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      Resolving the Phoma enigma

      research-article
      Author(s): 1 , 1 , 2 , 1 , ∗ , 3 , 4 , 5
      Publication date (Electronic):
      Journal: Studies in Mycology
      Publisher: CBS Fungal Biodiversity Centre
      Keywords: Ascochyta, Didymella, Multi-locus phylogeny, Phoma, Taxonomy, Microsphaeropsidaceae Q. Chen, L. Cai & Crous, Allophoma Q. Chen & L. Cai, Calophoma Q. Chen & L. Cai, Heterophoma Q. Chen & L. Cai, Neoascochyta Q. Chen & L. Cai, Neodidymelliopsis Q. Chen & L. Cai, Nothophoma Q. Chen & L. Cai, Paraboeremia Q. Chen & L. Cai, Phomatodes Q. Chen & L. Cai, Xenodidymella Q. Chen & L. Cai, Ascochytamedicaginicola var. medicaginicola Q. Chen & L. Cai, Didymellasenecionicola Q. Chen & L. Cai, Allophomanicaraguensis Q. Chen & L. Cai, Phomaneerlandica Q. Chen & L. Cai, Stagonosporopsishelianthi Q. Chen & L. Cai, Allophomalabilis (Sacc.) Q. Chen & L. Cai, All. minor (Aveskamp et al.) Q. Chen & L. Cai, All. piperis (Tassi) Q. Chen & L. Cai, All. tropica (R. Schneid. & Boerema) Q. Chen & L. Cai, All. zantedeschiae (Dippen.) Q. Chen & L. Cai, Ascochytaherbicola (Wehm.) Q. Chen & L. Cai, As. medicaginicola var. macrospora (Boerema et al.) Q. Chen & L. Cai, As. nigripycnidia (Boerema et al.) Q. Chen & L. Cai, As. phacae (Corbaz) Q. Chen & L. Cai, As. versabilis (Boerema et al.) Q. Chen & L. Cai, Boeremialilacis (Sacc.) Q. Chen & L. Cai, Calophomaaquilegiicola (M. Petrov) Q. Chen & L. Cai, Ca. clematidina (Thüm.) Q. Chen & L. Cai, Ca. clematidis-rectae (Petr.) Q. Chen & L. Cai, Ca. complanata (Tode) Q. Chen & L. Cai, Ca. glaucii (Brunaud) Q. Chen & L. Cai, Ca. vodakii (E. Müll.) Q. Chen & L. Cai, Didymellaacetosellae (A.L. Sm. & Ramsb.) Q. Chen & L. Cai, D. aliena (Fr.) Q. Chen & L. Cai, D. americana (Morgan-Jones & J.F. White) Q. Chen & L. Cai, D. anserina (Marchal) Q. Chen & L. Cai, D. aurea (Gruyter et al.) Q. Chen & L. Cai, D. bellidis (Neerg.) Q. Chen & L. Cai, D. boeremae (Gruyter) Q. Chen & L. Cai, D. calidophila (Aveskamp et al.) Q. Chen & L. Cai, D. chenopodii (P. Karst. & Har.) Q. Chen & L. Cai, D. coffeae-arabicae (Aveskamp et al.) Q. Chen & L. Cai, D. curtisii (Berk.) Q. Chen & L. Cai, D. dactylidis (Aveskamp et al.) Q. Chen & L. Cai, D. dimorpha (Aveskamp et al.) Q. Chen & L. Cai, D. eucalyptica (Sacc.) Q. Chen & L. Cai, D. gardeniae (S. Chandra & Tandon) Q. Chen & L. Cai, D. glomerata (Corda) Q. Chen & L. Cai, D. heteroderae (Boerema et al.) Q. Chen & L. Cai, D. longicolla (Aveskamp et al.) Q. Chen & L. Cai, D. mascrostoma (Mont.) Q. Chen & L. Cai, D. maydis (Arny & R.R. Nelson) Q. Chen & L. Cai, D. microchlamydospora (Aveskamp & Verkley) Q. Chen & L. Cai, D. molleriana (G. Winter) Q. Chen & L. Cai, D. musae (P. Joly) Q. Chen & L. Cai, D. negriana (Thüm.) Q. Chen & L. Cai, D. nigricans (P.R. Johnst. & Boerema) Q. Chen & L. Cai, D. pedeiae (Aveskamp et al.) Q. Chen & L. Cai, D. pinodella (L.K. Jones) Q. Chen & L. Cai, D. pomorum (Thüm.) Q. Chen & L. Cai, D. protuberans (Lév.) Q. Chen & L. Cai, D. rhei (Ellis & Everh.) Q. Chen & L. Cai, D. rumicicola (Boerema & Loer.) Q. Chen & L. Cai, D. sancta (Aveskamp et al.) Q. Chen & L. Cai, D. subglomerata (Boerema et al.) Q. Chen & L. Cai, D. subherbarum (Gruyter et al.) Q. Chen & L. Cai, D. viburnicola (Oudem.) Q. Chen & L. Cai, Epicoccumbrasiliense (Aveskamp et al.) Q. Chen & L. Cai, E. draconis (Berk. ex Cooke) Q. Chen & L. Cai, E. henningsii (Sacc.) Q. Chen & L. Cai, E. huancayense (Turkenst.) Q. Chen & L. Cai, E. plurivorum (P.R. Johnst.) Q. Chen & L. Cai, Heterophomaadonidis (Moesz) Q. Chen & L. Cai, H. nobilis (Kabát & Bubák) Q. Chen & L. Cai, H. novae-verbascicola (Aveskamp et al.) Q. Chen & L. Cai, H. poolensis (Taubenh.) Q. Chen & L. Cai, H. sylvatica (Sacc.) Q. Chen & L. Cai, Neoascochytadesmazieri (Cavara) Q. Chen & L. Cai, Neoa. europaea (Punith) Q. Chen & L. Cai, Neoa. exitialis (Morini) Q. Chen & L. Cai, Neoa. graminicola (Punith.) Q. Chen & L. Cai, Neoa. paspali (P.R. Johnst.) Q. Chen & L. Cai, Neodidymelliopsiscannabis (Aa & Boerema) Q. Chen & L. Cai, Neod. polemonii (Cooke) Q. Chen & L. Cai, Neod. xanthina (Sacc.) Q. Chen & L. Cai, Nothophomaanigozanthi (Tassi) Q. Chen & L. Cai, No. arachidis-hypogaeae (V.G. Rao) Q. Chen & L. Cai, No. gossypiicola (Gruyter) Q. Chen & L. Cai, No. infossa (Ellis & Everh.) Q. Chen & L. Cai, No. quercina (Syd.) Q. Chen & L. Cai, Paraboeremiaadianticola (Aa & Boerema) Q. Chen & L. Cai, Pa. putaminum (Speg.) Q. Chen & L. Cai, Pa. selaginellae (Sacc.) Q. Chen & L. Cai, Phomatodesaubrietiae (Moesz) Q. Chen & L. Cai, Phomat. nebulosa (Pers.) Q. Chen & L. Cai, Xenodidymellaapplanata (Niessl) Q. Chen & L. Cai, X. asphodeli ( E. Müll.) Q. Chen & L. Cai, X. catariae (Cooke & Ellis) Q. Chen & L. Cai, X. humicola (J.C. Gilman & E.V. Abbott) Q. Chen & L. Cai

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          Abstract

          The Didymellaceae was established in 2009 to accommodate Ascochyta, Didymella and Phoma, as well as several related phoma-like genera. The family contains numerous plant pathogenic, saprobic and endophytic species associated with a wide range of hosts. Ascochyta and Phoma are morphologically difficult to distinguish, and species from both genera have in the past been linked to Didymella sexual morphs. The aim of the present study was to clarify the generic delimitation in Didymellaceae by combing multi-locus phylogenetic analyses based on ITS, LSU, rpb2 and tub2, and morphological observations. The resulting phylogenetic tree revealed 17 well-supported monophyletic clades in Didymellaceae, leading to the introduction of nine genera, three species, two nomina nova and 84 combinations. Furthermore, 11 epitypes and seven neotypes were designated to help stabilise the taxonomy and use of names. As a result of these data, Ascochyta, Didymella and Phoma were delineated as three distinct genera, and the generic circumscriptions of Ascochyta, Didymella, Epicoccum and Phoma emended. Furthermore, the genus Microsphaeropsis, which is morphologically distinct from the members of Didymellaceae, grouped basal to the Didymellaceae, for which a new family Microsphaeropsidaceae was introduced.

