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Abstract The debates over the requirement of the International Code of Nomenclature for algae, fungi, and plants (ICNafp) for a viable specimen to represent the name-bearing type material for a species or infraspecific taxon have a long history. Taxonomy of fungi commonly studied as living cultures exemplified by yeasts and moulds, strongly depend on viable reference material. The availability of viable cultures is also particularly useful for several groups of filamentous and dimorphic fungi. While the preservation of metabolically inactive cultures is permitted and recommended by the ICNafp, there is room for improvement. Below, we review the history and current status of cultures as the name-bearing type material under the Code. We also present a roadmap with tasks to be achieved in order to establish a stable nomenclatural system that properly manages taxa typified by viable specimens. Furthermore, we propose setting up rules and defining the nomenclatural status of ex-type cultures under Chapter F, the section of the ICNafp that includes provisions specific to names of fungi.

A rare smut fungus, Anthracoidea pseudofoetidae, is reported for the first time from Russia, based on a specimen from the Altai Republic. Carex enervis is a new host record for A. pseudofoetidae. Two new records for Greece, Microbotryum duriaeanum and M. moenchiae-manticae, and a new record for Morocco, Microbotryum moehringiae, are also presented. Microbotryum moehringiae is recorded for the first time from Africa. Cerastium brachypetalum subsp. roeseri is a new host record for Microbotryum duriaeanum. New molecular data are provided for these smut fungi. Updated phylogenetic trees for Anthracoidea and Microbotryum are also given.

Fungal communities in above-ground tree tissues are hyperdiverse and are influenced by biotic interactions with other organisms living in or on these tissues. These biotic interactions are, however, still poorly understood. In this study, we aimed to understand how insect-associated gall formation on Eucalyptus foliage correlates with the diversity of foliar fungal communities in surrounding healthy leaf tissue, as well as the co-occurrence patterns among the members of the fungal community. We used ITS metabarcoding to characterise the foliar fungal communities of 179 individual E. grandis trees. These trees were assigned to infestation levels of the wasp Leptocybe invasa (Eulophidae: Hymenoptera), which causes gall formation on shoot tips and leaves of its host. Fungal community networks were calculated using a Pearson correlation coefficient. The composition and diversity of fungal communities were influenced by the severity of L. invasa infestations. We identified potential Eucalyptus pathogens with high sequence abundance at all disease severity levels, but network analysis indicated that the co-occurrence of potential pathogens between no to mild and medium to heavy infestation differed significantly. A better understanding of microbial interactions, especially the role of pathogens, can be useful for controlling disease- and beneficial host-associated microbial communities.

After examination of specimens, mainly from the herbarium (MA) and the mycological collection (MA-Fungi) of the Royal Botanic Garden of Madrid, we report several novelties on smut fungi within Europe. Two species of smut fungi, Sporisorium egyptiacum and Tilletia viennotii, are reported for the first time from Europe. A finding of Sphacelotheca polygoni-serrulati represents a second record for Europe. Six species of smut fungi, Moreaua kochiana, Schizonella elynae, Sporisorium egyptiacum, Thecaphora thlaspeos, Tilletia viennotii, and Ustanciosporium majus, are recorded for the first time from the Iberian Peninsula. Five species of smut fungi, Moreaua kochiana, Schizonella elynae, Sporisorium egyptiacum, Thecaphora thlaspeos, and Ustanciosporium majus, are newly recorded from Spain. Three species, Moreaua kochiana, Sphacelotheca polygoni-serrulati, and Tilletia viennotii, are new for Portugal. A specimen of Moreaua kochiana represents a new record for France. Arabis serpillifolia is reported as a new host of Thecaphora thlaspeos. New distribution records from the Iberian Peninsula are given for Anthracoidea arenariae, Microbotryum minuartiae, M. silenes-saxifragae, and Tranzscheliella sparti. We also include a phylogenetic analysis of DNA sequences of Moreaua kochiana, generated in this study, to understand this species’ relationships within its genus.

Foliar fungi, especially endophytic fungi, constitute an important part of the microbiome of plants. Yet little is known about the composition of these communities. In this study, we isolated fungi from leaf tissues of the desert plant Welwitschia mirabilis to determine the culturable diversity of the foliar fungal community. The isolated fungal taxa, which grouped into 17 distinct lineages, were identified by sequencing elongation factor 1 alpha, beta-tubulin 1, beta-tubulin 2 and the internal transcribed spacer region. The culturable community was mainly composed of cosmopolitan fungal genera despite the unique taxonomic position of the plant and its geographic isolation. To test for endemism in two of the common fungal genera, Alternaria and Aureobasidium, we built haplotype networks using a global data set. Even this broad data set showed little evidence for specialisation within this unique host or its geographical location. The data suggest that the culturable members of communities of leaf-associated fungi in habitats with little plant coverage, such as the Namib Desert, are mainly established by long-distance aerially distributed fungal inocula and few of these taxa co-evolve with the host within the habitat. Significance: • The culturable members of fungal communities associated with an ecological and evolutionary isolated plant have not co-speciated with their hosts, but to a large extent are composed of globally distributed fungal species. • Harsh environmental conditions and the geographic isolation of host plants seem to favour ubiquitous fungal species over more specialist fungal species. Open data sets*: https://www.ncbi.nlm.nih.gov/genbank *see accession numbers in text

Fungi represent a common and diverse part of the microbial communities that associate with plants. They also commonly colonise various plant parts asymptomatically. The molecular mechanisms of these interactions are, however, poorly understood. In this study we use transcriptomic data from Eucalyptus grandis, to demonstrate that RNA-seq data are a neglected source of information to study fungal−host interactions, by exploring the fungal transcripts they inevitably contain. We identified fungal transcripts from E. grandis data based on their sequence dissimilarity to the E. grandis genome and predicted biological functions. Taxonomic classifications identified, amongst other fungi, many well-known pathogenic fungal taxa in the asymptomatic tissue of E. grandis. The comparison of a clone of E. grandis resistant to Chrysoporthe austroafricana with a susceptible clone revealed a significant difference in the number of fungal transcripts, while the number of fungal taxa was not substantially affected. Classifications of transcripts based on their respective biological functions showed that the fungal communities of the two E. grandis clones associate with fundamental biological processes, with some notable differences. To shield the greater host defence machinery in the resistant E. grandis clone, fungi produce more secondary metabolites, whereas the environment for fungi associated with the susceptible E. grandis clone is more conducive for building fungal cellular structures and biomass growth. Secreted proteins included carbohydrate active enzymes that potentially are involved in fungal−plant and fungal−microbe interactions. While plant transcriptome datasets cannot replace the need for designed experiments to probe plant−microbe interactions at a molecular level, they clearly hold potential to add to the understanding of the diversity of plant−microbe interactions.

