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-Phylogenetic analysis of combined data of the 5 0-end of the 28S rRNA gene (including domains D1 and D2) and ITS region for 34 sequences from Erysiphe gracilis s. lat. and three outgroup sequences. This tree is a single parsimonious tree with 143 steps, which were found using a heuristic search. Horizontal branch lengths are proportional to the number of substitutions that were inferred to have occurred along a particular branch of the tree. BS (!70%) values by the maximum parsimony (MP) and maximum likelihood (ML) methods were shown on the respective branches.
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Erysiphe gracilis is a powdery mildew species that occurs on evergreen oak species belonging to Quercus subgen. Cyclobalanopsis in East Asia (China and Japan). In a previous report, we found that E. gracilis var. gracilis is divided into four genotypes each of them forming a separate clade with strong bootstrap support. In this study, we further in...
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... Three se- quences from E. japonica (S. Ito & Hara) C.T. Wei were used as outgroup. The alignment data matrix consisted of 37 sequences and 1369 characters, of which 118 (8.6%) characters were variable and 97 (7.1%) characters were informative for parsimony analysis. A single most parsimonious tree with 143 steps was constructed by the MP analysis (Fig. 2). BS supports by ML analysis were shown on the MP tree. The IGS data set consisted of 62 sequences and 391 characters, of which 73 (18.7%) characters were variable and 60 (15.3%) characters were informative for parsimony analysis. Two sequences from E. japonica were used as outgroup. A total of 47,131 equally parsimonious trees with 94 ...
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... Circinate, branched, and clavate appendages were defined as complex, whereas mycelioid appendages were defined as simple. The appendages of Erysiphe uncinuloides and E. longiappendiculata are mentioned as "uncinate-circinate to helicoid at the apex" in the original description (Siahaan et al. 2018). However, photographs of their chasmothecia show the appendages as long undulating mycelia, rather than setiform, and they are estimated to be functionally almost identical to mycelioid. ...
... In certain species, appendages exhibit intermediate or ambiguous morphologies, which makes it difficult to deduce their functions during overwintering. For example, E. astragali, E. bremeri, E. hyperici, E. intermedia, E. longiappendiculata, E. russellii, E. trifoliorum, and E. uncinuloides exhibit intermediate states between complex and simple appendages (Braun and Cook 2012;Siahaan et al. 2018). These species may represent transitional stages in which appendage simplifications on herb/evergreen hosts are in progress. ...
Evolutionary relationships between the morphological and ecological traits of fungi are poorly understood. The appendages of chasmothecia, which are sexual reproductive organs of Erysiphaceae, are considered to play a crucial role in the overwintering strategies of these fungi on host plants. Previous studies suggested that both the host type and appendage morphology evolved at the same nodes and transitioned from complex appendages on deciduous hosts to simple appendages on herb/evergreen hosts. However, the evolutionary dependence between host type and appendage morphology remains unproven owing to the limited species data used in analyses. To elucidate the evolutionary relationship between host type and appendage morphology, we used phylogenetic comparative methods (PCMs) to investigate the state transition, ancestral state, evolutionary dependence, and contingent evolution within Erysipheae, the largest and most diverse tribe in Erysiphaceae. Our PCMs, based on a comprehensive data set of Erysipheae, revealed that the most ancestral states were deciduous host types and complex appendages. From these ancestral states, convergent evolution toward the herb/evergreen host types and simple appendages occurred multiple times at the same nodes. For the first time in Erysiphaceae, we detected an evolutionary dependence between host type and appendage morphology. This is one of the few examples in which evolutionary dependence between host phenology and morphological traits in plant-parasitic fungi was demonstrated using PCMs. Appendage simplification on herb/evergreen hosts and complications on deciduous hosts can be reasonably explained by the functional advantages of each appendage type in different overwintering strategies. These expected appendage functions can explain approximately 90% of host type and appendage morphology combinations observed in the analyzed taxa. However, our results also highlighted the occurrence of evolutionary shifts that deviate from the expected advantages of each appendage morphology. These seemingly irrational shifts might be interpretable from the flexibility of overwintering strategies and quantification of appendage functions.
