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Best tree of the Maximum Likelihood analysis of caryophyllaceous anther smuts in the genus Microbotryum based on a concatenated ITS and LSU dataset. The tree was rooted with M. anomalum (J. Kunze ex G. Winter) Vánky (based on Kemler & al. 2006). Bootstrap values ≥ 50 are indicated above branches.
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... The position and branch lengths of the molecular phylogenetic analysis of the Greek specimen of Microbotryum on Stellaria media (Fig. 1, Table 1) indicates that this specimen belongs to a distinct species. The phylogenetic analysis confirmed previous phylogenetic relationships within the genus Microbotryum (Denchev et al. 2019(Denchev et al. , 2020b(Denchev et al. , 2023. The Greek specimen falls within the group of caryophyllaceous seed parasites. ...
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.
... Phylogenetic studies have shown that the presentation of spores in the anthers, reflecting adaptation of the pathogen to vector-based transmission, is a trait derived several times in the genus Microbotryum (Kemler et al., 2006). However, some species of Microbotryum within the clade that predominantly infects Silene and its relatives have secondarily evolved sporulation in the whole flower (Denchev et al., 2019;Kemler et al., 2020), suggesting a possible reversion to aerial transmission. Further studies are needed to assess if evolutionary changes in transmission mode occur within contrasting host populations of D. pavonius, and if there are "trade-offs" (negative genetic correlations) between effective transmission by the two modes, for example, as might be determined by morphological features of spores. ...
Pathogen transmission mode is a key determinant of epidemiological outcomes. Theory shows that host density can influence the spread of pathogens differentially depending on their mode of transmission. Host density could therefore play an important role in determining the pathogen transmission mode. We tested theoretical expectations using floral arrays of the alpine carnation Dianthus pavonius in field experiments of spore dispersal of the anther‐smut fungus, Microbotryum, by vector (pollinator)‐based floral transmission and passive aerial transmission at a range of host densities. Pollinators deposited fewer spores per plant at high host density than at lower density (ranging from a 0.2–2 m spacing between plants), and vector‐based spore deposition at higher densities declined more steeply with distance from diseased plant sources. In contrast, while aerial spore deposition declined with distance from the diseased source, the steepness of this decline was independent of host density. Our study indicates that the amount and distance of vector‐based transmission are likely to be a nonmonotonic function of host density as a result of vector behavior, which is not readily encapsulated by fixed dispersal functions. We conclude that the spatial spread of pathogens by vectors is likely to be greater at lower and intermediate densities, whereas the spatial spread of aerially transmitted pathogens would be greater at high densities. These contrasting patterns could lead to differential importance of each transmission mode in terms of its contribution to subsequent infections across host densities.
... Microbotryum Lév. Notes: In its broad circumscription, Microbotryum comprises 99 species (Denchev and Denchev 2011;Vánky 2011;Piątek et al. 2012Piątek et al. , 2013Denchev et al. 2019;Ziegler et al. 2018;Kemler et al. 2020). During an examination of Polygonum specimens in the herbarium of the Meise Botanic Garden, Belgium (BR), a smut fungus belonging to Microbotryum was found on a specimen of Polygonum polycnemoides (BR, s.n.) from Turkey. ...
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. quercofocculosus, Coryneum fagi, Cruentomycena uttarakhandina, Cryptocoryneum rosae, Cyathus uniperidiolus, Cylindrotorula indica, Diaporthe chamaeropicola, Didymella azollae, Diplodia alanphillipsii, Dothiora coronicola, Efbula 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, Tubeufa 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 fci, 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 favovirens, E. lata, Favolus septatus, Fusarium atrovinosum, F. clavum, Helicosporium luteosporum, Hermatomyces nabanheensis, Hermatomyces sphaericoides, Longipedicellata aquatica, Lophiostoma caudata, L. clematidisvitalbae, Lophiotrema hydei, L. neoarundinaria, Marasmiellus palmivorus, Megacapitula villosa, Micropsalliota globocystis, M. gracilis, Montagnula thailandica, Neohelicosporium irregulare, N. parisporum, Paradictyoarthrinium difractum, Phaeoisaria aquatica, Poaceascoma taiwanense, Saproamanita manicata, Spegazzinia camelliae, Submersispora variabilis, Thyronectria caudata, T. mackenziei, Tubeufa 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.
... Microbotryomycetes comprises seven orders (Heitmaniales, Heterogastridiales, Kriegeriales, Leucosporidiales, Microbotryales, Rosettozymales, Sporidiobolales), eleven families (Camptobasidiaceae, Chrysozymaceae, Colacogloeaceae, Heitmaniaceae, Heterogastridiaceae, Kriegeriaceae, Leucosporidiaceae, Microbotryaceae, Rosettozymaceae, Sporidiobolaceae, Ustilentylomataceae), 44 genera, and about 300 species (Wang et al. 2015c;Begerow et al. 2018;Denchev et al. 2019Denchev et al. , 2020He et al. 2019;Kemler et al. 2020;Li et al. 2021). It is a diverse class, containing mycoparasites, plant parasites, and saprotrophic yeasts with largely distinct ecological, morphological, and ultrastructural features. ...
