Trichothecenes are terpenoid toxins produced by species in 10 fungal genera, including species of Trichoderma. The trichothecene biosynthetic gene (tri) cluster typically includes the tri5 gene, which encodes a terpene synthase that catalyzes formation of trichodiene, the parent compound of all trichothecenes. The two Trichoderma species, Trichoderma arundinaceum and T. brevicompactum, that have been examined are unique in that tri5 is located outside the tri cluster in a genomic region that does not include other known tri genes. In the current study, analysis of 35 species representing a wide range of the phylogenetic diversity of Trichoderma revealed that 22 species had tri5, but only 13 species had both tri5 and the tri cluster. tri5 was not located in the cluster in any species. Using complementation analysis of a T. arundinaceum tri5 deletion mutant, we demonstrated that some tri5 homologs from species that lack a tri cluster are functional, but others are not. Phylogenetic analyses suggest that Trichoderma tri5 was under positive selection following its divergence from homologs in other fungi but before Trichoderma species began diverging from one another. We propose two models to explain these diverse observations. One model proposes that the location of tri5 outside the tri cluster resulted from loss of tri5 from the cluster in an ancestral species followed by reacquisition via horizontal transfer. The other model proposes that in species that have a functional tri5 but lack the tri cluster, trichodiene production provides a competitive advantage.
Based on molecular phylogenetic analyses of a multigene matrix of partial nuSSU-ITS-LSU rDNA, RPB1, RPB2 and TEF1 sequences and by morphological evidence, the genus Mycosphaerangium is shown to be the closest relative of Neomelanconium, and confirmed to be a member of the Cenangiaceae (Leotiomycetes). While Mycosphaerangium and Neomelanconium share many traits like similar conidia, conidiogenesis, asci and ascospores, their apothecia differ particularly in excipular features and are therefore recognized as distinct genera. Mycosphaerangium tiliae, described from North America, is excluded from the genus but shown to represent the sexual morph of the European Neomelanconium gelatosporum, and it is therefore synonymized with the latter. Based on morphology, Neomelanconium deightonii is assumed to be congeneric with Neomelanconium gelatosporum, and it is lectotypified. Dermatea tetraspora and Phaeangium magnisporum, the basionyms of Mycosphaerangium tetrasporum and M. magnisporum, respectively, are lectotypified as well, and for M. tetrasporum, the asexual morph is recorded for the first time. Mycosphaerangium quercinum sp. nov. is described as a new species from various Quercus hosts in Europe, where it is shown to be widely distributed. It morphologically and ecologically closely resembles the North American M. tetrasporum, but differs in paraphysis and ascospore morphology and by croziers at its ascus base. The three accepted species of Mycosphaerangium and the two of Neomelanconium are described and illustrated. Mycosphaerangium magnisporum, M. quercinum and M. tetrasporum are recorded to be constantly associated with species of Coryneum, indicating a fungicolous habit, but no evidence for fungal associations has been found in Neomelanconium deightonii and N. gelatosporum.
The genus Melanconis (Melanconidaceae, Diaporthales) in the strict sense is here re-evaluated regarding phylogenetic structure, taxonomy, distribution and ecology. Using a matrix of sequences from ITS, LSU, ms204, rpb2, tef1 and tub2, eight species are recognised and their phylogenetic positions are determined. Based on phylogenetic, morphological and geographical differentiation, Melanconis marginalis is subdivided into four subspecies. Melanconis italica is reduced to a subspecies of Melanconis marginalis. The two species Melanconis larissae from Betula sp. and M. pacifica from Alnus rubra are described as new. Melanconis alni and M. stilbostoma are lectotypified and M. alni, M. marginalis and M. stilbostoma are epitypified. All GenBank sequences deposited as Melanconis alni are shown to actually represent M. marginalis and those as M. marginalis belong to the newly described M. pacifica. Currently, Alnus and Betula are the sole host genera of Melanconis. All species and subspecies are (re-)described and illustrated. In addition, the neotypification of Melanconium pterocaryae is here validated.