... Quercus species almost worldwide are infected by powdery mildew species in the genus Erysiphe. Erysiphe species occurring on oaks have been described morphologically and, in part, based on phylogenetic analyses (Takamatsu et al. 2007;Braun and Cook 2012;Siahaan et al. 2018). Erysiphe alphitoides, E. quercicola, and E. hypophylla are the most common and widespread species reported on oaks. ...
... In the interim, we recommend LC271391.1 as a reference sequence. This sequence stems from the comprehensive phylogenetic-taxonomic revision of the Erysiphe gracilis complex published by Siahaan et al. (2018). ...
... Holotype: TNS-F-198227 (Japan, on Quercus acuta). Epitype (designated by Siahaan et al. 2018 Darsaraei et al. (2021). Notes: Yamaguchi et al. (2021) disentangled Erysiphe salmonii s. lat. ...
This is the fourth contribution within an ongoing series dedicated to the phylogeny and taxonomy of powdery mildews. This particular installment undertakes a comprehensive evaluation of a group previously referred to as the "Uncinula lineage" within Erysiphe. The genus Erysiphe is too large to be assessed in a single paper; thus, the treatment of Erysiphe is split into three parts, according to phylogenetic lineages. The first paper, presented here, discusses the most basal lineage of Erysiphe and its relationship to allied basal genera within tribe Erysipheae (i.e., Brasiliomyces and Salmonomyces). ITS+28S analyses are insufficient to resolve the basal assemblage of taxa within the Erysipheae. Therefore, phylogenetic multilocus examinations have been carried out to better understand the evolution of these taxa. The results of our analyses favor maintaining Brasiliomyces, Bulbomicroidium, and Salmonomyces as separate genera, at least for the interim, until further phylogenetic multilocus data are available for additional basal taxa within the Erysipheae. The current analyses also confirmed previous results that showed that the "Uncinula lineage" is not exclusively composed of Erysiphe species of sect. Uncinula but also includes some species that morphologically align with sect. Erysiphe, as well as species that had previously been assigned to Californiomyces and Typhulochaeta. Numerous sequences of Erysiphe species from the "Uncinula lineage" have been included in the present phylogenetic analyses and were confirmed by their position in well-supported species clades. Several species have been sequenced for the first time, including Erysiphe clintonii, E. couchii, E. geniculata, E. macrospora, and E. parvula. Ex-type sequences are provided for 16 taxa including E. nothofagi, E. trinae, and E. variabilis. Epitypes are designated and ex-epitype sequences are added for 18 taxa including Erysiphe carpophila, E. densa, and U. geniculata var. carpinicola. The new species Erysiphe canariensis is described, and the new names E. hosagoudarii and E. pseudoprunastri and the new combination E. ampelopsidis are introduced. ARTICLE HISTORY
... Powdery mildews infecting Quercus species are common and widespread across the globe. There are an estimated 23 Erysiphe species described morphologically and, in part through phylogenetic analyses (Braun & Cook, 2012;Siahaan et al., 2018;Takamatsu et al., 2007). In Europe, the most comprehensively studied, and common species, are Erysiphe alphitoides, E. quercicola and E. hypophylla. ...
Powdery mildew, caused by Erysiphe spp., on oak has been shown to have serious ecological consequences on Quercus hosts. Erysiphe alphitoides and E. quercicola are two of the most heavily studied and common powdery mildews known to occur on Quercus species. In recent years, these species have been noted throughout the world on a range of hosts within and outside the Quercus genus. Reports that E. alphitoides was absent in European herbaria before 1921 and the discovery of the holotype of E. alphitoides from 1911 in an American herbarium (FH) led to the current study in which we genetically analysed six specimens of E. alphitoides s. lat including, most importantly, the holotype of E. alphitoides from France collected in 1911. The results of our analyses revealed that: (1) The sequence of the E. alphitoides holotype falls within the E. quercicola clade, confirming that E. alphitoides did not spread to Europe until ~1921. (2) E. alphitoides var. chenii forms a monophyletic clade with E. epigena and should be reduced to synonymy with that species and (3) through sequence analyses E. alphitoides and E. quercicola are confirmed to have spread to North America. The sequencing results of the E. alphitoides holotype have severe nomenclatural‐taxonomic consequences. A proposal was submitted simultaneously with the present manuscript to conserve the name E. alphitoides so that the traditional usage of the names E. alphitoides and E. quercicola could be maintained. The sequences obtained for the current study provide new insight into the taxonomy and spread of these ecologically significant, globally distributed species. The present study highlights the importance of sequencing specimens from type material, above all when morphological similar species are involved.
