Journal Articles: 2022-present

2024
Bradshaw, M., et al., 2024. Phylogeny and taxonomy of the genera of Erysiphaceae, part 5: Erysiphe (the “Microsphaera lineage” part 1). Mycologia , 116 (1) , pp. 106-147. Publisher's VersionAbstract
In this contribution, we offer the fifth installment of a series focusing on the phylogeny and taxonomy of powdery mildews. This paper is the second segment evaluating the genus Erysiphe. The first treatment of Erysiphe focused on phylogenetically basal species in the “Uncinula lineage.” This research presents a phylogenetic-taxonomic assessment of species that form the group previously referred to as the “Microsphaera lineage.” Given the size of the group, we split the treatment of this lineage of Erysiphe species into two parts based on their phylogenetic placement. Phylogenetic trees based on ITS+28S data are supplemented by sequences of additional markers (CAM, GADPH, GS, RPB2, and TUB). Included in the analysis of the Microsphaera lineage is the “Erysiphe aquilegiae complex” (group, clade, cluster), which encompasses sequences obtained from an assemblage of Erysiphe species with insufficient resolution in rDNA analyses. Attempts have been made to resolve this group at the species level by applying a multilocus approach. A detailed discussion of the “Erysiphe aquilegiae complex” is provided. Sequences are provided for the first time for several species, particularly North American species, such as Erysiphe aggregata, E. erineophila, E. parnassiae, and E. semitosta. Ex-type sequences for Microsphaera benzoin and M. magnusii have been retrieved. Alphitomorpha penicillata, Microsphaera vanbruntiana, and M. symphoricarpi are epitypified with ex-epitype sequences. The new species Erysiphe alnicola, E. deutziana, E. cornigena, E. lentaginis, and E. sambucina are described, the new combinations E. lauracearum, E. passiflorae, and E. sambucicola are introduced, and the new name E. santali is proposed.
2023
Du, X.H., et al., 2023. Cytological studies reveal high variation in ascospore number and shape and conidia produced directly from ascospores in Morchella galilaea. Frontiers in Microbiology. Publisher's VersionAbstract
Spores are important as dispersal and survival propagules in fungi. In this study we investigated the variation in number, shape, size and germination mode of ascospores in Morchella galilaea, the only species of the genus Morchella known to fruit in the autumn. Based on the observation of five samples, we first discovered significant variation in the shape and size of ascospores in Morchella. One to sixteen ascospores were found in the asci. Ascospore size correlated negatively with ascospore number, but positively with ascus size, and ascus size was positively correlated with ascospore number. We noted that ascospores, both from fresh collections and dried specimens, germinated terminally or laterally either by extended germ tubes, or via the production of conidia that were formed directly from ascospores at one, two or multiple sites. The direct formation of conidia from ascospores takes place within asci or after ascospores are discharged. Using laser confocal microscopy, we recorded the number of nuclei in ascospores and in conidia produced from ascospores. In most ascospores of M. galilaea, several nuclei were observed, as is typical of species of Morchella. However, nuclear number varied from zero to around 20 in this species, and larger ascospores harbored more nuclei. One to six nuclei were present in the conidia. Nuclear migration from ascospores to conidia was observed. Conidia forming directly from ascospores has been observed in few species of Pezizomycetes; this is the first report of the phenomenon in Morchella species. Morphological and molecular data show that conidial formation from ascospores is not found in all the specimens of this species and, hence, is not an informative taxonomic character in M. galilaea. Our data suggest that conidia produced from ascospores and successive mitosis within the ascus may contribute to asci with more than eight spores. The absence of mitosis and/or nuclear degeneration, as well as cytokinesis defect, likely results in asci with fewer than eight ascospores. This study provides new insights into the poorly understood life cycle of Morchella species and more broadly improves knowledge of conidia formation and reproductive strategies in Pezizomycetes.
