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Background and Aims Pinus cembra represent a typical and important tree species growing in European subalpine and alpine habitats. The ectomycorrhizal (ECM) fungal communities associated to this tree under natural conditions are largely unknown. Methods In this study, we investigated the ECM fungal abundance and composition at four high-altitude sites (two northern-exposed and two southern-exposed habitats) in South Tyrol (Italy), and included also two different age classes of P. cembra. The ECM partners were characterized morphologically, and identified by rDNA ITS sequence analysis. Results The degree of mycorrhization in adult P. cembra was typically 100% in these natural habitats, with a total species diversity of 20 ECM species. The four high-altitude sites were similar concerning their species richness and mycobiont diversity, but they differed significantly in ECM species composition. Young P. cembra had a mycorrhization degree of 100% and a total of 10 species were observed. All mycorrhizal partners of naturally regenerated young P. cembra were only detected in one specific location, with the exception of Cenococcum sp. and Amphinema sp. which were detected at two sites. Young trees harbour a distinct ectomycorrhizal fungal diversity, which is clearly lower than the diversity detected in adult P. cembra trees. The P. cembra bolete (Suillus plorans) is the most important symbiotic partner of P. cembra at Southern Tyrolean high-altitude sites and is known for its strict, species-specific host association. Conclusions The ectomycorrhizal fungal community composition strongly depends on geographic region and on the slope exposure (north or south) of the site.

Alpine habitats are one of the most vulnerable ecosystems to environmental change, however, little information is known about the drivers of plant-fungal interactions in these ecosystems and their resilience to climate change. We investigated the influence of the main drivers of ectomycorrhizal (EM) fungal communities along elevation and environmental gradients in the alpine zone of the European Alps and measured their degree of specialisation using network analysis. We sampled ectomycorrhizas of Dryas octopetala, Bistorta vivipara and Salix herbacea, and soil fungal communities at 28 locations across five countries, from the treeline to the nival zone. We found that: (1) EM fungal community composition, but not richness, changes along elevation, (2) there is no strong evidence of host specialisation, however, EM fungal networks in the alpine zone and within these, EM fungi associated with snowbed communities, are more specialised than in other alpine habitats, (3) plant host population structure does not influence EM fungal communities, and (4) most variability in EM fungal communities is explained by fine-scale changes in edaphic properties, like soil pH and total nitrogen. The higher specialisation and narrower ecological niches of these plant-fungal interactions in snowbed habitats make these habitats particularly vulnerable to environmental change in alpine ecosystems.

Aboveground species richness patterns of vascular plants, aphyllophoroid macrofungi, bryophytes and lichens were compared along an altitudinal gradient (80&ndash, 310 m a.s.l.) on the Slantsevaya mountain at the eastern macroslope of the Polar Urals (Russia). Five altitudinal levels were included in the study: (1) Northern boreal forest with larch-spruce in the Sob&rsquo, river valley habitats, (2&ndash, 3) two levels of closed, northern boreal, larch-dominated forests on the slopes, (4) crook-stemmed forest, (5) tundra habitats above the timberline. Vascular plant or bryophyte species richness was not affected by altitudinal levels, but lichen species richness significantly increased from the river valley to the tundra. For aphyllophoroid macrofungi, species richness was highest at intermediate and low altitudes, and poorest in the tundra. These results indicate a positive ecotone effect on aphyllophoroid fungal species richness. The species richness of aphyllophoroid fungi as a whole was neither correlated to mortmass stocks, nor to species richness of vascular plants, but individual ecological or morphological groups depended on these parameters. Poroid fungal species richness was positively correlated to tree age, wood biomass and crown density, and therefore peaked in the middle of the slope and at the foot of the mountain. In contrast, clavarioid fungal species richness was negatively related to woody bio- and mortmass, and therefore peaked in the tundra. This altitudinal level was characterized by high biomass proportions of lichens and mosses, and by high litter mortmass. The proportion of corticoid fungi increased with altitude, reaching its maximum at the timberline. Results from the different methods used in this work were concordant, and showed significant patterns. Tundra communities differ significantly from the forest communities, as is also confirmed by nonmetric multidimensional scaling (NMDS) analyses based on the spectrum of morphological and ecological groups of aphyllophoroid fungi.