The Kingdom Fungi adds substantially to the diversity of life, but due to their cryptic morphology and lifestyle, tremendous diversity, paucity of formally described specimens, and the difficulty in isolating environmental strains into culture, fungal communities are difficult to characterize. This is especially true for endophytic communities of fungi living in healthy plant tissue. The developments in next generation sequencing technologies are, however, starting to reveal the true extent of fungal diversity. One of the promising new technologies, namely semiconductor sequencing, has thus far not been used in fungal diversity assessments. In this study we sequenced the internal transcribed spacer 1 (ITS1) nuclear encoded ribosomal RNA of the endophytic community of the economically important tree, Eucalyptus grandis, from South Africa using the Ion Torrent Personal Genome Machine (PGM). We determined the impact of various analysis parameters on the interpretation of the results, namely different sequence quality parameter settings, different sequence similarity cutoffs for clustering and filtering of databases for removal of sequences with incomplete taxonomy. Sequence similarity cutoff values only had a marginal effect on the identified family numbers, whereas different sequence quality filters had a large effect (89 vs. 48 families between least and most stringent filters). Database filtering had a small, but statistically significant, effect on the assignment of sequences to reference sequences. The community was dominated by Ascomycota, and particularly by families in the Dothidiomycetes that harbor well-known plant pathogens. The study demonstrates that semiconductor sequencing is an ideal strategy for environmental sequencing of fungal communities. It also highlights some potential pitfalls in subsequent data analyses when using a technology with relatively short read lengths.

BACKGROUND: Yeast-like fungi inhabit soils throughout all climatic zones in a great abundance. While recent estimations predicted a plethora of prokaryotic taxa in one gram of soil, similar data are lacking for fungi, especially yeasts. METHODOLOGY/PRINCIPAL FINDINGS: We assessed the diversity of soil yeasts in different forests of central Germany using cultivation-based techniques with subsequent identification based on rDNA sequence data. Based on experiments using various pre-cultivation sample treatment and different cultivation media we obtained the highest number of yeasts by analysing mixed soil samples with a single nutrient-rich medium. Additionally, several species richness estimators were applied to incidence-based data of 165 samples. All of them predicted a similar range of yeast diversity, namely 14 to 16 species. Randomized species richness curves reached saturation in all applied estimators, thus indicating that the majority of species is detected after approximately 30 to 50 samples analysed. CONCLUSIONS/SIGNIFICANCE: In this study we demonstrate that robust species identification as well as mathematical approaches are essential to reliably estimate the sampling effort needed to describe soil yeast communities. This approach has great potential for optimisation of cultivation techniques and allows high throughput analysis in the future.

The debates over the requirement of the International Code of Nomenclature for algae, fungi, and plants (ICNafp) for a viable specimen to represent the name-bearing type material for a species or infraspecific taxon have a long history. Taxonomy of fungi commonly studied as living cultures exemplified by yeasts and moulds, strongly depend on viable reference material. The availability of viable cultures is also particularly useful for several groups of filamentous and dimorphic fungi. While the preservation of metabolically inactive cultures is permitted and recommended by the ICNafp, there is room for improvement. Below, we review the history and current status of cultures as the name-bearing type material under the Code. We also present a roadmap with tasks to be achieved in order to establish a stable nomenclatural system that properly manages taxa typified by viable specimens. Furthermore, we propose setting up rules and defining the nomenclatural status of ex-type cultures under Chapter F, the section of the ICNafp that includes provisions specific to names of fungi. [ABSTRACT FROM AUTHOR]

The smut fungal genus Anthracoidea contains more than 100 species that parasitize hosts predominantly in the sedge genus Carex. Anthracoidea species are mainly found in the boreal zones of the Northern Hemisphere and many species have an arctic-alpine distribution. Recent re-organization of the taxonomy of the main host genus Carex questions current understanding of host associations in Anthracoidea. Host specificity for many of the species in this genus is considered to be quite broad and a host spectrum of over 10 host species is common. One aim of the study is to understand the potential influence that host taxonomy has on the evolutionary patterns of Anthracoidea. Additionally, by including more specimens, we clarify host specificity and species delimitation in Anthracoidea sempervirentis, a prevalent species occurring on different host species in different Carex subgroups using molecular data. Host colonization patterns within Anthracoidea are complex, and different subclades of Carex have been colonized several times independently, whereas clades of related Anthracoidea species often occur on Carex species from the same host clade. Parasites previously thought to be Anthracoidea sempervirentis occurring on the different Carex host are shown to be at least four distinct species that are restricted to individual host species. Three new species, Anthracoidea ferrugineae on Carex ferruginea from the Alps and the Carpathians, A. firmae on Carex firma from the Alps, and A. kitaibelianae on Carex kitaibeliana from mountains in the Balkan Peninsula, are described and illustrated. An emended description of Anthracoidea sempervirentis is also provided. Anthracoidea sempervirentis in its emended circumscription consists of two clades that correspond to respective clades within Carex sempervirens. The study shows that host colonization in Anthracoidea is more complex than current host taxonomy suggests. Further, including several specimens per host species results in a much higher diversity within Anthracoidea than previously assumed. [ABSTRACT FROM AUTHOR]

Basidiomycota is one of the major phyla in the fungal tree of life. The outline of Basidiomycota provides essential taxonomic information for researchers and workers in mycology. In this study, we present a time-framed phylogenomic tree with 487 species of Basidiomycota from 127 families, 47 orders, 14 classes and four subphyla; we update the outline of Basidiomycota based on the phylogenomic relationships and the taxonomic studies since 2019; and we provide notes for each order and discuss the history, defining characteristics, evolution, justification of orders, problems, significance, and plates. Our phylogenomic analysis suggests that the subphyla diverged in a time range of 443–490 Myr (million years), classes in a time range of 312–412 Myr, and orders in a time range of 102–361 Myr. Families diverged in a time range of 50–289 Myr, 76–224 Myr, and 62–156 Myr in Agaricomycotina, Pucciniomycotina, and Ustilaginomycotina, respectively. Based on the phylogenomic relationships and divergence times, we propose a new suborder Mycenineae in Agaricales to accommodate Mycenaceae. In the current outline of Basidiomycota, there are four subphyla, 20 classes, 77 orders, 297 families, and 2134 genera accepted. When building a robust taxonomy of Basidiomycota in the genomic era, the generation of molecular phylogenetic data has become relatively easier. Finding phenotypical characters, especially those that can be applied for identification and classification, however, has become increasingly challenging. [ABSTRACT FROM AUTHOR]