Bradshaw, M., et al., 2023. The Erysiphe alphitoides complex (powdery mildews) – unravelling the phylogeny and taxonomy of an intricate assemblage of species. New Zealand Journal of Botany . Publisher's VersionAbstract

Powdery mildews on oaks, caused by Erysiphe species, have serious ecological consequences on a range of Quercus hosts. In addition to Erysiphe quercicolaE. alphitoides is one of the most common and widespread species of Erysiphe having a wide host range among oak species, and a clear economic significance in applied ecology, forestry, and forest pathology. There are many publications addressing these important tree pathogens. Previous phylogenetic examinations have shown that E. alphitoides refers to a complicated species assemblage with insufficient taxonomic resolution in ITS + 28S analyses; the associated sequences form an insufficiently resolved species complex. The majority of species within the E. alphitoides complex cannot be unequivocally identified based solely on ITS + 28S analyses. Most of the additional species of the E. alphitoides complex are distributed in Asia, with a concentration in Japan. The question posed is whether there is a single widespread powdery mildew species, E. alphitoides, or an assemblage of closely allied species. To answer this question, specimens of related recognized species, particularly those from Japan, have been subjected to phylogenetic multilocus examinations, including CAMGAPDHGS, ITS + 28S, RPB2, and TUB sequences. An analysis of the concatenated sequences resulted in the confirmation of several distinct species. These species form highly supported clades that include E. alphitoidesE. aucubaeE. euonymicolaE. ipomoeaeE. menispermi var. dahuricaE. orixaeE. pseudoloniceraeE. sinomenii and E. wallrothiiErysiphe akebiae as well as the relationship between Japanese and North American collections requires further examinations.

Bradshaw, M., Braun, U. & Pfister, D.H., 2023. Phylogeny and taxonomy of the genera of Erysiphaceae, part 4: Erysiphe (the “Uncinula lineage”). Mycologia. Publisher's VersionAbstract
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.
Kuhar, F., et al., 2023. Paedomorphosis and Evolution of Sequestrate Basidiomycetes. In Evolution of Fungi and Fungal-Like Organisms . Springer International Publishing. Publisher's VersionAbstract
Current theories on the evolution of sequestrate (enclosed) basidiomes explain the origin of these forms in a gradualist adaptational process led by selective forces, such as drought and animal consumption. Paedomorphosis (the retention of juvenile traits) has been invoked as the phenomenon underlying sequestration, but many consequences of this process have not yet been explored. Our present interpretation of sequestrate morphologies, in light of Stephen Jay Gould’s characterization of neoteny (retention of juvenile features in an adult stage with mature reproductive structures) and progenesis (the onset of sexual maturity in a morphologically immature stage that does not reach the mature morphology observed in the ancestral form), both involved in paedomorphosis, implies that the origin of sequestrate basidiomes might constitute two distinct evolutionary processes. These two processes could be recognized among fungi by contrasting their morphological plasticity, phylogenetic diversification, and ecological patterns. The hypotheses discussed here provide new insights for interpreting and studying the evolution of sequestrate fungi.
Bradshaw, M., et al., 2023. Extensive intragenomic variation in the internal transcribed spacer (ITS) region of fungi. iScience , 26. Publisher's VersionAbstract
Fungi are among the most biodiverse organisms in the world. Accurate species identification is imperative for studies on fungal ecology and evolution. The internal transcribed spacer (ITS) rDNA region has been widely accepted as the universal barcode for fungi. However, several recent studies have uncovered intragenomic sequence variation within the ITS in multiple fungal species. Here, we mined the genome of 2414 fungal species to determine the prevalence of intragenomic variation and found that the genomes of 641 species, about one-quarter of the 2414 species examined, contained multiple ITS copies. Of those 641 species, 419 ( 65%) contained variation among copies revealing that intragenomic variation is common in fungi. We proceeded to show how these copies could result in the erroneous description of hundreds of fungal species and skew studies evaluating environmental DNA (eDNA) especially when making diversity estimates. Additionally, many genomes were found to be contaminated, especially those of unculturable fungi.
Bradshaw, M., et al., 2023. Phylogeny and taxonomy of the genera of Erysiphaceae, part 3: Cystotheca. Mycologia.Abstract
This contribution is part of a series devoted to the phylogeny and taxonomy of powdery mildews, with an emphasis on North American taxa. An overview of Cystotheca species is given, including references to ex-type sequences or, if unavailable, proposals for representative reference sequences for phylogenetic-taxonomic purposes. The new species C. mexicana is described, based on Mexican collections on Quercus glaucoides × Quercus microphylla and Quercus liebmannii × Q. microphylla. Cystotheca lanestris is reported for the first time worldwide on Quercus laceyi (Collected in Mexico) and on Q. toumeyi (collected in Arizona, USA). Cystotheca lanestris on Q. agrifolia and on Q. cerris is reported for the first time in Mexico. Epitypes with ex-epitype sequences are designated for Cystotheca wrightii, Lanomyces tjibodensis (= C. tjibodensis), Sphaerotheca kusanoi, and S. lanestris (C. lanestris).
