CBS-KNAW Fungal Biodiversity Centre, P.O. Box 85167, 3508 AD Utrecht, The Netherlands ; Wageningen University and Research Centre (WUR), Laboratory of Phytopathology, Droevendaalsesteeg 1, 6708 PB Wageningen, The Netherlands.
Stud Mycol. 2013 Jun 30;75(1):1-36. doi: 10.3114/sim0004.
The anamorphic genus Phoma was subdivided into nine sections based on morphological characters, and included teleomorphs in Didymella, Leptosphaeria, Pleospora and Mycosphaerella, suggesting the polyphyly of the genus. Recent molecular, phylogenetic studies led to the conclusion that Phoma should be restricted to Didymellaceae. The present study focuses on the taxonomy of excluded Phoma species, currently classified in Phoma sections Plenodomus, Heterospora and Pilosa. Species of Leptosphaeria and Phoma section Plenodomus are reclassified in Plenodomus, Subplenodomus gen. nov., Leptosphaeria and Paraleptosphaeria gen. nov., based on the phylogeny determined by analysis of sequence data of the large subunit 28S nrDNA (LSU) and Internal Transcribed Spacer regions 1 & 2 and 5.8S nrDNA (ITS). Phoma heteromorphospora, type species of Phoma section Heterospora, and its allied species Phoma dimorphospora, are transferred to the genus Heterospora stat. nov. The Phoma acuta complex (teleomorph Leptosphaeria doliolum), is revised based on a multilocus sequence analysis of the LSU, ITS, small subunit 18S nrDNA (SSU), β-tubulin (TUB), and chitin synthase 1 (CHS-1) regions. Species of Phoma section Pilosa and allied Ascochyta species were determined to belong to Pleosporaceae based on analysis of actin (ACT) sequence data. Anamorphs that are similar morphologically to Phoma and described in Ascochyta, Asteromella, Coniothyrium, Plectophomella, Pleurophoma and Pyrenochaeta are included in this study. Phoma-like species, which grouped outside the Pleosporineae based on a LSU sequence analysis, are transferred to the genera Aposphaeria, Paraconiothyrium and Westerdykella. The genera Medicopsis gen. nov. and Nigrograna gen. nov. are introduced to accommodate the medically important species formerly known as Pyrenochaeta romeroi and Pyrenochaeta mackinnonii, respectively.
New genera: Medicopsis Gruyter, Verkley & Crous, Nigrograna Gruyter, Verkley & Crous, Paraleptosphaeria Gruyter, Verkley & Crous, Subplenodomus Gruyter, Verkley & Crous. New species: Aposphaeria corallinolutea Gruyter, Aveskamp & Verkley, Paraconiothyrium maculicutis Verkley & Gruyter. New combinations: Coniothyrium carteri (Gruyter & Boerema) Verkley & Gruyter, C. dolichi (Mohanty) Verkley & Gruyter, C. glycines (R.B. Stewart) Verkley & Gruyter, C. multiporum (V.H. Pawar, P.N. Mathur & Thirum.) Verkley & Gruyter, C. telephii (Allesch.) Verkley & Gruyter, Heterospora (Boerema, Gruyter & Noordel.) Gruyter, Verkley & Crous, H. chenopodii (Westend.) Gruyter, Aveskamp & Verkley, H. dimorphospora (Speg.) Gruyter, Aveskamp & Verkley, Leptosphaeria errabunda (Desm.) Gruyter, Aveskamp & Verkley, L. etheridgei (L.J. Hutchison & Y. Hirats.) Gruyter, Aveskamp & Verkley, L. macrocapsa (Trail) Gruyter, Aveskamp & Verkley, L. pedicularis (Fuckel) Gruyter, Aveskamp & Verkley, L. rubefaciens (Togliani) Gruyter, Aveskamp & Verkley, L. sclerotioides (Sacc.) Gruyter, Aveskamp & Verkley, L. sydowii (Boerema, Kesteren & Loer.) Gruyter, Aveskamp & Verkley, L. veronicae (Hollós) Gruyter, Aveskamp & Verkley, Medicopsis romeroi (Borelli) Gruyter, Verkley & Crous, Nigrograna mackinnonii (Borelli) Gruyter, Verkley & Crous, Paraconiothyrium flavescens (Gruyter, Noordel. & Boerema) Verkley & Gruyter, Paracon. fuckelii (Sacc.) Verkley & Gruyter, Paracon. fusco-maculans (Sacc.) Verkley & Gruyter, Paracon. lini (Pass.) Verkley & Gruyter, Paracon. tiliae (F. Rudolphi) Verkley & Gruyter, Paraleptosphaeria dryadis (Johanson) Gruyter, Aveskamp & Verkley, Paralept. macrospora (Thüm.) Gruyter, Aveskamp & Verkley, Paralept. nitschkei (Rehm ex G. Winter) Gruyter, Aveskamp & Verkley, Paralept. orobanches (Schweinitz: Fr.) Gruyter, Aveskamp & Verkley, Paralept. praetermissa (P. Karst.) Gruyter, Aveskamp & Verkley, Plenodomus agnitus (Desm.) Gruyter, Aveskamp & Verkley, Plen. biglobosus (Shoemaker & H. Brun) Gruyter, Aveskamp & Verkley, Plen. chrysanthemi (Zachos, Constantinou & Panag.) Gruyter, Aveskamp & Verkley, Plen. collinsoniae (Dearn. & House) Gruyter, Aveskamp & Verkley, Plen. confertus (Niessl ex Sacc.) Gruyter, Aveskamp & Verkley, Plen. congestus (M.T. Lucas) Gruyter, Aveskamp & Verkley, Plen. enteroleucus (Sacc.) Gruyter, Aveskamp & Verkley, Plen. fallaciosus (Berl.) Gruyter, Aveskamp & Verkley, Plen. hendersoniae (Fuckel) Gruyter, Aveskamp & Verkley, Plen. influorescens (Boerema & Loer.) Gruyter, Aveskamp & Verkley, Plen. libanotidis (Fuckel) Gruyter, Aveskamp & Verkley, Plen. lindquistii (Frezzi) Gruyter, Aveskamp & Verkley, Plen. lupini (Ellis & Everh.) Gruyter, Aveskamp & Verkley, Plen. pimpinellae (Lowen & Sivan.) Gruyter, Aveskamp & Verkley, Plen. tracheiphilus (Petri) Gruyter, Aveskamp & Verkley, Plen. visci (Moesz) Gruyter, Aveskamp & Verkley, Pleospora fallens (Sacc.) Gruyter & Verkley, Pleo. flavigena (Constantinou & Aa) Gruyter & Verkley, Pleo. incompta (Sacc. & Martelli) Gruyter & Verkley, Pyrenochaetopsis pratorum (P.R. Johnst. & Boerema) Gruyter, Aveskamp & Verkley, Subplenodomus apiicola (Kleb.) Gruyter, Aveskamp & Verkley, Subplen. drobnjacensis (Bubák) Gruyter, Aveskamp & Verkley, Subplen. valerianae (Henn.) Gruyter, Aveskamp & Verkley, Subplen. violicola (P. Syd.) Gruyter, Aveskamp & Verkley, Westerdykella capitulum (V.H. Pawar, P.N. Mathur & Thirum.) de Gruyter, Aveskamp & Verkley, W. minutispora (P.N. Mathur ex Gruyter & Noordel.) Gruyter, Aveskamp & Verkley. New names: Pleospora angustis Gruyter & Verkley, Pleospora halimiones Gruyter & Verkley.
变形属 Phoma 根据形态特征分为九个部分,包括 Didymella、Leptosphaeria、 Pleospora 和 Mycosphaerella 中的有性型,表明该属的多系性。最近的分子、系统发育研究得出结论,Phoma 应仅限于 Didymellaceae。本研究重点关注排除的 Phoma 物种的分类学,目前这些物种被归类为 Phoma 节 Plenodomus、Heterospora 和 Pilosa。属于 Leptosphaeria 和 Phoma 节 Plenodomus 的物种被重新分类为 Plenodomus、Subplenodomus gen. nov.、Leptosphaeria 和 Paraleptosphaeria gen. nov.,这是基于大核糖体亚基 28S nrDNA(LSU)、内部转录间隔区 1 和 2 以及 5.8S nrDNA(ITS)序列数据分析得出的系统发育。Phoma heteromorphospora,Phoma 节 Heterospora 的模式种,及其相关种 Phoma dimorphospora,被转移到新的属 Heterospora stat. nov.。基于 LSU、ITS、小核糖体亚基 18S nrDNA(SSU)、β-微管蛋白(TUB)和几丁质合成酶 1(CHS-1)区域的多基因序列分析,对 Phoma acuta 复合种(有性型 Leptosphaeria doliolum)进行了修订。Phoma 节 Pilosa 和相关的 Ascochyta 种被确定为 Pleosporaceae,这是基于肌动蛋白(ACT)序列数据分析得出的。与 Phoma 和描述在 Ascochyta、Asteromella、Coniothyrium、Plectophomella、Pleurophoma 和 Pyrenochaeta 中相似形态的无性型也包括在本研究中。基于 LSU 序列分析,与 Pleosporineae 无关的 Phoma 样种被转移到新属 Aposphaeria、Paraconiothyrium 和 Westerdykella。新的属 Medicopsis Gruyter、Verkley 和 Crous 和 Nigrograna Gruyter、Verkley 和 Crous 被引入,以容纳以前称为 Pyrenochaeta romeroi 和 Pyrenochaeta mackinnonii 的医学上重要的物种。