The boletoid genera Butyriboletus and Exsudoporus have recently been suggested by some researchers to constitute a single genus, and Exsudoporus was merged into Butyriboletus as a later synonym. However, no convincing arguments have yet provided significant evidence for this congeneric placement. In this study, we analyze material from Exsudoporus species and closely related taxa to assess taxonomic and phylogenetic boundaries between these genera and to clarify species delimitation within Exsudoporus. Outcomes from a multilocus phylogenetic analysis (ITS, nrLSU, tef1-α and rpb2) clearly resolve Exsudoporus as a monophyletic, homogenous and independent genus that is sister to Butyriboletus. An accurate morphological description, comprehensive sampling, type studies, line drawings and a historical overview on the nomenclatural issues of the type species E. permagnificus are provided. Furthermore, this species is documented for the first time from Israel in association with Quercus calliprinos. The previously described North American species Exsudoporus frostii and E. floridanus are molecularly confirmed as representatives of Exsudoporus, and E. floridanus is epitypified. The eastern Asian species Leccinum rubrum is assigned here to Exsudoporus based on molecular evidence, and a new combination is proposed. Sequence data from the original material of the Japanese Boletus kermesinus were generated, and its conspecificity with L. rubrum is inferred as formerly presumed based on morphology. Four additional cryptic species from North and Central America previously misdetermined as either B. frostii or B. floridanus are phylogenetically placed but remain undescribed due to the paucity of available material. Boletus weberi (syn. B. pseudofrostii) and Xerocomus cf. mcrobbii cluster outside of Exsudoporus and are herein assigned to the recently described genus Amoenoboletus. Biogeographic distribution patterns are elucidated, and a dichotomous key to all known species of Exsudoporus worldwide is presented.

Ectomycorrhizae (EM) are important for the survival of seedlings and trees, but how they will react to global warming or changes in soil fertility is still in question. We tested the effect of soil temperature manipulation and nitrogen fertilization on EM communities in a high-altitude Pinus cembra afforestation. The trees had been inoculated in the 1960s in a nursery with a mixture of Suillus placidus, S. plorans and S. sibircus. Sampling was performed during the third year of temperature manipulation in June and October 2013. Root tips were counted, sorted into morphotypes, and sequenced. Fungal biomass was measured as ergosterol and hyphal length. The EM potential of the soil was assessed with internal transcribed spacers (ITS) clone libraries from in-growth mesh bags (MB). Temperature manipulation of ± 1 °C had no effect on the EM community. A total of 33 operational taxonomic units (OTUs) were identified, 20 from the roots, 13 from MB. The inoculated Suillus spp. colonized 82% of the root tips, thus demonstrating that the inoculation was sustainable. Nitrogen fertilization had no impact on the EM community, but promoted depletion in soil organic matter, and caused a reduction in soil fungal biomass.

Apple replant disease (ARD) is a frequently occurring plant disease, which causes retarded growth and mortality of young apple trees in replanted orchards. The aetiology is not well understood, but soil-borne micro-organisms are often discussed as primary causal agents of the replant problem. A greenhouse study was conducted in Laimburg, Italy, with orchard soils from the region, with the aim of obtaining information about the influence of soil biotic and abiotic factors on the aetiology of the disease. Apple rootstocks (M9) were planted into soils cultivated with apple trees that were either fumigated with chloropicrin or not fumigated, as well as mixtures of fumigated and non-fumigated soils. In addition, uncultivated soils (from the inter-row, from a fallow plot and from a meadow) were taken as controls. Various parameters were measured after 62 days in a controlled pot assay. Soils fumigated with chloropicrin resulted in higher apple shoot growth and lower microbial biomass carbon than non-fumigated soils. Uncultivated soils had generally the highest microbial biomass carbon and the highest ergosterol contents. No considerable differences between basal respiration, ergosterol content, pH, electrical conductivity, and most nutrient and metal contents were observed between fumigated and non-fumigated soils. Denaturing gradient gel electrophoresis gels of DNA extracted from the soils revealed differences in the fungal, bacterial and actinobacterial communities of the different soils, indicating significant shifts in microbial community composition after chloropicrin treatment. This study indicates biotic factors in soil to be a causal agent of apple replant disease.

