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Fallopia japonica and Impatiens glandulifera are major plant invaders on a global scale that often become dominant in riparian areas. However, little is known about how these species affect interactions in soil–plant systems. The aim of this study was to investigate the impact of both species on abiotic and biotic soil properties, with a special focus on fungi. We investigated eight sites along small streams invaded by F. japonica and I. glandulifera, respectively, and compared each with nearby sites dominated by the native species Urtica dioica. Three different types of samples were collected: bulk soil, rhizosphere soil and roots from invasive and native stands at each site. Bulk soil samples were analysed for soil physicochemical, microbial properties (soil microbial respiration and ergosterol) and soil arthropod abundance (Acari and Collembola). Soil respiration was also evaluated in rhizosphere samples. The fungal community composition of both bulk soil and roots were analysed using a metabarcoding approach. Soil physicochemical properties as well as soil microbial activity, fungal biomass and soil fungal operational unit taxonomic unit (OTU) richness did not differ between invaded and native riparian habitats, indicating only minor belowground impacts of the two invasive plant species. Soil microbial activity, fungal biomass and soil fungal OTU richness were rather related to the soil physicochemical properties. In contrast, Acari abundance decreased by 68% in the presence of F. japonica, while Collembola abundance increased by 11% in I. glandulifera sites. Moreover, root-associated fungal communities differed between the invasive and native plants. In F. japonica roots, fungal OTU richness of all investigated ecological groups (mycorrhiza, endophytes, parasites, saprobes) were lower compared to U. dioica. However, in I. glandulifera roots only the OTU richness of mycorrhiza and saprobic fungi was lower. Overall, our findings show that F. japonica and I. glandulifera can influence the abundance of soil arthropods and are characterized by lower OTU richness of root-associated fungi.

Phylogenetic analyses based on a three-locus nuclear data set (ITS1-5.8S-ITS2, the 5′ end of the 28S, and the largest subunit of RNA polymerase I) supported the pagoda fungus (Podoserpula, Amylocor...

While mineral fertilization increases agricultural yields, it also bears the risk of contaminating non-target ecosystems and negatively affecting soil chemical parameters and microbial communities. This calls for alternative and more sustainable agricultural practices that reduce the use of fertilizers. Flood pulse irrigation could be an alternative to mineral fertilization of hay meadows, since it increases the yield with little or no application of fertilizer. However, the positive and negative implications of flood pulse irrigation on soil chemical parameters and particularly soil microbial communities are still largely unknown. In this study, we assessed shifts in soil microbial communities (SMC) as a response to changes in soil chemical parameters after flood pulse irrigation and/or fertilization of meadows. We determined soil chemical (Corg, Ntot, water extractable N, P, K, pH) and microbial (phospholipid-derived fatty acids, PLFA) parameters of 12 meadows in a 2 × 2 factorial design, comprising flood pulse irrigation and fertilization. Corg, Ntot, and water content as well as microbial biomass were higher in flood-irrigated than in non-flooded soils. Soil microbial biomass positively correlated with Corg, Ntot, and water extractable N. Gram-negative bacteria significantly increased, whereas the fungi/bacteria ratio significantly decreased in flood-irrigated soils compared to non-flooded soils. Arbuscular mycorrhizal fungi were positively correlated with soil pH. Flood pulse irrigation seemed to promote the build-up of a larger soil carbon and nitrogen pool as well as higher water content and microbial biomass. By this, it potentially mitigated negative mineral fertilization effects such as changed soil pH and reduced carbon use efficiency. We conclude that flood pulse irrigation may represent a sustainable alternative to mineral fertilization.

Fungi belonging to the Entorrhizales (Entorrhizomycota) comprise biotrophic pathogens associated with roots of the Cyperaceae and Juncaceae plant species. They are nearly globally distributed but rarely studied due to a hidden lifestyle without causing visible effects on host plants. Therefore, the evolutionary origin and phylogenetic relationships of the group are still poorly understood and it is not known whether species diversification was the result of co-evolution with their hosts or the result of host jumps. To infer hypotheses about the evolutionary history of the Entorrhizales, divergence times were estimated and plant-fungal tanglegrams calculated. Relaxed molecular clock analyses suggest that the Entorrhizomycota originated around the Neoproterozoic-Palaeozoic and diverged during the Late Cretaceous-Paleogene into the extant orders Entorrhizales and Talbotiomycetales. The split of the major lineages within the Entorrhizales took place in the Eocene, somewhat later than the divergence of the host families Cyperaceae and Juncaceae. Topology- and distance-based co-phylogenetic analyses of the fungi and their hosts revealed a large number of co-speciation and lineage sorting events in early fungal speciation, which resulted in a phylogenetic split corresponding to species infecting Cyperaceae or Juncaceae. Given that this split is congruent with spore differences, Entorrhiza s. str. is emended for species infecting hosts in the Cyperaceae, and a new genus Juncorrhiza is described for species restricted to hosts in the Juncaceae. Additionally, three new species are described: Entorrhiza fuirenae, Juncorrhiza maritima and J. oxycarpi.