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.
... However, as genes responsible for host adaptation are not necessarily involved in sexual reproduction, the way in which host adaptation plays a role in reproductive isolation and speciation of smut fungal populations on different hosts is still an active field of research. Phylogenetic studies support the hypothesis that host adaptation and speciation are linked, as the vast majority of smut fungi on different host species form evolutionary independent lineages (e.g., Begerow et al. 2002;Lutz et al. 2005;Stoll et al. 2005;Piątek et al. 2013;Denchev et al. 2019). On the other hand, work on smut fungal populations shows ongoing gene flow between closely related parasite species (Petit et al. 2017) and experimental work clearly indicates that even distantly related species are able to hybridize (e.g. ...
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.
... The Caryophyllaceae is a large family of dicotyledonous plants (Greenberg & Donoghue 2011), and its species are hosts for many plant-parasitic microfungi, among them at least 38 species of smut fungi assigned to the genera Microbotryum Lév. and Thecaphora Fingerh. (Vánky (2012), with additional species in Denchev et al. (2009) ;Denchev & Denchev (2011); Piątek et al. (2012Piątek et al. ( , 2013; Denchev et al. (2019); Kemler et al. (2020)), which form their sori in the floral organs. Although there are similarities in the appearance of the infections and the affected hosts, these two genera are not closely related; Microbotryum is in the Microbotryaceae within the subphylum Pucciniomycotina, and Thecaphora is the only g en u s i n t h e G lo m os p o ri ac e ae wi th in t he Ustilaginomycotina. ...
Thecaphora melandrii (Syd.) Vánky & M.Lutz infects species in the Caryophyllacaeae forming sori with spore balls in the floral organs. We report new finds from Britain, supported by phylogenetic analysis, that confirm its occurrence on Silene uniflora Roth. We review published and web accessible records and note the relatively few records of this smut, its sparse distribution, confined to Europe but scattered predominantly from central to eastern Europe. Analysis of the rDNA ITS and 28S sequences demonstrates little variability among specimens, even those parasitising different host genera, which suggests that the species has evolved relatively recently. Some
Microbotryum species infect the same host plants, and we found two species, M. lagerheimii Denchev and M. silenes-inflatae (DC. ex Liro) G.Deml & Oberw., in the same locations as T. melandrii, identified by morphology and molecular phylogenetic analysis. These species may form a stable multi-species community of parasites of Silene uniflora.
... The fungal genus Microbotryum, with 98 described species (Denchev and Denchev 2011;Vánky 2012;Piątek et al. 2012Piątek et al. , 2013Ziegler et al. 2018;Denchev et al. 2019), is mainly known for containing the agents of anther smut disease in hosts of the Caryophyllaceae. In this group of parasites, it is assumed that the production of teliospores in the anthers is most likely an adaptation to host pollinators that increases parasite dispersal. ...
... Caryophyllaceae species evolved only once. Following this single colonization event, a major radiation led to the formation of Microbotryum species on many members of the Caryophyllaceae (Lutz et al. 2005(Lutz et al. , 2008; Le Gac et al. 2007;Refrégier et al. 2008;Piątek et al. 2012Piątek et al. , 2013Smith et al. 2017;Denchev et al. 2019;Tang et al. 2019). Anther smut infections are widespread throughout host distributions (Hood et al. 2010). ...
... Research on caryophyllaceous anther smuts has shown that there exists a high number of often host-species-specific parasite species, many of which have been described only in recent years based on molecular data, host information, and sometimes also on morphological differences. Currently, 24 species are recognized (Lutz et al. 2005(Lutz et al. , 2008Denchev 2007a, b;Denchev et al. 2009Denchev et al. , 2019Denchev and Denchev 2011;Vánky 2012;Piątek et al. 2012Piątek et al. , 2013. Based on morphological species concepts prevalent in the existing literature, species diversity and host specificity seem lower in ovule/seed parasites (ten species), but our phylogenies indicate that host specific lineages exist in this group, e.g., the smut fungi infecting different Cerastium spp. ...
The smut fungal genus Microbotryum (Microbotryales, Pucciniomycotina) contains species that parasitize plants from many
different lineages of euasterids, with host specificity of individual parasite species in general being exceptionally high.
