Ash dieback caused by the mitosporic ascomycete Chalara fraxinea is a novel disease of major concern affecting Fraxinus excelsior and Fraxinus angustifolia in large parts of Europe. Recently, its teleomorph was detected and assigned to Hymenoscyphus albidus, which has been known from Europe since 1851. In this study, we present molecular evidence for the existence of two morphologically very similar taxa, H. albidus, which is lectotypified and Hymenoscyphus pseudoalbidus sp. nov. Differences were found between the species in the loci calmodulin, translation elongation factor 1-a and the internal transcribed spacers of the rDNA genes, and strong differentiation was obtained with ISSR markers. It is likely that H. albidus is a non-pathogenic species, whereas H. pseudoalbidus is a virulent species causing ash dieback. Genotyping herbarium specimens showed that H. pseudoalbidus has been present in Switzerland for at least 30 years prior to the outbreak of the epidemic.

Dark septate endophytes (DSE), a diverse group of ascomycetes, are dominant root colonizers in many ecosystems. The most frequent DSE in natural forest ecosystems in the Northern hemisphere belong to the Phialocephala fortinii s.l. – Acephala applanata species complex (PAC). Recently, species rank was assigned to seven cryptic species (CSP) of P. fortinii s.l.: Phialocephala fortinii s. str. C.J.K. Wang & H.E. Wilcox, Phialocephala europaea C.R. Grünig et T.N. Sieber, Phialocephala helvetica C.R. Grünig et T.N. Sieber, Phialocephala letzii C.R. Grünig et T.N. Sieber, Phialocephala subalpina C.R. Grünig et T.N. Sieber, Phialocephala turiciensis C.R. Grünig et T.N. Sieber, and Phialocephala uotolensis C.R. Grünig et T.N. Sieber. PAC species occur on all parts of the root system of trees, from mycorrhizal root tips to the stem base. Up to 80% of fine roots in forest stands can be colonized by them, and up to eight species occur sympatrically. The present work is a mixture of review and reconsideration of published work in the light of the subdivision of P. fortinii s.l. into several species. We review the current knowledge related to taxonomy, geographical distribution, population biology, and ecology of PAC species. We identified strains of P. fortinii s.l. from previously published studies to species level. The reassessment of earlier studies indicates that PAC species behave in a versatile manner along the mutualism-parasitism continuum and lifestyle designation is complicated by the use of different experimental systems. Finally we define the most promising research areas, which will contribute to elucidate the ecological role of root endophytes in general and PAC species in particular.

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Root endophytes are common and genetically highly diverse suggesting important ecological roles. Yet, relative to above-ground endophytes, little is known about them. Dark septate endophytic fungi of the Phialocephala fortinii s.l.-Acephala applanata species complex (PAC) are ubiquitous root colonizers of conifers and Ericaceae, but their ecological function is largely unknown. Responses of Norway spruce seedlings of two seed provenances to inoculations with isolates of four PAC species were studied in vitro. In addition, isolates of Phialocephala subalpina from two populations within and one outside the natural range of Norway spruce were also included to study the effect of the geographic origin of P. subalpina on host response. The interaction of PAC with Norway spruce ranged from neutral to highly virulent and was primarily isolate-dependent. Variation in virulence was much higher within than among species, nonetheless only isolates of P. subalpina were highly virulent. Disease caused by P. subalpina genotypes from the native range of Norway spruce was more severe than that induced by genotypes from outside the natural distribution of Norway spruce. Virulence was not correlated with the phylogenetic relatedness of the isolates but was positively correlated with the extent of fungal colonization as measured by quantitative real-time PCR.

AimThe biogeography of microbes is poorly understood and there is an open debate regarding if and how microbial biodiversity is structured. At the beginning of the 20th century, Baas Becking laid the foundations for the biogeography of microbes by stating that 'Everything is everywhere, but the environment selects' (the EisE hypothesis). This hypothesis remained dogma for almost a century. However, the recognition that microbial 'species' are often assemblages of reproductively isolated lineages challenged the EisE hypothesis, leading to the now common assumption that microbial communities possess cryptic biogeographic structures. We tested the presence of a cryptic biogeographical structure for a well-characterized fungal species complex (the Phialocephala fortinii s.l.-Acephala applanata species complex, PAC) using precise molecular species resolution. In addition, we analysed factors that could govern PAC community assembling.Locations Forty-four study sites in temperate and boreal forests across the Northern Hemisphere were included. Methods(1) The distance-decay relationship among PAC communities was calculated and a resampling procedure was applied to analyse the effect of sampling intensity and geographic distances among PAC communities. (2) Factors shaping PAC communities (e.g. climatic factors and tree species composition) were studied.(3) We tested PAC communities for random composition. ResultsWe found that the similarity of species assemblages did not decrease with increasing geographical distance. Moreover, species diversity did not increase by expanding the area sampled. Instead, species diversity increased by increasing the sampling effort. Community composition correlated neither with tree species composition nor climate, and no association among species was observed. Main conclusionsWe could not discover any cryptic biogeographic structure even after applying refined species assignment but we demonstrate the importance of sampling effort for understanding the biogeography of microorganisms. Moreover, we show that primarily stochastic effects are responsible for the species composition of PAC communities.