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Entyloma bullosum on Ranunculus paludosus. a. Macroscopic symptoms of infection; b-d. spores, as seen in light microscopy, median and superficial views (from neotype).-Scale bars = 10 µm. 

Entyloma bullosum on Ranunculus paludosus. a. Macroscopic symptoms of infection; b-d. spores, as seen in light microscopy, median and superficial views (from neotype).-Scale bars = 10 µm. 

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Plant pathogenic smut fungi in the broader sense can be divided into the Ustilaginomycetes, which cause classical smut symptoms with masses of blackish spores being produced in a variety of angiosperms, and the Exobasidiomycetes, which are often less conspicuous, as many do not shed large amounts of blackish spores. The leaf-spot causing members of...

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... bullosum (Sacc.) J. Kruse, M. Lutz, Piątek & Thines, comb. nov. -MycoBank MB823957; Fig. 4 Basionym. Caeoma bullosum Sacc., Nuovo Giorn. Bot. Ital., n.s. 22: 32. ...

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... Species of Entyloma (Fungi: Ustilaginomycotina) infect dicotyledonous host plants where they form leaf spots (called sori) filled with smooth, rarely verruculose spores. The leaf spots are often associated with an external entylomella-like asexual morph composed of hyaline conidiophores and conidia (Aime et al., 2018;Kruse et al., 2018;Vánky, 2012). There are a few saprobic Entyloma species that colonize apples causing white haze disorder (Boekhout et al., 2006;Richter et al., 2019). ...
... Eschscholzia and Dendromecon are closely related genera of the family Papaveraceae (Hoot et al., 2015). In the light of high host specificity reported for Entyloma species (Begerow et al., 2002;Kruse et al., 2018;Vánky, 2012), it is uncertain whether Entyloma on Dendromecon sp. and on E. californica are the same species. The main host, E. californica, popularly called California poppy, is widespread in western United States and Mexico (Clark, 1997) and is commonly cultivated in North America and Europe as an ornamental plant. ...
... In order to attain an ITS + LSU sampling as complete as possible and to make sure that, for each species, both regions were from the same specimen, we sequenced specimens of an additional 29 Entyloma species. GenBank accession numbers of the sequences used are given in Figure 1 (Begerow et al., 1997(Begerow et al., , 2002Boekhout et al., 1995Boekhout et al., , 2006Denchev et al., 2021;Hamamoto et al., 2000;Hawkes et al., 2011;Henricot et al., 2013;Kemler et al., 2007;Kruse et al., 2018;Kruse & Thines, 2019;Lutz & Piątek, 2016;Marchese et al., 2021;Rooney-Latham et al., 2017;Savchenko et al., 2014Savchenko et al., , 2015Vánky & Lutz, 2010;Vu et al., 2019;Wang et al., 2015). ...
Article
Entyloma includes pathogenic and saprobic species that infect or colonize dicotyledonous host plants. Although most Entyloma species are known only from native areas of occurrence, some species were introduced with their host plants and spread outside their natural areas. The identification of introduced species is important for detection and management of invasive species. In this study, the morphology, phylogeny and species boundaries of Entyloma eschscholziae, recently introduced from North America to Europe, are revisited. Morphology was similar among the type and other specimens of E. eschscholziae analysed on Eschscholzia californica. Both asexual and sexual morphs were observed. The rDNA ITS1-5.8S-ITS2 sequences of the E. eschscholziae specimens from Europe and New Zealand and the environmental sequence obtained from grassland soil in California, United States, were identical. Morphological and molecular analyses confirm that the causative agents of white smut on E. californica in native (North America) and introduced (Europe, New Zealand) areas belong to the same species. DNA barcodes obtained in this study (especially ITS sequence from the designated epitype specimen) could be used to facilitate its molecular identification. Specimens on Dendromecon rigida, previously assigned to E. eschscholziae, are morphologically distinct. An attempt to obtain DNA barcode data from degraded holotype material was not successful and no more recent material is available. However, based on the morphological differences and high host specificity found in Entyloma spp., it is appropriate to describe a new species, Entyloma dendromeconis, for this smut pathogen.