... The current results strongly suggest that there is a strict host specificity in these Fraxinus powdery mildew species. Recent molecular phylogenetic analyses revealed that powdery mildews include many cryptic species distinguished by their host plants (e.g., Braun et al., 2006;Scholler, Schmidt, Siahaan, Takamatsu, & Braun, 2016;Meeboon, Siahaan, Fujioka, & Takamatsu, 2017;Abasova et al., 2018;Siahaan, Sakamoto, Shinoda, & Takamatsu, 2018;Meeboon, Takamatsu, & Braun, 2020). This is especially conspicuous in the Uncinula-lineage of Erysiphe to which the Fraxinus powdery mildews belong. ...
... For example, the E. carpinicola species complex from Carpinus species was divided into five species differentiated by their host species (Braun et al., 2006). Similarly, the E. gracilis species complex was divided into six species in line with their host species (Siahaan et al., 2018). Shin (2000) reported E. fraxinicola (as U. fraxini) from Korea, but Heluta et al. (2017) suggested that this might be a mis-identification of E. salmonii based on the drawing in Shin (2000). ...
The genus Fraxinus (Oleaceae), known as ash trees, currently comprises 43 recognized species that are distributed in temperate and subtropical regions of the northern hemisphere. Two Erysiphe (sect. Uncinula) species have been known on Fraxinus spp. so far. In this study, Fraxinus powdery mildews from different areas of the world were collected to make molecular and morphological analyses. These specimens are divided into three distinct molecular phylogenetic groups, which are distinguishable by their morphology and/or host preference. The powdery mildew occurring on F. apertisquamifera and F. lanuginosa is described as a new species, E. fraxinea. Epitypes are designated for E. fraxinicola and E. salmonii. Applying previous traditional species delimitations, various hosts were shared by E. fraxinicola as well as E. salmonii, but the current analyses strongly suggest strict host specificity among these three powdery mildew species. Evolutionary timing calculated by molecular clock analysis suggests co-evolution of powdery mildews with their Fraxinus hosts.
... revealed a high degree of diversity and cryptic speciation, and most species examined were shown to be phylogenetically and taxonomically heterogeneous. These results are similar to those of previous studies that analyzed complex powdery mildew species, such as those that evaluated E. carpinicola (Braun et al. 2006) and E. gracilis (Siahaan et al. 2018). Erysiphe corylacearum, a name based on Microsphaera hommae, was originally introduced for powdery mildew on Corylus spp. ...
Erysiphe species (powdery mildews) on Corylus and Ostrya hosts (Betulaceae subfam. Coryloideae) in Asia and North America are widespread pathogens on these economically and ecologically valuable nut crops. An improved understanding of their phylogeny and taxonomy is of ecological and applied importance. Phylogenetic analyses and morphological reexaminations conducted in this study revealed a higher degree of diversity and cryptic speciation than reflected in earlier species concepts. North American collections on C. cornuta, which were previously assigned to E. corylacearum, proved to constitute a species of its own and are herein introduced as E. cornutae, sp. nov. Two additional North American species, E. coryli-americanae, sp. nov. and E. ostryae, sp. nov., have been detected on C. americana and O. virginiana and are described. They are morphologically similar to E. cornutae, but genetically distinct. Based on phylogenetic analyses, E. corylacearum is an Asian species confined to various Asian Corylus species. Sequence data retrieved from Japanese type material of E. corylicola revealed that this species clusters with sequences from E. elevata on Catalpa species, distant from all other Erysiphe species on Corylus. Morphologically similar, yet distinct, specimens on C. sieboldiana, which were previously assigned to E. corylicola, form a distinct, distant clade. The species involved is described herein as E. pseudocorylacearum, sp. nov. Additionally, an unusual infection of C. sieboldiana in Japan by E. syringae has been shown by means of sequence data. The phylogeny and taxonomy of Erysiphe species belonging to the Corylioideae are discussed in detail, and a key to the species concerned is provided.