The group of seed-destroying Microbotryum species on caryophyllaceous hosts includes ten species, among which only M. alsines, on Stellaria nitens from USA, is known on Stellaria. Recently, a specimen of Stellaria media from Greece, infected by a seed-destroying Microbotryum species, was found in a phanerogams herbarium. Based on distinct morphology and phylogenetic evidence, this fungus was considered to represent an unknown species which is described and illustrated as Microbotryum graecum. It can be distinguished from M. alsines by having smaller spores, fewer meshes per spore diameter, a different spore ornamentation, wider meshes, and higher spore wall muri. [ABSTRACT FROM AUTHOR]

Context: Land use change can significantly affect plant-fungal interactions. Objectives: We assessed how fungal endophytes within African wild olive (Olea europaea subsp. cuspidata) twigs are influenced by different levels of land use change and differences in surrounding vegetation types. Methods: Twigs were sampled in the Western Cape Province (South Africa) and their fungal endophyte assemblages were characterised using culture-independent DNA metabarcoding. We assessed the effects of land use change (natural, semi-natural and planted (completely transformed)) and differences in surrounding vegetation types (grasses/low-growing plants versus shrubs/trees versus other olives) using fungal endophyte alpha and beta diversity measures. Co-occurrence networks were constructed to assess assemblage connectivity under different scenarios and to identify OTUs of potential ecological significance. Results: OTU richness, but not abundance, was significantly influenced by both land use change and differences in the surrounding vegetation types. Planted African olives and those surrounded by heterospecific trees harboured the highest OTU richness. Only levels of land use change significantly influenced fungal endophyte assemblage composition. Specifically, fungal assemblages from natural habitats were distinct from those in planted and semi-natural habitats, which were similar to each other. Co-occurrence network analyses revealed that cohesive and species rich networks could only be maintained within the natural habitats. Conclusion: These findings suggest that although the African olive is widespread, the identity and composition of their associated fungal assemblages are particularly sensitive to land use change. This study highlights the importance of conserving natural habitats, not just for the plants, but also for the maintenance of their associated fungal endophytes. [ABSTRACT FROM AUTHOR]

Novel species of fungi described in this study include those from various countries as follows: Argentina, Neocamarosporium halophilum in leaf spots of Atriplex undulata. Australia, Aschersonia merianiae on scale insect (Coccoidea), Curvularia huamulaniae isolated from air, Hevansia mainiae on dead spider, Ophiocordyceps poecilometigena on Poecilometis sp. Bolivia, Lecanora menthoides on sandstone, in open semi-desert montane areas, Sticta monlueckiorum corticolous in a forest, Trichonectria epimegalosporae on apothecia of corticolous Megalospora sulphurata var. sulphurata, Trichonectria puncteliae on the thallus of Punctelia borreri. Brazil, Catenomargarita pseudocercosporicola (incl. Catenomargarita gen. nov.) hyperparasitic on Pseudocercospora fijiensis on leaves of Musa acuminata, Tulasnella restingae on protocorms and roots of Epidendrum fulgens. Bulgaria, Anthracoidea umbrosae on Carex spp. Croatia, Hymenoscyphus radicis from surface-sterilised, asymptomatic roots of Microthlaspi erraticum, Orbilia multiserpentina on wood of decorticated branches of Quercus pubescens. France, Calosporella punctatispora on dead corticated twigs of Acer opalus. French West Indies (Martinique), Eutypella lechatii on dead corticated palm stem. Germany, Arrhenia alcalinophila on loamy soil. Iceland, Cistella blauvikensis on dead grass (Poaceae). India, Fulvifomes maritimus on living Peltophorum pterocarpum, Fulvifomes natarajanii on dead wood of Prosopis juliflora, Fulvifomes subazonatus on trunk of Azadirachta indica, Macrolepiota bharadwajii on moist soil near the forest, Narcissea delicata on decaying elephant dung, Paramyrothecium indicum on living leaves of Hibiscus hispidissimus, Trichoglossum syamviswanathii on moist soil near the base of a bamboo plantation. Iran, Vacuiphoma astragalicola from stem canker of Astragalus sarcocolla. Malaysia, Neoeriomycopsis fissistigmae (incl. Neoeriomycopsidaceae fam. nov.) on leaf spots on flower Fissistigma sp. Namibia, Exophiala lichenicola lichenicolous on Acarospora cf. luederitzensis. Netherlands, Entoloma occultatum on soil, Extremus caricis on dead leaves of Carex sp., Inocybe pseudomytiliodora on loamy soil. Norway, Inocybe guldeniae on calcareous soil, Inocybe rupestroides on gravelly soil. Pakistan, Hymenagaricus brunneodiscus on soil. Philippines, Ophiocordyceps philippinensis parasitic on Asilus sp. Poland, Hawksworthiomyces ciconiae isolated from Ciconia ciconia nest, Plectosphaerella vigrensis from leaf spots on Impatiens noli-tangere, Xenoramularia epitaxicola from sooty mould community on Taxus baccata. Portugal, Inocybe dagamae on clay soil. Saudi Arabia, Diaporthe jazanensis on branches of Coffea arabica. South Africa, Alternaria moraeae on dead leaves of Moraea sp., Bonitomyces buffelskloofinus (incl. Bonitomyces gen. nov.) on dead twigs of unknown tree, Constrictochalara koukolii on living leaves of Itea rhamnoides colonised by a Meliola sp., Cylindromonium lichenophilum on Parmelina tiliacea, Gamszarella buffelskloofina (incl. Gamszarella gen. nov.) on dead insect, Isthmosporiella africana (incl. Isthmosporiella gen. nov.) on dead twigs of unknown tree, Nothoeucasphaeria buffelskloofina (incl. Nothoeucasphaeria gen. nov.), on dead twigs of unknown tree, Nothomicrothyrium beaucarneae (incl. Nothomicrothyrium gen. nov.) on dead leaves of Beaucarnea stricta, Paramycosphaerella proteae on living leaves of Protea caffra, Querciphoma foliicola on leaf litter, Rachicladosporium conostomii on dead twigs of Conostomium natalense var. glabrum, Rhamphoriopsis synnematosa on dead twig of unknown tree, Waltergamsia mpumalanga on dead leaves of unknown tree. Spain, Amanita fulvogrisea on limestone soil, in mixed forest, Amanita herculis in open Quercus forest, Vuilleminia beltraniae on Cistus symphytifolius. Sweden, Pachyella pulchella on decaying wood on sand-silt riverbank. Thailand, Deniquelata cassiae on dead stem of Cassia fistula, Stomiopeltis thailandica on dead twigs of Magnolia champaca. Ukraine, Circinaria podoliana on natural limestone outcrops, Neonematogonum carpinicola (incl. Neonematogonum gen. nov.) on dead branches of Carpinus betulus. USA, Exophiala wilsonii water from cooling tower, Hygrophorus aesculeticola on soil in mixed forest, and Neocelosporium aereum from air in a house attic. Morphological and culture characteristics are supported by DNA barcodes. [ABSTRACT FROM AUTHOR]

