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Morphophylogenetic analyses revealed that Podosphaera tridactyla constitutes a species complex
Mycologia
Abstract
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.
... The taxonomic list of powdery mildew species, which included the collections of the authors and literature data, was compiled using the fundamental taxonomic manual of Erysiphaceae (Braun, 2012) with additions for the several taxonomic revisions of the species complexes: Blumeria graminis (Liu et al., 2021;Heluta, 2022), Erysiphe adunca (Darsaraei et al., 2021), E. lonicerae (Bradshaw et al., 2021), Golovinomyces biocellatus (Scholler et al., 2016), G. cynoglossi (Braun et al., 2018), G. orontii , Podosphaera tridactyla (Meeboon et al., 2020;Smith et al., 2021), P. cerasi and P. prunicola (Moparthi et al., 2019); Leveillula taurica (Khodaparast et al., 2001;2012). All names of host plant species are given by the open database "Catalogue of life" (Catalogue of Life Database, 2022). ...
... Over the past 10 years, the list of powdery mildew fungi has increased by another 20 species, which can be considered as new for the Ile-Alatau due to taxonomic revisions of the species complexes: Blumeria graminis (Liu et al., 2021;Heluta, 2022), Erysiphe adunca (Darsaraei et al., 2021), E. lonicerae (Bradshaw et al., 2021), Golovinomyces biocellatus (Scholler et al., 2016), G. cynoglossi (Braun et al., 2018), G. orontii , Podosphaera tridactyla (Meeboon et al., 2020;Smith et al., 2021), P. cerasi and P. prunicola (Moparthi et al., 2019); Leveillula taurica (Khodaparast et al., 2001;2012). ...
... Note: The species is rare in the Ile-Alatau. Note: Based on morphological studies and analysis of molecular sequences, some authors (Meeboon et al., 2020) propose to divide the complex species P. tridactyla s. lat. for 10 species, including 7 new species (P. ...
... In other cases, phylogenetic revisions of traditional powdery mildew species revealed that these 'species' are complexes, including undescribed cryptic species, such as in the case of Podosphaera tridactyla s. lat. (Meeboon et al. 2020). Such recently published taxonomic revisions with introduced new species names require re-examinations of the species complexes concerned in the respective countries. ...
... As this species is currently spreading in Europe, it is likely that it may already have invaded further European countries. (Cunnington et al. 2005), Meeboon et al. (2020) carried out a comprehensive phylogenetic-taxonomic revision of this complex and revealed that Po. tridactyla constitutes a species complex. Numerous new species were described, including Po. ampla on Prunus cerasifera, P. domestica, and P. spinosa, distributed in the Caucasus region (Armenia, Georgia) and Europe (Belarus, Bulgaria, Estonia, Germany, Greece, Lithuania, Montenegro, Slovakia, Sweden, Switzerland, Ukraine). ...
... In autumn 2022, Po. ampla was recorded several times in North Tyrol, Austria (occurrences on three different specimens of Prunus cerasifera) and Lombardia (Brescia, Italy, one sample on Prunus spinosa and one on Prunus cerasifera), which are the first documented occurrences in Austria and Italy. The morphology of the examined specimens agrees perfectly with the description given in Meeboon et al. (2020). This species is characterised by having few (up to 6) stiff appendages with variable length, formed in parallel bundles (fascicles) on the uppermost part of the chasmothecium and a typical branching pattern of the branched apex, with up to six times dichotomous branched apices, which is a combination unknown from any other Podosphaera species infecting Prunus spp. ...
... de Bary and P. pruni-avium Meeboon, S. Takam. & U. Braun, respectively, have been reported to infect sweet cherries; however, these fungi have not been reported in the U.S. (Blumer 1960;Choi et al. 2015;Meeboon et al. 2020). ...
... form a separate well-supported species clade in phylogenic analyses (Takamatsu et al. 2010). Meeboon et al. (2020) conducted morphological reexaminations on P. tridactyla s. lat. and it was concluded that P. tridactyla primarily infects Prunus subgen. ...
... In contrast, P. tridactyla (s. str.) chasmothecia from Prunus padus, the type, host of this species, measured (60-)70-105(-120) μm in diameter and had 2-6 appendages that were 3-6 times dichotomously branched (Meeboon et al. 2020). P. pruniavium chasmothecia were 80-110 μm in diameter with 2-5 stiff or slightly curved appendages that were arranged in the upper half (Meeboon et al. 2020). ...
... de Bary and P. pruni-avium Meeboon, S. Takam. & U. Braun, respectively, have been reported to infect sweet cherries; however, these fungi have not been reported in the U.S. (Blumer 1960;Choi et al. 2015;Meeboon et al. 2020). ...
... form a separate well-supported species clade in phylogenic analyses (Takamatsu et al. 2010). Meeboon et al. (2020) conducted morphological reexaminations on P. tridactyla s. lat. and it was concluded that P. tridactyla primarily infects Prunus subgen. ...
... In contrast, P. tridactyla (s. str.) chasmothecia from Prunus padus, the type, host of this species, measured (60-)70-105(-120) μm in diameter and had 2-6 appendages that were 3-6 times dichotomously branched (Meeboon et al. 2020). P. pruniavium chasmothecia were 80-110 μm in diameter with 2-5 stiff or slightly curved appendages that were arranged in the upper half (Meeboon et al. 2020). ...
Sweet cherry is a high-value fresh market crop whose commercial value is directly related to fruit quality. Morphological studies and phylogenetic analyses have shown that a distinct powdery mildew species, Podosphaera cerasi, infects sweet cherry in orchards, nurseries, and greenhouses for stock production. The fungal species P. clandestina, which was thought to cause powdery mildew infection in sweet cherry, is now confined to the powdery mildew that infects Crataegus spp. The intricate nomenclatural-taxonomic history of sweet cherry powdery mildew is briefly outlined. This review has been written to serve production managers as well as applied and basic research regarding control of this pathogen. The economic impact of this powdery mildew, environmental aspects contributing to its epidemiology, management options including both conventional chemical and biopesticide products, as well as mycoviruses associated with P. cerasi are discussed. Additionally, an overview of host–pathogen resistance based on cultivar, fungicide resistance, the role of chasmothecia in epidemic initiation, and susceptible stages of fruit to infection are discussed.
... tridactyla complex ( Figure 2). Both data sets included sequences generated from this study and sequences of Podosphaera obtained from GenBank that were selected from published studies [11,16,17] and NCBI searches of Podosphaera spp. ITS and ITS+28S sequences. ...
... ITS and ITS+28S sequences. Phylogenetic analyses of these two data sets used Cystotheca lanestris (GenBank accessions AF011289 and AF011288) as the outgroup taxon, in line with Meeboon et al. [11]. The phylogenetic analysis of host matK sequences included sequences from Prunus species subgenera Cerasus, Padus and Prunus obtained from Chin et al. [4] and the outgroup was Oemleria cerasiformis (matK AF288110). ...
... ampla clade, including the Australian sequences, were as follows: ITS (BS 80, BI 1) and 28S (BS 100, BI 1) ( Figures S2 and S3). In the ITS tree, two Australian sequences (AY833656 and AF154321) published by Cunnington et al. [10] and labelled as Podosphaera sp. by Meeboon et al. [11] fell into the Po. ampla clade. ...
The Podosphaera tridactyla species complex is highly variable morphologically and causes powdery mildew on a wide range of Prunus species, including stone fruit. A taxonomic revision of the Po. tridactyla species complex in 2020 identified 12 species, seven of which were newly characterised. In order to clarify which species of this complex are present in Australia, next generation sequencing was used to isolate the fungal ITS+28S and host matK chloroplast gene regions from 56 powdery mildew specimens of stone fruit and ornamental Prunus species accessioned as Po. tridactyla or Oidium sp. in Australian reference collections. The specimens were collected in Australia, Switzerland, Italy and Korea and were collected from 1953 to 2018. Host species were confirmed using matK phylogenetic analysis, which identified that four had been misidentified as Prunus but were actually Malusprunifolia. Podosphaera species were identified using ITS+28S phylogenetic analysis, recognising three Podosphaera species on stone fruit and related ornamental Prunus hosts in Australia. These were Po.pannosa, the rose powdery mildew, and two species in the Po. tridactyla species complex: Po. ampla, which was the predominant species, and a previously unidentified species from peach, which we describe here as Po. cunningtonii.
... These discoveries stem from intensified global explorations as well as numerous recently published phylogenetic-taxonomic revisions of species complexes. In most cases, such revisions have resulted in the identification of previously unknown cryptic species (e.g., Bradshaw et al. 2020Bradshaw et al. , 2021cDarsaraei et al. 2021;Liu et al. 2021;Meeboon et al. 2020). Additionally, the use of sequence analyses has made it easier to identify anamorphic collections by enabling clarification of their generic affinity and taxonomic status. ...