Additionally, it has been shown that the location of spore production in some species is related to spore dispersal. In this
phylogenetic study based on ITS and LSU rDNA data of 57 Microbotryum spp., host spectra and sorus location are mapped
on the phylogeny of Microbotryum species in order to understand the macroevolutionary patterns of these two traits.We find that
monophyletic parasite clades correspond well with monophyletic host clades and also that monophyletic parasite groups in
general produce their spores in the same plant organ. Ancestral state reconstruction inferred the most probable ancestral trait for
sorus location being leaves and the most probable ancestral host family for the genus Microbotryum as being the Polygonaceae.
According to molecular analyses, a newly sequenced specimen of Ustilago ducellieri, a seed parasite on Arenaria leptoclados,
previously treated as synonym of Microbotryum duriaeanum, belongs to a lineage distinct from specimens of M. duriaeanum. A
new combination, Microbotryum ducellieri, is accordingly proposed. Taxonomic implications of the presented analyses for the
genera Bauhinus and Haradaea are briefly discussed.
... It is a group of seventeen, highly host specific fungi. Regarding the sorus morphology, they may be divided into two groups: (i) species causing typical anther infection, with sori restricted to the anthers (four species, M. arcticum, M. lagerheimii, M. silenes-acaulis, and M. stellariae, in Greenland); and (ii) species causing atypical infection, with sori usually formed not only in the anthers but also in the filaments, and causing formation of swollen and deformed flowers, completely filled with spore mass (one species, M. savilei, potentially occurring in Greenland) (Denchev et al. 2019). ...
... Microbotryum arcticum on Silene uralensis subsp. arctica was recently described from the High Arctic of Greenland and the Canadian Arctic Archipelago (Denchev et al. 2019). Four specimens from North Greenland (including the holotype) and a specimen from East Greenland are listed in the protologue, as examined. ...
... Other six specimens from East Greenland -reported by Hagen (1947) as 'U. violacea' on 'Melandrium apetalum', but not found in the herbarium in Oslo -were also considered as belonging to M. arcticum (Denchev et al. 2019). ...
The first taxonomic treatment of the smut fungi in Greenland is provided. A total of 43 species in 11 genera are treated and illustrated by photographs of sori, microphotographs of spores in LM and SEM, and distribution maps. Two species, Anthracoidea pseudofoetidae and Urocystis tothii, are recorded as new from North America. Thirteen species, Anthracoidea altera, A. capillaris, A. limosa, A. liroi, A. pseudofoetidae, A. scirpoideae, A. turfosa, Microbotryum lagerheimii, M. stellariae, Schizonella elynae, Stegocintractia luzulae, Urocystis fischeri, and U. tothii, are reported for the first time from Greenland. Three new fungus-host combinations, Anthracoidea capillaris on Carex boecheriana, Anthracoidea pseudofoetidae on Carex maritima, and Urocystis tothii on Juncus biglumis, are given. Five plant species are reported as new hosts of smut fungi in Greenland, namely, Carex nigra for Anthracoidea heterospora, C. canescens for Anthracoidea karii, C. fuliginosa subsp. misandra for Anthracoidea misandrae, C. maritima for Orphanomyces arcticus, and C. fuliginosa subsp. misandra for Schizonella melanogramma. Three species, Microbotryum violaceum s. str. (recorded as ‘Ustilago violacea’), Urocystis anemones, and U. junci, which were previously reported from Greenland, are considered wrongly identified. Additional distribution records are given for 12 species from Greenland: Anthracoidea bigelowii, A. caricis, A. elynae, A. lindebergiae, A. misandrae, A. nardinae, A. rupestris, A. scirpi, Schizonella melanogramma, Stegocintractia hyperborea, Urocystis agropyri, and U. sorosporioides. The most numerous distribution groups are the following: circumpolar–alpine and Arctic–alpine species – 14; circumboreal–polar species – 10; and circumpolar and Arctic species – 6. The most widely distributed smut fungi in Greenland were Anthracoidea bigelowii, A. elynae, Microbotryum bistortarum, and M. vinosum. Most species were found in the High Arctic zone (29 species), while from the Low Arctic zone and the Subarctic zone, 26 and 19 species were known, respectively. Ten species, Anthracoidea bigelowii, A. capillaris, A. elynae, Microbotryum bistortarum, M. koenigiae, M. pustulatum, M. silenes-acaulis, M. vinosum, Schizonella elynae, and Urocystis sorosporioides, were recorded from all three zones. Only plants belonging to six families, Cyperaceae, Poaceae, Juncaceae, Ranunculaceae, Caryophyllaceae, and Polygonaceae, out of a total of 55 in the flora of Greenland, hosted smut fungi. Cyperaceae was the plant family with most host species (23). Carex was the genus with the highest number of host species (22). The total number of the host plants (45 species) was 8.5 % out of a total of 532 vascular plants in the flora of Greenland. A new combination in Carex, C. macroprophylla subsp. subfilifolia, is proposed for Kobresia filifolia subsp. subfilifolia.
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.