This article is the 14th in the Fungal Diversity Notes series, wherein we report 98 taxa distributed in two phyla, seven classes, 26 orders and 50 families which are described and illustrated. Taxa in this study were collected from Australia, Brazil, Burkina Faso, Chile, China, Cyprus, Egypt, France, French Guiana, India, Indonesia, Italy, Laos, Mexico, Russia, Sri Lanka, Thailand, and Vietnam. There are 59 new taxa, 39 new hosts and new geographical distributions with one new combination. The 59 new species comprise Angustimassarinakunmingense, Asterinalopi, Asterinabrigadeirensis, Bartaliniabidenticola, Bartaliniacaryotae, Buelliapruinocalcarea, Coltriciainsularis, Colletotrichumflexuosum, Colletotrichumthasutense, Coniochaetacaraganae, Coniothyriumyuccicola, Dematipyriformaaquatic, Dematipyriformaglobispora, Dematipyriformanilotica, Distoseptisporabambusicola, Fulvifomesjawadhuvensis, Fulvifomesmalaiyanurensis, Fulvifomesthiruvannamalaiensis, Fusariumpurpurea, Gerronemaatrovirens, Gerronemaflavum, Gerronemakeralense, Gerronemakuruvense, Grammotheletaiwanensis, Hongkongmyceschangchunensis, Hypoxyloninaequale, Kirschsteiniotheliaacutisporum, Kirschsteiniotheliacrustaceum, Kirschsteiniotheliaextensum, Kirschsteiniotheliaseptemseptatum, Kirschsteiniotheliaspatiosum, Lecanoraimmersocalcarea, Lepiotasubthailandica, Lindgomycesguizhouensis, Martheasmiuspallidoaurantiacus, Marasmiustangerinus, Neovaginatisporamangiferae, Pararamichloridiumaquisubtropicum, Pestalotiopsispiraubensis, Phacidiumchinaum, Phaeoisariagoiasensis, Phaeoseptumthailandicum, Pleurotheciumaquisubtropicum, Pseudocercosporavernoniae, Pyrenophoraverruculosa, Rhachomycescruralis, Rhachomyceshyperommae, Rhachomycesmagrinii, Rhachomycesplatyprosophi, Rhizomarasmiuscunninghamietorum, Skeletocutiscangshanensis, Skeletocutissubchrysella, Sporisoriumanadelphiae-leptocomae, Tetraploadashaoensis, Tomentellaexiguelata, Tomentellafuscoaraneosa, Tricholomopsislechatii, Vaginatisporaflavispora and Wetmoreanablastidiocalcarea. The new combination is Torulasundara. The 39 new records on hosts and geographical distribution comprise Apiosporaguiyangensis, Aplosporellaartocarpi, Ascochytamedicaginicola, Astrocystisbambusicola, Atheliarolfsii, Bambusicolabambusae, Bipolarisluttrellii, Botryosphaeriadothidea, Chlorophyllumsquamulosum, Colletotrichumaeschynomenes, Colletotrichumpandanicola, Coprinopsiscinerea, Corylicolaitalica, Curvulariaalcornii, Curvulariasenegalensis, Diaporthefoeniculina, Diaporthelongicolla, Diaporthephaseolorum, Diatrypellaquercina, Fusariumbrachygibbosum, Helicomaaquaticum, Lepiotametulispora, Lepiotapongduadensis, Lepiotasubvenenata, Melanconiellameridionalis, Monotosporellaerecta, Nodulosphaeriadigitalis, Palmiascomagregariascomum, Periconiabyssoides, Periconiacortaderiae, Pleopunctumellipsoideum, Psilocybekeralensis, Scedosporiumapiospermum, Scedosporiumdehoogii, Scedosporiummarina, Spegazziniadeightonii, Torulafici, Wiesneriomyceslaurinus and Xylariavenosula. All these taxa are supported by morphological and multigene phylogenetic analyses. This article allows the researchers to publish fungal collections which are important for future studies. An updated, accurate and timely report of fungus-host and fungus-geography is important. We also provide an updated list of fungal taxa published in the previous fungal diversity notes. In this list, erroneous taxa and synonyms are marked and corrected accordingly. [ABSTRACT FROM AUTHOR]

Fungi are eukaryotes that play essential roles in ecosystems. Among fungi, Basidiomycota is one of the major phyla with more than 40,000 described species. We review species diversity of Basidiomycota from five groups with different lifestyles or habitats: saprobic in grass/forest litter, wood-decaying, yeast-like, ectomycorrhizal, and plant parasitic. Case studies of Agaricus, Cantharellus, Ganoderma, Gyroporus, Russula, Tricholoma, and groups of lichenicolous yeast-like fungi, rust fungi, and smut fungi are used to determine trends in discovery of biodiversity. In each case study, the number of new species published during 2009–2020 is analysed to determine the rate of discovery. Publication rates differ between taxa and reflect different states of progress for species discovery in different genera. The results showed that lichenicolous yeast-like taxa had the highest publication rate for new species in the past two decades, and it is likely this trend will continue in the next decade. The species discovery rate of plant parasitic basidiomycetes was low in the past ten years, and remained constant in the past 50 years. We also found that the establishment of comprehensive and robust taxonomic systems based on a joint global initiative by mycologists could promote and standardize the recognition of taxa. We estimated that more than 54,000 species of Basidiomycota will be discovered by 2030, and estimate a total of 1.4–4.2 million species of Basidiomycota globally. These numbers illustrate a huge gap between the described and yet unknown diversity in Basidiomycota. [ABSTRACT FROM AUTHOR]