... It was introduced by de Bary [ 1 ] and is typified with E. microsporum (Unger) J. Schröt. on Ranunculus repens L. Entyloma is a large genus, with 189 recognised species on host plants belonging to 27 families of eudicots [2][3][4][5][6][7][8][9][10][11][12]. The species of Entyloma form sori in vegetative organs of plants, mostly in leaves, rarely in stems or roots, usually forming few to numerous spots, sometimes swellings or galls formed by hypertrophic growth of host tissue. ...
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Three smut fungi are reported for the first time from Bulgaria: Entyloma fergussonii on Myosotis orbelica, E. pastinacae on Pastinaca sativa subsp. urens, and E. thalictri on Thalictrum minus, extending the known species of Entyloma to 11 in this country. Brief descriptions and illustrations are provided for the newly recorded smut fungi. Myosotis orbelica is a new host plant for Entyloma fergussonii.
... Reliable identification of new species requires a substantial investment of time and resources to document diverse properties such as morphology, physiology, life-cycle, host relationships, and nucleotide sequences not restricted to the common DNA-barcode, ribosomal ITS. High numbers of predicted but yet undescribed species needs the mycological Bezerra et al. 2018;Brewer et al. 2014;Cabañes et al. 2011Cabañes et al. , 2016Cao et al. 2018;Chamnanpa et al. 2013;Crous et al. 2019;, 2012, 2016cDenchev et al. 20102012, 2020bFrancesca et al. 2016;Guo and Xu 2013;He et al. 2011;Honnavar et al. 2016;Kennedy et al. 2012;Aime 2016, 2017;Kruse and Thines 2019;Kruse et al. 2018aKruse et al. , 2018bKruse et al. , 2018cKruse et al. , 2020Li et al. 2014bLi et al. , 2017aLi et al. , 2017bLi et al. , 2019Li et al. , 2020Limtong et al. 2017;Lorch et al. 2018;Lutz et al. 2012;Macedo et al. 2016;McTaggart et al. 2020;Mekha et al. 2014;Nasr et al. 2014Nasr et al. , 2017Nasr et al. , 2019Oliveira et al. 2013Oliveira et al. , 2014Piątek 2014;Piątek and Shivas 2011;Piątek et al. , 2012aPiątek et al. ,b, 2013aPiątek et al. ,b, 2015aPiątek et al. ,b, 2016Piepenbring et al. 2012Piepenbring et al. , 2020Richter et al. 2019;Riess et al. 2019;Rooney-Latham et al. 2017;Rush and Aime 2013;Rush et al. 2020;Savchenko 2015;Savchenko and Carris 2017;Savchenko et al. 2013Savchenko et al. , 2014aShivas et al. 2011Shivas et al. , 2012Singh et al. 2020;Sipiczky 2020;Stajsic et al. 2018;Sun et al. 2018;Thanh et al. 2012Thanh et al. , 2013Vánky 2011bVánky , 2013Vánky and Abbasi 2011a,b;Vánky and Salo 2011;Ziegler et al. 2018 community to speed-up publication rates to formally document diversity in a race against biodiversity loss (Pearce et al. 2020). ...
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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.
... The polyphasic approach involves using multiple data types and methods when defining species boundaries for cryptic fungal species or species complexes (Hudler et al. 1998). Most of the studies suggest an integrative taxonomic approach combining evidence from genealogical, biological, phenotypic and phylogenetic approaches to include different aspects of fungal autecology, physiology, and biochemistry to resolve cryptic species or species complexes (Yang and Rannala 2010;Padial et al. 2010;Udayanga et al. 2014b;Haelewaters et al. 2018;Kruse et al. 2018;Bhunjun et al. 2020;Lücking et al. 2020). When defining species boundaries for cryptic species or for species complexes, it is recommended to apply genealogical concordance phylogenetic species recognition, coalescentbased or distance-based techniques together with other types of data such as morphological and ecological data. ...