European species of the genus Dendrostoma (Erythrogloeaceae, Diaporthales) occurring on Castanea sativa and Quercus spp. based on freshly collected material are presented. Using a matrix of sequences from ITS, LSU, rpb2, and tef1, five species are recognized, and their phylogenetic positions are determined. Four species are added to the 14 described species of Dendrostoma. Dendrostoma atlanticum on Castanea sativa, D. creticum on Quercus coccifera and D. istriacum on Q. ilex are described as new species, Valsa castanea is combined in Dendrostoma, and D. leiphaemia is redescribed and illustrated. A key to the European species of Dendrostoma is provided.
The new species Stilbocrea walteri is described and illustrated from Quercus ilex collected in Portugal. Phylogenetic analyses of LSU rDNA, rpb1, rpb2 and tef1 sequence matrices place S. walteri in the Bionectriaceae, Hypocreales, within a clade of specimens morphologically identified as Stilbocrea macrostoma, the generic type of Stilbocrea. Stilbocrea walteri differs from S. macrostoma in dark olive green to blackish ascomata basally immersed in a stroma, KOH+ and LA+ ascomata and the lack of a stilbella-like asexual morph on natural substrate and pure culture. A simple phialidic asexual morph is formed in pure culture. To enable a morphological comparison, Stilbocrea macrostoma is illustrated.
Based on molecular phylogenetic analyses of a multigene matrix of partial nuSSU-ITS-LSU rDNA, cal, his, ms204, rpb1, rpb2, tef1 and tub2 sequences, recent European and Iranian collections of Melanconium pterocaryae from the type host, Pterocarya fraxinifolia, are shown to be distinct from the Japanese Melanconis pterocaryae from Pterocarya rhoifolia, and both are confirmed as closely related members of the recently described genus Juglanconis. Therefore, the new name Juglanconis japonica is proposed for Melanconis pterocaryae. As no type collection could be traced, Melanconium pterocaryae (syn. J. pterocaryae) is neotypified, described and illustrated, and it is recorded for Europe for the first time. During field surveys in natural stands of P. fraxinifolia in Guilan province (Iran), Juglanconis pterocaryae was consistently isolated from tissues affected by branch and trunk cankers, twig dieback and wood necrosis, indicating that it is the causal agent of these diseases. The external and internal symptoms associated with these trunk diseases are described and illustrated.
Phylogenetic analyses of a combined DNA data matrix containing ITS, LSU, rpb2 and tub2 sequences of representative Xylariales revealed that the genus Barrmaelia is a well-defined monophylum, as based on four of its described species (B. macrospora, B. moravica, B. oxyacanthae, B. rhamnicola) and the new species B. rappazii. The generic type of Entosordaria, E. perfidiosa, is revealed as the closest relative of Barrmaelia, being phylogenetically distant from the generic type of Clypeosphaeria, C. mamillana, which belongs to Xylariaceae sensu stricto. Entosordaria and Barrmaelia are highly supported and form a distinct lineage, which is recognised as the new family Barrmaeliaceae. The new species E. quercina is described. Barrmaelia macrospora, B. moravica and B. rhamnicola are epitypified and E. perfidiosa is lecto- and epitypified. Published sequences of Anthostomella and several Anthostomella-like species from the genera Alloanthostomella, Anthostomelloides, Neoanthostomella, Pseudoanthostomella and Pyriformiascoma are evaluated, demonstrating the necessity of critical inspection of published sequence data before inclusion in phylogenies. Verified isolates of several species from these genera should be re-sequenced to affirm their phylogenetic affinities. In addition, the generic type of Anthostomella should be sequenced before additional generic re-arrangements are proposed.
Two new species of Pleosporales, Anteaglonium rubescens (Anteagloniaceae) and Atrocalyx asturiensis (Lophiotremataceae), are described. Phylogenetic placement was determined by combined analyses of a DNA data matrix containing ITS, LSU, SSU, rpb2, and tef1. Anteaglonium rubescens is a stromatic fungus characterized by brown didymospores disarticulating within asci, and by the production of a red-orange to pink pigment produced in nature and in artificial culture. Atrocalyx asturiensis has massive ascomatal crests and brown phragmospores.