The order Ceraceosorales (Ustilaginomycotina) currently includes the single genus Ceraceosorus, with one species, Ceraceosorus bombacis, parasitic on Bombax ceiba in India. The diversity, biogeography, evolution, and phylogenetic relationships of this order are still relatively unknown. Here, a second species of Ceraceosorus is described from West Africa as a novel species, Ceraceosorus africanus, infecting Bombax costatum in Benin, Ghana, and Togo. This species produces conspicuous fructifications, similar to corticioid basidiomata when mature, but sorus-like in early stages of ontogenetic development. The fructifications cover much of the leaf surface and resemble leaf blight. This contrasts with the inconspicuous fructifications of C. bombacis comprising small spots scattered over the lower leaf surface that resemble leaf spot. Both species of Ceraceosorus differ in several micromorphological traits, infect different host plant species in widely separated geographical areas, and are separated by a considerable genetic distance in 28S rDNA and RPB2 genes. The distinct corticioid fructification of C. africanus is a unique morphological trait within the Ustilaginomycotina. Molecular phylogenetic analyses of a single gene dataset (D1/D2 28S rDNA) supported the monophyly of the two Ceraceosorus species and the Ceraceosorales and their placement within the Ustilaginomycotina. Molecular phylogenetic analyses of a multigene dataset (18S/5.8S/28S rDNA/RPB2/TEF1) revealed Exobasidium rhododendri (Exobasidiales) as the closest relative of Ceraceosorus, both clustering together with Entyloma calendulae (Entylomatales), indicating affinities to the Exobasidiomycetes. This phylogenetic placement is in agreement with ultrastructural characteristics (presence of local interaction zone and interaction apparatus) reported for the Ceraceosorales, Entylomatales, and Exobasidiales.

Ceraceosorus bombacis is an early-diverging lineage of smut fungi and a pathogen of cotton trees (Bombax ceiba). To study the evolutionary genomics of smut fungi in comparison with other fungal and oomycete pathogens, the genome of C. bombacis was sequenced and comparative genomic analyses were performed. The genome of 26.09 Mb encodes for 8,024 proteins, of which 576 are putative-secreted effector proteins (PSEPs). Orthology analysis revealed 30 ortholog PSEPs among six Ustilaginomycotina genomes, the largest groups of which are lytic enzymes, such as aspartic peptidase and glycoside hydrolase. Positive selection analyses revealed the highest percentage of positively selected PSEPs in C. bombacis compared with other Ustilaginomycotina genomes. Metabolic pathway analyses revealed the absence of genes encoding for nitrite and nitrate reductase in the genome of the human skin pathogen Malassezia globosa, but these enzymes are present in the sequenced plant pathogens in smut fungi. Interestingly, these genes are also absent in cultivable oomycete animal pathogens, while nitrate reductase has been lost in cultivable oomycete plant pathogens. Similar patterns were also observed for obligate biotrophic and hemi-biotrophic fungal and oomycete pathogens. Furthermore, it was found that both fungal and oomycete animal pathogen genomes are lacking cutinases and pectinesterases. Overall, these findings highlight the parallel evolution of certain genomic traits, revealing potential common evolutionary trajectories among fungal and oomycete pathogens, shaping the pathogen genomes according to their lifestyle.