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This is the opening paper in the special issue of Fungal Diversity, which collates the data on defining species. Defining and recognizing species has long been a controversial issue. Since Darwin's proposed origin of species, over 30 species criteria have been brought forth and used to define species boundaries. In recent times, phylogenetic analyses based on multiple loci have been extensively used as a method to define species boundaries. However, only a few mycologists are aware that phylogenetic species criteria can mask discordances among fungal groups, leading to inaccurately defined species boundaries. In the current review, we discuss species recognition criteria, how and where these criteria can be applied along with their limitations and derived alternatives. In order to delimit fungal species, authors need to take into account not only the phylogenetic and phenotypic coherence, but also the timing of events that lead to fungal speciation and subsequent diversifications. Variations in the rate of phenotypic diversifications and convergent fungal evolution make it difficult to establish a universal species recognition criterion. The best practice can only be defined in the context of each fungal group. In this review, we provide a set of guidelines, encouraging an integrative taxonomic approach for species delimitation that can be used to define fungal species boundaries in the future. The other papers in this special issue deal with fungal speciation in Ascomycota, Dothideomycetes, Basidiomycota, basal fungi, lichen-forming fungi, plant pathogenic fungi, and yeasts.
... Entyloma is a large genus in the Entylomataceae, with 188 recognized species on host plants belonging to 27 families (Vánky 2011;Denchev & al. 2013;Savchenko & al. 2014Savchenko & al. , 2016Rossman & al. 2016;Rooney-Latham & al. 2017;Savchenko & Carris 2017;Kruse & al. 2018;Kruse & Thines 2019;Richter & al. 2019). Entyloma species form sori in vegetative organs of plants from many different lineages of eudicots, mostly in leaves, rarely in stems, occasionally in roots, usually forming few to numerous spots, sometimes swellings or galls formed by hypertrophic growth of host tissue. ...
... Liro 1938, Lindeberg 1959, Vánky 2011) applied narrower species concepts, considering Entyloma species as infecting one or more hosts from the same host genus or occasionally a few closely related host genera. During the last two decades with the application of molecular methods, it became evident that members of Entyloma exhibit a far higher host specificity, parasitizing a single or only a few closely related host species (Begerow & al. 2002, Vánky & Lutz 2010, Savchenko & al. 2014, Kruse & al. 2018, Kruse & Thines 2019. It also became evident that much higher diversity than currently recognized in Entyloma should be expected (Kruse & al. 2018). ...
... During the last two decades with the application of molecular methods, it became evident that members of Entyloma exhibit a far higher host specificity, parasitizing a single or only a few closely related host species (Begerow & al. 2002, Vánky & Lutz 2010, Savchenko & al. 2014, Kruse & al. 2018, Kruse & Thines 2019. It also became evident that much higher diversity than currently recognized in Entyloma should be expected (Kruse & al. 2018). From Uzbekistan, 16 different Entyloma species are known, among which three are recorded on hosts in Ranunculaceae: the E. ranunculi-repentis complex on R. polyanthemos L., E. thalictri J. Schröt. ...
Article
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A new smut fungus, Entyloma eranthidis on Eranthis longistipitata from Uzbekistan, revealed by molecular, morphological, and ecological evidence, is described and illustrated. It differs from all other species of Entyloma by host specialization on Eranthis and by having longer (≤35(–38) μm) spores and thicker (≤10(–12) μm) spore walls. ITS rDNA sequence analysis indicates that the new species does not cluster with other species of Entyloma on Ranunculaceae.