Based on phylogenetic analyses of an ITS-LSU-SSU-rpb2-tef1 sequence data matrix three taxa once classified in Cucurbitaria are referred to the Melanommataceae. Cucurbitaria rhododendri, also known as Melanomma rhododendri, is not congeneric with the generic type of Melanomma, M. pulvis-pyrius and thus classified in the new genus Alpinaria. Cucurbitaria piceae, known as Gemmamyces piceae, the cause of the Gemmamyces bud blight of Picea spp., belongs also to the Melanommataceae. The name Gemmamyces is conserved. For Cucurbitaria obducens, also known as Teichospora obducens, the new genus Praetumpfia is described, as it cannot be accommodated in any known genus. All species are redescribed and epitypified. Based on sequence data and morphology, Blastostroma, Mycodidymella and Xenostigmina are synonyms of Petrakia. The genus Petrakia is emended. We also provide sequences of additional markers for Beverwykella pulmonaria, Melanomma pulvis-pyrius, Petrakia echinata and Pseudotrichia mutabilis.
Stigmatodiscus pruni (Stigmatodiscaceae, Stigmatodiscales) is described and illustrated from corticated dead twigs of Prunus spinosa collected in Austria and France. It is characterised by hysteriform ascomata with two lateral black lips, which are erumpent through the periderm of the host, and a black disc in combination with two-celled, asymmetric brown verruculose ascospores with a distinct sheath. Phylogenetic analyses of a multigene matrix containing a representative selection of Dothideomycetes from four genes (nuc 18S rDNA, nuc 28S rDNA, rpb2 and tef1) revealed a highly supported placement within Stigmatodiscales as sister species to Stigmatodiscus enigmaticus. Micromorphology of the sexual and asexual morph matches the genus Stigmatodiscus, except for the hysteriform shape of the ascomata and the two-celled ascospores.
During a survey on corticolous Dothideomycetes, several collections with ascospores matching the genera Asteromassaria and Stigmatomassaria (Pleomassariaceae, Pleosporales) were revealed from dead corticated twigs of Acer, Carpinus and Tamarix. Closer morphological examination showed that their ascomata were apothecial, with a hamathecium consisting of septate, branched paraphyses, which are apically swollen at maturity. Several collections were cultured and sequenced, and a Blast search of their nuc 28S rDNA sequences revealed dothideomycetous affiliation, but without a close match to a specific family or order. Phylogenetic analyses of a multigene matrix containing a representative selection of Dothideomycetes from four genes (nuc 18S rDNA, nuc 28S rDNA, rpb2 and tef1) revealed placement within Dothideomycetes but without a supported familial or ordinal affiliation. Based on the phylogenetic analyses and morphological investigations, the new genera Asterodiscus and Stigmatodiscus, with the two new species A. tamaricis and S. enigmaticus, are described and illustrated, and placed in the new family Stigmatodiscaceae and new order Stigmatodiscales.
With the advance to one scientific name for each fungal species, the generic names in the class Sordariomycetes typified by sexual and asexual morphs are evaluated based on their type species to determine if they compete with each other for use or protection. Recommendations are made for which of the competing generic names should be used based on criteria such as priority, number of potential names changes, and frequency of use. Some recommendations for well-known genera include Arthrinium over Apiospora, Colletotrichum over Glomerella, Menispora over Zignoëlla, Microdochium over Monographella, Nigrospora over Khuskia, and Plectosphaerella over Plectosporium. All competing generic names are listed in a table of recommended names along with the required action. If priority is not accorded to sexually typified generic names after 2017, only four names would require formal protection: Chaetosphaerella over Oedemium, Diatrype over Libertella, Microdochium over Monographella, and Phaeoacremonium over Romellia and Togninia. Concerning species in the recommended genera, one replacement name (Xylaria benjaminii nom. nov.) is introduced, and the following new combinations are made: Arthrinium sinense, Chloridium caesium, C. chloroconium, C. gonytrichii, Corollospora marina, C. parvula, C. ramulosa, Juncigena fruticosae, Melanospora simplex, Seimatosporium massarina, Sporoschisma daemonoropis, S. taitense, Torpedospora mangrovei, Xylaria penicilliopsis, and X. termiticola combs. nov.