Fungal communities in above-ground tree tissues are hyperdiverse and are influenced by biotic interactions with other organisms living in or on these tissues. These biotic interactions are, however, still poorly understood. In this study, we aimed to understand how insect-associated gall formation on Eucalyptus foliage correlates with the diversity of foliar fungal communities in surrounding healthy leaf tissue, as well as the co-occurrence patterns among the members of the fungal community. We used ITS metabarcoding to characterise the foliar fungal communities of 179 individual E. grandis trees. These trees were assigned to infestation levels of the wasp Leptocybe invasa (Eulophidae: Hymenoptera), which causes gall formation on shoot tips and leaves of its host. Fungal community networks were calculated using a Pearson correlation coefficient. The composition and diversity of fungal communities were influenced by the severity of L. invasa infestations. We identified potential Eucalyptus pathogens with high sequence abundance at all disease severity levels, but network analysis indicated that the co-occurrence of potential pathogens between no to mild and medium to heavy infestation differed significantly. A better understanding of microbial interactions, especially the role of pathogens, can be useful for controlling disease- and beneficial host-associated microbial communities. [ABSTRACT FROM AUTHOR]

This article is the 13th contribution in the Fungal Diversity Notes series, wherein 125 taxa from four phyla, ten classes, 31 orders, 69 families, 92 genera and three genera incertae sedis are treated, demonstrating worldwide and geographic distribution. Fungal taxa described and illustrated in the present study include three new genera, 69 new species, one new combination, one reference specimen and 51 new records on new hosts and new geographical distributions. Three new genera, Cylindrotorula (Torulaceae), Scolecoleotia (Leotiales genus incertae sedis) and Xenovaginatispora (Lindomycetaceae) are introduced based on distinct phylogenetic lineages and unique morphologies. Newly described species are Aspergillus lannaensis, Cercophora dulciaquae, Cladophialophora aquatica, Coprinellus punjabensis, Cortinarius alutarius, C. mammillatus, C. quercoflocculosus, Coryneum fagi, Cruentomycena uttarakhandina, Cryptocoryneum rosae, Cyathus uniperidiolus, Cylindrotorula indica, Diaporthe chamaeropicola, Didymella azollae, Diplodia alanphillipsii, Dothiora coronicola, Efibula rodriguezarmasiae, Erysiphe salicicola, Fusarium queenslandicum, Geastrum gorgonicum, G. hansagiense, Helicosporium sexualis, Helminthosporium chiangraiensis, Hongkongmyces kokensis, Hydrophilomyces hydraenae, Hygrocybe boertmannii, Hyphoderma australosetigerum, Hyphodontia yunnanensis, Khaleijomyces umikazeana, Laboulbenia divisa, Laboulbenia triarthronis, Laccaria populina, Lactarius pallidozonarius, Lepidosphaeria strobelii, Longipedicellata megafusiformis, Lophiotrema lincangensis, Marasmius benghalensis, M. jinfoshanensis, M. subtropicus, Mariannaea camelliae, Melanographium smilaxii, Microbotryum polycnemoides, Mimeomyces digitatus, Minutisphaera thailandensis, Mortierella solitaria, Mucor harpali, Nigrograna jinghongensis, Odontia huanrenensis, O. parvispina, Paraconiothyrium ajrekarii, Parafuscosporella niloticus, Phaeocytostroma yomensis, Phaeoisaria synnematicus, Phanerochaete hainanensis, Pleopunctum thailandicum, Pleurotheciella dimorphospora, Pseudochaetosphaeronema chiangraiense, Pseudodactylaria albicolonia, Rhexoacrodictys nigrospora, Russula paravioleipes, Scolecoleotia eriocamporesi, Seriascoma honghense, Synandromyces makranczyi, Thyridaria aureobrunnea, Torula lancangjiangensis, Tubeufia longihelicospora, Wicklowia fusiformispora, Xenovaginatispora phichaiensis and Xylaria apiospora. One new combination, Pseudobactrodesmium stilboideus is proposed. A reference specimen of Comoclathris permunda is designated. New host or distribution records are provided for Acrocalymma fici, Aliquandostipite khaoyaiensis, Camarosporidiella laburni, Canalisporium caribense, Chaetoscutula juniperi, Chlorophyllum demangei, C. globosum, C. hortense, Cladophialophora abundans, Dendryphion hydei, Diaporthe foeniculina, D. pseudophoenicicola, D. pyracanthae, Dictyosporium pandanicola, Dyfrolomyces distoseptatus, Ernakulamia tanakae, Eutypa flavovirens, E. lata, Favolus septatus, Fusarium atrovinosum, F. clavum, Helicosporium luteosporum, Hermatomyces nabanheensis, Hermatomyces sphaericoides, Longipedicellata aquatica, Lophiostoma caudata, L. clematidis-vitalbae, Lophiotrema hydei, L. neoarundinaria, Marasmiellus palmivorus, Megacapitula villosa, Micropsalliota globocystis, M. gracilis, Montagnula thailandica, Neohelicosporium irregulare, N. parisporum, Paradictyoarthrinium diffractum, Phaeoisaria aquatica, Poaceascoma taiwanense, Saproamanita manicata, Spegazzinia camelliae, Submersispora variabilis, Thyronectria caudata, T. mackenziei, Tubeufia chiangmaiensis, T. roseohelicospora, Vaginatispora nypae, Wicklowia submersa, Xanthagaricus necopinatus and Xylaria haemorrhoidalis. The data presented herein are based on morphological examination of fresh specimens, coupled with analysis of phylogenetic sequence data to better integrate taxa into appropriate taxonomic ranks and infer their evolutionary relationships. [ABSTRACT FROM AUTHOR]

Fungi are ubiquitous in nature; that is, they are present everywhere on the planet; understanding the active state and functional capacity of airborne microbes associated with health of human, animal, and plant is critical for biosafety management. Here, we firstly and directly proved that there were about 40% active fungi in the air via rRNA amplicon sequencing and imaging flow cytometry simultaneously. Amplicon sequencing analysis showed differences between structures of active and total fungal community; Ascomycota were dominant in the active community, while Basidiomycota have low transcriptional activity across all samples. Notably, plant pathogenic fungi were predominant in the air, and more than 50% were active, including not only several common plant pathogens but also biotrophic fungi (Erysiphe sp. and Microbotryum sp.) and host-specific pathogens, which were generally considered to be inactive after leaving the host. Putative plant pathogens of eight genera were found active across the sampling season, indicating their superior ability to obtain nutrients even in barren nutrient environments. Interestingly, we detected several potentially active unrecorded fungi in China (Diatrype prominens , Septofusidium herbarum , Pseudomicrostroma glucosiphilum , and Uromycladium tepperianum), which suggested that they spread over a long distance by air and may cause diseases under favorable conditions. Our results suggested that maintaining transmission in air is an essential feature of many fungi including plant pathogens regardless of being a biotrophic, hemibiotrophic, or necrotrophic group. Moreover, two potentially active human pathogens and one animal pathogen were captured, which indicated their potential risks. This study provided a new perspective for more comprehensive understanding of airborne fungi, including their multidimensional lifestyle, state, functioning, and potential pathogenic risk. It also laid the foundation for further prediction and management of airborne microbial communities, which would be of interest for public health and agriculture. [ABSTRACT FROM AUTHOR]

