redisposition of Phoma-like anamorphs in Pleosporales - CBS - KNAW

available online at www.studiesinmycology.org 

Studies in Mycology 75: 1–36. 

Redisposition of Phoma-like anamorphs in Pleosporales 

J. de Gruyter 1–3* , J.H.C. Woudenberg 1 , M.M. Aveskamp 1 , G.J.M. Verkley 1 , J.Z. Groenewald 1 and P.W. Crous 1,3,4 

1 

CBS-KNAW Fungal Biodiversity Centre, P.O. Box 85167, 3508 AD Utrecht, The Netherlands; 2 National Reference Centre, National Plant Protection Organization, P.O. Box 9102, 

6700 HC Wageningen, The Netherlands; 3 Wageningen University and Research Centre (WUR), Laboratory of Phytopathology, Droevendaalsesteeg 1, 6708 PB Wageningen, 

The Netherlands; 4 Microbiology, Department of Biology, Utrecht University, Padualaan 8, 3584 CH Utrecht, The Netherlands 

*Correspondence: Hans de Gruyter, j.de.gruyter@minlnv.nl 

Abstract: 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. 

Studies in Mycology 

Key words: coelomycetes, Coniothyriaceae, Cucurbitariaceae, Leptosphaeriaceae, Melanommataceae, molecular phylogeny, Montagnulaceae, Phaeosphaeriaceae, 

Pleosporaceae, Sporormiaceae, taxonomy, Trematosphaeriaceae. 

Taxonomic novelties: 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. 

Published online: 15 May 2012; doi:10.3114/sim0004. 

Introduction 

The anamorphic genus Phoma includes many important plant 

pathogens. The taxonomy of Phoma has been studied intensively 

in the Netherlands for more than 40 years resulting in the 

development of a generic concept as an outline for identification 

of Phoma species (Boerema 1997). In this concept species of 

the genus Phoma are classified based on their morphological 

Copyright CBS-KNAW Fungal Biodiversity Centre, P.O. Box 85167, 3508 AD Utrecht, The Netherlands. 

characters into nine sections: Phoma, Heterospora, Macrospora, 

Paraphoma, Peyronellaea, Phyllostictoides, Pilosa, Plenodomus 

and Sclerophomella (Boerema 1997). The species placed in each 

of the sections were systematically described culminating in the 

publication of the “Phoma Identification Manual” (Boerema et al. 

2004), which contained the descriptions of 223 specific and infraspecific 

taxa of Phoma, and more than 1000 synonyms in other 

coelomycetous genera. The classification of the Phoma species in 

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1

De Gruyter et al. 

sections based on morphology is artificial (Boerema et al. 2004), 

and several species can be classified in more than one section as 

they reveal multiple “section-specific” characters. 

A large, well-studied Phoma culture collection that includes 

more than 1100 strains of Phoma resulted from the extensive 

morphological studies conducted on Phoma in The Netherlands. 

That culture collection is the basis of an intensive molecular 

phylogenetic study of the genus Phoma, which commenced in 

2006. Molecular studies of species of Phoma prior to the onset of 

this project concentrated on the development of molecular detection 

methods for specific, important plant pathogenic Phoma species, 

such as Ph. macdonaldii, Ph. tracheiphila, Stagonosporopsis 

cucurbitacearum (as Ph. cucurbitacearum) and Boeremia foveata 

(as Ph. foveata) (Aveskamp et al. 2008). The phylogeny of the 

type species of the nine Phoma sections and morphologically 

similar coelomycetes was determined utilising the sequence data 

of the large subunit 28S nrDNA (LSU) and the small subunit 18S 

nrDNA (SSU) regions (de Gruyter et al. 2009). Results of that study 

demonstrated that the type species of the nine Phoma sections 

all grouped in Pleosporales. The type species of five Phoma 

sections, Phoma, Phyllostictoides, Sclerophomella, Macrospora 

and Peyronellaea and similar genera, grouped in a distinct clade 

in Didymellaceae. The type species of the remaining four Phoma 

sections, Heterospora, Paraphoma, Pilosa and Plenodomus, 

clustered in several clades outside Didymellaceae based on the 

LSU and SSU sequence analysis leading to the conclusion that 

these species should be excluded from Phoma (de Gruyter et al. 

2009, Aveskamp et al. 2010). 

The molecular phylogeny of the Phoma species in Didymellaceae 

was determined in a subsequent study (Aveskamp et al. 2010) 

and, as the phylogenetic placement of the sectional type species 

already suggested, included species mainly from sections Phoma, 

Phyllostictoides, Sclerophomella, Macrospora and Peyronellaea. 

The molecular phylogeny of 11 Phoma species classified in Phoma 

section Paraphoma based on their setose pycnidia was investigated 

using LSU and SSU sequences (de Gruyter et al. 2010) and this 

section was highly polyphyletic, with species clustering mainly in 

Phaeosphaeriaceae and Cucurbitariaceae. 

