these results and reanalysis of subcellular data, current knowledge of the evolution of septal features of fungal hyphae is synthesized, and a preliminary reassessment of ascomal evolution is presented. Based on previously unpublished data and sequences from GenBank, this study provides a phylogenetic synthesis for the Fungi and a framework for future phylogenetic studies on fungi.     

Organic chemical and nutrient dynamics in decomposing beech leaf litter in relation to fungal ingrowth and succession during 3-year decomposition processes in a cool temperate deciduous forest in Japan

Decomposition processes of beech leaf litter were studied over a 3-year period in a cool temperate deciduous forest in Japan. Organic chemical and nutrient dynamics, fungal biomass and succession were followed on upper (Moder) and lower (Mull) of a forest slope. Litter decomposition rates were similar between the sites. Nutrient dynamics of the decomposing litter was categorized into two types: weight changes in nitrogen and phosphorus showed two phases, the immobilization (0–21 months) and the mobilization phase (21–35 months), while those in potassium, calcium and magnesium showed only the mobilization phase. The rate of loss of organic chemical constituents was lignin < holocellulose < soluble carbohydrate < polyphenol in order. The changes in lignocellulose index (LCI), the ratio of holocellulose in lignin and holocellulose, were significantly correlated to the changes in nitrogen and phosphorus concentrations during the decomposition. During the immobilization phase, increase in total fungal biomass contributed to the immobilization of nitrogen and phosphorus. The percentage of clamp-bearing fungal biomass (biomass of the Basidiomycota) to total fungal biomass increased as the decomposition proceeded and was significantly correlated to LCI. Two species in the xylariaceous Ascomycota were dominantly isolated by the surface sterilization method from decomposing litter collected in the 11th month. The organic chemical, nitrogen and phosphorus dynamics during the decomposition were suggested to be related to the ingrowth, substrate utilization and succession of the Xylariaceae and the Basidiomycota. Twenty-one species in the other Ascomycota and the Zygomycota isolated by the washing method were classified into three groups based on their occurrence patterns: primary saprophytes, litter inhabitants and secondary sugar fungi. These species showed different responses to LCI and soluble carbohydrate concentration of the litter between the groups.