Three closely related species in Alnicola sect. Alnicola, Alnicola, are described, illustrated, and compared. Alnicola badia is revised based on the type collection and recent collections made under Alnus alnobetula. Two species (A. longicystis, A. xanthophylla) associated with Alnus glutinosa and A. incana and locally frequent in western and central Europe are described as new.

Fungal pure cultures identified with both classical morphological methods and through barcoding sequences are a basic requirement for reliable reference sequences in public databases. Improved techniques for an accelerated DNA barcode reference library construction will result in considerably improved sequence databases covering a wider taxonomic range. Fast, cheap, and reliable methods for obtaining DNA sequences from fungal isolates are, therefore, a valuable tool for the scientific community. Direct colony PCR was already successfully established for yeasts, but has not been evaluated for a wide range of anamorphic soil fungi up to now, and a direct amplification protocol for hyphomycetes without tissue pre-treatment has not been published so far. Here, we present a colony PCR technique directly from fungal hyphae without previous DNA extraction or other prior manipulation. Seven hundred eighty-eight fungal strains from 48 genera were tested with a success rate of 86%. PCR success varied considerably: DNA of fungi belonging to the genera Cladosporium, Geomyces, Fusarium, and Mortierella could be amplified with high success. DNA of soil-borne yeasts was always successfully amplified. Absidia, Mucor, Trichoderma, and Penicillium isolates had noticeably lower PCR success.

Several collections of Amanita species from section Vaginatae have been reported to be strictly associated with Helianthemum plants growing in grasslands, a still largely under-explored ectomycorrhizal habitat. The main aim of this study was to investigate the taxonomic status and phylogenetic position of strictly Helianthemum -associated Amanita sect. Vaginatae taxa, informally named Amanita “ helianthemicola ” . Collections from Italy, England and France were included in this study. The morphologically closely related species complex A. lividopallescens was also examined. Analyses were carried out based on both morphological and molecular data (phylogenetic analysis of the nrITS sequences). All investigated Amanita collections, which are strictly associated with Helianthemum nummularium, turned out to be conspecific with A . simulans. Amanita simulans was recently described from Sardinia (Italy) from Populus nigra habitats based on morphological characters only. As the holotype of A. simulans was lost, a neotype is designated here based on a voucher from the original collecting area. Amanita simulans is re-described, and an extensive discussion on the morphological variability, host species range, distribution and related taxa is provided. Amanita lividopallescens was confirmed as a good species, and re-delimited based on our phylogenetic analysis; moreover, it was epitypified with a recent and well-documented collection from Corse (France) . Amanita stenospora is a synonym of A . lividopallescens, being a colour form of the latter without taxonomic value.

In this paper we have attempted to clarify the taxonomy and nomenclature of thirteen taxa of the genus Cortinarius subgenus Telamonia (sections Hydrocybe, Fraternii) well represented in the southwestern Mediterranean area of Europe (C. atrocoeruleus, C. bombycinus, C casimiri, C. contrarius, C. decipiens, C. fraternus, C. gallurae, C. hoffmannii, C. petroselineus, C. sertipes, C. subturibulosus, C. urdaibaiensis and C. vernus). To this end we have performed a combined study of morphological and molecular data (rDNA ITS sequences). The morphological analysis was carried out on 114 collections and the molecular analysis involved 31 of the 114 collections, including 11 type collections (types for C. casimiri and C. fraternus were not available). In addition, a study of spores under field emission scanning electron microscopy (FESEM) was conducted. The results of the combined analysis allowed us to asign the studied material to five species (C. casimiri s.l., C. decipiens s.l., C. gallurae, C. subturibulosus s.l. and C. vernus s.l.). Thus, all collections from more continental areas, which were originally identified as six different taxa (C. atrocoeruleus, C. contrarius, C. decipiens, C. fraternus, C. sertipes, C. flexipes fo. sertipes) corresponded to C. decipiens sensu lato, a widely distributed, genetically and morphologically variable species. Cortinarius casimiri is also found in such habitats, but it is confirmed as distinct taxon. Collections from Mediterranean sclerophyllous communities correspond to C. gallurae, C. vernus sensu lato and C. subturibulosus sensu lato. Due to close phylogenetic relationships we propose the new combinations C. casimiri var. hoffmannii (=C. decipiens var. hoffmannii non C. hoffmannii) and C. subturibulosus var. bombycinus (=C. bombycinus), and the new variety C. vernus var. nevadavernus (=C. vernus H. Lindstr. & Melot sensu auct.).