Quijada, L., Baral, H.O. & Pfister, D.H., 2023. The Unexpected Identity of Tympanis vagabunda. Life , 13 (3). Publisher's VersionAbstract
Tympanis species (Leotiales) are plant pathogens distributed mostly in northern temperate ecosystems. The diversity and identity of some species remains unclear. Tympanis vagabunda, found in Sicilia (Italy) on dry twigs of RosaRubus, and Pistacia, is one example of an obscure and poorly known species. During the study of its type specimen in S, which contained one twig with a wood anatomy fitting neither of the three mentioned hosts, the microanatomic structures indicated that it belongs to the genus Rutstroemia (Helotiales). To investigate its identity, the types of R. fruticetiR. juniperiR. urceolus, and R. longiasca were studied for comparison. The species for which molecular data were available were included in a dataset that contained identified species of Rutstroemia, along with other select species from the families Rutstroemiaceae and Sclerotiniaceae. R. fruticeti, a saprobe frequently reported from Rubus fruticosus in Europe, is found to be a later synonym of T. vagabunda, and the combination Rutstroemia vagabunda is proposed. R. juniperi is an infrequently reported European species on twigs of Juniperus and is morphologically hard to distinguish from R. vagabunda; available molecular data support its recognition as a distinct species. R. longiasca differs from R. vagabunda in its black apothecia, smaller asci, and narrower ascospores. R. urceolus differs from R. vagabunda in having black apothecia and smaller inamyloid asci, and excipulum at the flanks and margin is composed of dark-walled hyphae.
Van Caenegem, W., et al., 2023. Hesperomyces (Fungi, Ascomycota) associated with Hyperaspis ladybirds (Coleoptera, Coccinellidae): Rethinking host specificity. Frontiers in Fungal Biology.Abstract
Laboulbeniales (Ascomycota, Laboulbeniomycetes) are biotrophic microfungi always attached to the exoskeleton of their arthropod hosts. They do not form hyphae or a mycelium; instead, they undergo determinate growth, developing from a two-celled ascospore to form a multicellular thallus. Hesperomyces virescens has been reported on over 30 species of ladybirds (Coleoptera, Coccinellidae); in reality, it represents a complex of species, presumably segregated by host genus association. In this study, we report on Hesperomyces thalli on Hyperaspis vinciguerrae from the Canary Islands and compare them with the Hesperomyces hyperaspidis described on Hyperaspis sp. from Trinidad. We generated the sequences of the internal transcribed spacer (ITS) region, the large subunit (LSU) nuclear ribosomal RNA gene, and the minichromosome maintenance complex component 7 (MCM7) protein-coding gene. Our phylogenetic reconstruction of Hesperomyces based on a concatenated ITS–LSU–MCM7 dataset revealed Hesperomyces sp. ex Hy. vinciguerrae as a member of the He. virescens species complex distinct from He. virescens sensu stricto (s.s.). It also revealed that the Hesperomyces sp. ex Chilocorus bipustulatus from Algeria is different from He. virescens s.s., which is associated with Chilocorus stigma from the USA. This suggests that the species of Hesperomyces are not solely segregated by host association, but that there is also a biogeographical component involved. Based on these data, we refrained from referring our material from Hy. vinciguerrae to He. hyperaspidis. Finally, we discuss the usefulness of MCM7 as a useful marker for species delimitation in Hesperomyces.
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2022
Quijada, L., et al., 2022. A review of Hyphodiscaceae. Studies in Mycology , 103 (1). Publisher's VersionAbstract
In a recently published classification scheme for Leotiomycetes, the new family Hyphodiscaceae was erected; unfortunately, this study was rife with phylogenetic misinterpretations and hampered by a poor understanding of this group of fungi. This manifested in the form of an undiagnostic familial description, an erroneous familial circumscription, and the redescription of the type species of an included genus as a new species in a different genus. The present work corrects these errors by incorporating new molecular data from this group into phylogenetic analyses and examining the morphological features of the included taxa. An emended description of Hyphodiscaceae is provided, notes and descriptions of the included genera are supplied, and keys to genera and species in Hyphodiscaceae are supplied. Microscypha cajaniensis is combined in Hyphodiscus, and Scolecolachnum nigricans is a taxonomic synonym of Fuscolachnum pteridis. Future work in this family should focus on increasing phylogenetic sampling outside of Eurasia and better characterising described species to help resolve outstanding issues.