... The powdery mildew monograph by Braun and Cook (2012) serves as the general basis for the accepted names listed. This monograph was supplemented by several recent phylogenetic-taxonomic studies focusing on specific groups within the Erysiphaceae (e.g., Bradshaw et al. 2020Bradshaw et al. , 2021aBradshaw et al. , b, c, 2022aDarsaraei et al. 2021;Liu et al. 2021;Meeboon et al. 2020). The phylogenetic revisions of powdery mildews are ongoing, and comprehensive treatments for phylogenetically heterogeneous species such as Podosphaera aphanis and P. erigerontis-canadensis, as recognized in Braun and Cook (2012), are currently unavailable. ...
Powdery mildews are highly destructive fungal plant pathogens that have a significant economic impact on both agricultural and ecological systems worldwide. The intricate relationship between powdery mildews and their host plants has led to co-speciation. In this study, we conducted an extensive evaluation of powdery mildew hosts to provide an updated understanding of the host ranges and distributions of these fungi. The "United States National Fungus Collections Fungus-Host Dataset" is the primary source of information for our analyses. The analysis of the dataset demonstrated the worldwide prevalence of powdery mildews; the data contained over 72,000 reports of powdery mildews, representing ~8.7% of all host-fungal records. We have updated the taxonomy and nomenclature of powdery mildews. In total, powdery mildews infect ~10,125 host taxa belonging to 205 families of flowering plants accounting for 1,970 genera in 200 countries across six continents. Furthermore, we estimate that powdery mildews infect approximately 2.9% of described angiosperm species. Our study underscores the need for regular updates on powdery mildew host information due to the continuously evolving taxonomy and the discovery of new host taxa; since 1986 we estimate an additional 1,866 host taxa, 353 genera, and 36 families have been reported. Additionally, the identification of powdery mildew hosts provides valuable insights into the co-evolutionary dynamics between the fungi and their plant hosts. Overall, this updated list provides valuable insights into the taxonomy and geographic distribution of powdery mildew species, building upon the previous work of Amano in 1986. Discerning the geographic spread and host range of economically significant plant pathogens is vital for biosecurity measures and identifying the origins and expansion of potentially harmful pathogens.
... Magnicellulatae (Ito and Takamatsu 2010), North American Podosphaera species on Prunus spp. (Moparthi et al. 2019), phylogenetic revision of the P. tridactyla complex (Meeboon et al. 2020). ...
... This hypothesis was supported by the ITS phylogenetic tree, on which the samples of Po. tridactyla were grouped into 2-4 clades, representing several separate phylogenetic species . A comprehensive phylogenetic-taxonomic revision of Po. tridactyla s. lat., carried out by Meeboon et al. (2020), confirmed this assumption and led to a splitting of Po. tridactyla s. lat. into numerous species. ...
Powdery mildew diseases on crops and ornamental plants are common and cause significant economic losses, by reducing the yield and quality of crops and downgrading the value of ornamentals. In surveys published annually in the Canadian Plant Disease Survey, powdery mildews were reported frequently. In recent years, the taxonomy of powdery mildew fungi has changed tremendously as the result of the increased application of molecular phylogenetic analyses. Several generic concepts have been re-adjusted. Many traditionally considered species turned out to be a complex of several phylogenetic species. In order to facilitate effective communication in the scientific community, it is important to apply the current correct names for field identifications. In this document, we summarize the powdery mildew fungi reported in the Canadian Plant Disease Survey from 2000 to 2019, address the taxonomic and nomenclatural issues associated with the listed species, and tentatively suggest correct names based on host plants wherever possible. Nevertheless, a reliable identification of the powdery mildew must depend on DNA sequence analyses and morphological examinations.
... There are three powdery mildew species in the genus Podosphaera that infect cherry: P. cerasi, P. prunicola, and P. pruniavium (a newly named species within the P. tridactyla complex) (Meeboon et al., 2020;Moparthi et al., 2019). In addition, P. clandestina was formerly applied in a much wider sense that included powdery mildews with cherry as a host, such as by Braun (1987). ...
... obtained from GenBank that were selected from published studies (Meeboon et al., 2020;Moparthi et al., 2019;Smith et al., 2020) and ...
A strain of Podosphaera clandestina has been highlighted as a priority pest threat to the Australian cherry industry. Australia currently has no records of powdery mildew on cherry (Prunus avium). P. clandestina is reported to cause disease on a range of Rosaceae genera including Crataegus and Prunus and in Australia, P. clandestina has only been recorded on Crataegus. A recent species revision identified P. cerasi on P. avium as a separate species from P. clandestina. Therefore, a revision of which powdery mildew species is present in Australia on Crataegus is required to inform Australian plant biosecurity. Reference collection specimens from the Victorian Plant Pathology Herbarium (VPRI) recorded as Podosphaera spp. collected between 1889 to 2008 on cherry and three other host plant genera from Australia and overseas were sampled for DNA extraction and next‐generation sequencing (NGS). Sequence data from preserved specimens were successfully mapped to internal transcribed spacer (ITS) sequences of P. clandestina in the strict sense, P. cerasi, and P. prunicola, and chloroplast matK sequences were used to identify plant hosts. Australian specimens on Crataegus hosts were P. clandestina in the strict sense and specimens on Prunus from the USA were identified as P. cerasi and P. prunicola. The outcome of this study confirmed the powdery mildew on Australian Crataegus specimens to be P. clandestina and none of the cherry powdery mildews (P. cerasi, P. pruni‐avium, or P. prunicola) are present on Australian specimens in the VPRI collection, which suggests they are not present in Australia.
... Anm.: In Thiel et al. (2023) ist dieser Pilz nur von Prunus avium angegeben. Dies basiert auf den Mitteilungen zu dieser Pilz-Wirt-Kombination für Deutschland inMeeboon et al. (2020), die als weiteren Wirt nur noch Prunus laurocerasus aufführen. Die Zuordnung der Podosphaera-Befälle auf P. ...
Abstract: Some interesting records of plant parasitic microfungi of the Erysiphaceae, Peronosporomycetes, Pucciniomycotina, Ustilaginomycotina, Chytridiomycota and other Ascomycota from Germany, Austria and Switzerland are reported.
Podosphaera pruni-lusitanicae on cultivated Prunus lusitanica is documented in Germany for the first time. Podosphaera cerasi on Prunus cerasus was confirmed to occur in Germany based on molecular genetic data. Uromyces proëminens on Euphorbia prostrata is new for Germany. Investigations have shown that the species has been present in Germany since 2011. A find from Switzerland on the host mentioned above is a new fungus-host combination for the fungus, which is still rare in Switzerland. Uromyces sparsus was rediscovered after 50 years in Germany on Spergularia marina. Literature references to Spergularia rubra as host for this fungus are incorrect. Thecaphora lathyri on Lathyrus pratensis, which has not been reported in Germany since 1963, was found again in 2021 and has since been detected in three other federal states. Urocystis ornithogali on Ornithogalum umbellatum agg. was rediscovered in Germany after around 140 years. Exobasidium schinzianum was found on Saxifraga rotundifolia in Austria, probably the only recent record for a long time. As new hosts for Germany Allium x proliferum for Uromyces ambiguus, Phaseolus coccineus for
Podosphaera xanthii and Symphytum caucasicum for Golovinomyces asperifoliorum were identified. The finds of Anthemis ruthenica with Golovinomyces macrocarpus and Leontodon incanus with Ramularia inaequalis are probably matrices novae.
In addition other interesting records of phytopathogenic microfungi are listed at the end of the publication, e.g. new records for federal states of Germany, Austria and Switzerland.
Keywords: Podosphaera spp., Prunus spp., Thecaphora lathyri, Urocystis ornithogali, Uromyces proëminens, Uromyces sparsus, Germany, Austria, Switzerland, matrix nova, new records, sequences
... Наприклад, значні зміни було внесено у видовий склад таких родів як Blumeria Golovin ex Speer (Liu et al., 2021), Golovinomyces (U. Braun) V.P. Heluta (Scholler et al., 2016;Braun et al., 2018Braun et al., , 2019 та Podosphaera Kunze (Moparthi et al., 2019;Meeboon et al., 2020). Отже, настав час здійснити узагальнення видового складу борошнисторосяних грибів України, розробити й опублікувати ключі для їхнього визначення. ...