The purpose of the present study was to clarify the molecular 

phylogeny of the Phoma species currently classified in sections 

Plenodomus and Pilosa, along with Phoma species which were 

determined to be distantly related to the generic type species 

Ph. herbarum in previous molecular studies. Additionally, Phomalike 

isolates of coelomycetes currently classified in Ascochyta 

and Coniothyrium and clustering outside the Didymellaceae (de 

Gruyter et al. 2009, Aveskamp et al. 2010) are included in this study 

along with a number of Phoma-like species that do not belong to 

Pleosporineae. 

In the present study, the initial focus was to determine the 

molecular phylogeny of Phoma betae (teleom. Pleospora betae) 

and Ph. lingam (teleom. Leptosphaeria maculans), type species of 

the Phoma sections Pilosa and Plenodomus, respectively, at the 

generic rank based on the sequence data of the LSU and the SSU 

regions. In a subsequent study, the sequence data of both the LSU 

and the ITS regions were used for a revised classification of the 

Phoma species currently classified in Phoma section Plenodomus. 

Only a limited number of the species currently classified in this 

section have a confirmed Leptosphaeria teleomorph. 

The Phoma acuta species complex was subject of a more 

detailed study. The teleomorph of Ph. acuta is Leptosphaeria 

doliolum, type species of the genus Leptosphaeria. A multilocus 

analysis of sequence data of the SSU, LSU, ITS, β-tubulin (TUB), 

and chitin synthase 1 (CHS-1) regions was performed. The 

phylogeny of Phoma species of section Pilosa, with a Pleospora 

teleomorph (Pleosporaceae) was studied utilising actin (ACT) 

sequence data. 

Phoma-like species currently attributed to the genera 

Aposphaeria, Asteromella, Coniothyrium, Phoma, Plenodomus, 

Pleurophoma and Pyrenochaeta, which could not be classified in the 

Pleosporineae based on their molecular phylogeny, were included 

in a LSU sequence analysis. All Phoma taxa that are unrelated to 

Didymellaceae and treated in this paper are redisposed to other 

genera. 

A further aim of this study was to establish a single nomenclature 

for well-resolved anamorph–teleomorph relationships as discussed 

by Hawksworth et al. (2011). In cases where one anamorphteleomorph 

generic relation is involved in a monophyletic lineage, 

one generic name was chosen based on priority and the other 

named teleomorph or anamorph state is treated as a synonym. 

Similar approaches towards single nomenclature have been 

employed in Botryosphaeriales (Crous et al. 2006, 2009a, b, Phillips 

et al. 2008), Pleosporales (Aveskamp et al. 2010), and Hypocreales 

(Lombard et al. 2010a–c, Chaverri et al. 2011, Gräfenhan et al. 

2011, Schroers et al. 2011). 

MATERIALS AND METHODS 

Isolate selection, culture studies and DNA extraction 

The generic abbreviations used in this study are: Ascochyta 

(A.), Coniothyrium (C.), Heterospora (H.), Leptosphaeria (L.), 

Paraconiothyrium (Paracon.), Paraleptosphaeria (Paralep.), Phoma 

(Ph.), Plenodomus (Plen.), Pleospora (Pleo.), Pyrenochaeta (Py.), 

Subplenodomus (Subplen.) and Westerdykella (W.). The isolates 

included in this study were obtained from the culture collections 

of the Centraalbureau voor Schimmelcultures, Utrecht, The 

Netherlands (CBS-KNAW) and the Dutch National Plant Protection 

Organization, Wageningen, The Netherlands (PD) (Table 1). 

The freeze-dried isolates were revived overnight in 2 mL malt/ 

peptone (50 % / 50 %) liquid medium and subsequently transferred 

and maintained on oatmeal agar (OA) (Crous et al. 2009c). The 

isolates, which were stored at -196 °C, were directly transferred 

to OA. Cultures growing on OA and malt extract agar (MEA) 

(Crous et al. 2009c) were studied morphologically as described in 

detail by Boerema et al. (2004). The genomic DNA isolation was 

performed using the Ultraclean Microbial DNA isolation kit (Mo Bio 

Laboratories, Carlsbad, California) according to the instructions of 

the manufacturer. All DNA extracts were diluted 10 × in milliQ water 

and stored at 4 °C before use. 

PCR and sequencing 

For nucleotide sequence comparisons, partial regions of SSU, LSU 

and ITS, as well as part of the ACT, TUB and CHS-1 genes were 

amplified. The SSU region was amplified with the primers NS1 and 

NS4 (White et al. 1990) and the LSU region was amplified with 

the primers LR0R (Rehner & Samuels 1994) and LR7 (Vilgalys 

& Hester 1990). The ITS and TUB regions were amplified as 

described by Aveskamp et al. (2009) using the primer pair V9G (de 

Hoog & Gerrits van den Ende 1998) and ITS4 (White et al. 1990) 

for the ITS and the BT2Fw and BT4Rd primer pair (Woudenberg 

et al. 2009) for the TUB locus. The ACT and CHS-1 regions 

2

Phoma sections Plenodomus, Pilosa 

were amplified using the primer pairs ACT-512F / ACT-783R and 

CHS-354R / CHS-79F (Carbone & Kohn 1999). The amplification 

reactions were performed and analysed as described by de Gruyter 

et al. (2009). 