Species delimitation is one of the most fundamental processes in biology. Biodiversity undertakings, for instance, require explicit species concepts and criteria for species delimitation in order to be relevant and translatable. However, a perfect species concept does not exist for Fungi. Here, we review the species concepts commonly used in Basidiomycota, the second largest phylum of Fungi that contains some of the best known species of mushrooms, rusts, smuts, and jelly fungi. In general, best practice is to delimitate species, publish new taxa, and conduct taxonomic revisions based on as many independent lines of evidence as possible, that is, by applying a so-called unifying (or integrative) conceptual framework. However, the types of data used vary considerably from group to group. For this reason we discuss the different classes of Basidiomycota, and for each provide: (i) a general introduction with difficulties faced in species recognition, (ii) species concepts and methods for species delimitation, and (iii) community recommendations and conclusions. [ABSTRACT FROM AUTHOR]

Here we review how evolving species concepts have been applied to understand yeast diversity. Initially, a phenotypic species concept was utilized taking into consideration morphological aspects of colonies and cells, and growth profiles. Later the biological species concept was added, which applied data from mating experiments. Biophysical measurements of DNA similarity between isolates were an early measure that became more broadly applied with the advent of sequencing technology, leading to a sequence-based species concept using comparisons of parts of the ribosomal DNA. At present phylogenetic species concepts that employ sequence data of rDNA and other genes are universally applied in fungal taxonomy, including yeasts, because various studies revealed a relatively good correlation between the biological species concept and sequence divergence. The application of genome information is becoming increasingly common, and we strongly recommend the use of complete, rather than draft genomes to improve our understanding of species and their genome and genetic dynamics. Complete genomes allow in-depth comparisons on the evolvability of genomes and, consequently, of the species to which they belong. Hybridization seems a relatively common phenomenon and has been observed in all major fungal lineages that contain yeasts. Note that hybrids may greatly differ in their post-hybridization development. Future in-depth studies, initially using some model species or complexes may shift the traditional species concept as isolated clusters of genetically compatible isolates to a cohesive speciation network in which such clusters are interconnected by genetic processes, such as hybridization. [ABSTRACT FROM AUTHOR]

Cyclocybe parasitica is a wood-destroying parasitic edible mushroom growing on diverse broad-leafed trees in New Zealand and other Pacific areas. Recent molecular systematics of European Cyclocybe aegerita, a newly delimited Asian phylum and of related species, corroborated the distinction of the chiefly saprobic cultivated edible mushroom C. aegerita from C. parasitica. Here, we show that C. parasitica exhibits a morpho-physiological trait characteristic to its European cousin, i.e., monokaryotic fruiting sensu stricto (basidiome formation without mating). Monokaryotic fruiting structures formed by C. parasitica ICMP 11668-derived monokaryons were categorized into four phenotypes. One of them displays ulcer-like structures previously reported from bracket fungi. Histology of dikaryotic and monokaryotic C. parasitica fruiting structures revealed anatomical commonalities and differences between them, and towards monokaryotic fruiting structures of C. aegerita. Mating experiments with C. parasitica strains representative of each fruiting phenotype identified compatible sibling monokaryons. Given reports on hypothetically monokaryotic basidiome field populations of ‘C. aegerita sensu lato’, it seems worthwhile to prospectively investigate whether monokaryotic fruiting s.str. occurs in nature. Sampling from such populations including karyotyping, comparative -omics, and competition assays may help to answer this question and provide evidence whether this trait may confer competitive advantages to a species capable of it. [ABSTRACT FROM AUTHOR]

The species of Anthracoidea on Carex are host-specific smut fungi restricted to sedges belonging to the same or closely related sections. They are characterized by sori that form black, hard bodies around aborted nuts of their hosts. In Carex sect. Hallerianae, only one species, C. halleriana, is known as a host of Anthracoidea. The taxonomic status of this smut fungus was problematic due to a lack of molecular data. It has been reported under different names, mainly as "A. caricis" or "A. irregularis". A comparative morphological study and molecular phylogenetic analysis, using LSU (large subunit) nuclear rDNA sequences, supported the recognition of a distinct species, A. hallerianae. The new species is described and illustrated based on material from Central Europe, the Iberian Peninsula, the Balkan Peninsula, the Mediterranean area and Transcaucasia. Citation: Denchev T. T., Denchev C. M., Koopman J., Begerow D. & Kemler M. 2021: Host specialization and molecular evidence support a distinct species of smut fungus, Anthracoidea hallerianae (Anthracoideaceae), on Carex halleriana (Cyperaceae). – Willdenowia 51: 57–67. Version of record first published online on 9 March 2021 ahead of inclusion in April 2021 issue. [ABSTRACT FROM AUTHOR]

The Basidiomycota constitutes a major phylum of the kingdom Fungi and is second in species numbers to the Ascomycota. The present work provides an overview of all validly published, currently used basidiomycete genera to date in a single document. An outline of all genera of Basidiomycota is provided, which includes 1928 currently used genera names, with 1263 synonyms, which are distributed in 241 families, 68 orders, 18 classes and four subphyla. We provide brief notes for each accepted genus including information on classification, number of accepted species, type species, life mode, habitat, distribution, and sequence information. Furthermore, three phylogenetic analyses with combined LSU, SSU, 5.8s, rpb1, rpb2, and ef1 datasets for the subphyla Agaricomycotina, Pucciniomycotina and Ustilaginomycotina are conducted, respectively. Divergence time estimates are provided to the family level with 632 species from 62 orders, 168 families and 605 genera. Our study indicates that the divergence times of the subphyla in Basidiomycota are 406–430 Mya, classes are 211–383 Mya, and orders are 99–323 Mya, which are largely consistent with previous studies. In this study, all phylogenetically supported families were dated, with the families of Agaricomycotina diverging from 27–178 Mya, Pucciniomycotina from 85–222 Mya, and Ustilaginomycotina from 79–177 Mya. Divergence times as additional criterion in ranking provide additional evidence to resolve taxonomic problems in the Basidiomycota taxonomic system, and also provide a better understanding of their phylogeny and evolution. [ABSTRACT FROM AUTHOR]