The previously compiled list of species of powdery mildew fungi of Ukraine was published more than 30 years ago. That list included 108 species belonging to 12 genera. However, over the past three decades, numerous changes have been made in the system of the former order Erysiphales, in particular, in its generic composition; moreover, dozens of new species have been described. At the same time, many species of powdery mildew fungi were introduced to Europe, including Ukraine, in particular, from East Asia and North America. Thus, the species composition of powdery mildew fungi of Ukraine has undergone significant changes and requires a critical revision. Therefore, a series of relevant articles is planned to be published. This article, the first in the series, deals with the small genera Arthrocladiella and Blumeria. A list of species of these genera recorded in Ukraine (Arthrocladiella mougeotii, Blumeria avenae, B. bulbigera, B. dactylidis, B. graminicola, B. graminis, and B. hordei), their distribution by regions of Ukraine and a key for identification are provided. Arthrocladiella mougeotii was found to occur occasionally only in nine regions, mainly in the southern part of Ukraine, primarily in the forest-steppe and steppe zones and on the South Coast of Crimea. Blumeria avenae was recorded only in two regions, on the South Coast of Crimea and in Central Polissya and should be also considered rare in Ukraine. The remaining species were observed quite often and registered in 14–24 regions, while B. graminicola and B. graminis are regarded as the most common ones.
... According to the above cited authors, within the well known morphologic species of powdery mildews there are usually many subspecies, forms, varieties, biotypes or races which cannot be distinguished morphologically or which can be distinguished by only slight morphological differences. More recently the study on the phylogeny of the P. tridactyla, carried out by Meeboon et al. (2020) suggested that P. tridactyla is a complex consisting of at least 12 different species. Based on detailed morphological examinations and molecular sequence analyses, they have proposed dividing P. tridactyla into 10 species, encompassing 7 new species. ...
The rootstock Myrobalan 29C (Prunus cerasifera Ehrh.) is largely used as rootstocks of plum and apricot planting material. In May 2020, symptoms of powdery mildew were observed on leaves and young stems of Myrobalan 29C rootstocks. On the adaxial and abaxial leaf surface white colonies with white superficial mycelium and conidia were present. Chasmothecia appeared in some injured leaf tissues. Microscopic examination showed globose chasmothecia with appendages arising from the upper half of the fruiting bodies. Usually, the appendages were one to four times longer than the chasmothecial diameter and had two to four dichotomous branches. A single ascus observed in chasmothecium was broadly ellipsoid-ovoid and contained six to eight ascospores. The shape of the ascospores was ellipsoid-ovoid, too. Conidia formed in true chains on erect conidiophores, which were ellipsoid, hyaline and measured 24.6-33.6 × 10.8-20.8 µm (average 29.6 × 15.8 µm). Pathogenicity was confirmed by inoculation of healthy Myrobalan 29C, apricot (P. armeniaca L.) and cherry (P. avium L.) plants. Disease signs on all of the inoculated experimental species appeared as white powdery coating very similar to the powdery coating on natural infected Myrobalan 29C. Based on the morphological characteristics of the conidia, chasmothecia and results of the pathogenicity tests, the causative agent of the symptoms observed was presumed to belong to Podosphaera sp. (Ascomycetes). To the best of our knowledge, previously no detailed characterization of Podosphaera sp. in P. cerasifera was made in Bulgaria.
... For the selection of sequences and determination of host types, we used the following criteria: (i) reliable species identification based on both morphological characteristics and DNA sequences and (ii) originating from the host type on which the chasmothecial formation had been confirmed. The trait data of fungi were determined according to the descriptions in previous studies (e.g., Braun and Cook 2012;Moparthi et al. 2019;Meeboon et al. 2020). In the trait data set, the host types were classified into three categories: deciduous broad-leaved trees (deciduous), evergreen broad-leaved trees (evergreen), and herbaceous plants (herb). ...
The chasmothecial appendages of Erysiphaceae are considered to function in the overwintering strategy and evolve morphologically in line with transitions of different host type. However, the evolutionary patterns and relationships of these traits have not yet been verified using statistical models based on phylogenetic information. We aimed to clarify the evolutionary process of host type and appendage morphology in Cystotheceae using phylogenetic comparative methods (PCMs) and to evaluate the evolutionary relationship of these traits. The ancestral state estimation of host types showed that the deciduous type is the most ancestral in Cystotheceae, and the herb or evergreen types evolved secondarily four times and twice, respectively. Branched-or circinate-type appendages were estimated to be the most ancestral, and the mycelioid and rudimentary types evolved secondarily thrice and once, respectively. The results of the random forest analysis showed that the host type was predictable from the phylogeny and appendage morphology. The ancestral state estimation suggested that simultaneous transitions of the host type and appendage morphology occurred at several ancestral nodes. These results suggest some functional relationships between host type and appendage morphology, but there was no statistical support for an overall trend in evolutionary dependence between these traits. Our results demonstrate the utility of PCMs in the study of trait evolution in Cystotheceae, which can be applied to a broader phylogeny of powdery mildews to elucidate the evolutionary relationship and functional causality of phenotypic traits.
... Developing the specific molecular-phylogenetic tech-niques have also essentially contributed to the precise identification, construction of phylogeny and study of evolution of this group of fungi (Hirata & Takamatsu 1996, Takamatsu & Kano 2001, Bradshaw & Tobin 2020. The modern investigations have defined taxonomic status of many cryptic species and complex constitutes of powdery mildews (Scholler et al. 2016;Braun et al. 2018Braun et al. , 2019Meeboon et al. 2020;Qiu et al. 2020a, b) and comprehensive information about pathogenic features of different species has been obtained. Currently, powdery mildews are allocated within the order Helotiales according to the results of recent molecularphylogenetic studies (Johnston et al. 2019). ...
The present paper summarizes the taxonomic studies of Erysiphaceae (Helotiales, Ascomycota) in Azerbaijan. It represents a checklist of powdery mildews, which was prepared by combining all data collected from various works dealing with mycobiota of the country since mid of the last century. Consequently, 133 taxa belonging to the currently recognised genera – Arthrocladiella, Blumeria, Erysiphe, Golovinomyces, Leveillula, Neoërysiphe, Phyllactinia, Podosphaera, Sawadaea of the family Erysiphaceae have been reported. Host plants include angiosperms from 51 families and 227 genera. Current systematic position of each powdery mildew fungus was given with detailed information on their examined specimens.
... Qiu et al. (2020) revisited the heterogeneous species complex within the genus Golovinomyces by a multi-locus phylogenetic analysis. Meeboon et al. (2020) clarified that Podosphaera tridactyla represents a species complex consisting of at least 12 different species based on a phylogenetic analysis of combined ITS and 28S rDNA sequences. Phylogenetic analyses also played a key role in the recent study of Acalypha powdery mildews. ...
Powdery mildew on various Acalypha spp. has previously been assigned to Erysiphe acalyphae (≡ Uncinula acalyphae). In a recently published phylogenetic-taxonomic reassessment, it has been revealed that sequences retrieved from this Acalypha powdery mildew form a separate lineage representing a genus of its own for which the old genus name Salmonomyces was reinstated, with the new combination Salmonomyces acalyphae. Pseudoidium javanicum, described from Indonesia on Acalypha javanica, was reduced to synonymy with Salmonomyces acalyphae. Attempts to obtain sequence data from type material of Uncinula acalyphae, described from Kunming, Yunnan Province, China, on Acalypha supera (= Acalypha brachystachya), and efforts to collect fresh Chinese samples for epitypification purposes failed during the course of the works for the abovementioned studies. Therefore, the name Salmonomyces acalyphae was applied to all collections on Acalypha, first and foremost on the basis of the congruence of the morphological characteristics of the asexual morphs on Acalypha supera and other Acalypha spp. Meanwhile, attempts to trace new collections of this powdery mildew on Acalypha supera in Kunming, Yunnan Province, were successful and allow epitypification of Uncinula acalyphae with ex-epitype sequence data. Phylogenetic analyses surprisingly revealed the hidden diversity that two sister species are involved within Salmonomyces, viz., Salmonomyces acalyphae with both anamorph and teleomorph, confined to Acalypha supera in South China as host, and a second species, so far only known as anamorph, with wider host range and distribution, for which the new combination Salmonomyces javanicus, based on Pseudoidium javanicum, being introduced.
... ITS, SSU [234] Podosphaera aphanis var. aphanis Fragaria ananassa ITS [235] Podosphaera caricae-papayae Carica papaya ITS [236] Podosphaera caricicola Carica papaya ITS [145] Podosphaera pannosa Rosa hybrida ITS [235] Podosphaera pruni-cerasoidis Prunus cerasoides ITS, LSU [228] Podosphaera xanthii Justicia comata ITS [235] Gymnema inodorum ITS [235] Bidens pilosa ITS [235] Dahlia sp. ITS [235] Galinsoga parviflora ITS [235] Spilanthes iabadicensis ITS [235] Impatiens balsamina ITS [235] Millingtonia hortensis ITS [235] Heliotropium indicum ITS [235] Cleome rutidosperma ITS [235] Cucumis sativus ITS [235] Cucurbita pepo ITS [235] Momordica charantia ITS [235] Euphorbia hirta ITS [235] Crotalaria pallida ITS [235] Macroptilium atropurpureum ITS [235] Phaseolus vulgaris ITS [235] Senna occidentalis ITS [235] Vigna radiata ITS [235] Vigna unguiculata subsp. ...