A multigene analysis of a combined ITS, LSU, SSU, rpb2 and tef1 sequence data matrix was applied to infer the phylogenetic position of the genus Teichospora in the Pleosporales, based on isolates from freshly collected material of the generic type T. trabicola and several additional species. Phylogenetic analyses revealed that Misturatosphaeria and Floricola are synonyms of Teichospora. All species of these genera and several species recently described in the genus Curreya belong to Teichospora and are thus combined in this genus. Also, Melanomma radicans and Ramusculicola thailandica are combined in Teichospora. The new name Teichospora parva is established for Misturatosphaeria minima. Three new species, T. melanommoides, T. pusilla and T. rubriostiolata, are described, and an expanded description of T. mariae is given. The family Teichosporaceae is currently confined to Teichospora, which can be phylogenetically clearly separated from Lophiostoma, the type genus of the Lophiostomataceae. The family name Floricolaceae is a synonym of Teichosporaceae. All species described here form apically free paraphyses among immature asci. This finding contradicts the current general dogma that apically free paraphyses are absent in the Pleosporales and questions the wide use of the term pseudoparaphysis.
The genus Allantonectria is synonymised with Thyronectria, based on morphological and molecular phylogenetic considerations. Investigations of types and fresh collections revealed that Tubercularia concentrica is an earlier name for Sphaeria (syn. Allantonectria) miltina and is thus combined in Thyronectria. Allantonectria yuccae is recognised as a distinct species and transferred to Thyronectria, as well as the recently described A. zangii. Descriptions and illustrations are provided for T. concentrica and T. yuccae. A recent collection of the North American Thyronectroidea chrysogramma, the generic type of Thyronectroidea, was studied with respect to morphology of its sexual morph in fresh condition and of its asexual morph produced in pure culture. Molecular phylogenies based on six loci (ITS and LSU regions of nuc rDNA, act1, rpb1, rpb2, tef1 and tub2 genes) place T. chrysogramma within Thyronectria, confirming synonymy of Thyronectroidea with Thyronectria, but remarkably a relationship to European species with green to brown spores (former genus Mattirolia) receives no support, and its closest relatives remain unclear. Thyronectria caraganae is described as a new species from herbarium specimens of Caragana arborescens collected in the Ukraine. It is characterised by morphology of the sexual morph and by DNA sequence data, which place it within the T. austroamericana - T. rhodochlora clade with high support. This is also supported by its morphology, specifically ascomata partly embedded within a stroma, muriform ascospores not budding within the ascus, becoming yellowish to rosy at maturity.
The Calosphaeriales is revisited with new collection data, living cultures, morphological studies of ascoma centrum, secondary structures of the internal transcribed spacer (ITS) rDNA and phylogeny based on novel DNA sequences of five nuclear ribosomal and protein-coding loci. Morphological features, molecular evidence and information from predicted RNA secondary structures of ITS converged upon robust phylogenies of the Calosphaeriales and Togniniales. The current concept of the Calosphaeriales includes the Calosphaeriaceae and Pleurostomataceae encompassing five monophyletic genera, Calosphaeria, Flabellascus gen. nov., Jattaea, Pleurostoma and Togniniella, strongly supported by Bayesian and Maximum Likelihood methods. The structural elements of ITS1 form characteristic patterns that are phylogenetically conserved, corroborate observations based on morphology and have a high predictive value at the generic level. Three major clades containing 44 species of Phaeoacremonium were recovered in the closely related Togniniales based on ITS, actin and ß-tubulin sequences. They are newly characterized by sexual and RNA structural characters and ecology. This approach is a first step towards understanding of the molecular systematics of Phaeoacremonium and possibly its new classification. In the Calosphaeriales, Jattaea aphanospora sp. nov. and J. ribicola sp. nov. are introduced, Calosphaeria taediosa is combined in Jattaea and epitypified. The sexual morph of Phaeoacremonium cinereum was encountered for the first time on decaying wood and obtained in vitro. In order to achieve a single nomenclature, the genera of asexual morphs linked with the Calosphaeriales are transferred to synonymy of their sexual morphs following the principle of priority, i.e. Calosphaeriophora to Calosphaeria, Phaeocrella to Togniniella and Pleurostomophora to Pleurostoma. Three new combinations are proposed, i.e. Pleurostoma ochraceum comb. nov., P. repens comb. nov. and P. richardsiae comb. nov. The morphology-based key is provided to facilitate identification of genera accepted in the Calosphaeriales.