Sequencing of the PCR amplicons was conducted using the 

same primer combinations, although the primer LR5 (Vilgalys & 

Hester 1990) was used as an additional internal sequencing primer 

for LSU. The sequence products were purified using Sephadex 

columns (Sephadex G-50 Superfine, Amersham Biosciences, 

Roosendaal, Netherlands) and analysed with an ABI Prism 3730XL 

Sequencer (Applied Biosystems) according to the manufacturer’s 

instructions. Consensus sequences were computed from both 

forward and reverse sequences using the Bionumerics v. 4.61 

software package (Applied Maths, Sint-Martens-Latem, Belgium) 

and were lodged with GenBank. All sequences of reference isolates 

included in this study were obtained from GenBank (Table 1). 

Phylogenetic analyses 

To determine the phylogeny of Phoma betae and Ph. lingam at 

rank, the SSU and LSU sequence data of two isolates were aligned 

with the sequences of 46 reference isolates in the Pleosporales that 

were obtained from GenBank (Table 1), 14 of which were classified 

in the Pleosporaceae or Leptosphaeriaceae. The phylogeny of 

Phoma section Plenodomus was determined with the combined 

data set of LSU and ITS sequences of 87 isolates, including 

53 isolates currently classified in Leptosphaeria and Phoma 

section Plenodomus. Phoma apiicola, Ph. dimorphospora, Ph. 

heteromorphospora, Ph. lupini, Ph. valerianae, Ph. vasinfecta and 

Ph. violicola classified in Phoma sections Phoma or Heterospora 

(Boerema et al. 2004) grouped in previous molecular phylogenetic 

studies outside Didymellaceae (de Gruyter et al. 2009, Aveskamp 

et al. 2010), and are therefore treated here. 

In the study of the Leptosphaeria doliolum complex, that includes 

the subspecies of Ph. acuta, viz. subsp. acuta, errabunda and also 

Ph. acuta subsp. acuta f. sp. phlogis, a phylogenetic analysis was 

performed utilising the ITS, ACT, TUB, CHS-1 sequences of 18 

isolates. Phoma macrocapsa, Ph. sydowii and Ph. veronicicola 

being closely related to this species complex were included. 

The species concept of Phoma-like anamorphs in Pleosporaceae 

was determined by alignments of the ACT sequences of 15 isolates 

and five reference isolates. Phoma fallens, Ph. glaucispora and 

Ph. flavigena were also included. These species were originally 

classified in Phoma sect. Phoma (de Gruyter & Noordeloos 1992, 

de Gruyter et al. 1998). However, a molecular phylogenetic study 

demonstrated that these species grouped in a clade representing 

Leptosphaeriaceae and Pleosporaceae (Aveskamp et al. 2010). 

Sequence data were compared with those of isolates currently 

classified in the genera Phoma, Ascochyta and Coniothyrium, as 

well as isolates of Leptosphaeria clavata and the generic type 

species Pleospora herbarum. Phoma incompta is the only species 

classified in Phoma section Sclerophomella, which proved to be 

unrelated to Didymellaceae (Aveskamp et al. 2010). 

The Phoma-like species that could not be attributed to 

Pleosporineae (Zhang et al. 2009) were studied with the LSU 

sequences of 40 isolates, including 20 reference isolates 

representing the anamorph genera Beverwykella, Neottiosporina, 

Paraconiothyrium, as well as the teleomorph genera Byssothecium, 

Falciformispora, Herpotrichia, Massaria, Melanomma, 

Paraphaeosphaeria, Pleomassaria, Preussia, Roussoella, 

Sporormiella, Thyridaria, Trematosphaeria and Westerdykella. 

Four Phoma species were included which are currently described in 

Phoma section Phoma, viz. Ph. capitulum, Ph. flavescens, Ph. lini, 

and Ph. minutispora (de Gruyter & Noordeloos 1992, de Gruyter et 

al. 1993). In addition, the human pathogens Pyrenochaeta romeroi 

and Py. mackinnonii, which could not be classified in a recent study 

dealing with Phoma-like species with setose pycnidia (de Gruyter 

et al. 2010), were included. 

The multiple alignments were automatically calculated by 

the BioNumerics software package, but manual adjustments for 

improvement were made by eye where necessary. For multilocus 

alignments, the phylogenetic analyses were done for each dataset 

individually, and where similar tree topologies were obtained, an 

analysis was performed on the combined alignment of all the 

gene regions in the multilocus alignment. Neighbour-Joining (NJ) 

distance analyses were conducted using PAUP (Phylogenetic 

Analysis Using Parsimony) v. 4.0b10 (Swofford 2003) with 

the uncorrected “p”, Jukes-Cantor and Kimura 2-parameter 

substitution models. The robustness of the trees obtained was 

evaluated by 1000 bootstrap replications. A Bayesian analysis was 

conducted with MrBayes v. 3.1.2 (Huelsenbeck & Ronqvist 2001) 