European aspen is one of the most widely distributed trees in Central Europe and is a typical early colonizer of poor and disturbed soils. However, little is known about ectomycorrhizal (ECM) fungi in these ecosystems. We examined the ECM community of European aspen growing on a heavily contaminated site in southern Austria by analysing ECM roots, sorting them into morphotypes, subjecting them to DNA extraction, PCR, and DNA sequencing. ECM root symbionts were sampled two times in 2004. During this time, the below-ground community structure was relatively stable; we found no evidence of taxa adapted to summer or autumn conditions and only two species varied widely in occurrence between soil horizons. The ECM fungal community was diverse (54 species), rich in Basidiomycota (43 species), and dominated by Cenococcum geophilum and fungi with corticoid basidiomes (e.g. Thelephoraceae).

Anamika gen. nov. is described from southern India with one species, A. indica sp. nov. The genus is characterized by spores with a strongly cavernous ornamentation due to cavernae in the epitunica (perispore), and the possession of cheilo-, pleuro-, and caulocystidia. It seems to be associated with Hopea species (Dipterocarpaceae). Molecular data suggest a close relationship to Hebeloma and Naucoria, but indicate that it represents an independent line of evolution.

Gymnopilus turficola sp. nov. is described. This species is characterized by its habitat in palsa mires growing on peat in subarctic areas, by greenish lamellae in young state and the development of an iodoform odour when kept closed for some hours. Psilocin, psilocybin or related compounds could not be detected. G. turficola seems closely related to G. odini. According to the analysis of ITS sequences this species falls into a well supported clade of closely related taxa with G. odini, G. decipiens and G. penetrans.

Among the numerous items of equipment with the ‘Iceman’, who died more than 5000 years ago on an alpine glacier, were three fungal objects: two different shaped fruitbody pieces of the polypore Piptoporus betulinus, each mounted separately on a leather thong, and, found in his girdle bag, a relatively large quantity of tinder material prepared from the ‘true tinder bracket’ Fomes fomentarius. A full description of these items and a chronological report on their identification is given. The question about the possible use of the fungi is discussed on the basis of a comprehensive collection of ethnomycological and pharmacological literature data.

Mycophiles forage for and pick vast quantities of a wide variety of wild mushroom species. As a result, mushroom intoxications are comparatively frequent in such countries with mycophiles. Thus, national governments are forced to release guidelines or enact legislation in order to ensure the safe commerce of wild mushrooms due to food safety concerns. It is in these guidelines and laws that one can observe whether a country is indeed mycophobic or mycophilic. Furthermore, these laws and guidelines provide valuable information on mushroom preferences and on the consumption habits of each country. As such we were interested in the questions as to whether mushroom consumption behaviour was different within Europe, and if it was possible to discover the typical or distinctive culinary preferences of Slavic or Romanic speaking people, people from special geographical regions or from different zones. This work is based on the analysis of edible mushroom lists available in specific guidelines or legislation related to the consumption and commerce of mushrooms in 27 European countries. The overall diversity of edible mushrooms authorised to be commercialised in Europe is very high. However, only 60 out of a total 268 fungal species can be cultivated. This highlights the importance of guidelines or legislation for the safe commerce of wild mushrooms. The species richness and composition of the mushrooms listed for commerce is very heterogeneous within Europe. The consumption behaviour is not only languagefamily- related, but is strongly influenced by geographical location and neighbouring countries. Indicator species were detected for different European regions ; most of them are widespread fungi, and thus prove culture-specific preferences for these mushrooms. Our results highlight tradition and external input such as trade and cultural exchange as strong factors shaping mushroom consumption behaviour.

The leading agaricologist Meinhard M. Moser died on 30 September 2002. He was a Centenary Fellow of the British Mycological Society, and the mentor of numerous students and colleagues throughout the world.