Ascomycota/classification , DNA, Ribosomal Spacer/chemistry , Genes, Fungal/genetics , Phylogeny , RNA, Fungal/genetics , RNA, Ribosomal, 16S/genetics , Ascomycota/genetics , Bayes Theorem , DNA, Ribosomal Spacer/genetics , Evolution, Molecular , Likelihood Functions , Nucleic Acid Conformation
Two new species of Thyronectria growing in Mediterranean vegetation are described from southern Spain; they are T. giennensis from Quercus ilex ssp. rotundifolia and T. pistaciae from Pistacia lentiscus. Both species are characterized by morphology of sexual and asexual morphs and by DNA data. They have olivaceous to green-brown muriform ascospores and are closely related to T. asturiensis and T. roseovirens, as determined by multigene phylogenetic analyses of a matrix containing six loci (ITS and 28S regions of nuc rDNA, ACT1, RPB1, RPB2, TEF1 and TUB2 genes). We also report that Cucurbitaria bicolor is a synonym of Thyronectria rhodochlora, the type species of Thyronectria.
Ascomycota/classification , Ascomycota/isolation & purification , Phylogeny , Ascomycota/genetics , Ascomycota/growth & development , Fungal Proteins/genetics , Mediterranean Region , Molecular Sequence Data , Peptide Elongation Factor 1/genetics , RNA Polymerase II/genetics , Spores, Fungal/classification , Spores, Fungal/genetics , Spores, Fungal/growth & development , Spores, Fungal/isolation & purification , Tubulin/genetics
The present paper represents the second contribution in the Genera of Fungi series, linking type species of fungal genera to their morphology and DNA sequence data, and where possible, ecology. This paper focuses on 12 genera of microfungi, 11 of which the type species are neo- or epitypified here: Allantophomopsis (A. cytisporea, Phacidiaceae, Phacidiales, Leotiomycetes), Latorua gen. nov. (Latorua caligans, Latoruaceae, Pleosporales, Dothideomycetes), Macrodiplodiopsis (M. desmazieri, Macrodiplodiopsidaceae, Pleosporales, Dothideomycetes), Macrohilum (M. eucalypti, Macrohilaceae, Diaporthales, Sordariomycetes), Milospium (M. graphideorum, incertae sedis, Pezizomycotina), Protostegia (P. eucleae, Mycosphaerellaceae, Capnodiales, Dothideomycetes), Pyricularia (P. grisea, Pyriculariaceae, Magnaporthales, Sordariomycetes), Robillarda (R. sessilis, Robillardaceae, Xylariales, Sordariomycetes), Rutola (R. graminis, incertae sedis, Pleosporales, Dothideomycetes), Septoriella (S. phragmitis, Phaeosphaeriaceae, Pleosporales, Dothideomycetes), Torula (T. herbarum, Torulaceae, Pleosporales, Dothideomycetes) and Wojnowicia (syn. of Septoriella, S. hirta, Phaeosphaeriaceae, Pleosporales, Dothideomycetes). Novel species include Latorua grootfonteinensis, Robillarda africana, R. roystoneae, R. terrae, Torula ficus, T. hollandica, and T. masonii spp. nov., and three new families: Macrodiplodiopsisceae, Macrohilaceae, and Robillardaceae. Authors interested in contributing accounts of individual genera to larger multi-authored papers to be published in IMA Fungus, should contact the associate editors listed for the major groups of fungi on the List of Protected Generic Names for Fungi (www.generaoffungi.org).