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          Alternaria redefined

          J.H.C. Woudenberg, J.Z. Groenewald, M. Binder … (2013)
          Alternaria is a ubiquitous fungal genus that includes saprobic, endophytic and pathogenic species associated with a wide variety of substrates. In recent years, DNA-based studies revealed multiple non-monophyletic genera within the Alternaria complex, and Alternaria species clades that do not always correlate to species-groups based on morphological characteristics. The Alternaria complex currently comprises nine genera and eight Alternaria sections. The aim of this study was to delineate phylogenetic lineages within Alternaria and allied genera based on nucleotide sequence data of parts of the 18S nrDNA, 28S nrDNA, ITS, GAPDH, RPB2 and TEF1-alpha gene regions. Our data reveal a Pleospora/Stemphylium clade sister to Embellisia annulata, and a well-supported Alternaria clade. The Alternaria clade contains 24 internal clades and six monotypic lineages, the assemblage of which we recognise as Alternaria. This puts the genera Allewia, Brachycladium, Chalastospora, Chmelia, Crivellia, Embellisia, Lewia, Nimbya, Sinomyces, Teretispora, Ulocladium, Undifilum and Ybotromyces in synonymy with Alternaria. In this study, we treat the 24 internal clades in the Alternaria complex as sections, which is a continuation of a recent proposal for the taxonomic treatment of lineages in Alternaria. Embellisia annulata is synonymised with Dendryphiella salina, and together with Dendryphiella arenariae, are placed in the new genus Paradendryphiella. The sexual genera Clathrospora and Comoclathris, which were previously associated with Alternaria, cluster within the Pleosporaceae, outside Alternaria s. str., whereas Alternariaster, a genus formerly seen as part of Alternaria, clusters within the Leptosphaeriaceae. Paradendryphiella is newly described, the generic circumscription of Alternaria is emended, and 32 new combinations and 10 new names are proposed. A further 10 names are resurrected, while descriptions are provided for 16 new Alternaria sections. Taxonomic novelties: New combinations - Alternaria abundans (E.G. Simmons) Woudenb. & Crous, Alternaria alternariae (Cooke) Woudenb. & Crous, Alternaria atra (Preuss) Woudenb. & Crous, Alternaria bornmuelleri (Magnus) Woudenb. & Crous, Alternaria botrytis (Preuss) Woudenb. & Crous, Alternaria caespitosa (de Hoog & C. Rubio) Woudenb. & Crous, Alternaria cantlous (Yong Wang bis & X.G. Zhang) Woudenb. & Crous, Alternaria caricis (E.G. Simmons) Woudenb. & Crous, Alternaria cinerea (Baucom & Creamer) Woudenb. & Crous, Alternaria didymospora (Munt.-Cvetk.) Woudenb. & Crous, Alternaria fulva (Baucom & Creamer) Woudenb. & Crous, Alternaria hyacinthi (de Hoog & P.J. Mull. bis) Woudenb. & Crous, Alternaria indefessa (E.G. Simmons) Woudenberg & Crous, Alternaria leptinellae (E.G. Simmons & C.F. Hill) Woudenb. & Crous, Alternaria lolii (E.G. Simmons & C.F. Hill) Woudenb. & Crous, Alternaria multiformis (E.G. Simmons) Woudenb. & Crous, Alternaria obclavata (Crous & U. Braun) Woudenb. & Crous, Alternaria obovoidea (E.G. Simmons) Woudenb. & Crous, Alternaria oudemansii (E.G. Simmons) Woudenb. & Crous, Alternaria oxytropis (Q. Wang, Nagao & Kakish.) Woudenb. & Crous, Alternaria penicillata (Corda) Woudenb. & Crous, Alternaria planifunda (E.G. Simmons) Woudenb. & Crous, Alternaria proteae (E.G. Simmons) Woudenb. & Crous, Alternaria scirpinfestans (E.G. Simmons & D.A. Johnson) Woudenb. & Crous, Alternaria scirpivora (E.G. Simmons & D.A. Johnson) Woudenb. & Crous, Alternaria septospora (Preuss) Woudenb. & Crous, Alternaria slovaca (Svob.