in two parallel runs, using the default settings but with the following 

adjustments: the GTR model (trees 1–3, 5) with gamma-distributed 

rate and the HKY+ γ-model (tree 4) were selected for the partitions 

using the Findmodel freeware (http://hcv.lanl.gov/content/hcv-db/ 

findmodel/findmodel.html), and a MCMC heated chain was set 

with a “temperature” value of 0.05. The number of generations and 

sample frequencies were set at 5 million and 10 (trees 3–5) or 100 

(trees 1, 2) respectively and the run was automatically stopped 

as soon as the average standard deviation of split frequencies 

reached below 0.01. The resulting trees were printed with TreeView 

v. 1.6.6 (Page 1996) and alignments and trees were deposited into 

TreeBASE (www.treebase.org). 

RESULTS 

The data for the aligned sequence matrices for the trees obtained in 

the different studies are provided below. In the case that alignments 

of multiple loci are involved, the topologies of the obtained trees 

for each locus were compared by eye to confirm that the overall 

tree topology of the individual datasets were similar to each other 

and to that of the tree obtained from the combined alignment. The 

NJ analyses with the three substitution models showed similar tree 

topologies and were congruent to those obtained in the Baysian 

analyses. The results of the molecular phylogenetic analyses are 

supplied below; the summarised additional ecology and distribution 

data of the taxa involved were adopted from Boerema et al. (2004), 

where the references to original literature are provided. 

Phylogeny of Phoma lingam and Ph. betae, the type 

species of Phoma sections Plenodomus and Pilosa 

(Pleosporineae) 

The aligned sequence matrix obtained for the SSU and LSU 

regions had a total length of 2 671 nucleotide characters, 1 367 and 

1 304 respectively. In the alignment, an insertion in the SSU at the 

positions 478–832 was observed for the cultures CBS 216.75, CBS 

165.78, CBS 138.96, CBS 331.37 and CBS 674.75. This insertion 

was excluded from further phylogenetic analyses. The combined 

dataset used in the analyses included 48 taxa and contained 2 316 

characters with 101 and 213 unique site patterns for SSU and LSU, 

www.studiesinmycology.org 

3


Table 1. Isolates used in this study and their GenBank accession numbers. Name changes and newly generated sequences are indicated in bold. 

Species name, final identification Former identification CBS no. Other no. ITS SSU LSU ACT TUB CHS-1 Host, substrate Country 

Aposphaeria corallinolutea sp. nov. Pleurophoma sp. CBS 131286 PD 83/367 JF740329 Kerria japonica (Rosaceae) Netherlands 

Pleurophoma sp. CBS 131287 PD 83/831 JF740330 Fraxinus excelsior 

(Oleaceae) 

Aposphaeria populina CBS 543.70 EU754130 Populus canadensis 

(Salicaceae) 

Pyrenochaeta sp. CBS 350.82 JF740265 Picea abies (Pinaceae) Germany 

Netherlands 

Netherlands 

Pleurophoma sp. CBS 130330 PD 84/221 JF740328 Cornus mas (Cornaceae) Netherlands 

Beverwykella pulmonaria CBS 283.53 ATCC 32983, IFO 6800 GU301804 Fagus sylvatica (Fagaceae) Netherlands 

Byssothecium circinans CBS 675.92 ATCC 52767, ATCC 52678, 

IMI 266220 

AY016357 Medicago sativa 

(Fabaceae) 

Chaetodiplodia sp. Chaetodiplodia sp. CBS 453.68 JF740115 Halimione portulacoides 

(Chenopodiaceae) 

Chaetosphaeronema hispidulum CBS 216.75 EU754045 EU754144 Anthyllis vulneraria 

(Fabaceae) 

Cochliobolus sativus DAOM 226212 DQ677995 DQ678045 (Poaceae) Unknown 

USA 

Netherlands 

Germany 

Coniothyrium carteri comb. nov. Phoma carteri CBS 101633 PD 84/74 JF740180 GQ387593 Quercus sp. Fagaceae) Netherlands 

Phoma carteri CBS 105.91 JF740181 GQ387533 GQ387594 Quercus robur (Fagaceae) Germany 

Coniothyrium dolichi comb. nov. Pyrenochaeta dolichi CBS 124143 IMI 217261 JF740182 GQ387610 Dolichos biforus 

(Fabaceae) 

Pyrenochaeta dolichi CBS 124140 IMI 217262 JF740183 GQ387550 GQ387611 Dolichos biforus 

(Fabaceae) 

Coniothyrium glycines comb. nov. Phoma glycinicola CBS 124455 IMI 294986 JF740184 GQ387536 GQ387597 Glycine max (Fabaceae) Zambia 

India 

India 

Phoma glycinicola CBS 124141 PG-1 JF740185 GQ387598 Glycine max (Fabaceae) Zimbabwe 

Coniothyrium multiporum comb. nov. Phoma multipora CBS 501.91 PD 83/888 JF740186 GU238109 Unknown Egypt 

Phoma multipora CBS 353.65 IMI 113689, ATCC 16207, 

HACC 164 

JF740187 JF740268 Saline soil India 

Coniothyrium palmarum CBS 400.71 AY720708 EU754054 EU754153 Chamaerops humilis 

(Arecaceae) 

Coniothyrium telephii comb. nov. Phoma septicidalis CBS 188.71 JF740188 GQ387538 GQ387599 Air Finland 

Italy 

Phoma septicidalis CBS 856.97 JF740189 GQ387539 GQ387600 Mineral wool Finland 

Phoma septicidalis CBS 101636 PD 86/1186 JF740190 GQ387540 GQ387601 Glycine max (Fabaceae) Zimbabwe 

Cucurbitaria berberidis, anam. 

