Kasun M. Thambugala

Families of Dothideomycetes

Dothideomycetes comprise a highly diverse range of fungi characterized mainly by asci with two wall layers (bitunicate asci) and often with fissitunicate dehiscence. Many species are saprobes, with many asexual states comprising important plant pathogens. They are also endophytes, epiphytes, fungicolous, lichenized, or lichenicolous fungi. They occur in terrestrial, freshwater and marine habitats in almost every part of the world. We accept 105 families in Dothideomycetes with the new families Anteagloniaceae, Bambusicolaceae, Biatriosporaceae, Lichenoconiaceae, Muyocopronaceae, Paranectriellaceae, Roussoellaceae, Salsugineaceae, Seynesiopeltidaceae and Thyridariaceae introduced in this paper. Each family is provided with a description and notes, including asexual and asexual states, and if more than one genus is included, the type genus is also characterized. Each family is provided with at least one figure-plate, usually illustrating the type genus, a list of accepted genera, including asexual genera, and a key to these genera. A phylogenetic tree based on four gene combined analysis add support for 64 of the families and 22 orders, including the novel orders, Dyfrolomycetales, Lichenoconiales, Lichenotheliales, Monoblastiales, Natipusillales, Phaeotrichales and Strigulales. The paper is expected to provide a working document on Dothideomycetes which can be modified as new data comes to light. It is hoped that by illustrating types we provide stimulation and interest so that more work is carried out in this remarkable group of fungi.

The Faces of Fungi database: fungal names linked with morphology, phylogeny and human impacts

Taxonomic names are key links between various databases that store information on different organisms. Several global fungal nomenclural and taxonomic databases (notably Index Fungorum, Species Fungorum and MycoBank) can be sourced to find taxonomic details about fungi, while DNA sequence data can be sourced from NCBI, EBI and UNITE databases. Although the sequence data may be linked to a name, the quality of the metadata is variable and generally there is no corresponding link to images, descriptions or herbarium material. There is generally no way to establish the accuracy of the names in these genomic databases, other than whether the submission is from a reputable source. To tackle this problem, a new database (FacesofFungi), accessible at www.facesoffungi.org (FoF) has been established. This fungal database allows deposition of taxonomic data, phenotypic details and other useful data, which will enhance our current taxonomic understanding and ultimately enable mycologists to gain better and updated insights into the current fungal classification system. In addition, the database will also allow access to comprehensive metadata including descriptions of voucher and type specimens. This database is user-friendly, providing links and easy access between taxonomic ranks, with the classification system based primarily on molecular data (from the literature and via updated web-based phylogenetic trees), and to a lesser extent on morphological data when molecular data are unavailable. In FoF species are not only linked to the closest phylogenetic representatives, but also relevant data is provided, wherever available, on various applied aspects, such as ecological, industrial, quarantine and chemical uses. The data include the three main fungal groups (Ascomycota, Basidiomycota, Basal fungi) and fungus-like organisms. The FoF webpage is an output funded by the Mushroom Research Foundation which is an NGO with seven directors with mycological expertise. The webpage has 76 curators, and with the help of these specialists, FoF will provide an updated natural classification of the fungi, with illustrated accounts of species linked to molecular data. The present paper introduces the FoF database to the scientific community and briefly reviews some of the problems associated with classification and identification of the main fungal groups. The structure and use of the database is then explained. We would like to invite all mycologists to contribute to these web pages.

Naming and outline of Dothideomycetes-2014 including proposals for the protection or suppression of generic names

Article 59.1, of the International Code of Nomenclature for Algae, Fungi, and Plants (ICN; Melbourne Code), which addresses the nomenclature of pleomorphic fungi, became effective from 30 July 2011. Since that date, each fungal species can have one nomenclaturally correct name in a particular classification. All other previously used names for this species will be considered as synonyms. The older generic epithet takes priority over the younger name. Any widely used younger names proposed for use, must comply with Art. 57.2 and their usage should be approved by the Nomenclature Committee for Fungi (NCF). In this paper, we list all genera currently accepted by us in Dothideomycetes (belonging to 23 orders and 110 families), including pleomorphic and non-pleomorphic genera. In the case of pleomorphic genera, we follow the rulings of the current ICN and propose single generic names for future usage. The taxonomic placements of 1261 genera are listed as an outline. Protected names and suppressed names for 34 pleomorphic genera are listed separately. Notes and justifications are provided for possible proposed names after the list of genera. Notes are also provided on recent advances in our understanding of asexual and sexual morph linkages in Dothideomycetes. A phylogenetic tree based on four gene analyses supported 23 orders and 75 families, while 35 families still lack molecular data.