Thailand is rich in fungal diversity and new taxa are continuously introduced. In the last decade there has been a vast improvement on the taxonomy, phylogeny and classification of fungi, including in Thailand. Earlier studies were solely based on morphology while modern research uses morphology and molecular phylogeny. There is a huge gap between currently accepted species numbers and previously described species based on morphology. Hence, clarification of previously introduced species using molecular data and providing missing molecular data for those species, is as important as introducing novel taxa. Correct identification of fungi is important in plant pathology and for quarantine purposes. Some introduced fungal species have been discovered to be species complexes based on modern molecular phylogeny. The adoption of "one fungus one name" has necessitated the updating of previous classifications. There are also many taxa yet to be discovered from Thailand. Thus it is important to recollect, sequence, re-describe and resolve the taxonomic ambiguities. It is important to document all fungal taxa as well as provide information on their life modes and to update plant quarantine checklists. In this review we discuss the status of phytopathogenic fungal identification, evolution of the fungal studies through some well-known examples, modern technologies that can be used, and prospects. Most importantly we provide a checklist of plant associated fungi of Thailand, for which there is molecular data available.
... Most powdery mildew species are host specific, with limited host ranges (Takamatsu et al. 2007) and rarely occurring on a wide range of hosts to form a species complex (Kiss et al. 2020, Meeboon et al. 2020. Podosphaera sect. ...
Podosphaera filipendulensis, a new teleomorph of powdery mildew was discovered on living leaves of Filipendula vestita (Rosaceae). It is the first species of powdery mildew that causes wilting and ultimately death of apical infected leaves of F. vestita. It is described, illustrated and compared with other Podosphaera species reported on Rosaceae and a key to all species reported on Rosaceae is provided. The phylogeny of this species has been inferred from internal transcribed spacer (ITS) rDNA sequence data. Phylogenetically, P. filipendulensis is close to P. ferruginea var. ferruginea and P. macularis but differs in having larger chasmothecia, smaller peridium cells, shorter, narrower and aseptate appendages, 8-spored asci and smaller ascospores.
... The diversity of powdery mildews in Japan based on modern taxonomic approaches and additional collections has been explored in recent years (e.g. Meeboon, Siahaan, Fujioka, & Takamatsu, 2017;Takamatsu, Katsuyama, & Shinoda, 2017;Meeboon, Takamatsu, & Braun, 2020). During the present study, we found powdery mildew specimens on Actinidia polygama (Siebold et Zucc.) ...
Erysiphe actinidiicola on Actinidia polygama is described based on morphological and molecular data. Erysiphe actinidiicola is distinguished from E. actinidiae var. actinidiae by having irregularly to dichotomously branched chasmothecial appendages, larger chasmothecia sizes and numbers of asci per chasmothecium. Molecular analyses indicated that this species forms a clade separated from E. actinidiae var. actinidiae. An epitype is proposed for E. actinidiae var. actinidiae with ex-epitype sequences. A powdery mildew found on Limonium tetragonum is tentatively described as Erysiphe sp. This species is distinguished from E. limonii, a powdery mildew on Limonium spp., based on the DNA sequence differences in the 28S rDNA and internal transcribed spacer region as well as the morphological differences in the length of the conidiophores. This is the first record of powdery mildew on L. tetragonum in the world.
... 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. ...
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.
... These sequences were aligned with the reference sequences of Erysiphe species shown in Table 1 by MUSCLE implemented in MEGA 7.0 (Kumar et al. 2016). Phylogenetic trees were constructed by maximum parsimony (MP) and maximum likelihood (ML) methods in PAUP* v. 4.0 (Swofford 2003) and raxmlGUI v. 1.3 (Silvestro & Michalak 2012), respectively, according to the procedures of Meeboon et al. (2020). All sites were treated as unordered and unweighted, with gaps treated as missing data. ...
The phylogeny and taxonomy of powdery mildews, belonging to the genus Erysiphe, on Lonicera species
throughout the world are examined and discussed. Phylogenetic analyses revealed that sequences retrieved from
Erysiphe lonicerae, a widespread powdery mildew species distributed in the Northern Hemisphere on a wide range of
Lonicera spp., constitutes a complex of two separate species, viz., E. lonicerae (s. str.) and Erysiphe ehrenbergii comb. nov.
Erysiphe lonicerae occurs on Lonicera spp. belonging to Lonicera subgen. Lonicera (= subgen. Caprifolium and subgen.
Periclymenum), as well as L. japonica. Erysiphe ehrenbergii comb. nov. occurs on Lonicera spp. of Lonicera subgen.
Chamaecerasus. Phylogenetic and morphological analyses have also revealed that Microsphaera caprifoliacearum (≡
Erysiphe caprifoliacearum) should be reduced to synonymy with E. lonicerae (s. str.). Additionally, Erysiphe lonicerina sp.
nov. on Lonicera japonica in Japan is described and the new name Erysiphe flexibilis, based on Microsphaera lonicerae
var. flexuosa, is introduced. The phylogeny of Erysiphe ehrenbergii and E. lonicerae as well as other Erysiphe species on
honeysuckle is discussed, and a survey of all species, including a key to the species concerned, is provided.
... were intensively studied by Cunnington et al. (2005c) based on specimens collected in Australia and elsewhere. Recently, the taxonomy of this group was revised (Moparthi et al., 2019;Meeboon et al., 2020), therefore the identity of the Australian specimens should be revisited. ...
In contrast to Eurasia and North America, powdery mildews (Ascomycota, Erysiphales) are understudied in Australia. There are over 900 species known globally, with fewer than currently 60 recorded from Australia. Some of the Australian records are doubtful as the identifications were presumptive, being based on host plant-pathogen lists from overseas. The goal of this study was to provide the first comprehensive catalog of all powdery mildew species present in Australia. The project resulted in (i) an up-to-date list of all the taxa that have been identified in Australia based on published DNA barcode sequences prior to this study; (ii) the precise identification of 117 specimens freshly collected from across the country; and (iii) the precise identification of 30 herbarium specimens collected between 1975 and 2013. This study confirmed 42 species representing 10 genera, including two genera and 13 species recorded for the first time in Australia. In Eurasia and North America, the number of powdery mildew species is much higher. Phylogenetic analyses of powdery mildews collected from Acalypha spp. resulted in the transfer of Erysiphe acalyphae to Salmonomyces, a resurrected genus. Salmonomyces acalyphae comb. nov. represents a newly discovered lineage of the Erysiphales. Another taxonomic change is the transfer of Oidium ixodiae to Golovinomyces. Powdery mildew infections have been confirmed on 13 native Australian plant species in the genera Acacia, Acalypha, Cephalotus, Convolvulus, Eucalyptus, Hardenbergia, Ixodia, Jagera, Senecio, and Trema. Most of the causal agents were polyphagous species that infect many other host plants both overseas and in Australia. All powdery mildews infecting native plants in Australia were phylogenetically closely related to species known overseas. The data indicate that Australia is a continent without native powdery mildews, and most, if not all, species have been introduced since the European colonization of the continent.
This is an updated version of the book published in 2019 and contains new records and almost a hundred new host/powdery mildew associations. An attempt has been made to revise species names following on from taxonomic revisions based on recent research and DNA methodology.
Powdery mildews are important obligate biotrophic ectoparasites that cause severe damage to various plant species in Southeast Asia. So far, 14 genera and 96 species of powdery mildew have been reported in Southeast Asia, with Thailand having the most number of recorded species. Of the 14 genera reported, Oidium sp. had the most recorded host, including several important species of vegetables, fruits, vines, herbs, field crops, ornamentals, and tree species. This paper briefly reviews the knowledge of powdery mildew disease in Southeast Asia, focusing on the pathogens' host range, impact, and some sources of resistance to powdery mildew.
Powdery mildews are a monophyletic group of obligate plant pathogenic fungi in the family Erysiphaceae. Powdery mildews are economically important in that they cause damage to many agriculturally significant crops and plants in ecologically important habitats. In this contribution, we introduce a new series of publications focusing on the phylogeny and taxonomy of this group, with an emphasis on specimens collected from North America. The first part of the series focuses on the genus Golovinomyces and includes a section detailing the powdery mildew species concept. We conducted analyses of Golovinomyces spp. with available rDNA sequence data from GenBank and supplemented the data set with rDNA (ITS, 28S, IGS) as well as protein-coding (GAPDH) data from 94 North American collections. Many of the species evaluated are included in phylogenetic and morphological analyses for the first time, including the American species G. americanus, G. brunneopunctatus, G. californicus, G. greeneanus, G. hydrophyllacearum, and G. sparsus. A special emphasis was placed on acquiring ex-type or ex-epitype sequences or presenting reference sequences for phylogenetic-taxonomic purposes. Three new species, G. eurybiarum, G. galiorum, and G. malvacearum, are described, and the new combinations G. fuegianus, G. mutisiae, and G. reginae are introduced. Ex-holotype sequences of Erysiphe sparsa (≡ G. sparsus) reveal that it should be reduced to synonymy with G. ambrosiae, and ex-epitype sequences of G. valerianae reveal that it should be reduced to synonymy with G. orontii. Multiple epitypes are designated with ex-epitype sequences.