Pyrenochaeta berberidis 

CBS 363.93 JF740191 GQ387545 GQ387606 Berberis vulgaris 

(Berberidaceae) 

Didymella exigua CBS 183.55 EU754056 EU754155 Rumex arifolius 

(Polygonaceae) 

Netherlands 

France 

4


Table 1. (Continued). 


Netherlands 

Didymella lycopersici, anam. Boeremia 

lycopersici 

CBS 378.67 JF740097 GU237950 Lycopersicon esculentum 

(Solanaceae) 

Falcisormispora lignatilis BCC 21118 GU371827 Elaeis guineensis 

(Arecaceae) 

Herpotrichia juniperi CBS 200.31 DQ678080 Juniperus nana 

(Cupressaceae) 

Heterospora chenopodii comb. nov. Phoma heteromorphospora CBS 448.68 FJ427023 EU754088 EU754187 Chenopodium album 


Phoma heteromorphospora CBS 115.96 PD 94/1576 JF740227 EU754188 Chenopodium album 


Heterospora dimorphospora comb. nov. Phoma dimorphospora CBS 345.78 PD 76/1015 JF740203 GU238069 Chenopodium quinoa 


Leptosphaeria conoidea Leptosphaeria conoidea, 

anam. Phoma doliolum 

Phoma dimorphospora CBS 165.78 PD 77/884 JF740204 JF740098 JF740281 Chenopodium quinoa 


Leptosphaeria conoidea, 

anam. Phoma doliolum 

Leptosphaeria doliolum Leptosphaeria doliolum 

subsp. doliolum var. 

doliolum, anam. Phoma 

acuta subsp. acuta 

Leptosphaeria doliolum 

subsp. errabunda, anam. 

Phoma acuta subsp. 

errabunda 

Phoma acuta subsp. acuta 

f.sp. phloxis 

Phoma acuta subsp. acuta 

f.sp. phloxis 









Leptosphaeria errabunda comb. nov. Leptosphaeria doliolum 



errabunda 

CBS 616.75 ATCC 32813, IMI 199777, 

PD 74/56 

JF740201 JF740099 JF740279 Lunaria annua 

(Brassicaceae) 

Thailand 

Switzerland 

Netherlands 

Netherlands 

Peru 

Peru 

Netherlands 

CBS 125977 PD 82/888 JF740202 JF740280 Senecio sp. (Asteraceae) Netherlands 

CBS 505.75 PD 75/141 JF740205 GQ387515 GQ387576 JF740126 JF740144 JF740162 Urtica dioica (Urticaceae) Netherlands 

CBS 541.66 PD 66/221 JF740206 JF740284 JF740127 JF740145 JF740163 Rudbeckia sp. (Asteraceae) Netherlands 

CBS 155.94 PD 77/80 JF740207 JF740282 JF740128 JF740146 JF740164 Phlox paniculata 

(Polemoniaceae) 

CBS 125979 PD 78/37 JF740208 JF740283 JF740129 JF740147 JF740165 Phlox paniculata 


Netherlands 

Netherlands 

CBS 504.75 PD 74/55 JF740209 JF740130 JF740148 JF740166 Urtica dioica (Urticaceae) Netherlands 

CBS 130000 PD 82/701 JF740210 JF740131 JF740149 JF740167 Urtica dioica (Urticaceae) Netherlands 

CBS 617.75 ATCC 32814, IMI 199775, 

PD 74/201 

JF740216 JF740289 JF740132 JF740150 JF740168 Solidago sp. (hybrid) 

(Asteraceae) 

Netherlands 


5




Netherlands 




errabunda 




errabunda 




errabunda 




errabunda 

CBS 125978 PD 74/61 JF740217 JF740290 JF740133 JF740151 JF740169 Delphinium sp. 

(Ranunculaceae) 

CBS 129999 PD 78/569 JF740218 JF740134 JF740152 JF740170 Aconitum sp. 