The group of the anthericolous Microbotryum species on Silene consists of narrowly host-specialized fungi. Despite intensive taxonomic and phylogenetic studies in this group over the past two decades, the actual species richness has not yet been fully uncovered. Thirteen of these species cause typical anther infection, with soral development restricted to the anthers. Three other species cause atypical infection, with soral development resulting in swollen and deformed flowers (completely filled with spores) and affecting both the anthers and the filaments. A comparative morphological study and molecular phylogenetic analyses, using ITS and LSU rDNA sequences, revealed a new species, Microbotryum arcticum, causing anther infection of Silene uralensis subsp. arctica. Microbotryum arcticum is described and illustrated on the basis of material from Greenland and the eastern Canadian Arctic. An emended description of M. savilei, which causes atypical infection of the same host plant in the eastern Canadian Arctic, is also given. Morphological characters of healthy flowers of S. uralensis subsp. arctica are compared with those of flowers with anther infection and with those of flowers with atypical infection. [ABSTRACT FROM AUTHOR]

Trees in the genus Vachellia (previously Acacia) are commonly infected by the gall-inducing rusts Ravenelia macowaniana and R. evansii. Rust galls bearing aecial infections and relating uredinial and telial infections on the leaves of nine Vachellia species not previously recorded to be infected by Ravenelia spp. have recently been collected in South Africa. The rust fungi causing these infections were characterised using molecular phylogenetic analyses of DNA sequence data of the LSU and ITS rDNA regions as well as morphological examinations. The host range of R. macowaniana and R. evansii was thus re-assessed and extended from four to nine species and from one to three species, respectively. Application of Principal Component Analyses (PCA) of telial morphological characters provided evidence of an effect of the host species on the teliospore morphology in R. evansii, but only minor effects in R. macowaniana. A novel gall-inducing Ravenelia sp. closely related to R. macowaniana, was found on Vachellia xanthophloea and it is described here as R. xanthophloeae. [ABSTRACT FROM AUTHOR]

The increasing number and duration of inundations is reported to be a consequence of climate change and may severely compromise non-adapted macroorganisms. The effect of flooding events on terrestrial and aquatic microbial communities is, however, less well understood. They may respond to the changed abiotic properties of their native habitat, and the native community may change due to the introduction of alien species. We designed an experiment to investigate the effect of five different flooding durations on the terrestrial and aquatic communities of eukaryotic microorganism, using the AquaFlow mesocosms. With amplicon sequencing of the small subunit (SSU) and internal transcribed spacer (ITS) rRNA gene regions, we analyzed community compositions directly before and after flooding. Subsequently, they were monitored for another 28 days, to determine the sustainability of community changes. Our results revealed a temporary increase in similarity between terrestrial and aquatic communities according to OTU composition (operational taxonomic unit, serves as a proxy for species). Increased similarity was mainly caused by the transmission of OTUs from water to soil. A minority of these were able to persist in soil until the end of the experiment. By contrast, the vast majority of soil OTUs was not transmitted to water. Flooding duration affected the community structure (abundance) more than composition (occurrence). Terrestrial communities responded immediately to flooding and the flooding duration influenced the community changes. Independent from flooding duration, all terrestrial communities recovered largely after flooding, indicating a remarkable resilience to the applied disturbances. Aquatic communities responded immediately to the applied inundations too. At the end of the experiment, they grouped according to the applied flooding duration and the amount of ammonium and chloride that leached from the soil. This indicates a sustained long-term response of the aquatic communities to flooding events. [ABSTRACT FROM AUTHOR]

Host specialization is a key evolutionary process for the diversification and emergence of new pathogens. However, the molecular determinants of host range are poorly understood. Smut fungi are biotrophic pathogens that have distinct and narrow host ranges based on largely unknown genetic determinants. Hence, we aimed to expand comparative genomics analyses of smut fungi by including more species infecting different hosts and to define orphans and positively selected genes to gain further insights into the genetics basis of host specialization. We analyzed nine lineages of smut fungi isolated from eight crop and non-crop hosts: maize, barley, sugarcane, wheat, oats, Zizania latifolia (Manchurian rice), Echinochloa colona (a wild grass), and Persicaria sp. (a wild dicot plant). We assembled two new genomes: Ustilago hordei (strain Uhor01) isolated from oats and U. tritici (strain CBS 119.19) isolated from wheat. The smut genomes were of small sizes, ranging from 18.38 to 24.63 Mb. U. hordei species experienced genome expansions due to the proliferation of transposable elements and the amount of these elements varied among the two strains. Phylogenetic analysis confirmed that Ustilago is not a monophyletic genus and, furthermore, detected misclassification of the U. tritici specimen. The comparison between smut pathogens of crop and non-crop hosts did not reveal distinct signatures, suggesting that host domestication did not play a dominant role in shaping the evolution of smuts. We found that host specialization in smut fungi likely has a complex genetic basis: different functional categories were enriched in orphans and lineage-specific selected genes. The diversification and gain/loss of effector genes are probably the most important determinants of host specificity. [ABSTRACT FROM AUTHOR]

Botryosphaeria dothidea is the type species of Botryosphaeria (Botryosphaeriaceae, Botryosphaeriales). Fungi residing in this order are amongst the most widespread and important canker and dieback pathogens of trees worldwide, with B. dothidea one of the most common species on a large number of hosts. Its taxonomic circumscription has undergone substantial change in the past decade, making it difficult to interpret the large volume of literature linked to the name B. dothidea. This pathogen profile synthesizes the current understanding of B. dothidea pertaining to its distribution, host associations and role as a pathogen in managed and natural woody environments. The prolonged latent infection or endophytic phase is of particular importance, as it implies that the fungus can easily pass undetected by quarantine systems in traded living plants, fruits and other plant parts. Infections typically become obvious only under conditions of host stress, when disease symptoms develop. This study also considers the knowledge emerging from the recently sequenced B. dothidea genome, elucidating previously unknown aspects of the species, including mating and host infection strategies. Despite more than 150 years of research on B. dothidea, there is clearly much to be learned regarding this global tree pathogen. This is increasingly important given the stresses imposed on various woody hosts as a result of climate change. Taxonomy Botryosphaeria dothidea (Moug. ex Fr) Ces. & De Not, 1863. Kingdom Fungi, Phylum Ascomycota, Class Dothideomycetes, Order Botryosphaeriales, Family Botryosphaeriaceae, Genus Botryosphaeria, Species dothidea. Host range Confirmed on more than 24 host genera, including woody plants, such as Acacia (= Vachellia), Eucalyptus, Vitis and Pistachio. Disease symptoms Associated with twig, branch and stem cankers, tip and branch dieback, fruit rot, blue stain and plant death. Useful websites The Botryosphaeria site for detailed morphological descriptions (/ );, Systematic Mycology and Microbiology Laboratory Fungal Database for all literature and associated hosts (/ );, TreeBASE link for the combined ITS and TEF-1α tree (:S18906);, DOE Joint Genome Institute, JGI Mycocosm for the Botryosphaeria dothidea genome (). [ABSTRACT FROM AUTHOR]