Based on the results of the authors' research and revisions of previous information, modern species diversity has been established and the first annotated list of powdery fungi on woody plants in urban habitats of Sverdlovsk Region (Ekaterinburg and several cities of the region) has been compiled. Totally, 29 species of Erysiphaceae have been recorded, among which the majority belong to the genus Erysiphe (17 species), and other belong to the genera Podosphaera (7), Phyllactinia (3) and Sawadaea (2). Six species were first found in Sverdlovsk Region: Erysiphe ehrenbergii, E. euonymi, E. lonicerae, E. viburni, Podosphaera myrtillina, and P. spiraeae. Almost half of all identified species (14 of 29) are alien to Sverdlovsk Region, while 6 species can be considered as invasive, of which two species originate from North America (Erysiphe necator and Podosphaera mors-uvae) and four species from East Asia (Erysiphe alphitoides, E. corylacearum, E. palczewskii, and E. vanbruntiana), and another eight species can be considered as unintentionally introduced from neighboring regions together with their host plants. The growth of most species occurs in the period from mid-June to the end of September. The majority of recorded Erysiphaceae species (25 out of 29) form both anamorph and teleomorph, and only four species (Erysiphe aquilegiae, E. necator, Podosphaera pannosa и P. spiraeae) are recorded in the anamorphic stage only.
2021) Noteworthy records of phytopathogenic micromycetes (16). Zeitschrift für Mykologie 87(2):229-329. Abstract: Some interesting records of plant parasitic microfungi of the Erysiphaceae, Pero-nosporomycetes, Pucciniomycotina, Ustilaginomycotina and anamorphic Ascomycetes collected in Germany and Austria are reported. A record of Entyloma on Scorzonera hispanica belongs to a new species that has not yet been described. The plant species was not known as a host for the genus Entyloma before. The North American Microbotryum alsines was found for the first time in Europe on Stellaria media. A record of Erysiphe syringae on Fraxinus pennsylvanica is the first record of this powdery mildew on the genus Fraxinus in Europe. The host species is a matrix nova for this fungus. Puccinia scirpi was rediscovered in Germany after 80 years on Nymphoi-des peltata. Exobasidium schinzianum on Saxifraga rotundifolia, Peronospora ruegeriae on Onobry-chis viciifolia and Urocystis bolivarii on Lolium perenne are reported as new for Germany. Three recorded host species are matrices novae: Cornus mas for Erysiphe pulchra, Eutrema japonicum for Erysiphe cruciferarum and Origanum vulgare for Peronospora stigmaticola. In addition other interesting records are listed, including formerly unknown host species in Germany, e. g.
Erysiphe adunca s. lat. is a common powdery mildew species, distributed almost worldwide, on hosts of Populus and Salix species. Based on its wide host range and distribution as well as its strong morphological variations, E. adunca s. lat. was previously divided into several species or, alternatively, into varieties. However, comprehensive phylogenetic examinations on the putative monophyly of this species or the existence of a possible E. adunca species complex have not been conducted based on combined morphological and molecular approaches. In this study, we retrieved rDNA (ITS and LSU) sequences of 22 powdery mildew specimens on Populus and Salix spp. collected in China, Germany, Iran, Japan, Russia, and the United States and conducted morphological examinations of these and additional specimens. The morphological and molecular data obtained clearly indicate that E. adunca is a species complex comprised of at least four species., viz., E. adunca s. str. (on Populus spp. in Asia, Europe, and North America), E. capreae, including E. pseudoregularis as a new synonym (on Salix spp., particularly on species in Salix subgen. Vetrix sect. Vetrix in Asia, Europe, and North America), the reinstated E. salicis (on Salix spp. belonging to various subgenera of Salix in Asia and Europe), and E. mandshurica (which has been confirmed as a species of its own on Populus spp., in Asia). In addition to the clear phylogenetic differentiation, the morphology of the asexual and sexual morphs of E. adunca s. str., E. capreae and E. salicis is easily distinguishable. Sequences retrieved from collections on Salix gracilistyla (Uncinula salicis-gracilistylae), a species that was previously considered a variety of Uncinula adunca and Erysiphe adunca, formed a small subclade within the E. salicis clade and are now referred to as Erysiphe salicis var. salicis-gracilistylae comb. nov. Alphitomorpha adunca; Erysiphe capreae and E. salicis are epitypified, and a neotype is designated for Alphitomorpha obtusata, with ex-epitype sequences and, in the latter case, an ex-neotype sequence.
A first annotated checklist of powdery mildews (Erysiphaceae) recorded from the Fergana Valley (within Uzbekistan) was compiled on the base of original long-term research, revisions of previously collected specimens deposited at Tashkent Mycological Herbarium of the Institute of Botany of Academy of Sciences of Uzbekistan (TASM), and earlier publications of other mycologists and phytopathologists. In total, 67 erysiphaceous species of ten genera on 153 angiosperm host plant species of 100 genera and 37 families have been recorded, including 21 species and 1 variety found for the first time in the Fergana Valley. The most species-rich genera in the investigated area are Leveillula (16 species), Erysiphe (15), Podosphaera (12), Golovinomyces (10), and Phyllactinia (8), whereas the genera Arthrocladiella, Blumeria, and Sawadaea are only represented by a single species per genus. Such a particular taxonomic composition of powdery mildews can be considered typical for Central Asia, especially with regard to the prevailing position of Leveillula. The present checklist contains detailed information about all particular findings for each powdery mildew species: recorded host plant species, typical habitats, collection locations, collection dates, collectors’ names and TASM herbarium specimen numbers. The most common species are Blumeria graminis, Erysiphe cruciferarum, E. trifoliorum, E. polygoni, Golovinomyces artemisiae, Leveillula alhagi, L. cylindrospora, L. lappae, L. papilionacearum, L. picridis, L. taurica, Podosphaera clandestina, P. pannosa, P. xanthii, Phyllactinia babayanii, P. fraxini, and P. moricola; whereas all other powdery mildew species are relatively rare
New records of powdery mildews (Erysiphaceae) from Switzerland and adjacent countries are listed and annotated, including first records of multiple host plants worldwide. The collections concerned are described, illustrated, discussed, and some identifications have been confirmed by results of sequencing (ITS + 28S rDNA). The new species, Erysiphe abeliana (on Abelia ×grandiflora) and Phyllactinia cruchetii (on Passiflora coerulea), are described. Some previous records of powdery mildews
from Switzerland based only on morphology have been confirmed by sequence data. Type specimens of Erysiphe chifengensis and E. potentillae have been sequenced, and new sequences have been generated from collections of E. malvae and E. punicae. Microsphaera abeliae (Erysiphe abeliicola) isepitypified with an ex-epitype sequence.
We present the latest version of the Molecular Evolutionary Genetics Analysis (MEGA) software, which contains many sophisticated
methods and tools for phylogenomics and phylomedicine. In this major upgrade, MEGA has been optimized for use on 64-bit computing
systems for analyzing bigger datasets. Researchers can now explore and analyze tens of thousands of sequences in MEGA. The
new version also provides an advanced wizard for building timetrees and includes a new functionality to automatically predict
gene duplication events in gene family trees. The 64-bit MEGA is made available in two interfaces: graphical and command line.
The graphical user interface (GUI) is a native Microsoft Windows application that can also be used on Mac OSX. The command
line MEGA is available as native applications for Windows, Linux, and Mac OSX. They are intended for use in high-throughput
and scripted analysis. Both versions are available from www.megasoftware.net free of charge.
The internal transcribed spacer (ITS) DNA marker is routinely used for fungal identification but gives a clear result for only three out of four powdery mildew samples. A search for new markers indicates that some genes offer enhanced identification in comparison with ITS. Others fail due to amplification and sequencing difficulties and lack of informative variability. Powdery mildews (Ascomycota, Erysiphales) are biotrophic, fungal plant pathogens that commonly occur worldwide on a wide range of host plants. They are unsightly and greatly reduce the vigor of their hosts and have major impacts on crop and other cultivated plants. Species within this order are straightforward to spot, but difficult to identify. A citizen science scheme was run in 2013–2016 in the UK to gather a wide array of samples on which identification methods could be developed. Current techniques for identification and phylogenetic reconstruction show scope for improvement. In this paper, we review genes used in other fungal groups for discrimination at species level. Working protocols for amplification and sequencing of seven genes (actin, β-tubulin, calmodulin,Chs, elongation factor 1-α [EF1-α], Mcm7, and Tsr1) are developed with varying success; Mcm7 proves to be the most useful at differentiation between closely related, phylogenetically young powdery mildew species for phylogenetic reconstruction when used separately and in tandem with ITS. We therefore propose this as the most appropriate candidate gene to be used commonly in powdery mildew diagnostics alongside the ITS; furthermore, this could be transferred to similarly troublesome fungal clades.