Netherlands 

CBS 129998 PD 84/462 JF740219 JF740135 JF740153 JF740171 Gailardia (Asteraceae) Netherlands 

CBS 129997 PD 78/631 JF740220 JF740136 JF740154 JF740172 Achillea millefolium 

(Apiaceae) 

Leptosphaeria etheridgei comb. nov. Phoma etheridgei CBS 125980 DAOM 216539, PD 95/1483 JF740221 JF740291 Populus tremuloides 

(Salicaceae) 

Leptosphaeria macrocapsa comb. nov. Phoma macrocapsa CBS 640.93 PD 78/139 JF740237 JF740304 JF740138 JF740156 JF740174 Mercurialis perennis 

(Euphorbiaceae) 

Leptosphaeria pedicularis comb. nov. Phoma pedicularis CBS 126582 PD 77/710 JF740223 JF740293 Gentiana punctata 

(Gentianaceae) 

Phoma pedicularis CBS 390.80 PD 77/711 JF740224 JF740294 JF740137 JF740155 JF740173 Pedicularis sp. 

(Scrophulariaceae) 

Netherlands 

Canada 

Netherlands 

Switzerland 

Switzerland 

Leptosphaeria rubefaciens comb. nov. Phoma rubefaciens CBS 387.80 IMI 248432, ATCC 42533, 

PD 78/809 

JF740242 JF740311 Tilia (x) europea 

(Malvaceae) 

Netherlands 

Phoma rubefaciens CBS 223.77 JF740243 JF740312 Quercus sp. (Fagaceae) Switzerland 

Leptosphaeria sclerotioides comb. nov. Phoma sclerotioides CBS 144.84 CECT 20025, PD 82/1061 JF740192 JF740269 Medicago sativa 

(Fabaceae) 

Leptosphaeria slovacica Leptosphaeria slovacica, 

anam. Phoma leonuri 

Phoma sclerotioides CBS 148.84 PD 80/1242 JF740193 JF740270 Medicago sativa 

(Fabaceae) 

Leptosphaeria slovacica, 

anam. Phoma leonuri 

CBS 389.80 PD 79/171 JF740247 JF740101 JF740315 Balota nigra (Lamiaceae) Netherlands 

CBS 125975 PD 77/1161 JF740248 JF740316 Balota nigra (Lamiaceae) Netherlands 

Leptosphaeria sydowii comb. nov. Phoma sydowii CBS 385.80 PD 74/477 JF740244 JF740313 JF740139 JF740157 JF740175 Senecio jacobaea 

(Asteraceae) 

Canada 

Canada 

UK 

Phoma sydowii CBS 125976 PD 84/472 JF740245 JF740314 JF740140 JF740158 JF740176 Senecio jacobaea 

(Asteraceae) 

Netherlands 

6




Phoma sydowii CBS 297.51 JF740246 JF740141 JF740159 JF740177 Papaver rhoeas 

Switzerland 

(Papaveraceae) 

Leptosphaeria veronicae comb. nov. Phoma veronicicola CBS 145.84 CECT 20059, PD 78/273 JF740254 JF740320 JF740142 JF740160 JF740178 Veronica chamaedryoides Netherlands 


Phoma veronicicola CBS 126583 PD 74/227 JF740255 JF740321 JF740143 JF740161 JF740179 Veronica ‘Shirley Blue’ Netherlands 


Massaria platani CBS 221.37 DQ678013 DQ678065 Platanus occidentalis USA 

(Platanaceae) 

Massarina eburnea H 3953, HHUF 26621, JCM 

AB521718 AB521735 Fagus sylvatica (Fagaceae) UK 

14422 

Massarina eburnea CBS 473.64 ETH 2945 GU296170 GU301840 Fagus sylvatica (Fagaceae) Switzerland 

Medicopsis romeroi comb. nov. Pyrenochaeta romeroi CBS 252.60 ATCC 13735, FMC 151, 

UAMH 10841 

EU754108 EU754207 Human, maduromycosis Venezuela 

Pyrenochaeta romeroi CBS 122784 PD 84/1022 EU754208 Hordeum vulgare 

(Gramineae) 

Melanomma pulvis-pyrius CBS 371.75 GU301845 Wood France 

Unknown 

CBS 400.97 DQ678020 DQ678072 Fagus sp. (Fagaceae) Belgium 

Neophaeosphaeria filamentosa CBS 102202 BPI 802755 JF740259 GQ387516 GQ387577 Yucca rostrata (Agavaceae) Mexico 

Neosetophoma samarorum CBS 138.96 PD 82/653 GQ387517 GQ387578 Phlox paniculata 


Neottiosporina paspali CBS 331.37 EU754073 EU754172 Paspalum notatum 

(Poaceae) 

Nigrogana mackinnonii comb. nov. Pyrenochaeta mackinnonii CBS 674.75 FMC 270 

GQ387613 Human, black grain 

Venezuela 

GQ387552 

mycetoma 

Pyrenochaeta mackinnonii CBS 110022 GQ387614 Human, mycetoma Mexico 

Netherlands 

USA 

Paraconiothyrium flavescens comb. nov. Phoma flavescens CBS 178.93 PD 82/1062 GU238075 Soil Netherlands 

Paraconiothyrium fuckelii comb. nov. Coniothyrium fuckelii CBS 797.95 GU238204 GU237960 Rubus sp. (Rosaceae) Denmark 

Paraconiothyrium fusco-maculans comb. 

nov. 