Bradshaw, M.J., Braun, U. & Pfister, D.H., 2022. Phylogeny and taxonomy of the genera of Erysiphaceae, part 1: Golovinomyces. Mycologia , pp. 1-30. Publisher's VersionAbstract
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.
Bradshaw, M.J., et al., 2022. Phylogeny and taxonomy of the genera of Erysiphaceae, part 2: Neoerysiphe. Mycologia , pp. 1-14. Publisher's VersionAbstract
The second contribution to a new series devoted to the phylogeny and taxonomy of powdery mildews is presented. An overview of Neoerysiphe species is given, including references to ex-type sequences or, if unavailable, representative reference sequences for phylogenetic-taxonomic purposes are provided. The new species N. stachydis is described, and Striatoidium jaborosae is reduced to synonymy with Neoerysiphe macquii. Epitypes with ex-epitype sequences are designated for Alphitomorpha ballotae, A. labiatarum, Erysiphe galii, E. chelones, and E. galeopsidis. Based on phylogenetic analyses, it has been demonstrated that Neoerysiphe cumminsiana is confined to its type host, Roldana hartwegii (= Senecio seemannii), and other North and South American parasites on Asteraceae hosts, previously assigned to this species, pertain to N. macquii. The first record of N. macquii from Europe (Germany) on cultivated Bidens aurea was confirmed by sequencing. Sequence analysis of type material of N. rubiae reveals that this species should be excluded from Neoerysiphe; however, the true affinity of this taxon is not yet clear.
Bradshaw, M.J., et al., 2022. More Than Just Plants: Botanical Gardens Are an Untapped Source of Fungal Diversity . HortScience , 57 (10) , pp. 1289-1293. Publisher's VersionAbstract
Botanical gardens have extensive spatial databases of their plant specimens; however,
the fungi occurring in them are generally unstudied. Botanical gardens, with their great plant diversity, undoubtedly harbor a wide range of symbiotic fungi, including those that are plant-pathogenic. One such group of fungi is powdery mildews (Erysiphaceae). The powdery mildews are among the most prevalent and economically important plant pathogens in the world, with an estimated 906 species in 19 genera. They are known to infect more than 10,000 species of flowering plants and although some species occur across a range of hosts, many are associated with specific plants. Powdery mildews have undergone a long and dynamic coevolution with their host plants, resulting
in co-speciation. Botanical gardens provide a living laboratory in which to study these fungi, leading to a wealth of undiscovered fungal diversity. Furthermore, monitoring pathogens in botanical gardens has led to important ecological findings related to the plant sciences and plant protection. Between 2018 and 2022, a collaborative citizen science project was established with 10 botanical gardens in the United States and Mexico. A total of more than 300 powdery mildew specimens were collected on 220 different host taxa. We sequenced the entire internal transcribed spacer
(ITS) and large subunit (LSU) rDNA loci and phylogenetically and morphologically analyzed these collections revealing 130 species, of which 31 are likely unknown to science. This research highlights the importance of botanical gardens as a reservoir of fungal diversity. Future research will further elucidate the coevolutionary relationship between powdery mildews and their hosts and extend the current study to evaluate other plant pathogens and fungi in botanical gardens.
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Bradshaw, M., et al., 2022. Secondary DNA Barcodes (CAM, GAPDH, GS, and RpB2) to Characterize Species Complexes and Strengthen the Powdery Mildew Phylogeny. Frontiers in Ecology and Evolution. Publisher's VersionAbstract
Powdery mildews are a group of economically and ecologically important plant pathogens. In the past 25 years the use of ribosomal DNA (rDNA) in the powdery mildews has led to major taxonomic revisions. However, the broad scale use of rDNA has also revealed multiple species complexes that cannot be differentiated based on ITS + LSU data alone. Currently, there are only two powdery mildew taxonomic studies that took a multi-locus approach to resolve a species complex. In the present study, we introduce primers to sequence four additional regions (CAM, GAPDH, GS, and RPB2) that have the potential to improve support values in both broad and fine scale phylogenetic analyses. The primers were applied to a broad set of powdery mildew genera in China and the United States, and phylogenetic analyses included some of the common complexes. In taxa with nearly identical ITS sequences the analyses revealed a great amount of diversity. In total 154 non-rDNA sequences from 11 different powdery mildew genera were deposited in NCBI’s GenBank, laying the foundation for secondary barcode databases for powdery mildews. The combined and single loci phylogenetic trees constructed generally followed the previously defined species/genus concepts for the powdery mildews. Future research can use these primers to conduct in depth phylogenetic, and taxonomic studies to elucidate the evolutionary relationships of species and genera within the powdery mildews.