The effect of various media formulations on in vitro ectomycorrhizal synthesis of identified fungal strains with European aspen (Populus tremula L.) was tested in Petri dishes. Pre-grown seedlings were transferred to various nutrient media and inoculated with Paxillus involutus isolates using modified sandwich techniques. Mycorrhiza formation was evaluated macroscopically and further confirmed by microscopic examination of semi-thin sections for anatomical features of the mantle and the Hartig net. Standard media formulations did not support successful ectomycorrhiza formation because of either very poor plant survival (below 20%) or impaired fungal growth. The inclusion of micronutrients and vitamins in a Melin Norkrans (MMN)-based medium increased plant survival rate to above 60% and supported successful mycorrhizal synthesis. P. involutus isolates formed mycorrhizas with a characteristic Hartig net restricted to the epidermis. Mantle density and thickness varied depending on the isolate. In a follow-up experiment, the adapted medium supported successful ectomycorrhiza formation by various Laccaria and Hebeloma isolates. Our results show that an exogenous supply of vitamins and micronutrients in the medium was a prerequisite for successful mycorrhization of P. tremula in vitro in Petri dishes.

The bog sedge Kobresia myosuroides is among the first ectomycorrhizal (EM) plants forming dense pads on receding glacier forefronts of the Austrian Alps. This is the only Cyperaceae species known to form EM. To date, little is known about fungal species involved in these EM associations. Therefore, the main aim of this study was to detect EM fungal communities of K. myosuroides (1) by describing mycorrhizal morphotypes (MT) and (2) by identifying the mycobionts by rDNA internal transcribed spacer (ITS) sequencing. Furthermore, seasonal dynamics of Kobresia mycobionts were investigated. Sampling was performed in all four seasons (also under snow cover) during the years 2005 and 2006 at the Rotmoos glacier forefront, a well-characterized alpine primary successional habitat in the Austrian alps (2,300 m above sea level). The degree of EM infection of K. myosuroides roots was high (95%). Ten MTs were described and sequences of 18 fungal taxa were obtained. This was the highest mycobiont diversity ever reported for this plant. Cenococcum geophilum was the most abundant mycobiont (37–46%) and shared dominancy with Sebacina incrustans (16–44%) and Tomentella spp. (7–37%). Tomentella (including Thelephora) was the most species-rich mycobiont genus with five taxa, followed by Cortinarius, Inocybe, and Sebacina with two taxa each and one Hebeloma species. Other ascomycete mycobionts beside C. geophilum were Helvella sp., Lecythophora sp., and one Pezizales species. Due to high interannual differences in the EM fungal community, no significant seasonal changes could be detected. The importance of fungal mycobionts in alpine habitats is discussed.

Dwarf willows (e.g. Salix herbacea) are among the earliest ectomycorrhizal (EM) plants colonising primary successional sites such as glacier forefronts in the Tyrolean Alps. EM of S. herbacea were sampled at the Rotmoos glacier forefront (Otz Valley, Austria) three times a year during the growing season and once a year during winter when plants were covered with snow in 2005 and 2006. EM were investigated using morphological methods and by sequencing the rDNA ITS region. The degree of EM mycorrhization was high throughout both years (93%). We distinguished 21 EM morphotypes and identified 19 fungal species. Cenococcum geophilum, Sebacina spp., Tomentella spp. and Cortinarius spp. dominated the mycobiont community of S. herbacea. The observed species richness in this about 150-year-old soil was at least 59% of the estimated species richness. Fungal communities differed significantly between consecutive years, and spatial heterogeneity was high. These differences made it difficult to detect seasonal impacts. Abundances of C. geophilum EM increased throughout the 2-year sampling period. Sebacina incrustans EM were very abundant in 2005, but nearly disappeared in 2006, whilst its fruitbodies were still frequent in the sampling area. This suggests that the mycorrhizae were displaced from the roots by an outcompeting species, whereas the mycelium was still present in the soil.