Fungal diversity notes 1–110: taxonomic and phylogenetic contributions to fungal species

This paper is a compilation of notes on 110 fungal taxa, including one new family, 10 new genera, and 76 new species, representing a wide taxonomic and geographic range. The new family, Paradictyoarthriniaceae is introduced based on its distinct lineage in Dothideomycetes and its unique morphology. The family is sister to Biatriosporaceae and Roussoellaceae. The new genera are Allophaeosphaeria (Phaeosphaeriaceae), Amphibambusa (Amphisphaeriaceae), Brunneomycosphaerella (Capnodiales genera incertae cedis), Chaetocapnodium (Capnodiaceae), Flammeascoma (Anteagloniaceae), Multiseptospora (Pleosporales genera incertae cedis), Neogaeumannomyces (Magnaporthaceae), Palmiascoma (Bambusicolaceae), Paralecia (Squamarinaceae) and Sarimanas (Melanommataceae). The newly described species are the Ascomycota Aliquandostipite manochii, Allophaeosphaeria dactylidis, A. muriformia, Alternaria cesenica, Amphibambusa bambusicola, Amphisphaeria sorbi, Annulohypoxylon thailandicum, Atrotorquata spartii, Brunneomycosphaerella laburni, Byssosphaeria musae, Camarosporium aborescentis, C. aureum, C. frutexensis, Chaetocapnodium siamensis, Chaetothyrium agathis, Colletotrichum sedi, Conicomyces pseudotransvaalensis, Cytospora berberidis, C. sibiraeae, Diaporthe thunbergiicola, Diatrype palmicola, Dictyosporium aquaticum, D. meiosporum, D. thailandicum, Didymella cirsii, Dinemasporium nelloi, Flammeascoma bambusae, Kalmusia italica, K. spartii, Keissleriella sparticola, Lauriomyces synnematicus, Leptosphaeria ebuli, Lophiostoma pseudodictyosporium, L. ravennicum, Lophiotrema eburnoides, Montagnula graminicola, Multiseptospora thailandica, Myrothecium macrosporum, Natantispora unipolaris, Neogaeumannomyces bambusicola, Neosetophoma clematidis, N. italica, Oxydothis atypica, Palmiascoma gregariascomum, Paraconiothyrium nelloi, P. thysanolaenae, Paradictyoarthrinium tectonicola, Paralecia pratorum, Paraphaeosphaeria spartii, Pestalotiopsis digitalis, P. dracontomelon, P. italiana, Phaeoisaria pseudoclematidis, Phragmocapnias philippinensis, Pseudocamarosporium cotinae, Pseudocercospora tamarindi, Pseudotrichia rubriostiolata, P. thailandica, Psiloglonium multiseptatum, Saagaromyces mangrovei, Sarimanas pseudofluviatile, S. shirakamiense, Tothia spartii, Trichomerium siamensis, Wojnowicia dactylidicola, W. dactylidis and W. lonicerae. The Basidiomycota Agaricus flavicentrus, A. hanthanaensis, A. parvibicolor, A. sodalis, Cantharellus luteostipitatus, Lactarius atrobrunneus, L. politus, Phylloporia dependens and Russula cortinarioides are also introduced. Epitypifications or reference specimens are designated for Hapalocystis berkeleyi, Meliola tamarindi, Pallidocercospora acaciigena, Phaeosphaeria musae, Plenodomus agnitus, Psiloglonium colihuae, P. sasicola and Zasmidium musae while notes and/or new sequence data are provided for Annulohypoxylon leptascum, A. nitens, A. stygium, Biscogniauxia marginata, Fasciatispora nypae, Hypoxylon fendleri, H. monticulosum, Leptosphaeria doliolum, Microsphaeropsis olivacea, Neomicrothyrium, Paraleptosphaeria nitschkei, Phoma medicaginis and Saccotheciaceae. A full description of each species is provided with light micrographs (or drawings). Molecular data is provided for 90 taxa and used to generate phylogenetic trees to establish a natural classification for species.