Mitchell, J.K., et al., 2022. Species of the common discomycete genus Bisporella reassigned to at least four genera. Mycologia. Publisher's VersionAbstract
Bisporella as typically conceived is a genus of noticeable, bright yellow inoperculate discomycetes. This interpretation of the genus, however, is at odds with Bisporella pallescens, the current name of the type species of the genus; furthermore, the genus has been interpreted as including the unusual species Bisporella resinicola. By comparing morphological and molecular traits of species traditionally included in Bisporella, we show that the genus is polyphyletic, with many “typical” members of the genus belonging instead in Calycina in Pezizellaceae. Bisporella pallescens is conclusively linked with its asexual morph, Bispora antennata, and the genus Bisporella is abandoned as a later synonym of the monotypic genus Bispora (previously applied only to asexual fungi) and placed as sister to Hymenoscyphus in Helotiaceae. Bisporella resinicola is shown to represent an independent monotypic genus, Eustilbum, which so far is placed incertae sedis in Helotiales. Finally, “Bisporella” subpallida, like Bispora, belongs to Helotiaceae but is instead related to “Phaeohelotium” epiphyllum.
Pfister, D.H., et al., 2022. A reexamination and realignment of Peziza sensu lato (Pezizomycetes) species in southern South America/Un reexamen y revisión de las especies de Peziza sensu lato (Pezizomycetes) en el cono sur de Sudamérica. Darwiniana , 10 (1) , pp. 148-177. Publisher's VersionAbstract

In this study we review recent collections and historical records of epigeous members of the Pezizales formerly placed in the large, heterogenous genus Peziza from temperate southern South America. Recent analyses using molecular phylogenetic methods allow placement of these species in several previously described genera in recognition of the heterogeneity of Peziza. We include species in nine genera, describe one new species (Peziza gamundiae sp. nov.), and propose one new combination (Phylloscypha nothofageti comb. nov.). We also demonstrate that Pustularia microspora is a synonym of the previously described taxon Peziza pseudosylvestris. Our purpose is to draw attention to these taxa in order to promote their collection and study in a modern framework.

En este trabajo hemos revisado material fresco y colecciones históricas de Pezizales epigeos formalmente incluidos en el amplio y heterogéneo género Peziza provenientes de las zonas templadas del sur de Sudamérica. Estudios recientes basados en filogenias moleculares han permitido posicionar estas especies en varios géneros previamente descritos demostrando su heterogeneidad. Incluimos aquí especies de nueve géneros, una especie nueva para la ciencia (Peziza gamundiae sp. nov.) y proponemos una nueva combinación (Phylloscypha nothofageti comb. nov.). También demostramos que Pustularia microspora es sinónimo de un taxón previamente descrito como Peziza pseudosylvestris. Nuestro objetivo es llamar la atención sobre la presencia de estos taxa para promover su recolección y estudio en trabajos científicos modernos.

Nokes, L.F., Haelewaters, D. & ., P.D.H., 2022. Exploration of Marine Lichenized Fungi as Bioindicators of Coastal Ocean Pollution in the Boston Harbor Islands National Recreation Area. Rhodora , 122 (992) , pp. 251–273. Publisher's VersionAbstract
This preliminary exploration of marine lichenized fungi (lichens) as bioindicators of water pollution examined the distribution of intertidal lichen communities in the Boston Harbor Islands National Recreation Area with respect to recorded pollution throughout the harbor. We found significant negative associations between pollution measurements and the health of the lichen community based on cover and species richness. We also observed significant differences in species composition between areas of higher pollution and areas of lower pollution, though not enough data are available to establish the pollution sensitivity or tolerance of individual species. We note that difficulties in the collection and identification of marine lichens hamper efforts to use them broadly as bioindicators. This study suggests that marine lichens could prove useful as bioindicators, but more research is needed to understand the differential effects of pollution on individual species as well as to establish practical procedures both for quantifying marine lichen community health and for widespread bioindication using marine lichens. Finally, one species collected during this study, Verrucaria ceuthocarpa, represents a first report for the Boston Harbor Islands National Recreation Area.