Itajahya is a member of Phallales (Agaricomycetes), which, based on the presence of a calyptra and DNA sequence data for I. rosea, has recently been raised to generic status from a subgenus of Phallus. The type species of the genus, I. galericulata, is commonly known as the Jacaranda stinkhorn in Pretoria, South Africa, which is the only area where the fungus is known outside the Americas. The common name is derived from its association with the South American originating Jacaranda mimosifolia trees in the city. The aim of this study was to consider the unusual occurrence of the fungus in South Africa, to place it on the available Phallales phylogeny and to test whether it merits generic status. Fresh basidiomes were collected during the summer of 2015 and sequenced. Phylogenetic analyses were based on sequence data for the nuc-LSU-rDNA (LSU) and ATPase subunit 6 (ATP6) regions. The results showed that I. rosea and I. galericulata are phylogenetically related. They are also clearly distinguished from other members of Phallales such as Phallus spp. and Dictyophora spp., and so our new data supports the raising of Itajahya to the generic level. [ABSTRACT FROM AUTHOR]

A letter to the editor is presented in response to an article by S. Fares in volume 519 of the periodical which discussed the sustainable development of Europe's forests.

A letter to the editor is presented concerning an article by Silvano Fares and colleagues in issue 519 of "Nature" concerning management of European forests to promote carbon storage and resilience and suggests the suggestions run counter to guidelines for biodiversity sustainability.

A letter to the editor is presented in response to an article by M. Aloosh and A. Aloosh in volume 520 of the journal which discusses the impact of economic sanctions on public health in Iran.

A letter to the editor is presented in response to an article by Isabel Marques in volume 520 of the periodical in which she suggested reforms to botanical education that focused on a more molecular approach.

A letter to the editor is presented concerning the International Network for Alpine Research Catchment Hydrology which was launched in 2015 by the World Climate Research Programme's Global Energy and Water Exchanges project to improve the predictability of water supplies from mountainous areas.

The Kingdom Fungi adds substantially to the diversity of life, but due to their cryptic morphology and lifestyle, tremendous diversity, paucity of formally described specimens, and the difficulty in isolating environmental strains into culture, fungal communities are difficult to characterize. This is especially true for endophytic communities of fungi living in healthy plant tissue. The developments in next generation sequencing technologies are, however, starting to reveal the true extent of fungal diversity. One of the promising new technologies, namely semiconductor sequencing, has thus far not been used in fungal diversity assessments. In this study we sequenced the internal transcribed spacer 1 (ITS1) nuclear encoded ribosomal RNA of the endophytic community of the economically important tree, Eucalyptus grandis, from South Africa using the Ion Torrent Personal Genome Machine (PGM). We determined the impact of various analysis parameters on the interpretation of the results, namely different sequence quality parameter settings, different sequence similarity cutoffs for clustering and filtering of databases for removal of sequences with incomplete taxonomy. Sequence similarity cutoff values only had a marginal effect on the identified family numbers, whereas different sequence quality filters had a large effect (89 vs. 48 families between least and most stringent filters). Database filtering had a small, but statistically significant, effect on the assignment of sequences to reference sequences. The community was dominated by Ascomycota, and particularly by families in the Dothidiomycetes that harbor well-known plant pathogens. The study demonstrates that semiconductor sequencing is an ideal strategy for environmental sequencing of fungal communities. It also highlights some potential pitfalls in subsequent data analyses when using a technology with relatively short read lengths. [ABSTRACT FROM AUTHOR]

Anther smuts in the genus Microbotryum often show very high host specificity toward their caryophyllaceous hosts, but some of the larger host groups such as Dianthus are crucially undersampled for these parasites so that the question of host specificity cannot be answered conclusively. In this study we sequenced the internal transcribed spacer (ITS) region of members of the Microbotryum dianthorum species complex as well as their Dianthus hosts. We compared phylogenetic trees of these parasites including sequences of anther smuts from other Caryophyllaceae, mainly Silene, with phylogenies of Caryophyllaceae that are known to harbor anther smuts. Additionally we tested whether observed patterns in parasites are due to shared ancestry or if geographic separation is a factor that should be taken into consideration in delimitating species. Parasites on Dianthus showed mainly an arbitrary distribution on Dianthus hosts, whereas parasites on other Caryophyllaceae formed well-supported monophyletic clades that corresponded to restricted host groups. The same pattern was observed in the Caryophyllaceae studied: morphologically described Dianthus species did not correspond well with monophyletic clades based on molecular data, whereas other Caryophyllaceae mainly did. We suggest that these different patterns primarily result from different breeding systems and speciation times between different host groups as well as difficulties in species delimitations in the genus Dianthus. [ABSTRACT FROM AUTHOR]

The grass smuts comprise a speciose group of biotrophic plant parasites, so-called Ustilaginaceae, which are specifically adapted to hosts of sweet grasses, the Poaceae family. Mating takes a central role in their life cycle, as it initiates parasitism by a morphological and physiological transition from saprobic yeast cells to pathogenic filaments. As in other fungi, sexual identity is determined by specific genomic regions encoding allelic variants of a pheromone-receptor (PR) system and heterodimerising transcription factors. Both operate in a biphasic mating process that starts with PR-triggered recognition, directed growth of conjugation hyphae, and plasmogamy of compatible mating partners. So far, studies on the PR system of grass smuts revealed diverse interspecific compatibility and mating type determination. However, many questions concerning the specificity and evolutionary origin of the PR system remain unanswered. Combining comparative genetics and biological approaches, we report on the specificity of the PR system and its genetic diversity in 10 species spanning about 100 million years of mating type evolution. We show that three highly syntenic PR alleles are prevalent among members of the Ustilaginaceae, favouring a triallelic determination as the plesiomorphic characteristic of this group. Furthermore, the analysis of PR loci revealed increased genetic diversity of single PR locus genes compared to genes of flanking regions. Performing interspecies sex tests, we detected a high potential for hybridisation that is directly linked to pheromone signalling as known from intraspecies sex. Although the PR system seems to be optimised for intraspecific compatibility, the observed functional plasticity of the PR system increases the potential for interspecific sex, which might allow the hybrid-based genesis of newly combined host specificities. [ABSTRACT FROM AUTHOR]