Powdery mildew of Prunus spp. is a significant disease in most cherry growing regions of Washington, USA. Powdery mildews on Prunus virginiana and Pr. avium were previously assigned to Podosphaera clandestina s. lat. (= Po. oxyacanthae) or Po. prunicola. In this report, we confirm the presence of two distinct Podosphaera species on these hosts. Phylogenetic analyses of nuc rDNA sequences from the internal transcribed spacer region (ITS1-5.8S-ITS2 = ITS) and 28S subunit confirmed the presence of two distinct species. A morphological comparison with type material of Po. prunicola and additional collections demonstrated that the powdery mildew on Pr. virginiana (including var. demissa and var. melanocarpa) is in fact Po. prunicola. The powdery mildew on Pr. avium is genetically, morphologically, and biologically distinct from Po. prunicola and is described here as the new species Po. cerasi. Cross-inoculation experiments confirmed that these two species are host specific. Podosphaera prunicola was unable to colonize Pr. avium, whereas Po. cerasi was unable to colonize Pr. virginiana. Morphological reexamination of numerous specimens identified as Po. prunicola on a broad range of Prunus species suggests that Po. prunicola is probably confined to species in Prunus subgen.Padus (= Prunus subgen. Cerasus sect. Laurocerasus, including sect. Padus), with Pr. virginiana as the principal host. Podosphaera cerasi occurs on hosts in Prunus subgen. Cerasus, and our work confirms a newly described species of powdery mildew on Pr. avium. This work also includes the first documented and genetically proven European record of Po. prunicola on Pr. serotina and its widespread occurrence in the United States.
Golovinomyces orontii is a common plurivorous powdery mildew species with wide host range and worldwide distribution, usually occurring as asexual morph. Ascomata (chasmothecia) are rarely formed on most hosts. Phylogenetic analyses based on rDNA ITS and 28S rDNA data of a wide range of powdery mildew collections of G. orontii s. lat. suggested a high degree of genetic heterogeneity of this species, which is undoubtedly not monophyletic. This study revealed that sequences retrieved from numerous collections referred to as G. orontii (s. lat.) split into three distinct main clusters, previously classified as groups 1 to 3. These groups have been genetically and morphologically analyzed, circumscribed, and named. One cluster (group 2), including a sequence retrieved from powdery mildew on the type host of Erysiphe orontii, Misopates orontium, constitutes G. orontii s. str. G. tabaci comb. nov. (≡ E. tabaci) is introduced for the second cluster in group 1 that is genetically and morphologically clearly distinct from G. orontii s. str. The third assemblage of sequences (group 3) comprises powdery mildews on hosts of the composite tribe Cichorieae, including Cichorium and Lactuca spp., and a wide range of hosts belonging to various other plant families for which the name G. bolayi sp. nov. is proposed. Euoidium longipes and Euoidium lycopersici, two additional powdery mildew species on solanaceous hosts, are briefly discussed and, based on previous phylogenetic analyses, reallocated to Golovinomyces. Oidium lactucae-debilis on Ixeris japonica in Asia is tentatively reduced to synonymy with G. sonchicola, i.e., it does not pertain to the G. orontii s. lat. complex. G. orontii s. lat. on Vinca spp. (Apocynaceae), mostly known as asexual morph, represents a separate species only distantly related to G. orontii s. str., which is described herein as Golovinomyces vincae sp. nov. Some re-examined collections on Vinca major from Germany misidentified as G. orontii turned out to belong to Ramularia vincae and represent first records of this species for Germany. Golovinomyces spadiceus is a further plurivorous Golovinomyces species discussed in this work, which, however, does not belong to the G. orontii complex.
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 investigated genotype speciation in E. gracilis var. longissima occurring on Q. acuta and Q. sessilifolia, and found that this variety is also divided into two distinct genotypes. These results suggested that E. gracilis represents a species complex consisting of six different species. Based on detailed morphological examinations correlating with results of molecular sequence analyses, we propose to divide E. gracilis into six species, encompassing three new species (E. uncinuloides, E. pseudogracilis, and E. longiappendiculata), one new name (E. longifilamentosa), and two known species (E. gracilis s. str. and E. hiratae). A key to the species concerned is provided.
Two new species, Erysiphe azerbaijanica on Castanea sativa and E. linderae on Lindera praecox, both belonging to the Microsphaera lineage of the genus Erysiphe are described based on morphological and molecular data. Erysiphe azerbaijanica is distinguished from other Erysiphe species occurring on Castanea spp. by its cylindrical conidia with a length/width ratio of 2-3.6, longer conidiophore, and foot-cells. Molecular analyses indicated that this species forms a clade of its own, supporting the morphological observations. Molecular phylogenetic analyses showed that E. blasti s. lat. is divided into two genetically differentiated groups associated with different host species. Based on the sequence differences in the 28S rRNA gene and ITS region, connected with differences in the number and length of appendages, the fungus on L. praecox is described as a new species, E. linderae.
Parauncinula, a genus belonging to the Erysiphaceae (powdery mildew family), consisted of only two species, P. curvispora and P. septata, which are distributed only in China and Japan (endemic to East Asia). This genus split from other powdery mildew genera at an early stage of evolution, and thus has been considered as one of the most ancestral genera in the Erysiphales. Phylogenetic analyses using rDNA internal transcribed spacer (ITS) region and 28S rRNA gene sequences indicated that one of the species, P. septata, is divided into three distinct genotypes. The ITS sequence similarities between the three genotypes were 96% or less, suggesting species level differences among the genotypes. Combined with detailed morphological observations, P. septata s. lat. is divided into three species, viz. P. septata s. str., P. polyspora, and P. uncinata. These species are clearly defined by morphological characteristics such as the number of ascospores per ascus and morphology of appendages, as well as DNA sequences. A key to the species of Parauncinula is also presented.
Podosphaera salatai sp. nov., a new powdery mildew species on Cerasus incana (Rosaceae) in Georgia (Transcaucasia), is described, illustrated, compared with allied species and discussed.
A phylogenetic analysis of the Erysiphe with uncinuloid ascoma appendages (Erysiphe section Uncinula, Erysiphales, Ascomycota) on Carpinus spp. was done using sequences of the rDNA ITS regions and the D1/D2 domains of the 28S rDNA. These results, combined with morphological data, revealed a complex consisting of several distinct taxa. These included the already described Erysiphe
carpinicola on C. japonica distinguishable from the Erysiphe sp. on C. betulus and C. tschonoskii as well as the one on C. laxiflora. Thus, it was shown that Oidium
carpini, described from Europe on Carpinus betulus, the powdery mildew with uncinula-like ascomata, recently found in Europe on this host, as well as an Erysiphe on C. tschonoskii in Japan, described previously as E. carpinicola, all belong to a single new species, named E. arcuata in this paper. As the powdery mildew on C. laxiflora was also distinct from other known species, it is named E. carpini-laxiflorae in this paper. The already described E. pseudocarpinicola and Erysiphe sp. on Carpinus cordata are two additional taxa, which are morphologically and genetically distinguished from the other species of Erysiphe sect. Uncinula on Carpinus spp.
The very close relationship between the genera Erysiphe s.str. ( + Erysiphe sect. Erysiphe), Microsphaera, and Uncinula, which are connected by numerous intermediate taxa, is strongly supported by light as well as scanning electron microscopical features and has recently been confirmed by examinations of nucleotide sequences of rDNA internal transcribed spaces (ITS). Erysiphe s.str. and Microsphaera did not group into separate monophyletic lineage, but formed several small mixed clusters. It turned out that the formation of branched ascoma appendages does not have any taxonomic value at generic level. Therefore, Microsphaera is reduced to synonymy with Erysiphe s.str., and the new combinations, which are necessary, are introduced. Uncinula spp. form a basal subclade of the Erysiphe s.str./Microsphaera (Pseudoidium) clade, but since some species of Erysiphe s.str. belong in this subclade, it is necessary to reduce Uncinula to synonymy with the latter genus, too. Furthermore, examinations of nucleotide sequences of rDNA ITS regions of numerous taxa belonging to tribe Cystotheceae showed that Podosphaera and Sphaerotheca did not form separate monophyletic clades. Taxa of Sphaerotheca sect. Sphaerotheca and sect. Magnicellulatae grouped together with two different sets of Podosphaera spp. in two separate subclades of a single Podosphaera/Sphaerotheca clade. Based on these results and since Podosphaera and Sphaerotheca are morphologically only distinguished by having appendages which are branched and simple, respectively, it is proposed to merge the two genera. New combinations for Sphaerotheca spp. under Podosphaera are introduced.