... Iberian Peninsula are a subject of long-term studies (e.g., González Fragoso 1914, 1917, 1919, 1923, 1924a, b, 1926Unamuno 1928Unamuno , 1930aUnamuno , b, c, 1931Unamuno , 1934aUnamuno , b, 1942Ciferri 1933;Maire 1933Maire , 1943Losa España 1942, 1949, 1954Alcalde 1944;Cámara & Oliveira 1945;Cámara 1946;Guyot & al. 1955Guyot & al. , 1958Guyot & al. , 1960Guyot & al. , 1969Jørstad 1962;Durrieu 1966;Losa Quintana 1970;Llorens i Villagrasa 1985;Denchev 1995Denchev , 1997Almaraz & Durrieu 1997;Al-maraz 1998Al-maraz , 1999aAl-maraz , b, c, 2002Almaraz & Medina 1998;Almaraz & Telleria 1998;Vánky 2011;Kemler & al. , 2020Denchev & Denchev 2017;Kruse & al. 2018), their inventory is incomplete and no regional monographic study has been published yet. Based on the modern taxonomic arrangement of the smut fungi, all known records from the Iberian Peninsula and Balearic Islands can be referred to 164 species. ...
Article
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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.
... A remarkable case was recently resolved in the Entyloma ranunculi-repentis complex on multiple Ranunculus species. Here, one species, previously considered as one morphological species (Vánky, 2012), was split into nine species with each of them containing discrete but species-specific morphological characters as well as genetic and ecological differences (Kruse, Piątek et al., 2018). ...
Article
Chinese rhubarb (Rheum palmatum) is an important crop that has been used in traditional Chinese medicine for over 2000 years. The main cultivation areas are the Chinese provinces of Gansu, Qinhai, Tibet and Sichuan. Various fungal diseases threaten Chinese rhubarb, the most common of which is leaf smut. In literature, the causal agent of this disease is described as Thecaphora schwarzmaniana. We present morphological data and molecular phylogenetic data of two rDNA loci (ITS1‐5.8S‐ITS2 and 28S D1‐D2) which, however, justify the classification of the causal agent as a distinct species. Here we describe the new species Thecaphora dahuangis for the leaf smut pathogen of Chinese rhubarb.
... gestellte saprobiontisch lebende Sippen zu, die phylogenetisch innerhalb der Gattung Entyloma gruppiert sind und deshalb inzwischen als neue Entyloma-Arten beschrieben wurden (Boekhout et al. 2006, Baric et al. 2010, Richter et al. 2019. Nach derzeitigem Stand umfasst die Gattung Entyloma 195 Arten (Vánky 2012, Denchev et al. 2013, Savchenko et al. 2014, Rooney-Latham et al. 2017, Savchenko & Carris 2017, Kruse et al. 2018b, Richter et al. 2019, von denen fast die Hälfte nur auf Korbblütlern (Asteraceae) parasitieren (Vánky 2012 ...
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Ploch S, Kummer V (2020) Noteworthy records of phy-topathogenic micromycetes (14). Zeitschrift für Mykologie 86/2:319-385. Abstract: Some interesting records of plant parasitic microfungi of the Oomycota, Entorrhizo-mycota, Erysiphales, Pucciniomycotina and Ustilaginomycotina collected in Germany are reported. Entyloma podospermi on Scorzonera laciniata, Erysiphe pulchra on cultivated Cornus florida and Peronospora vincae on cultivated Vinca major are the first records of these fungi for Germany, Austria and Switzerland respectively. The first confirmed collections of Juncorrhiza mari-tima on Juncus ranarius in Germany are portrayed, including a discussion of possible older records. Information about the rare rust fungus Uromyces dianthi and its distribution in Ger-many including comments on its field identification is given. Furthermore, some corrections regarding Phyllactinia species occurring on different hosts in Germany, published in different herbaria or papers, are presented. This paper concludes with a list of some formerly unknown fungi and host species for different plant parasitic microfungi for the federal states of Germany. Zusammenfassung: Vorgestellt werden interessante Funde von Falschen Mehltauen (Oomy-cota), Wurzelgallpilzen (Entorrhizomycota), Echten Mehltaupilzen (Erysiphales), Rostpilzen (Pucciniomycotina) und Brandpilzen (Ustilaginomycotina) aus Deutschland. Entyloma podo-spermi auf Scorzonera laciniata, Erysiphe pulchra auf kultiviertem Cornus florida und Peronospora vincae auf kultiviertem Vinca major sind die ersten Nachweise für Deutschland, Österreich und die Schweiz. Die ersten gesicherten Nachweise von Juncorrhiza maritima auf Juncus ranarius in Deutschland werden präsentiert und es wird auf mögliche ältere Fundangaben in der Litera-tur hingewiesen. Es werden Informationen zur Häufigkeit und zum Befallsbild von dem in Deutschland seltenen Rostpilz Uromyces dianthi gegeben. Weiterhin werden einige Korrektu-ren zu bisher in Herbarien oder wissenschaftlicher Literatur publizierten, deutschlandbezo-genen Phyllactinia-Angaben präsentiert. Erfolgreiche Nachsuchen von bereits in dieser Reihe
... Ref.: Lit.: ; Kruse et al. (2018b) 1. Entyloma achilleae Magnus Wirt, Vork.: in Blättern von ▶ Achillea millefolium L. subsp. millefolium., stets mit Anamorphe Entylomella microstigma (Sacc.) ...