-Pol., L. Chmel & Bojan.) Woudenb. & Crous, Alternaria subcucurbitae (Yong Wang bis & X.G. Zhang) Woudenb. & Crous, Alternaria tellustris (E.G. Simmons) Woudenb. & Crous, Alternaria tumida (E.G. Simmons) Woudenb. & Crous, Paradendryphiella salina (G.K. Sutherl.) Woudenb. & Crous, Paradendryphiella arenariae (Nicot) Woudenb. & Crous. New names - Alternaria aspera Woudenb. & Crous, Alternaria botryospora Woudenb. & Crous, Alternaria brassicae-pekinensis Woudenb. & Crous, Alternaria breviramosa Woudenb. & Crous, Alternaria chlamydosporigena Woudenb. & Crous, Alternaria concatenata Woudenb. & Crous, Alternaria embellisia Woudenb. & Crous, Alternaria heterospora Woudenb. & Crous, Alternaria papavericola Woudenb. & Crous, Alternaria terricola Woudenb. & Crous. Resurrected names - Alternaria cetera E.G. Simmons, Alternaria chartarum Preuss, Alternaria consortialis (Thüm.) J.W. Groves & S. Hughes, Alternaria cucurbitae Letendre & Roum., Alternaria dennisii M.B. Ellis, Alternaria eureka E.G. Simmons, Alternaria gomphrenae Togashi, Alternaria malorum (Ruehle) U. Braun, Crous & Dugan, Alternaria phragmospora Emden, Alternaria scirpicola (Fuckel) Sivan. New sections, all in Alternaria - sect. Chalastospora Woudenb. & Crous, sect. Cheiranthus Woudenb. & Crous, sect. Crivellia Woudenb. & Crous, sect. Dianthicola Woudenb. & Crous, sect. Embellisia Woudenb. & Crous, sect. Embellisioides Woudenb. & Crous, sect. Eureka Woudenb. & Crous, sect. Infectoriae Woudenb. & Crous, sect. Japonicae Woudenb. & Crous, sect. Nimbya Woudenb. & Crous, sect. Phragmosporae Woudenb. & Crous, sect. Pseudoulocladium Woudenb. & Crous, sect. Teretispora Woudenb. & Crous, sect. Ulocladioides Woudenb. & Crous, sect. Ulocladium Woudenb. & Crous, sect. Undifilum Woudenb. & Crous. New genus - Paradendryphiella Woudenb. & Crous.
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            Highlights of the Didymellaceae: A polyphasic approach to characterise Phoma and related pleosporalean genera

            M.M. Aveskamp, J. de Gruyter, J.H.C. Woudenberg … (2010)
            Fungal taxonomists routinely encounter problems when dealing with asexual fungal species due to poly- and paraphyletic generic phylogenies, and unclear species boundaries. These problems are aptly illustrated in the genus Phoma. This phytopathologically significant fungal genus is currently subdivided into nine sections which are mainly based on a single or just a few morphological characters. However, this subdivision is ambiguous as several of the section-specific characters can occur within a single species. In addition, many teleomorph genera have been linked to Phoma, three of which are recognised here. In this study it is attempted to delineate generic boundaries, and to come to a generic circumscription which is more correct from an evolutionary point of view by means of multilocus sequence typing. Therefore, multiple analyses were conducted utilising sequences obtained from 28S nrDNA (Large Subunit - LSU), 18S nrDNA (Small Subunit - SSU), the Internal Transcribed Spacer regions 1 & 2 and 5.8S nrDNA (ITS), and part of the β-tubulin (TUB) gene region. A total of 324 strains were included in the analyses of which most belonged to Phoma taxa, whilst 54 to related pleosporalean fungi. In total, 206 taxa were investigated, of which 159 are known to have affinities to Phoma. The phylogenetic analysis revealed that the current Boeremaean subdivision is incorrect from an evolutionary point of view, revealing the genus to be highly polyphyletic. Phoma species are retrieved in six distinct clades within the Pleosporales, and appear to reside in different families. The majority of the species, however, including the generic type, clustered in a recently established family, Didymellaceae. In the second part of this study, the phylogenetic variation of the species and varieties in this clade was further assessed. Next to the genus Didymella, which is considered to be the sole teleomorph of Phoma s. str., we also retrieved taxa belonging to the teleomorph genera Leptosphaerulina and Macroventuria in this clade. Based on the sequence data obtained, the Didymellaceae segregate into at least 18 distinct clusters, of which many can be associated with several specific taxonomic characters. Four of these clusters were defined well enough by means of phylogeny and morphology, so that the associated taxa could be transferred to separate genera. Aditionally, this study addresses the taxonomic description of eight species and two varieties that are novel to science, and the recombination of 61 additional taxa.