Epitypification and neotypification: guidelines with appropriate and inappropriate examples

A review of phylogenetic studies carried out together with morphological ones shows that a major problem with most early studies is that they concentrated on techniques and used material or strains of fungi that in most cases were not carefully reference, and in a worrying number of cases wrongly named. Most classical species, particularly of microfungi, are not represented by adequate type material, or other authoritatively identified cultures or specimens, that can serve as DNA sources for phylogenetic study, or for developing robust identification systems. Natural classifications of fungi therefore suffer from the lack of reference strains in resultant phylogenetic trees. In some cases, epitypification and neotypification can solve this problem and these tools are increasingly used to resolve taxonomic confusion and stabilize the understanding of species, genera, families, or orders of fungi. This manuscript discusses epitypification and neotypification, describes how to epitypify or neotypify species and examines the importance of this process. A set of guidelines for epitypification is presented. Examples where taxa have been epitypified are presented and the benefits and problems of epitypification are discussed. As examples of epitypification, or to provide reference specimens, a new epitype is designated for Paraphaeosphaeria michotii and reference specimens are provided for Astrosphaeriella stellata, A. bakeriana, Phaeosphaeria elongata, Ophiobolus cirsii, and O. erythrosporus. In this way we demonstrate how to epitypify taxa and its importance, and also illustrate the value of proposing reference specimens if epitypification is not advisable. Although we provided guidelines for epitypification, the decision to epitypify or not lies with the author, who should have experience of the fungus concerned. This responsibility is to be taken seriously, as once a later typification is made, it may not be possible to undo that, particularly in the case of epitypes, without using the lengthy and tedious formal conservation and rejection processes.

Revision of Phaeosphaeriaceae

Phaeosphaeriaceae is a large and important family in the order Pleosporales which includes economically important plant pathogens. Species may also be endophytes or saprobes on plant hosts, especially on monocotyledons (e.g., Cannaceae, Cyperaceae, Juncaceae, Poaceae); some species have also been reported on dicotyledons. The family previously accommodated 35 sexual and asexual genera and comprised more than 300 species with a range of morphological characters. The morphological characters of taxa in this family are often ambiguous and can be confused with other taxa in Leptosphaeriaceae and Montagnulaceae. Fourteen specimens of the type genera of Phaeosphaeriaceae were loaned from herbaria worldwide and were re-examined and illustrated. Fresh collections were obtained from Italy and Thailand, characterized, examined, isolated into pure culture and used to obtain molecular data. The asexual state was induced where possible on sterile bamboo pieces placed on water agar. Multigene phylogenetic analyses of ITS, LSU, SSU, RPB2 and TEF1 sequence datasets were carried out using maximum likelihood, maximum parsimony and Bayesian analysis. Molecular analyses shows that 21 genera (Amarenomyces, Ampelomyces, Chaetosphaeronema, Dematiopleospora, Entodesmium, Loratospora, Neosetophoma, Neostagonospora, Nodulosphaeria, Ophiobolus, Ophiosphaerella, Paraphoma, Parastagonospora, Phaeosphaeria, Phaeosphaeriopsis, Sclerostagonospora, Setomelanomma, Setophoma, Vrystaatia, Wojnowicia and Xenoseptoria) belong in Phaeosphaeriaceae, while seven genera (Amarenographium, Bricookea, Dothideopsella, Eudarluca, Phaeostagonospora, Scolecosporiella and Tiarospora) are included based on morphological data. Amarenomyces is reinstated and Nodulosphaeria is confirmed in Phaeosphaeriaceae. Eudarluca is distinguished from Sphaerellopsis based on its morphological characters and is typical of Phaeosphaeriaceae. ITS gene phylogenetic analysis indicates that Sphaerellopsis belongs to Leptosphaeriaceae. Ophiobolus species form a clade within Phaeosphaeriaceae while Ophiosphaerella is shown to be polyphyletic. Phaeosphaeria sensu stricto is redefined. Two new species of Phaeosphaeria and one of Phaeosphaeriopsis are introduced while the asexual states of Phaeosphaeria chiangraina and Phaeosphaeriopsis dracaenicola are reported. Scolicosporium minkeviciusii forms a sister clade with Neostagonospora and Parastagonospora in Phaeosphaeriaceae. However, Scolicosporium minkeviciusii is not the type species. Thus, the placement of Scolicosporium sensu stricto in Phaeosphaeriaceae is questionable. Phylogenetic analysis of combined ITS and LSU genes, confirm the placement of Septoriella oudemansii in Phaeosphaeriaceae. However, it is not represented by the generic type, thus the placement of Septoriella is questionable. Setophaeosphaeria is excluded from Phaeosphariaceae as the type species, Sp. hemerocallidis forms a clade at the base of Cucurbitariaceae. Wilmia clusters in Didymosphaeriaceae and is synonymized under Letendraea. Barria, Chaetoplea, Hadrospora, Lautitia, Metameris, Mixtura and Pleoseptum are excluded from Phaeosphaeriaceae based on their morphological characters. The asexual genera Mycopappus and Xenostigmina are excluded from this family based on the phylogenetic evidence; these genera form a clade close to Melanommataceae.