In advancing to one name for fungi, this paper treats generic names competing for use in the order Diaporthales (Ascomycota, Sordariomycetes) and makes a recommendation for the use or protection of one generic name among synonymous names that may be either sexually or asexually typified. A table is presented that summarizes these recommendations. Among the genera most commonly encountered in this order, Cytospora is recommended over Valsa and Diaporthe over Phomopsis. New combinations are introduced for the oldest epithet of important species in the recommended genus. These include Amphiporthe tiliae, Coryneum lanciforme, Cytospora brevispora, C. ceratosperma, C. cinereostroma, C. eugeniae, C. fallax, C. myrtagena, Diaporthe amaranthophila, D. annonacearum, D. bougainvilleicola, D. caricae-papayae, D. cocoina, D. cucurbitae, D. juniperivora, D. leptostromiformis, D. pterophila, D. theae, D. vitimegaspora, Mastigosporella georgiana, Pilidiella angustispora, P. calamicola, P. pseudogranati, P. stromatica, and P. terminaliae.
The discovery of a second species of Bicornispora in Spain, B. seditiosa, which is closely related to B. exophiala but has smaller ascospores, narrower asci and different ecology, gave us the opportunity to culture and sequence the fungus. Phylogenetic analyses of rDNA regions including partial nuc 28S rDNA (28S) and ITS1-5.8S-ITS2 (ITS) supported a close relationship with species of the genus Lambertella (Rutstroemiaceae), placing Bicornispora, previously ascribed to Coryneliales, within Helotiales. This result confirmed an evolutionary linkage between certain inoperculate discomycetes such as Lambertella palmeri and derived cleistothecial forms (Bicornispora spp.). Based on analyses of morphological study and molecular phylogenetic analyses, a new combination Rutstroemia asphodeli is proposed for Ciboria asphodeli.
Ascomycota/classification , Ascomycota/isolation & purification , Ascomycota/genetics , Ascomycota/growth & development , DNA, Fungal/genetics , DNA, Ribosomal/genetics , Molecular Sequence Data , Phylogeny , Plants/microbiology , Spain , Spores, Fungal/classification , Spores, Fungal/genetics , Spores, Fungal/growth & development , Spores, Fungal/isolation & purification
Trichoderma harzianum is known as a cosmopolitan, ubiquitous species associated with a wide variety of substrates. It is possibly the most commonly used name in agricultural applications involving Trichoderma, including biological control of plant diseases. While various studies have suggested that T. harzianum is a species complex, only a few cryptic species are named. In the present study the taxonomy of the T. harzianum species complex is revised to include at least 14 species. Previously named species included in the complex are T. guizhouense, T. harzianum, and T. inhamatum. Two new combinations are proposed, T. lentiforme and T. lixii. Nine species are described as new, T. afarasin, T. afroharzianum, T. atrobrunneum, T. camerunense, T. endophyticum, T. neotropicale, T. pyramidale, T. rifaii and T. simmonsii. We isolated Trichoderma cultures from four commercial biocontrol products reported to contain T. harzianum. None of the biocontrol strains were identified as T. harzianum s. str. In addition, the widely applied culture 'T. harzianum T22' was determined to be T. afroharzianum. Some species in the T. harzianum complex appear to be exclusively endophytic, while others were only isolated from soil. Sexual states are rare. Descriptions and illustrations are provided. A secondary barcode, nuc translation elongation factor 1-α (TEF1) is needed to identify species in this complex.
Agricultural Inoculants/classification , Trichoderma/classification , Agricultural Inoculants/genetics , Agricultural Inoculants/growth & development , Agricultural Inoculants/isolation & purification , DNA, Fungal/genetics , DNA, Ribosomal/genetics , Fungal Proteins/genetics , Molecular Sequence Data , Peptide Elongation Factor 1/genetics , Pest Control, Biological/economics , Phylogeny , Soil Microbiology , Spores, Fungal/classification , Spores, Fungal/genetics , Spores, Fungal/growth & development , Spores, Fungal/isolation & purification , Trichoderma/genetics , Trichoderma/growth & development , Trichoderma/isolation & purification