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Phytoparasitic fungi occurring in the federal state Saxony-Anhalt, Germany, are mapped in context with their host plants all over the state. Distribution maps and annotations for each fungus-host combination are given. This volume contains Phytomyxea p. p. (Parasitische Schleimpilze), Oomycota p. p. (Falsche Mehltaue), Pucciniomycotina (Rostpilze i. w. S.), Ustilaginomycotina (Brandpilze i. e. S.), other Basidiomycota, Blastocladiomycota p. p., Chytridiomycota p. p. (Flagellatenpilze), totally 678 fungal species and 1058 host species. Annotations are given in German language.
... Integrative taxonomy attempts to combine as much evidence as possible from genealogical, biological, phenotypic and other approaches to delimit species (Aime 2004;Yang and Rannala 2010;Padial et al. 2010;Udayanga et al. 2014;Haelewaters et al. 2018;Kruse et al. 2018a). The different approaches are thereby not competitive but components of a holistic strategy. ...
Article
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True fungi (Fungi) and fungus-like organisms (e.g. Mycetozoa, Oomycota) constitute the second largest group of organisms based on global richness estimates, with around 3 million predicted species. Compared to plants and animals, fungi have simple body plans with often morphologically and ecologically obscure structures. This poses challenges for accurate and precise identifications. Here we provide a conceptual framework for the identification of fungi, encouraging the approach of integrative (polyphasic) taxonomy for species delimitation, i.e. the combination of genealogy (phylogeny), phenotype (including autecology), and reproductive biology (when feasible). This allows objective evaluation of diagnostic characters, either phenotypic or molecular or both. Verification of identifications is crucial but often neglected. Because of clade-specific evolutionary histories, there is currently no single tool for the identification of fungi, although DNA barcoding using the internal transcribed spacer (ITS) remains a first diagnosis, particularly in metabarcoding studies. Secondary DNA barcodes are increasingly implemented for groups where ITS does not provide sufficient precision. Issues of pairwise sequence similarity-based identifications and OTU clustering are discussed, and multiple sequence alignment-based phylogenetic approaches with subsequent verification are recommended as more accurate alternatives. In metabarcoding approaches, the trade-off between speed and accuracy and precision of molecular identifications must be carefully considered. Intragenomic variation of the ITS and other barcoding markers should be properly documented, as phylotype diversity is not necessarily a proxy of species richness. Important strategies to improve molecular identification of fungi are: (1) broadly document intraspecific and intragenomic variation of barcoding markers; (2) substantially expand sequence repositories, focusing on undersampled clades and missing taxa; (3) improve curation of sequence labels in primary repositories and substantially increase the number of sequences based on verified material; (4) link sequence data to digital information of voucher specimens including imagery. In parallel, technological improvements to genome sequencing offer promising alternatives to DNA barcoding in the future. Despite the prevalence of DNA-based fungal taxonomy, phenotype-based approaches remain an important strategy to catalog the global diversity of fungi and establish initial species hypotheses.