Plenodomus fuscomaculans 

CBS 116.16 EU754197 Malus sp. (Rosaceae) USA 

Paraconiothyrium lini comb. nov. Phoma lini CBS 253.92 PD 70/998 EU238093 Wisconsin tank Netherlands 

Paraconiothyrium maculicutis sp. nov. Pleurophoma pleurospora CBS 101461 IMI 320754, UTHSC 87-144 EU754200 Human, cutaneous lesions USA 

Paraconiothyrium minitans CBS 122788 PD 07/03486739 EU754074 EU754173 Unknown UK 

CBS 122786 PD 99/1064-1 EU754174 Clematis sp. 


Netherlands 

Paraconiothyrium tiliae comb. nov. Asteromella tiliae CBS 265.94 EU754139 Tilia platyphyllos (Tiliaceae) Austria 


7




Paraleptosphaeria dryadis comb. nov. Leptosphaeria dryadis CBS 643.86 JF740213 GU301828 Dryas octopetala 

(Rosaceae) 

Switzerland 

Norway 

Paraleptosphaeria macrospora comb. 

nov. 

Phoma macrospora CBS 114198 UPSC 2686 JF740238 JF740305 Rumex domesticus 


Paraleptosphaeria nitschkei comb. nov. Leptosphaeria nitschkei CBS 306.51 JF740239 JF740308 Cirsium spinosissimum 

(Asteraceae) 

Paraleptosphaeria orobanches comb. 

nov. 

Paraleptosphaeria praetermissa comb. 

nov. 

Phoma korfii CBS 101638 PD 97/12070 JF400230 JF740299 Epifagus virginiana 

(Orobanchaceae) 

Leptosphaeria praetermissa CBS 114591 JF740241 JF740310 Rubus idaeus (Rosaceae) Sweden 

Paraphaeosphaeria michoti CBS 652.86 ETH 9483 GQ387520 GQ387581 Typha latifolia (Typhaceae) Switzerland 

Switzerland 

USA 

Paraphoma radicina CBS 111.79 IMI 386094, PD 76/437 EU754092 EU754191 Malus sylvestris 

(Rosaceae) 

Phaeosphaeria nodorum CBS 110109 EU754076 EU754175 Lolium perenne 

(Gramineae) 

Phoma herbarum CBS 615.75 FJ427022 EU754087 EU754186 Rosa multiflora (Rosaceae) Netherlands 

Netherlands 

Denmark 

Phoma paspali CBS 560.81 PD 92/1569 GU238227 G238124 Paspalum dilatum 

(Poaceae) 

Plenodomus agnitus comb. nov. Leptosphaeria agnita, 

anam. Phoma agnita 

Leptosphaeria agnita, 

anam. Phoma agnita 

CBS 121.89 PD 82/903 JF740194 JF740271 Eupatorium cannabinum 

(Asteraceae) 

CBS 126584 PD 82/561 JF740195 JF740272 Eupatorium cannabinum 

(Asteraceae) 

Plenodomus biglobosus comb. nov. Leptosphaeria biglobosa CBS 119951 JF740198 JF740102 JF740274 Brassica rapa 


Plenodomus chrysanthemi comb. nov. Phoma vasinfecta, 

synanam. Phialophora 

chrysanthemi 

CBS 127249 DAOM 229269 JF740199 JF740275 Brassica juncea 


CBS 539.63 JF740253 GU238230 GU238151 Chrysanthemum sp. 

(Asteraceae) 

Plenodomus collinsoniae comb. nov. Leptosphaeria collinsoniae CBS 120227 JCM 13073, MAFF 239583 JF740200 JF740276 Vitis coignetiae (Vitaceae) Japan 

New 

Zealand 

Netherlands 

Netherlands 

Netherlands 

France 

Greece 

Plenodomus confertus comb. nov. Leptosphaeria conferta, 

anam. Phoma conferta 

Plenodomus congestus comb. nov. Leptosphaeria congesta, 

anam. Phoma congesta 

Plenodomus enteroleucus comb. nov. Phoma enteroleuca var. 

enteroleuca 

Phoma enteroleuca var. 

enteroleuca 

CBS 375.64 AF439459 JF740277 Anacyclus radiatus 

(Asteraceae) 

CBS 244.64 AF439460 JF740278 Erigeron canadensis 

(Asteraceae) 

CBS 142.84 PD 81/654, CECT20063 JF740214 JF740287 Catalpa bignonioides 

(Bignoniaceae) 

CBS 831.84 JF740215 JF740288 Triticum aestivum 

(Poaceae) 

Spain 

Spain 

Netherlands 

Germany 

8




Plenodomus fallaciosus comb. nov. Leptosphaeria fallaciosa CBS 414.62 ETH 2961 JF740222 JF740292 Satureia montana 

France 

(Lamiaceae) 

Plenodomus hendersoniae comb. nov. Phoma intricans CBS 113702 UPSC 1843 JF740225 JF740295 Salix cinerea (Salicaceae) Sweden 

Phoma intricans CBS 139.78 JF740226 JF740296 Pyrus malus (Rosaceae) Netherlands 

Plenodomus influorescens comb. nov. Phoma enteroleuca var. 