Fungal Planet description sheets: 281-319.

Novel species of fungi described in the present study include the following from South Africa: Alanphillipsia aloeicola from Aloe sp., Arxiella dolichandrae from Dolichandra unguiscati, Ganoderma austroafricanum from Jacaranda mimosifolia, Phacidiella podocarpi and Phaeosphaeria podocarpi from Podocarpus latifolius, Phyllosticta mimusopisicola from Mimusops zeyheri and Sphaerulina pelargonii from Pelargonium sp. Furthermore, Barssia maroccana is described from Cedrus atlantica (Morocco), Codinaea pini from Pinus patula (Uganda), Crucellisporiopsis marquesiae from Marquesia acuminata (Zambia), Dinemasporium ipomoeae from Ipomoea pes-caprae (Vietnam), Diaporthe phragmitis from Phragmites australis (China), Marasmius vladimirii from leaf litter (India), Melanconium hedericola from Hedera helix (Spain), Pluteus albotomentosus and Pluteus extremiorientalis from a mixed forest (Russia), Rachicladosporium eucalypti from Eucalyptus globulus (Ethiopia), Sistotrema epiphyllum from dead leaves of Fagus sylvatica in a forest (The Netherlands), Stagonospora chrysopyla from Scirpus microcarpus (USA) and Trichomerium dioscoreae from Dioscorea sp. (Japan). Novel species from Australia include: Corynespora endiandrae from Endiandra introrsa, Gonatophragmium triuniae from Triunia youngiana, Penicillium coccotrypicola from Archontophoenix cunninghamiana and Phytophthora moyootj from soil. Novelties from Iran include Neocamarosporium chichastianum from soil and Seimatosporium pistaciae from Pistacia vera. Xenosonderhenia eucalypti and Zasmidium eucalyptigenum are newly described from Eucalyptus urophylla in Indonesia. Diaporthe acaciarum and Roussoella acacia are newly described from Acacia tortilis in Tanzania. New species from Italy include Comoclathris spartii from Spartium junceum and Phoma tamaricicola from Tamarix gallica. Novel genera include (Ascomycetes): Acremoniopsis from forest soil and Collarina from water sediments (Spain), Phellinocrescentia from a Phellinus sp. (French Guiana), Neobambusicola from Strelitzia nicolai (South Africa), Neocladophialophora from Quercus robur (Germany), Neophysalospora from Corymbia henryi (Mozambique) and Xenophaeosphaeria from Grewia sp. (Tanzania). Morphological and culture characteristics along with ITS DNA barcodes are provided for all taxa.

A molecular phylogenetic reappraisal of the Didymosphaeriaceae (=Montagnulaceae).

The ascomycetous families, Didymosphaeriaceae and Montagnulaceae, have been treated in Pleosporales, Dothideomycetes, and both include saprobes, endophytes and pathogens associated with a wide variety of substrates worldwide. Didymosphaeriaceae was characterized by 1-septate ascospores and trabeculate pseudoparaphyses, mainly anastomosing above the asci, while species in Montagnulaceae had 1 to multi-septate ascospores and generally cellular pseudoparaphyses. In recent treatments, Bimuria, Didymocrea, Kalmusia, Karstenula, Montagnula, Paraphaeosphaeria, Paraconiothyrium and Letendraea were placed in Montagnulaceae, while only Didymosphaeria has been placed in Didymosphaeriaceae. New morphological and molecular data from recent collections have become available and thus the understanding of the families can be improved. Based on analyses of concatenated internal transcribed spacer (ITS) with LSU, SSU and β-tubulin gene sequences, the taxonomy of the genera classified in these families are re-assessed. Our phylogenetic analyses conclude that, the recently introduced Didymosphaeria rubi-ulmifolii with Paraconiothyrium brasiliense species complex, Alloconiothyrium, Bimuria, Deniquelata, Didymocrea, Kalmusia, Karstenula, Letendraea, Montagnula, Neokalmusia, Paraconiothyrium, Paraphaeosphaeria, Phaeodothis and Tremateia, forms a robust clade named here Didymosphaeriaceae. We therefore synonymize Montagnulaceae under Didymosphaeriaceae which is the oldest name and has priority and provide a update account of the family. Didymosphaeria is represented by Didymosphaeria rubi-ulmifolii. A new genus, Neokalmusia, is introduced in the family to accommodate the bambusicolous taxa Kalmusia brevispora and K. scabrispora. This introduction is based on molecular data coupled with morphology. One new species, Paraconiothyrium magnoliae and the sexual morph of Paraconiothyrium fuckelii, is also introduced. Julella and Barria are assigned to the family based on morphological similarity with Didymosphaeriaceae. Wilmia (previously referred to the Phaeosphaeriaceae) is synonymized under Letendraea (Didymosphaeriaceae). Furthermore, a new species, Letendraea cordylinicola, is introduced and placed in Didymosphaeriaceae based on phylogeny and morphology. The paraphyletic nature of Paraconiothyrium is partially resolved, but further sampling with fresh collections and molecular data of species in this genus are needed to obtain a natural classification.