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              A multi-gene phylogeny of Clavicipitaceae (Ascomycota, Fungi): identification of localized incongruence using a combinational bootstrap approach.

              Deuk Sung, Joseph Spatafora, Jace W. Jones … (2007)
              Multi-gene phylogenetic analyses were conducted to address the evolution of Clavicipitaceae (Ascomycota). Data are presented here for approximately 5900 base pairs from portions of seven loci: the nuclear ribosomal small and large subunit DNA (nrSSU and nrLSU), beta-tubulin, elongation factor 1alpha (EF-1alpha), the largest and second largest subunits of RNA polymerase II (RPB1 and RPB2), and mitochondrial ATP Synthase subunit 6 (mtATP6). These data were analyzed in a complete 66-taxon matrix and 91-taxon supermatrix that included some missing data. Separate phylogenetic analyses, with data partitioned according to genes, produced some conflicting results. The results of separate analyses from RPB1 and RPB2 are in agreement with the combined analyses that resolve a paraphyletic Clavicipitaceae comprising three well-supported clades (i.e., Clavicipitaceae clade A, B, and C), whereas the tree obtained from mtATP6 is in strong conflict with the monophyly of Clavicipitaceae clade B and the sister-group relationship of Hypocreaceae and Clavicipitaceae clade C. The distribution of relative contribution of nodal support for each gene partition was assessed using both partitioned Bremer support (PBS) values and combinational bootstrap (CB) analyses, the latter of which analyzed bootstrap proportions from all possible combinations of the seven gene partitions. These results suggest that CB analyses provide a more consistent estimate of nodal support than PBS and that combining heterogeneous gene partitions, which individually support a limited number of nodes, results in increased support for overall tree topology. Analyses of the 91-taxa supermatrix data sets revealed that some nodes were more strongly supported by increased taxon sampling. Identifying the localized incongruence of mtATP6 and analyses of complete and supermatrix data sets strengthen the evidence for rejecting the monophyly of Clavicipitaceae and much of the current subfamilial classification of the family. Although the monophyly of the grass-associated subfamily Clavicipitoideae (e.g., Claviceps, Balansia, and Epichloë) is strongly supported, the subfamily Cordycipitoideae (e.g., Cordyceps and Torrubiella) is not monophyletic. In particular, species of the genus Cordyceps, which are pathogens of arthropods and truffles, are found in all three clavicipitaceous clades. These results imply that most characters used in the current familial classification of Clavicipitaceae are not diagnostic of monophyly.
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                Author and article information

                Contributors
                L. Cai
                Journal
                Journal ID (nlm-ta): Stud Mycol
                Journal ID (iso-abbrev): Stud. Mycol
                Title: Studies in Mycology
                Publisher: CBS Fungal Biodiversity Centre
                ISSN (Print): 0166-0616
                ISSN (Electronic): 1872-9797
                Publication date PMC-release: 26 November 2015
                Publication date (Print): September 2015
                Publication date (Electronic): 26 November 2015
                Volume: 82
                Pages: 137-217
                Affiliations
                [1 ]State Key Laboratory of Mycology, Institute of Microbiology, Chinese Academy of Sciences, No. 1 West Beichen Rd, Chaoyang District, Beijing 100101, China
                [2 ]College of Agriculture and Biotechnology, China Agricultural University, No. 2 West Yuanmingyuan Rd, Haidian District, Beijing 100193, China
                [3 ]CBS-KNAW Fungal Biodiversity Centre, Uppsalalaan 8, 3584 CT Utrecht, The Netherlands
                [4 ]Department of Microbiology and Plant Pathology, Forestry and Agricultural Biotechnology Institute (FABI), University of Pretoria, Pretoria 0002, South Africa
                [5 ]Microbiology, Department of Biology, Utrecht University, Padualaan 8, 3584 CH Utrecht, The Netherlands
                Author notes
                [∗ ] Correspondence: L. Cai cail@ 123456im.ac.cn
                Article
                Publisher ID: S0166-0616(15)00017-2
                DOI: 10.1016/j.simyco.2015.10.003
                PMC ID: 4774273
                PubMed ID: 26955202
                SO-VID: 111f3d42-76cf-4322-b328-8c3329ea7272
                Copyright © Copyright © 2015, CBS-KNAW Fungal Biodiversity Centre. Production and hosting by ELSEVIER B.V.