Bradshaw, M., Braun, U. & Pfister, D.H., 2022. Powdery mildews on Quercus: A worldwide distribution and rediscovered holotype provide insights into the spread of these ecologically important pathogens. Forest Pathology. Publisher's VersionAbstract
Powdery mildew, caused by Erysiphe spp., on oak has been shown to have serious ecological consequences on Quercus hosts. Erysiphe alphitoides and Equercicola 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 Ealphitoides was absent in European herbaria before 1921 and the discovery of the holotype of Ealphitoides from 1911 in an American herbarium (FH) led to the current study in which we genetically analysed six specimens of Ealphitoides s. lat including, most importantly, the holotype of Ealphitoides from France collected in 1911. The results of our analyses revealed that: (1) The sequence of the Ealphitoides holotype falls within the Equercicola clade, confirming that Ealphitoides did not spread to Europe until ~1921. (2) Ealphitoides var. chenii forms a monophyletic clade with Eepigena and should be reduced to synonymy with that species and (3) through sequence analyses Ealphitoides and Equercicola are confirmed to have spread to North America. The sequencing results of the Ealphitoides holotype have severe nomenclatural-taxonomic consequences. A proposal was submitted simultaneously with the present manuscript to conserve the name Ealphitoides so that the traditional usage of the names Ealphitoides and Equercicola 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.
Quijada, L., et al., 2022. Apothecial Ancestry, Evolution, and Re-Evolution in Thelebolales (Leotiomycetes, Fungi). Biology , 11 (583) , pp. 1-28. Publisher's VersionAbstract
Closed cleistothecia-like ascomata have repeatedly evolved in non-related perithecioid and apothecioid lineages of lichenized and non-lichenized Ascomycota. The evolution of a closed, darkly pigmented ascoma that protects asci and ascospores is conceived as either an adaptation to harsh environmental conditions or a specialized dispersal strategy. Species with closed ascomata have mostly lost sterile hymenial elements (paraphyses) and the capacity to actively discharge ascospores. The class Leotiomycetes, one of the most speciose classes of Ascomycota, is mainly apothecioid, paraphysate, and possesses active ascospore discharge. Lineages with closed ascomata, and their morphological variants, have evolved independently in several families, such as Erysiphaceae, Myxotrichaceae, Rutstroemiaceae, etc. Thelebolales is a distinctive order in the Leotiomycetes class. It has two widespread families (Thelebolaceae, Pseudeurotiaceae) with mostly closed ascomata, evanescent asci, and thus passively dispersed ascospores. Within the order, closed ascomata dominate and a great diversity of peridia have evolved as adaptations to different dispersal strategies. The type genus, Thelebolus, is an exceptional case of ascomatal evolution within the order. Its species are the most diverse in functional traits, encompassing species with closed ascomata and evanescent asci, and species with open ascomata, active ascospore discharge, and paraphyses. Open ascomata were previously suggested as the ancestral state in the genus, these ascomata depend on mammals and birds as dispersal agents. In this scheme, species with closed ascomata, a lack of paraphyses, and passive ascospore discharge exhibit derived traits that evolved in adaptation to cold ecosystems. Here, we used morphological and phylogenetic methods, as well as the reconstruction of ancestral traits for ascomatal type, asci dehiscence, the presence or absence of paraphyses, and ascospore features to explore evolution within Thelebolales. We demonstrate the apothecial ancestry in Thelebolales and propose a new hypothesis about the evolution of the open ascomata in Thelebolus, involving a process of re-evolution where the active dispersal of ascospores appears independently twice within the order. We propose a new family, Holwayaceae, within Thelebolales, that retains the phenotypic features exhibited by species of Thelebolus, i.e., pigmented capitate paraphyses and active asci discharge with an opening limitation ring
Braun, U., Bradshaw, M. & Pfister, D.H., 2022. (2863) Proposal to conserve the name Golovinomyces against Euoidium (Ascomycota: Erysiphaceae). Taxon , 71 (2) , pp. 459. Publisher's Version

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