The teleomorph of Erysiphe takamatsui, a powdery mildew of lotus, was found in 2012 at a lotus pond of Niigata-shi, Japan, where this species was first found in 1974. This is the second record of teleomorph of E. takamatsui. Morphological and molecular analyses revealed that the asexual stages of the lotus powdery mildews found in Aichi, Osaka and Tokyo represent the anamorph of E. takamatsui. The nucleotide sequences of the internal transcribed spacer (ITS) and 28S rRNA gene were identical to those of E. aquilegiae, E. catalpae, and E. macleayae. These species formed a homogeneous clade together with E. sedi, Pseudoidium neolycopersici, and Pseudoidium spp. occurring on a wide range of plant families with only five base substitutions of ITS sequences in maximum. These results suggest that E. takamatsui is a fungus that appeared as a result of recent host expansion.
Almonds and peaches are among the most economically important fruit crops in subtropical and temperate regions. Questions regarding the origin and domestication of the crop species, interspecific relationships, and evolutionary patterns of fruits have remained unanswered due to the lack of phylogenetic data of Prunus subgenus Amygdalus, in which both almond and peach are classified. Twenty-two species of subg. Amygdalus were included in phylogenetic analyses of plastid ndhF-rpl32, rpL16, trnH-psbA, trnL-trnF, trnQ-5′rps16, and trnS-trnG and nuclear s6pdh genes to investigate the circumscription of subg. Amygdalus, phylogenetic relationships among the species, and implication for mesocarp and endocarp evolution. Prunus subg. Amygdalus is best circumscribed to include both almonds and peaches, comprising two sections, Amygdalus and Persica. Species traditionally placed close to almonds, such as P. tenella, P. petunnikowii, P. triloba, and P. pedunculata, should be excluded from subg. Amygdalus. Characteristics used to diagnose subg. Amygdalus, such as dry, splitting mesocarp and the ornamentation of endocarp evolved multiple times in Prunus; thus classification based solely on these morphological characters produces a taxonomic scheme that might not represent common ancestry. A dry, splitting mesocarp is likely to be ancestral for the common ancestor of subg. Amygdalus, and the fleshy, non-splitting mesocarp found in peaches was derived from the dry, splitting mesocarp, presumably by domestication and selection for a fleshy fruit in China. Very low genetic divergence among cultivated almond and its close relatives might be associated with gene flow and/or rapid speciation within the group.
The classification of the economically important genus Prunus L. sensu lato (s.l.) is controversial due to the high levels of convergent or the parallel evolution of morphological characters. In the present study, phylogenetic analyses of fifteen main segregates of Prunus s.l. represented by eighty-four species were conducted with maximum parsimony and Bayesian approaches using twelve chloroplast regions (atpB-rbcL, matK, ndhF, psbA-trnH, rbcL, rpL16, rpoC1, rps16, trnS-G, trnL, trnL-F and ycf1) and three nuclear genes (ITS, S6PDH and SBEI) to explore their infrageneric relationships. The results of these analyses were used to develop a new, phylogeny-based classification of Prunus s.l. Our phylogenetic reconstructions resolved three main clades of Prunus s.l. with strong supports. We adopted a broad-sensed genus, Prunus, and recognised three subgenera corresponding to the three main clades: subgenus Padus, subgenus Cerasus and subgenus Prunus. Seven sections of subgenus Prunus were recognised. The dwarf cherries, which were previously assigned to subgenus Cerasus, were included in this subgenus Prunus. One new section name, Prunus L. subgenus Prunus section Persicae (T. T. Yü & L. T. Lu) S. L. Zhou and one new species name, Prunus tianshanica (Pojarkov) S. Shi, were proposed.
With the increasing availability of molecular data, maximum likelihood approaches have gained a new central role in phylogenetic reconstructions. Extremely fast tree-search algorithms have been developed to handle data sets of ample size in reasonable time. In the past few years, RAxML has achieved great relevance in this field and obtained wide distribution among evolutionary biologists and taxonomists because of its high computational performance and accuracy. However, there are certain drawbacks with regard to its usability, since the program is exclusively command-line based. To overcome this problem, we developed raxmlGUI, a graphical user interface that makes the use of RAxML easier and highly intuitive, enabling the user to perform phylogenetic analyses of varying complexity. The GUI includes all main options of RAxML, and a number of functions are automated
or simplified. In addition, some features extend the standard use of RAxML, like assembling concatenated alignments with automatic partitioning. RaxmlGUI is an open source Python program, available in a cross-platform package that incorporates RAxML executables for the main operating systems. It can be downloaded from http://sourceforge.net/projects/raxmlgui/.
Keywords: Rapid bootstrap; Graphical user interface; Maximum likelihood; Phylogenetic analyses; Python; RAxML
Podosphaera is a genus of the powdery mildew fungi belonging to the tribe Cystotheceae of the Erysiphaceae. Among the host plants of Podosphaera, 86 % of hosts of the section Podosphaera and 57 % hosts of the subsection Sphaerotheca belong to the Rosaceae. In order to reconstruct the phylogeny of Podosphaera and to determine evolutionary relationships between Podosphaera and its host plants, we used 152 ITS sequences and 69 28S rDNA sequences of Podosphaera for phylogenetic analyses. As a result, Podosphaera was divided into two large clades: clade 1, consisting of the section Podosphaera on Prunus (P. tridactyla s.l.) and subsection Magnicellulatae; and clade 2, composed of the remaining member of section Podosphaera and subsection Sphaerotheca. Because section Podosphaera takes a basal position in both clades, section Podosphaera may be ancestral in the genus Podosphaera, and the subsections Sphaerotheca and Magnicellulatae may have evolved from section Podosphaera independently. Podosphaera isolates from the respective subfamilies of Rosaceae each formed different groups in the trees, suggesting a close evolutionary relationship between Podosphaera spp. and their rosaceous hosts. However, tree topology comparison and molecular clock calibration did not support the possibility of co-speciation between Podosphaera and Rosaceae. Molecular phylogeny did not support species delimitation of P. aphanis, P. clandestina, P. ferruginea, P. spiraeae and P. tridactyla in their current circumscriptions, which suggests the need for revision of these species.
The nucleotide sequences of the nuclear ribosomal DNA including 18S, 5.8S, and 28S rDNA and the internal transcribed spacer regions were determined for 33 powdery mildew taxa spanning 15 genera to infer the phylogenetic relationships for these fungi. Phylogenetic analyses revealed that Uncinula septata was placed in the primitive base to the large clade composed of all other powdery mildew taxa. Powdery mildews excluding Unc. septata were split into five major lineages. The Pseudoidium lineage consisted of six genera, i.e., Erysiphe, Microsphaera, Uncinula, Uncinuliella, Brasiliomyces, and Typhulochaeta, all of which are characterized by polyascal cleistothecia and Pseudoidium-type anamorph. The Eudoidium lineage consisted of three Erysiphe species (E. cichoracearum, E. orontii and E. galeopsidis) and Arthrocladiella mougeotii, which are characterized by polyascal cleistothecia and Euoidium-type anamorph. The endophytic lineage consisted of two Phyllactinia species (Phy. moricola and Phy. kakicola), Leveillula taurica and Pleochaeta shiraiana, which are characterized by the presence of endophytic or partly endophytic mycelia. The fibrosin lineage consisted of the genera Sawadaea, Cystotheca, Podosphaera, and Sphaerotheca, all of which are characterized by the presence of well-developed fibrosin bodies in the conidia and conidiophores. The monocot lineage consisted of a single species, Blumeria graminis, which is well characterized by being parasitic to monocots, especially to cereal plants. Character polarization was discussed on several major characteristics. The mycelioid appendage which has long been regarded as an ancestral character may be a derived character as a result of an event that may have occurred multiple times independently due to convergence. Euoidium-type anamorph and clavate appressorial germ tube are considered to be ancestral characters to Pseudoidium-type anamorph and lobed germ tube. Ectophytic nature and polyascal cleistothecia are also regarded as ancestral features to endophytic nature and monoascal cleistothecia.
The economically important plum or cherry genus (PRUNUS:) and the subfamily Amygdaloideae of the Rosaceae have a controversial taxonomic history due to the lack of a phylogenetic framework. Phylogenetic analysis using the ITS sequences of nuclear ribosomal DNA (nrDNA) was conducted to construct the evolutionary history and evaluate the historical classifications of PRUNUS: and the Amygdaloideae. The analyses suggest two major groups within the Amygdaloideae: (1) PRUNUS: s.l. (sensu lato) and MADDENIA:, and (2) EXOCHORDA:, Oemleria, and PRINSEPIA: The ITS phylogeny supports the recent treatment of including EXOCHORDA: (formerly in the Spiraeoideae) in the Amygdaloideae. MADDENIA: is found to be nested within PRUNUS: s.l. in the parsimony and distance analyses, but basal to PRUNUS: s.l. in the maximum likelihood analysis. Within PRUNUS:, two major groups are recognizable: (1) the AMYGDALUS:-PRUNUS: group, and (2) the CERASUS:-LAUROCERASUS:-PADUS: group. The clades in the ITS phylogeny are not congruent with most subgeneric groups in the widely used classification of PRUNUS: by Rehder. A broadly defined PRUNUS: is supported.