                License:

                This is an open access article under the CC BY-NC-ND license (http://creativecommons.org/licenses/by-nc-nd/4.0/).

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                Subject: Research Paper

                ScienceOpen disciplines: Plant science & Botany
                Keywords: ascochyta,didymella,multi-locus phylogeny,phoma,taxonomy,microsphaeropsidaceae q. chen, l. cai & crous,allophoma q. chen & l. cai,calophoma q. chen & l. cai,heterophoma q. chen & l. cai,neoascochyta q. chen & l. cai,neodidymelliopsis q. chen & l. cai,nothophoma q. chen & l. cai,paraboeremia q. chen & l. cai,phomatodes q. chen & l. cai,xenodidymella q. chen & l. cai,ascochytamedicaginicola var. medicaginicola q. chen & l. cai,didymellasenecionicola q. chen & l. cai,allophomanicaraguensis q. chen & l. cai,phomaneerlandica q. chen & l. cai,stagonosporopsishelianthi q. chen & l. cai,allophomalabilis (sacc.) q. chen & l. cai,all. minor (aveskamp et al.) q. chen & l. cai,all. piperis (tassi) q. chen & l. cai,all. tropica (r. schneid. & boerema) q. chen & l. cai,all. zantedeschiae (dippen.) q. chen & l. cai,ascochytaherbicola (wehm.) q. chen & l. cai,as. medicaginicola var. macrospora (boerema et al.) q. chen & l. cai,as. nigripycnidia (boerema et al.) q. chen & l. cai,as. phacae (corbaz) q. chen & l. cai,as. versabilis (boerema et al.) q. chen & l. cai,boeremialilacis (sacc.) q. chen & l. cai,calophomaaquilegiicola (m. petrov) q. chen & l. cai,ca. clematidina (thüm.) q. chen & l. cai,ca. clematidis-rectae (petr.) q. chen & l. cai,ca. complanata (tode) q. chen & l. cai,ca. glaucii (brunaud) q. chen & l. cai,ca. vodakii (e. müll.) q. chen & l. cai,didymellaacetosellae (a.l. sm. & ramsb.) q. chen & l. cai,d. aliena (fr.) q. chen & l. cai,d. americana (morgan-jones & j.f. white) q. chen & l. cai,d. anserina (marchal) q. chen & l. cai,d. aurea (gruyter et al.) q. chen & l. cai,d. bellidis (neerg.) q. chen & l. cai,d. boeremae (gruyter) q. chen & l. cai,d. calidophila (aveskamp et al.) q. chen & l. cai,d. chenopodii (p. karst. & har.) q. chen & l. cai,d. coffeae-arabicae (aveskamp et al.) q. chen & l. cai,d. curtisii (berk.) q. chen & l. cai,d. dactylidis (aveskamp et al.) q. chen & l. cai,d. dimorpha (aveskamp et al.) q. chen & l. cai,d. eucalyptica (sacc.) q. chen & l. cai,d. gardeniae (s. chandra & tandon) q. chen & l. cai,d. glomerata (corda) q. chen & l. cai,d. heteroderae (boerema et al.) q. chen & l. cai,d. longicolla (aveskamp et al.) q. chen & l. cai,d. mascrostoma (mont.) q. chen & l. cai,d. maydis (arny & r.r. nelson) q. chen & l. cai,d. microchlamydospora (aveskamp & verkley) q. chen & l. cai,d. molleriana (g. winter) q. chen & l. cai,d. musae (p. joly) q. chen & l. cai,d. negriana (thüm.) q. chen & l. cai,d. nigricans (p.r. johnst. & boerema) q. chen & l. cai,d. pedeiae (aveskamp et al.) q. chen & l. cai,d. pinodella (l.k. jones) q. chen & l. cai,d. pomorum (thüm.) q. chen & l. cai,d. protuberans (lév.) q. chen & l. cai,d. rhei (ellis & everh.) q. chen & l. cai,d. rumicicola (boerema & loer.) q. chen & l. cai,d. sancta (aveskamp et al.) q. chen & l. cai,d. subglomerata (boerema et al.) q. chen & l. cai,d. subherbarum (gruyter et al.) q. chen & l. cai,d. viburnicola (oudem.) q. chen & l. cai,epicoccumbrasiliense (aveskamp et al.) q. chen & l. cai,e. draconis (berk. ex cooke) q. chen & l. cai,e. henningsii (sacc.) q. chen & l. cai,e. huancayense (turkenst.) q. chen & l. cai,e. plurivorum (p.r. johnst.) q. chen & l. cai,heterophomaadonidis (moesz) q. chen & l. cai,h. nobilis (kabát & bubák) q. chen & l. cai,h. novae-verbascicola (aveskamp et al.) q. chen & l. cai,h. poolensis (taubenh.) q. chen & l. cai,h. sylvatica (sacc.) q. chen & l. cai,neoascochytadesmazieri (cavara) q. chen & l. cai,neoa. europaea (punith) q. chen & l. cai,neoa. exitialis (morini) q. chen & l. cai,neoa. graminicola (punith.) q. chen & l. cai,neoa. paspali (p.r. johnst.) q. chen & l. cai,neodidymelliopsiscannabis (aa & boerema) q. chen & l. cai,neod. polemonii (cooke) q. chen & l. cai,neod. xanthina (sacc.) q. chen & l. cai,nothophomaanigozanthi (tassi) q. chen & l. cai,no. arachidis-hypogaeae (v.g. rao) q. chen & l. cai,no. gossypiicola (gruyter) q. chen & l. cai,no. infossa (ellis & everh.) q. chen & l. cai,no. quercina (syd.) q. chen & l. cai,paraboeremiaadianticola (aa & boerema) q. chen & l. cai,pa. putaminum (speg.) q. chen & l. cai,pa. selaginellae (sacc.) q. chen & l. cai,phomatodesaubrietiae (moesz) q. chen & l. cai,phomat. nebulosa (pers.) q. chen & l. cai,xenodidymellaapplanata (niessl) q. chen & l. cai,x. asphodeli ( e. müll.) q. chen & l. cai,x. catariae (cooke & ellis) q. chen & l. cai,x. humicola (j.c. gilman & e.v. abbott) q. chen & l. cai
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                ScienceOpen disciplines: Plant science & Botany
                Keywords: ascochyta, didymella, multi-locus phylogeny, phoma, taxonomy, microsphaeropsidaceae q. chen, l. cai & crous, allophoma q. chen & l. cai, calophoma q. chen & l. cai, heterophoma q. chen & l. cai, neoascochyta q. chen & l. cai, neodidymelliopsis q. chen & l. cai, nothophoma q. chen & l. cai, paraboeremia q. chen & l. cai, phomatodes q. chen & l. cai, xenodidymella q. chen & l. cai, ascochytamedicaginicola var. medicaginicola q. chen & l. cai, didymellasenecionicola q. chen & l. cai, allophomanicaraguensis q. chen & l. cai, phomaneerlandica q. chen & l. cai, stagonosporopsishelianthi q. chen & l. cai, allophomalabilis (sacc.) q. chen & l. cai, all. minor (aveskamp et al.) q. chen & l. cai, all. piperis (tassi) q. chen & l. cai, all. tropica (r. schneid. & boerema) q. chen & l. cai, all. zantedeschiae (dippen.) q. chen & l. cai, ascochytaherbicola (wehm.) q. chen & l. cai, as. medicaginicola var. macrospora (boerema et al.) q. chen & l. cai, as. nigripycnidia (boerema et al.) q. chen & l. cai, as. phacae (corbaz) q. chen & l. cai, as. versabilis (boerema et al.) q. chen & l. cai, boeremialilacis (sacc.) q. chen & l. cai, calophomaaquilegiicola (m. petrov) q. chen & l. cai, ca. clematidina (thüm.) q. chen & l. cai, ca. clematidis-rectae (petr.) q. chen & l. cai, ca. complanata (tode) q. chen & l. cai, ca. glaucii (brunaud) q. chen & l. cai, ca. vodakii (e. müll.) q. chen & l. cai, didymellaacetosellae (a.l. sm. & ramsb.) q. chen & l. cai, d. aliena (fr.) q. chen & l. cai, d. americana (morgan-jones & j.f. white) q. chen & l. cai, d. anserina (marchal) q. chen & l. cai, d. aurea (gruyter et al.) q. chen & l. cai, d. bellidis (neerg.) q. chen & l. cai, d. boeremae (gruyter) q. chen & l. cai, d. calidophila (aveskamp et al.) q. chen & l. cai, d. chenopodii (p. karst. & har.) q. chen & l. cai, d. coffeae-arabicae (aveskamp et al.) q. chen & l. cai, d. curtisii (berk.) q. chen & l. cai, d. dactylidis (aveskamp et al.) q. chen & l. cai, d. dimorpha (aveskamp et al.) q. chen & l. cai, d. eucalyptica (sacc.) q. chen & l. cai, d. gardeniae (s. chandra & tandon) q. chen & l. cai, d. glomerata (corda) q. chen & l. cai, d. heteroderae (boerema et al.) q. chen & l. cai, d. longicolla (aveskamp et al.) q. chen & l. cai, d. mascrostoma (mont.) q. chen & l. cai, d. maydis (arny & r.r. nelson) q. chen & l. cai, d. microchlamydospora (aveskamp & verkley) q. chen & l. cai, d. molleriana (g. winter) q. chen & l. cai, d. musae (p. joly) q. chen & l. cai, d. negriana (thüm.) q. chen & l. cai, d. nigricans (p.r. johnst. & boerema) q. chen & l. cai, d. pedeiae (aveskamp et al.) q. chen & l. cai, d. pinodella (l.k. jones) q. chen & l. cai, d. pomorum (thüm.) q. chen & l. cai, d. protuberans (lév.) q. chen & l. cai, d. rhei (ellis & everh.) q. chen & l. cai, d. rumicicola (boerema & loer.) q. chen & l. cai, d. sancta (aveskamp et al.) q. chen & l. cai, d. subglomerata (boerema et al.) q. chen & l. cai, d. subherbarum (gruyter et al.) q. chen & l. cai, d. viburnicola (oudem.) q. chen & l. cai, epicoccumbrasiliense (aveskamp et al.) q. chen & l. cai, e. draconis (berk. ex cooke) q. chen & l. cai, e. henningsii (sacc.) q. chen & l. cai, e. huancayense (turkenst.) q. chen & l. cai, e. plurivorum (p.r. johnst.) q. chen & l. cai, heterophomaadonidis (moesz) q. chen & l. cai, h. nobilis (kabát & bubák) q. chen & l. cai, h. novae-verbascicola (aveskamp et al.) q. chen & l. cai, h. poolensis (taubenh.) q. chen & l. cai, h. sylvatica (sacc.) q. chen & l. cai, neoascochytadesmazieri (cavara) q. chen & l. cai, neoa. europaea (punith) q. chen & l. cai, neoa. exitialis (morini) q. chen & l. cai, neoa. graminicola (punith.) q. chen & l. cai, neoa. paspali (p.r. johnst.) q. chen & l. cai, neodidymelliopsiscannabis (aa & boerema) q. chen & l. cai, neod. polemonii (cooke) q. chen & l. cai, neod. xanthina (sacc.) q. chen & l. cai, nothophomaanigozanthi (tassi) q. chen & l. cai, no. arachidis-hypogaeae (v.g. rao) q. chen & l. cai, no. gossypiicola (gruyter) q. chen & l. cai, no. infossa (ellis & everh.) q. chen & l. cai, no. quercina (syd.) q. chen & l. cai, paraboeremiaadianticola (aa & boerema) q. chen & l. cai, pa. putaminum (speg.) q. chen & l. cai, pa. selaginellae (sacc.) q. chen & l. cai, phomatodesaubrietiae (moesz) q. chen & l. cai, phomat. nebulosa (pers.) q. chen & l. cai, xenodidymellaapplanata (niessl) q. chen & l. cai, x. asphodeli ( e. müll.) q. chen & l. cai, x. catariae (cooke & ellis) q. chen & l. cai, x. humicola (j.c. gilman & e.v. abbott) q. chen & l. cai

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