Polygonum viviparum is one of the first ectomycorrhizal (EM) plant species colonising primary successional sites at the Rotmoos glacier forefront (Tyrolean Alps, Austria). On a site with soil development of about 150 years (2,400 m above sea level), mycobionts of P. viviparum were identified by morphotyping and fungal ribosomal deoxyribonucleic acid internal transcribed spacer sequencing. For studying seasonal dynamics and spatial heterogeneity, ectomycorrhizae were sampled on five plots during all seasons. P. viviparum root tips were always EM. In total, 18 mycobiont taxa of the following genera were identified: Cenococcum (1), Cortinarius (2), Helvella (1), Inocybe (3), Russula (1), Sebacina (2), Thelephora (2) and Tomentella (6). All were non-specific EM partners of EM plants. As early as 2 weeks after spring snow melt, EM were well developed, vital and showed high mycobiont diversity. The relative abundance of senescent root tips was lowest in spring and increased throughout the year, with a maximum in winter (frozen soil). Thus, mycobiont growth and physiological activity obviously start when soil is still under snow cover: We speculate that water availability is one important initiation factor for mycorrhizal development under snow cover, when temperatures still range around the freezing point. Irrespectively of the season, most abundant mycobionts at this primary successional site belonged to the genera Tomentella, Sebacina and Cenococcum, also in frozen soil. Spatial heterogeneity was high when considering species composition and diversity indices. Overall mycobionts species richness was restricted at this site, probably because of the limited availability of fungal partners. We regard the presence/absence of fungal partner and limiting abiotic impacts of the environment as key factors for the symbiotic status of P. viviparum.

Species delimitation is still controversial in the Xerocomus chrysenteron complex. We have therefore established comprehensible and reliable species concepts based on statistical evaluation of morphological and ecological characters. We examined many collections from different geographical regions and different developmental stages within collections. Quantitative micromorphological characters (basidiospores, pileipellis end cells) were measured in statistically relevant numbers. The same material was used to generate 24 rDNA-LSU sequences, and the results of phylogenetic analyses clearly confirmed our species concepts: spore size and ornamentation, length of the pileipellis end cells and ‘pruinatus-hyphae’ are most valuable characters for the delimitation of species in this complex. Molecular data demonstrated that the X. chrysenteron complex is a monophyletic group. All the examined species (X. chrysenteron, X. cisalpinus, X. pruinatus, X. ripariellus, X. dryophilus, X. fennicus, X. porosporus and X. rubellus) represent independent lineages. The faintly striate spores, a key character characterising species of section Striatulispori , probably evolved independently. In addition, the ‘pruinatus-hyphae’ have multiple origins, and truncate spore apices are derived at least twice. Xerocomus cisalpinus sp. nov. is characterised by striate spores, the presence of ‘pruinatus-hyphae’ and a pileipellis strongly reminiscent of X. chrysenteron. For reasons of discussion, microscopical data are presented on Boletellus episcopalis for the first time. Xerocomus fennicus (Boletellus) comb. nov. is proposed. We provide descriptions to all included taxa. Our results once more demonstrate that reliably identified and characterised voucher collections are the basic requirement for meaningful phylogenetic studies.

Three new Cortinarius species, Cortinarius conopileus, C. keralensis, and C. phlegmophorus spp. nov., are described from Kerala State in southern India. This is the first record of ectomycorrhizal Cortinarius spp. in the tropical part of India. In addition to distinct morphological characters, the comparative analysis of rDNA ITS sequences of the collections from India and morphologically similar species support the recognition of these taxa as new species. Phylogenetic analyses demonstrate that the three Indian Cortinarius spp. belong to both larger subclades of the genus Cortinarius, clade/cortinarius and clade/telamonia. As supported by morphological and molecular data, C. phlegmophorus belongs to Cortinarius subgen. Myxacium sect. Defibulati. Based on classical morphological characters, both C. keralensis and C. conopileus are representatives of subgen. Telamonia. However, C. conopileus belongs to clade/obtusi, which is a well-supported subclade of clade/cortinarius. Thus, in contrast to classical taxonomy, the clade/obtusi represents an independent evolutionary origin of telamonioid taxa. This result is also reflected by the distinct morphological characters of taxa of clade/obtusi, namely the lamellar trama with ellipsoid inflated hyphae and the presence of cystidia. In contrast, C. keralensis is a typical member of clade/telamonia. Within/telamonia, only relationships of closely related taxa are resolved due to the low genetic divergence found in ITS sequences. Based on morphological and molecular criteria, C. keralensis is a distinct taxon of sect. Saturnini.