The recently-developed statistical method known as the "bootstrap" can be used to place confidence intervals on phylogenies. It involves resampling points from one's own data, with replacement, to create a series of bootstrap samples of the same size as the original data. Each of these is analyzed, and the variation among the resulting estimates taken to indicate the size of the error involved in making estimates from the original data. In the case of phylogenies, it is argued that the proper method of resampling is to keep all of the original species while sampling characters with replacement, under the assumption that the characters have been independently drawn by the systematist and have evolved independently. Majority-rule consensus trees can be used to construct a phylogeny showing all of the inferred monophyletic groups that occurred in a majority of the bootstrap samples. If a group shows up 95% of the time or more, the evidence for it is taken to be statistically significant. Existing computer programs can be used to analyze different bootstrap samples by using weights on the characters, the weight of a character being how many times it was drawn in bootstrap sampling. When all characters are perfectly compatible, as envisioned by Hennig, bootstrap sampling becomes unnecessary; the bootstrap method would show significant evidence for a group if it is defined by three or more characters.
— We studied sequence variation in 16S rDNA in 204 individuals from 37 populations of the land snail Candidula unifasciata (Poiret 1801) across the core species range in France, Switzerland, and Germany. Phylogeographic, nested clade, and coalescence analyses were used to elucidate the species evolutionary history. The study revealed the presence of two major evolutionary lineages that evolved in separate refuges in southeast France as result of previous fragmentation during the Pleistocene. Applying a recent extension of the nested clade analysis (Templeton 2001), we inferred that range expansions along river valleys in independent corridors to the north led eventually to a secondary contact zone of the major clades around the Geneva Basin. There is evidence supporting the idea that the formation of the secondary contact zone and the colonization of Germany might be postglacial events. The phylogeographic history inferred for C. unifasciata differs from general biogeographic patterns of postglacial colonization previously identified for other taxa, and it might represent a common model for species with restricted dispersal.
A phylogeny of Erysiphe sect. Uncinula on Carpinus spp. was reconstructed using the 28S rDNA sequences and a combined alignment of the 28S, ITS, and IGS rDNA sequences. The analysis was supplemented with morphological data obtained from examination of voucher specimens. A sequence of Erysiphe sect. Uncinula on C. cordata formed a distinct lineage separated from sequences of other Erysiphe species on Carpinus spp., indicating a cryptic species, which is described as E. paracarpinicola. The new species is genetically as well as morphologically most similar to E. carpinicola s. str., but differs in having fewer asci per chasmothecium (mainly 3–5 vs 4–10) and shorter chasmothecial appendages. A key to species of Erysiphe sect. Uncinula on Carpinus spp. is provided.
Phylogenetic relationships among 48 species of Prunus were investigated by means of maximum parsimony analysis of sequence polymorphism from nuclear ITS and chloroplast trnL-trnF spacer DNA. Several genera that have been implicated as close relatives of Prunus in previous studies were utilized as outgroups. The objective of this study was to reconstruct the phylogeny of Prunus with the purpose of reviewing previously described taxonomic relationships and providing a basis for studies of morphological evolution in the genus. Variability and homoplasy are higher in ITS than in trnL-trnF. Parsimony analysis of both markers, separately and in combination, supported a close relationship among Exochorda, Oemleria, and Prinsepia but did not support a sister relationship of this clade with Prunus. Prunus is monophyletic and divided into two groups. The first is a strongly supported clade composed of subgenera Padus, Laurocerasus, and Cerasus. None of these subgenera is monophyletic and the trait used to characterize Padus and Laurocerasus, flowers arranged in racemes, can be more broadly interpreted as a synapomorphy for the clade. The second group included the subgenera Amygdalus and Prunus, and sections Microcerasus (subgenus Cerasus), and Penarmeniaca, which has never been assigned to a subgenus. The exclusively North American subgenus Emplectocladus may be an early diverging taxon sister to the second group. Sections Prunus and Armeniaca are monophyletic with low bootstrap support. Section Penarmeniaca, from southwestern US, and P. besseyi (section Microcerasus) are sister taxa. A Eurasian origin of Prunus is supported. Communicating Editor: Richard Jensen
Sequences of the chloroplast ndhF gene and the nuclear ribosomal ITS regions are employed to reconstruct the phylogeny of Prunus (Rosaceae), and evaluate the classification schemes of this genus. The two data sets are congruent in that the genera Prunus s.l. and Maddenia form a monophyletic group, with Maddenia nested within Prunus. However, the ndhF data set is incongruent with the ITS data supporting two major groups within Prunus: one consisting of subgenera Laurocerasus (including Pygeum) and Padus as well as the genus Maddenia and another of subgenera Amygdalus, Cerasus, and Prunus. The ITS data, on the other hand, support a clade composed of subgenera Amygdalus and Prunus and Prunus sect. Microcerasus in addition to a paraphyletic grade of subgenera Laurocerasus and Padus (and the genus Maddenia) taxa. In general, the subgeneric classifications of Prunus s.l. are not supported. The ITS and ndhF phylogenies differ mainly in interspecific relationships and the relative position of the Padus/Laurocerasus group. Both ITS and ndhF data sets suggest that the formerly recognized genus Pygeum is polyphyletic and that the distinction of the subgenera Padus and Laurocerasus is not supported. The biogeographic interactions of the temperate and tropical members in the Padus/Laurocerasus/Maddenia alliance including Pygeum are shown to be highly dynamic and complex.
We have designed two taxon-selective primers for the internal transcribed spacer (ITS) region in the nuclear ribosomal repeat unit. These primers, ITS1-F and ITS4-B, were intended to be specific to fungi and basidiomycetes, respectively. We have tested the specificity of these primers against 13 species of ascomycetes, 14 of basidiomycetes, and 15 of plants. Our results showed that ITS4-B, when paired with either a ‘universal’ primer ITS1 or the fungal-specific primer ITS1-F, efficiently amplified DNA from all basidiomycetes and discriminated against ascomycete DNAs. The results with plants were not as clearcut. The ITS1-F/ITS4-B primer pair produced a small amount of PCR product for certain plant species, but the quantity was in most cases less than that produced by the ‘universal’ ITS primers. However, under conditions where both plant and fungal DNAs were present, the fungal DNA was amplified to the apparent exclusion of plant DNA. ITS1-F/ITS4-B preferential amplification was shown to be particularly useful for detection and analysis of the basidiomycete component in ectomycorrhizae and in rust-infected tissues. These primers can be used to study the structure of ectomycorrhizal communities or the distribution of rusts on alternate hosts.
We describe MUSCLE, a new computer program for creating multiple alignments of protein sequences. Elements of the algorithm
include fast distance estimation using kmer counting, progressive alignment using a new profile function we call the log‐expectation score, and refinement using tree‐dependent
restricted partitioning. The speed and accuracy of MUSCLE are compared with T‐Coffee, MAFFT and CLUSTALW on four test sets
of reference alignments: BAliBASE, SABmark, SMART and a new benchmark, PREFAB. MUSCLE achieves the highest, or joint highest,
rank in accuracy on each of these sets. Without refinement, MUSCLE achieves average accuracy statistically indistinguishable
from T‐Coffee and MAFFT, and is the fastest of the tested methods for large numbers of sequences, aligning 5000 sequences
of average length 350 in 7 min on a current desktop computer. The MUSCLE program, source code and PREFAB test data are freely
available at http://www.drive5. com/muscle.
Podosphaera tridactyla (Ascomycota: Erysiphales) is a morphologically variable species occurring on Prunus s. lat. In order to assess the genetic variation within this species, the rDNA ITS region was amplified from 29 specimens from a range of Prunus species collected in Australia, Switzerland, and Korea. RFLP analysis of the PCR products revealed six groups, and a comparison of sequences from representatives of these groups revealed three clades: Clade I contained all specimens from Prunus subgen. Prunus; Clade 2 specimens from a variety of Prunus subgenera, except subgen. Prunus, and Podosphaera longiseta was closely allied to this clade; and Clade 3 contained two specimens, from Japan and Korea. Phylogenetic analysis comparing P. tridactyla with of a range of Podosphaera species suggests that P. tridactyla is paraphyletic.
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6′. Chasmothecia with (1-)2-6(-8) appendages; on
Prunus (s. lat.) spp.............................................. 7
7. Chasmothecial appendages rather long, up to
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................................................................Po. longiseta
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species of Prunus (Po. tridactyla s. lat.)......... 8
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