... The sequences of E. bulbouncinula are indicated in boldface E. kissiana (Takamatsu, Siahaan, & Shinoda, 2015b) and 12 sequences for E. gracilis s.lat. (Siahaan, Sakamoto, Shinoda, & Takamatsu, 2018a). ...
... Mori et al., (2000) first reported that E. gracilis, although morphologically Erysiphelike, is phylogenetically more closely related to Erysiphe species with Uncinula-like morphology, assigned to E. sect. Uncinula (Braun & Cook, 2012 (Siahaan et al., 2018a). ...
... At present, DNA sequence data are available for approximately only one third of the total number of species in the Erysiphaceae, partly due to their unculturable nature on artificial media (Siahaan, Sakamoto, Shinoda, & Takamatsu, 2018b). They have initiated drastic changes in the taxonomy of powdery mildews not only at higher taxonomic levels (Johnston et al., 2019), but also at species level (Meeboon, Siahaan, Fujioka, & Takamatsu, 2017;Qiu et al., 2020;Siahaan et al., 2018a). Further researches are needed for better understanding of phylogenetic relationships and the taxonomy of this important fungal group. ...
Erysiphe bulbouncinula (≡ Bulbouncinula bulbosa ) on Koelreuteria paniculata is a powdery mildew native and endemic in China. Fresh samples of this species were collected on K. paniculata in Sichuan Province, China, in 2018. This is the fourth confirmed identification of this species. The original description in 1932 was based on one specimen from Hangzhou and then was described in Nanjing in 1990 and in Taiwan in 1993. The morphology of the new collections of this species is described and illustrated in detail, and agrees well with the original description of the type specimen except for the number of asci (8–16 asci per chasmothecia versus 7–9 asci in the type specimen). To explore the phylogenetic position of this poorly known species, three separate phylogenetic analyses, using partial 28S rDNA and the combination of internal transcribed spacer (ITS) and 28S rDNA sequences, were conducted. The results clearly showed that E. bulbouncinula form a monophyletic clade with strong bootstrap (BS) support (MP = 100%) constituting a sister clade to E. gracilis s. lat., form a subclade with Erysiphe sect. Californiomyces and then form a big group with sect. Typhulochaeta and sect. Uncinula . Phylogenetic results confirm that E. bulbouncinula is more closely allied to Erysiphe than to Phyllactinia . Thus, this species can be maintained in Erysiphe in its current circumscription, confirming Bulbouncinula as a synonym of Erysiphe .
... The classification system of Erysiphaceae has been developed based on the morphological features of sexual and asexual morphs (Braun, 1987). In recent years, taxonomic treatments at the genus (Braun & Cook, 2012;Braun & Takamatsu, 2000;Liberato, Barreto, Niinomi, & Takamatsu, 2006; (Marmolejo, Siahaan, Takamatsu, & Braun, 2018); Takamatsu, Braun, & Limkaisang, 2005;Takamatsu et al., 2005;To-anun et al., 2005) and species levels (Braun et al., 2006, (Braun et al., 2019; Siahaan, Sakamoto, Shinoda, & Takamatsu, 2018;Takamatsu et al., 2007) have been revised to reflect their molecular phylogenetic relationships. The species of Erysiphaceae are thought to have evolved while having close interactions with their hosts, and the phylogeography and diversification process of this fungal group have been discussed as compared to the evolutionary history of their host plants (Matsuda & Takamatsu, 2003;Niinomi, Takamatsu, & Havrylenko, 2008;Takamatsu, 2013;Takamatsu, Matsuda, Niinomi, & Havrylenko, 2006). ...
