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.
Once the disease is established, management is hardly possible. The occurrence of a small fraction of partially tolerant trees constitutes hope for resistance breeding in the future. Healthy-looking trees should be preserved.
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.
Understanding the genetic diversity and structure of invasive pathogens in source and in introduced areas is crucial to the revelation of hidden biological features of an organism, to the reconstruction of the course of invasions and to the establishment of effective control measures. Hymenoscyphus pseudoalbidus (anamorph: Chalara fraxinea) is an invasive and highly destructive fungal pathogen found on common ash Fraxinus excelsior in Europe and is native to East Asia. To gain insights into its dispersal mechanisms and history of invasion, we used microsatellite markers and characterized the genetic structure and diversity of H. pseudoalbidus populations at three spatial levels: (i) between Europe and Japan, (ii) in Europe and (iii) at the epidemic's front in Switzerland. Phylogenetic and network analysis demonstrated that individuals from both regions are conspecific. However, populations from Japan harboured a higher genetic diversity and were genetically differentiated from European ones. No evident population structure was found among the 1208 European strains using Bayesian and multivariate clustering analysis. Only the distribution of genetic diversity in space, pairwise population differentiation (GST) and the spatial analysis of principal components revealed a faint geographical pattern around Europe. A significant allele deficiency in most European populations pointed to a recent genetic bottleneck, whereas no pattern of isolation by distance was found. Our data suggest that H. pseudoalbidus was introduced just once by at least two individuals. The potential source region of H. pseudoalbidus is vast, and further investigations are required for a more accurate localization of the source population.
The recently described ascomycete fungus Hymenoscyphus pseudoalbidus (anamorph: Chalara fraxinea) causes the current dieback of ash (Fraxinus excelsior) in large parts of Europe. The origin of this species and its relation to the native cryptic species Hymenoscyphus albidus are still enigmatic. The spatiotemporal pattern of the epidemic is typical for an introduced invasive species. However, the presence of two cryptic species indicates that hybridization or mutation might have been involved in driving speciation in this case. In this study, we present a set of 18 polymorphic microsatellite markers to study these processes in more detail on a population genetic level. Markers were designed such that they can be amplified in three individual multiplex PCRs and analysed in two fragment analysis runs. We thoroughly tested the marker set for pairwise linkage among loci, selective neutrality and Mendelian inheritance. Additionally, the markers were applied on two large collections of isolates derived from study sites in Germany. Population genetic calculations suggested a low yet significant level of differentiation, a large genotypic diversity and a limited genetic diversity within populations. Furthermore, we present additional data concerning the phylogenetic relation between H. albidus and H. pseudoalbidus, which seems to be more distantly related to each other than expected previously.
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