Morphological and molecular discrepancies in the biodiversity of monophyletic groups are challenging. The intention of this study was to find out whether the high molecular diversity in Sebacinales can be verified by micromorphological characteristics. Therefore, we carried out molecular and morphological studies on all generic type species of Sebacinales and additional representative taxa. Our results encouraged us to disentangle some phylogenetic and taxonomic discrepancies and to improve sebacinalean classifications. This comprises generic circumscriptions and affiliations, as well as higher taxon groupings. At the family level, we redefined the Sebacinaceae, formerly the Sebacinales group A, and set it apart from the Sebacinales group B. For taxonomical purposes, it seems appropriate to refer Paulisebacina, Craterocolla, Chaetospermum, Globulisebacina, Tremelloscypha, and Sebacina to the Sebacinaceae and Piriformospora, and Serendipita to the Sebacinales group B. At the lower taxonomic level, we propose within the Sebacinaceae (1) to introduce Paulisebacina for Sebacina allantoidea, (2) to transfer Efibulobasidium rolleyi into a new monotypic genus, Globulisebacina, (3) to include Tremellostereum in Tremelloscypha, (4) to transfer Sebacina amesii into Tremelloscypha, (5) to combine S. helvelloides and S. concrescens in their own genus, Helvellosebacina, (6) to transfer Tremellodendron spp. into Sebacina, (7) to define S. epigaea s.str. without cystidia and flagelliform dikaryophyses, but with star-shaped resting spores, and (8) to separate S. cystidiata with simultaneously irregular germinating spores and inconspicuous cystidia, and S. flagelliformis with flagelliform dikaryophyses from S. epigaea s.str. Additional clades in Sebacina, based on molecular differences, cannot be distinguished morphologically at present.

A taxonomic re-evaluation of the Ceratobasidium-Rhizoctonia group suggests that Ceratobasidium contains only the type species C. calosporum, which deviates in micromorphological and ultrastructural characters from all other species so far included in that genus. Rhizoctonia species are compared with the type species of Ceratobasidium, Cejpomyces, Oncobasidium, Tofispora, Waitea, and Ypsilonidium. The micromorphology, ultrastructure, cellular interaction with the host, and molecular phylogeny of a Rhizoctonia species parasitic on needles and young shoots of Picea abies have been studied. The parasite has been known for a long time, but misinterpreted, and not named so far. Rhizoctonia butinii is described and compared with related species of the genus.

A historical retrospect and a taxonomic update will deal with Sebacina s.l. and s.str., Craterocolla, Efibulobasidium, Serendipita, Tremellodendron, Tremelloscypha, Tremellostereum, and Piriformospora, the Sebacinaceae, and the Sebacinales. Phylogenetic hypotheses for the order and subordinal taxa are discussed, including environmental sequence taxa. The cryptic biodiversity in Sebacinales is extensive but mostly unresolved with respect to the species involved. Trophic stages are manifold in Sebacinales but restricted to plant dependencies. Most of the species grow endophytically or form various mycorrhizae, but Craterocolla and Efibulobasidium species appear to be saprobic. The sebacinalean mycorrhizal diversity is unparalleled: ectomycorrhizae, ericoid and orchid mycorrhizae are frequent, both in autotrophs and heterotrophs, as well as mycothalli with Jungermanniales. Mycorrhizal community structures are difficult to evaluate in Sebacinales because of the high percentage of environmental sequence taxa lacking further characteristics. Nutritional requirements and exchanges have been studied extensively in Piriformospora indica, suggesting future possibilities for agricultural applications. The genomes of this species and of Sebacina vermifera have been sequenced recently, thus opening new fields in studying and understanding functional and evolutionary aspects.

Different distance-based threshold selection approaches were used to assess and compare use of the internal transcribed spacer (ITS) region to distinguish among 901 Cortinarius species represented by >3000 collections. Sources of error associated with genetic markers and selection approaches were explored and evaluated using MOTUs from genus and lineage based-alignments. Our study indicates that 1%-2% more species can be distinguished by using the full-length ITS barcode as compared to either the ITS1 or ITS2 regions alone. Optimal threshold values for different picking approaches and genetic marker lengths inferred from a subset of species containing major lineages ranged from 97.0% to 99.5% sequence similarity using clustering optimization and UNITE SH, and from 1% to 2% sequence dissimilarity with CROP. Errors for the optimal cutoff ranged from 0% to 70%, and these can be reduced to a maximum of 22% when excluding species lacking a barcode gap. A threshold value of 99% is suitable for distinguishing species in the majority of lineages in the genus using the entire ITS region but only 90% of the species could be identified using just the ITS1 or ITS2 region. Prior identification of species, lacking barcode gaps and their subsequent separate analyses, maximized the accuracy of threshold approaches.