新属:Medicopsis Gruyter、Verkley 和 Crous、Nigrograna Gruyter、Verkley 和 Crous、Paraleptosphaeria Gruyter、Verkley 和 Crous。新种:Aposphaeria corallinolutea Gruyter、Aveskamp 和 Verkley、Paraconiothyrium maculicutis Verkley 和 Gruyter。新组合:Coniothyrium carteri(Gruyter 和 Boerema)Verkley 和 Gruyter、C. dolichi(Mohanty)Verkley 和 Gruyter、C. glycines(R.B. Stewart)Verkley 和 Gruyter、C. multiporum(V.H. Pawar、P.N. Mathur 和 Thirum.)Verkley 和 Gruyter、C. telephii(Allesch.)Verkley 和 Gruyter、Heterospora(Boerema、Gruyter 和 Noordel.)Gruyter、Verkley 和 Crous、H. chenopodii(Westend.)Gruyter、Aveskamp 和 Verkley、H. dimorphospora(Speg.)Gruyter、Aveskamp 和 Verkley、Leptosphaeria errabunda(Desm.)Gruyter、Aveskamp 和 Verkley、L. etheridgei(L.J. Hutchison 和 Y. Hirats.)Gruyter、Aveskamp 和 Verkley、L. macrocapsa(Trail)Gruyter、Aveskamp 和 Verkley、L. pedicularis(Fuckel)Gruyter、Aveskamp 和 Verkley、L. rubefaciens(Togliani)Gruyter、Aveskamp 和 Verkley、L. sclerotioides(Sacc.)Gruyter、Aveskamp 和 Verkley、L. sydowii(Boerema、Kesteren 和 Loer.)Gruyter、Aveskamp 和 Verkley、L. veronicae(Hollós)Gruyter、Aveskamp 和 Verkley、Medicopsis romeroi(Borelli)Gruyter、Verkley 和 Crous、Nigrograna mackinnonii(Borelli)Gruyter、Verkley 和 Crous、Paraconiothyrium flavescens(Gruyter、Noordel. 和 Boerema)Verkley 和 Gruyter、Parac. fuckelii(Sacc.)Verkley 和 Gruyter、Parac. fusco-maculans(Sacc.)Verkley 和 Gruyter、Parac. lini(Pass.)Verkley 和 Gruyter、Parac. tiliae(F. Rudolphi)Verkley 和 Gruyter、Paraleptosphaeria dryadis(Johanson)Gruyter、Aveskamp 和 Verkley、Paralept. macrospora(Thüm.)Gruyter、Aveskamp 和 Verkley、Paralept. nitschkei(Rehm ex G. Winter)Gruyter、Aveskamp 和 Verkley、Paralept. orobanches(Schweinitz: Fr.)Gruyter、Aveskamp 和 Verkley、Paralept. praetermissa(P. Karst.)Gruyter、Aveskamp 和 Verkley、Plenodomus agnitus(Desm.)Gruyter、Aveskamp 和 Verkley、Plen. biglobosus(Shoemaker 和 H. Brun)Gruyter、Aveskamp 和 Verkley、Plen. chrysanthemi(Zachos、Constantinou 和 Panag.)Gruyter、Aveskamp 和 Verkley、Plen. collinsoniae(Dearn. 和 House)Gruyter、Aveskamp 和 Verkley、Plen. confertus(Niessl ex Sacc.)Gruyter、Aveskamp 和 Verkley、Plen. congestus(M.T. Lucas)Gruyter、Aveskamp 和 Verkley、Plen. enteroleucus(Sacc.)Gruyter、Aveskamp 和 Verkley、Plen. fallaciosus(Berl.)Gruyter、Aveskamp 和 Verkley、Plen. hendersoniae(Fuckel)Gruyter、Aveskamp 和 Verkley、Plen. influorescens(Boerema 和 Loer.)Gruyter、Aveskamp 和 Verkley、Plen. libanotidis(Fuckel)Gruyter、Aveskamp 和 Verkley、Plen. lindquistii(Frezzi)Gruyter、Aveskamp 和 Verkley、Plen. lupini(Ellis 和 Everh.)Gruyter、Aveskamp 和 Verkley、Plen. pimpinellae(Lowen 和 Sivan.)Gruyter、Aveskamp 和 Verkley、Plen. tracheiphilus(Petri)Gruyter、Aveskamp 和 Verkley、Plen. visci(Moesz)Gruyter、Aveskamp 和 Verkley、Pleospora fallens(Sacc.)Gruyter 和 Verkley、Pleo. flavigena(Constantinou 和 Aa)Gruyter 和 Verkley、Pleo. incompta(Sacc. 和 Martelli)Gruyter 和 Verkley、Pyrenochaetopsis pratorum(P.R. Johnst. 和 Boerema)Gruyter、Aveskamp 和 Verkley、Subplenodomus apiicola(Kleb.)Gruyter、Aveskamp 和 Verkley、Subplen. drobnjacensis(Bubák)Gruyter、Aveskamp 和 Verkley、Subplen. valerianae(Henn.)Gruyter、Aveskamp 和 Verkley、Subplen. violicola(P. Syd.)Gruyter、Aveskamp 和 Verkley、Westerdykella capitulum(V.H. Pawar、P.N. Mathur 和 Thirum.)de Gruyter、Aveskamp 和 Verkley、W. minutispora(P.N. Mathur ex Gruyter 和 Noordel.)Gruyter、Aveskamp 和 Verkley。新名:Pleospora angustis Gruyter 和 Verkley、Pleospora halimiones Gruyter 和 Verkley。