The classification system and evolutionary history of Erysiphaceae have been studied based on the results of molecular phylogenetic analyses. However, the sequence data used for these phylogenetic estimations have been limited to the nrDNA of ca., 50 taxa, and the relationships among higher taxonomic groups are not well understood. To provide a phylogenetic overview of Erysiphaceae, we performed phylogenetic estimations based on nrDNA and MCM7 sequences obtained from ca., 270 taxa. The phylogenetic tree showed a similar topology to the trees obtained in previous studies, although the branching order between Golovinomyceteae and Phyllactinieae was different and Phyllactinieae was not monophyletic. Phyllactinieae and Erysipheae were estimated to diversify after the divergence of Golovinomyceteae, suggesting an evolutionary trend in which non-catenate conidia + endoparasitic or non-catenate conidia + ectoparasitic lineages were derived from catenate conidia + ectoparasitic lineages. Phyllactinieae was divided into a clade of Phyllactinia + Leveillula and other clade(s) consisting of Pleochaeta and Queirozia. The phylogenetic hypothesis of Erysiphaceae was updated based on the largest dataset to date, but the higher-level phylogenetic relationships remain unclear. For a more robust phylogenetic hypothesis of Erysiphaceae, further sequence data, including protein coding regions, should be added to the dataset of nrDNA sequences.
... Molecular phylogenetic analyses often revealed the presence of cryptic species within traditional morphology-based species (complexes), and subsequent reevaluations of morphological traits frequently aligned with the molecular results. For example, Erysiphe carpinicola (Hara) U. Braun & S. Takam., previously considered a single species, was divided into four species(Braun et al. 2006;Meeboon and Takamatsu 2013), Erysiphe gracilis R.Y. Zheng & G. Q. Chen into six species(Siahaan et al. 2018), Parauncinula septata (E.S. Salmon) S. Takam. & U. Braun into three species ...
Podosphaera tridactyla (s. lat.) is a powdery mildew species occurring on a wide range of Prunus spp. almost worldwide. We have investigated the phylogeny of the Po. tridactyla complex, with special emphasis on potential aspects of cryptic speciation. The results suggested that Po. tridactyla represents a species complex consisting of at least 12 different species. Based on detailed morphological examinations and molecular sequence analyses, we propose dividing Po. tridactyla s. lat. into 10 species, encompassing 7 new species (Po. ampla, Po. pruni-avium, Po. pruni-cerasoidis, Po. prunigena, Po. pruni-lusitanicae, Po. prunina, and Po. pruni-japonicae) and 3 known species (Po. longiseta, Po. salatai, and Po. tridactyla s. str.). Oidium passerinii on Pr. laurocerasus is confirmed as a synonym of Po. tridactyla s. str. Epitypes are designated for Po. tridactyla and Oidium passerinii.
... Brasiliomyces malvastri and B. malachrae commonly infect host plants of the family Malvaceae, but the former species infects hosts of genus Malvastrum, and the latter occurs on Mala- chra species. Recent phylogenetic analyses often revealed that morphology-based species are actually species complex consisting of cryptic species confined to hosts of plant genera or species (Braun et al., 2006;Meeboon, Siahaan, Fujioka, & Takamatsu, 2017;Siahaan, Sakamoto, Shinoda, & Takamatsu, 2018; Takamatsu et al., 2007). Thus, the synonymy of these two species should be re- investigated by molecular analyses. ...
Brasiliomyces, comprising only four species, is a unique powdery mildew genus characterized by having small, semitransparent chasmothecia with a thin, single peridium layer. Brasiliomyces malachrae, a type species, is distributed in Central and South America and the morphology of this species, especially its asexual morph, is poorly known. This study was performed to describe the sexual and asexual morphs of B. malachrae on Malvastrum coromandelianum in detail, and to clarify the phylogenetic placement of this fungus. The asexual morph of this species is characterized by having hyaline ectophytic mycelium with lobed hyphal appressoria and catenescent conidia. Molecular phylogenetic analyses based on 18S, 5.8S, and 28S rRNA gene sequences revealed that B. malachrae is a sister to the genus Erysiphe and situated at the boundary between tribes Golovinomyceteae and Erysipheae, which supports its unique phylogenetic position. The evolutionary relationships among the tribes Golovinomyceteae, Erysipheae, and Phyllactinieae are discussed on the basis of the present phylogenetic analyses.