Towards a natural classification and backbone tree for Pleosporaceae

The family Pleosporaceae includes numerous saprobic, opportunistic human, and plant pathogenic taxa. The classification of genera and species Pleosporaceae has been a major challenge due to the lack of a clear understanding of the importance of the morphological characters used to distinguish taxa as well as the lack of reference strains. Recent treatments concluded that Pleospora and some other genera in Pleosporaceae are likely polyphyletic. In order to establish the evolutionary relationships and to resolve the polyphyletic nature of Pleospora and allied genera, we sequenced the 18S nrDNA, 28S nrDNA, ITS, GAPDH, RPB2 and TEF1-alpha gene regions of Pleosporaceae species and phylogenetically analysed this data. Multigene phylogenies strongly support the monophyletic nature of Pleosporaceae among the other families in Pleosporales, and the acceptance of the genera Alternaria, Bipolaris, Clathrospora, Comoclathris, Curvularia, Dactuliophora, Decorospora, Diademosa, Exserohilum, Extrawettsteinina, Gibbago, Neocamarosporium, Paradendryphiella, Platysporoides, Pleospora, Porocercospora, Pseudoyuconia and Pyrenophora. Austropleospora, Dendryphion, Edenia and Macrospora are excluded from the family based on morphology coupled with molecular data. Two novel species, Alternaria murispora in this paper and Comoclathris sedi are introduced. The sexual morph of Alternaria alternata is re-described and illustrated using modern concepts from fresh collections. The paraphyletic nature of Pleospora is resolved based on the available morpho-molecular data, but further sampling with fresh collections, reference or ex-type strains and molecular data are needed to obtain a natural classification of genera and the family.

Towards a natural classification and backbone tree for Lophiostomataceae , Floricolaceae , and Amorosiaceae fam. nov .

The genera Lophiostoma, Misturatosphaeria and several other allied taxa in Lophiostomataceae are revisited. Accounts of these taxa, including their history, morphology, and family placement, based on molecular phylogeny, are provided. Type or representative specimens of Lophiostoma and Misturatosphaeria were examined and fresh specimens were obtained from Germany, Italy, Japan and Thailand. A multi-gene phylogenetic analysis of the lophiostomataceous genera Floricola, Lophiostoma, Misturatosphaeria and related taxa is provided. Sixteen genera including Lophiostoma, Lophiohelichrysum, Dimorphiopsis, Platystomum and Vaginatispora, plus eleven newly introduced genera Biappendiculispora, Alpestrisphaeria, Capulatispora, Coelodictyosporium, Guttulispora, Lophiopoacea, Neotrematosphaeria, Paucispora, Pseudolophiostoma, Pseudoplatystomum and Sigarispora are accepted in Lophiostomataceae based on morphology and phylogeny. Lophiostoma caulium, Lophiostoma arundinis and Lophiostoma caudatum are accommodated in Sigarispora. Lophiostoma winteri and Lophiostoma fuckelii are placed in the genera Lophiopoacea and Vaginatispora respectively. Three Curreya species and Misturatosphaeria claviformis are transferred to a new genus, Neocurreya. All other Misturatosphaeria species except Misturatosphaeria aurantiinotata and M. uniseptata are separated in the new genera Asymmetrispora, Aurantiascoma, Magnibotryascoma, Pseudoaurantiascoma and Pseudomisturatosphaeria based on their morphological and phylogenetic affinities. Another new genus, Ramusculicola is introduced for a new collection from Thailand. These seven new genera are accommodated in a new family Floricolaceae, together with Floricola and Misturatosphaeria. Several massarina-like species clustered as a sister clade to Amorosia littoralis and are accommodated in a new genus Angustimassarina. A new family Amorosiaceae is proposed to accommodate the genera Amorosia and Angustimassarina. The putatively named species Decaisnella formosa and Thyridaria macrostomoides form a separate clade together with a new genus Lignosphaeria which is placed in Dothideomycetes, genera incertae sedis.