The recently described fungal phylum Entorrhizomycota was established solely for the genus Entorrhiza, species of which cause root-galls in Cyperaceae and Juncaceae. Talbotiomyces calosporus (incertae sedis) shares morphological characteristics and an ecological niche with species of Entorrhiza. We investigated the higher classification of T. calosporus to determine whether it belongs in Entorrhizomycota. Ribosomal DNA sequences showed Talbotiomyces to be a close relative of Entorrhiza and both taxa form a highly supported monophyletic group. Based on molecular phylogenetic analyses and in congruence with existing morphological and ecological data, Entorrhiza and Talbotiomyces represent a deep dichotomy within the Entorrhizomycota. While species of Entorrhiza are characterised by dolipores and occur on monocotyledons, members of Talbotiomyces are characterised by simple pores and are associated with eudicotyledons. This expands the host range of the recently described Entorrhizomycota from Poales to other angiosperms. Higher taxa, namely Talbotiomycetales ord. nov. and Talbotiomycetaceae fam. nov., are proposed here to accommodate Talbotiomyces.

Background Phylogenetic studies, particularly those based on rDNA sequences from plant roots and basidiomata, have revealed a strikingly high genetic diversity in the Sebacinales. However, the factors determining this genetic diversity at higher and lower taxonomic levels within this order are still unknown. In this study, we analysed patterns of genetic variation within two morphological species, Sebacina epigaea and S. incrustans, based on 340 DNA haplotype sequences of independent genetic markers from the nuclear (ITS + 5.8S + D1/D2, RPB2) and mitochondrial (ATP6) genomes for 98 population samples. By characterising the genetic population structure within these species, we provide insights into species boundaries and the possible factors responsible for genetic diversity at a regional geographic scale. Results We found that recombination events are relatively common between natural populations within Sebacina epigaea and S. incrustans, and play a significant role in generating intraspecific genetic diversity. Furthermore, we also found that RPB2 and ATP6 genes display higher levels of intraspecific synonymous polymorphism. Phylogenetic and demographic analyses based on nuclear and mitochondrial loci revealed three distinct phylogenetic lineages within of each of the morphospecies S. epigaea and S. incrustans: one major and widely distributed lineage, and two geographically restricted lineages, respectively. We found almost no differential morphological or ecological characteristics that could be used to discriminate between these lineages. Conclusions Our results suggest that recombination and negative selection have played significant roles in generating genetic diversity within these morphological species at small geographical scales. Concordance between gene genealogies identified lineages/cryptic species that have evolved independently for a relatively long period of time. These putative species were not associated with geographic provenance, geographic barrier, host preference or distinct phenotypic innovations.

To study the diversity and phylogenetic structure of Sebacinales communities from eight vegetation communities along an altitudinal gradient in the Bavarian Alps (Germany), we analysed 456 thalli or roots of plants. We detected 264 sebacinoid sequences, spanning the intergenic transcribed spacer region, 5.8S and D1/D2 regions of the nuclear rRNA gene, mostly using a nested PCR approach. Based on 97% sequence similarity, 73 Sebacinales molecular taxonomic units were found from 70 host species belonging to 44 plant families. Twenty-six molecular taxonomic units represented singletons, the most frequent of these being restricted exclusively to either wooded or grassland habitats. Although Sebacinales appear to occur in low abundance in plant roots, these microorganisms are phylogenetically diverse and widely spread in the ecosystems studied. Ordination analyses showed that land use, pH and humus content strongly influence the diversity and assembly of Sebacinales. In most cases, Sebacinales communities in ecosystems with extreme soil conditions or intensive land use exhibited significant phylogenetic clustering, whereas in undisturbed plant communities no trend was observed. These results suggest that ecosystem disturbance and environmental forces have an influence on the diversity and structure of Sebacinales community assembly over local and spatial scales.

International audience; Inconspicuous basidiomycetes from the order Sebacinales are known to be involved in a puzzling variety of mutualistic plant-fungal symbioses (mycorrhizae), which presumably involve transport of mineral nutrients. Recently a few members of this fungal order not fitting this definition and commonly referred to as ‘endophytes’ have raised considerable interest by their ability to enhance plant growth and to increase resistance of their host plants against abiotic stress factors and fungal pathogens. Using DNA-based detection and electron microscopy, we show that Sebacinales are not only extremely versatile in their mycorrhizal associations, but are also almost universally present as symptomless endophytes. They occurred in field specimens of bryophytes, pteridophytes and all families of herbaceous angiosperms we investigated, including liverworts, wheat, maize, and the non-mycorrhizal model plant Arabidopsis thaliana. They were present in all habitats we studied on four continents. We even detected these fungi in herbarium specimens originating from pioneering field trips to North Africa in the 1830s/40s. No geographical or host patterns were detected. Our data suggest that the multitude of mycorrhizal interactions in Sebacinales may have arisen from an ancestral endophytic habit by specialization. Considering their proven beneficial influence on plant growth and their ubiquity, endophytic Sebacinales may be a previously unrecognized universal hidden force in plant ecosystems.