influorescens 

Phoma enteroleuca var. 

influorescens 

CBS 143.84 PD 78/883, CECT 20064 JF400228 JF740297 Fraxinus excelsior 

(Oleaceae) 

Netherlands 

PD 73/1382 JF400229 JF740298 Lilium sp. (Liliaceae) Netherlands 

Plenodomus libanotidis comb. nov. Leptosphaeria libanotis CBS 113795 UPSC 2219 JF400231 JF740300 Seseli libanotis (Apiaceae) Sweden 

Plenodomus lindquistii comb. nov. Leptosphaeria lindquistii, 

anam. Phoma macdonaldii 

Leptosphaeria lindquistii, 

anam. Phoma macdonaldii 

Plenodomus lingam Leptosphaeria maculans, 

anam. Phoma lingam 

CBS 386.80 PD 77/336 JF400232 JF740301 Helianthus annuus 

(Asteraceae) 

CBS 381.67 JF400233 JF740302 Helianthus annuus 

(Asteraceae) 

CBS 275.63 MUCL 9901, UPSC 1025 JF400234 JF740103 JF740306 Brassica sp. (Brassicaceae) UK 

former 

Yugoslavia 

Canada 

Leptosphaeria maculans, 


CBS 260.94 PD 78/989 JF400235 JF740307 JF740116 Brassica oleracea 


Netherlands 

Leptosphaeria maculans, 


CBS 147.24 JF740117 Unknown Unknown 

Plenodomus lupini comb. nov. Phoma lupini CBS 248.92 PD 79/141 JF740236 JF740303 Lupinus mutabilis 

(Fabaceae) 

Plenodomus pimpinellae comb. nov. Leptosphaeria pimpinellae, 

anam. Phoma pimpinellae 

CBS 101637 PD 92/41 JF740240 JF740309 Pimpinella anisum 

(Apiaceae) 

Plenodomus tracheiphilus comb. nov. Phoma tracheiphila CBS 551.93 PD 81/782 JF740249 JF740104 JF740317 Citrus limonium (Rutaceae) Israel 

Peru 

Israel 

Phoma tracheiphila CBS 127250 PD 09/04597141 JF740250 JF740318 Citrus sp. (Rutaceae) Italy 

Plenodomus visci comb. nov. Plectophomella visci CBS 122783 PD 74/1021 JF740256 EU754096 EU754195 Viscum album (Viscaceae) France 

Plenodomus wasabiae Phoma wasabiae CBS 120119 FAU 559 JF740257 JF740323 Wasabia japonica 


Phoma wasabiae CBS 120120 FAU 561 JF740258 JF740324 Wasabia japonica 


Pleomassaria siparia CBS 279.74 AY004341 Betula verrucosa 

(Betulaceae) 

Pleospora angustis nom. nov. Leptosphaeria clavata CBS 296.51 JF740122 Unknown Switzerland 

Taiwan 

Taiwan 

Netherlands 

Pleospora betae Pleospora betae, anam. 

Phoma betae 

CBS 523.66 PD 66/270, IHEM 3915 EU754080 EU754179 JF740118 Beta vulgaris 


Netherlands 

Pleospora betae, anam. 

Phoma betae 

CBS 109410 PD 77/113 EU754178 JF740119 Beta vulgaris 


Netherlands 


9




Germany 

Pleospora calvescens Pleospora calvescens, 

anam. Ascochyta caulina 

CBS 246.79 PD 77/655 EU754032 EU754131 JF740120 Atriplex hastata 


Pleospora calvescens, 


CBS 343.78 JF740121 Atriplex hastata 


Netherlands 

Pleospora chenopodii Ascochyta hyalospora CBS 206.80 PD 74/1022 JF740095 JF740266 JF740109 Chenopodium quinoa 


Pleospora calvescens, 


CBS 344.78 PD 68/682 JF740110 Atriplex hastata 


Bolivia 

Netherlands 

Pleospora fallens comb. nov. Phoma fallens CBS 161.78 LEV 1131 JF740106 Olea europaea 

(Oleaeceae) 

Phoma glaucispora CBS 284.70 PD 97/2400 JF740107 Nerium oleander 

(Apocynaceae) 

Pleospora flavigena comb. nov. Phoma flavigena CBS 314.80 PD 91/1613 JF740108 Water Romania 

New 

Zealand 

Italy 

Pleospora halimiones nom. nov. Ascochyta obiones CBS 432.77 IMI 282137 JF740096 JF740267 JF740113 Halimione portulacoides 


Ascochyta obiones CBS 786.68 JF740114 Halimione portulacoides 


Pleospora herbarum CBS 191.86 IMI 276975 GU238232 GU238160 JF740123 Medicago sativa 

(Fabaceae) 

Pleospora incompta comb. nov. Phoma incompta CBS 467.76 JF740111 Olea europaea 

(Oleaeceae) 

Pleospora typhicola Pleospora typhicola, anam. 

Phoma typharum 

Phoma incompta CBS 526.82 JF740112 Olea europaea 

(Oleaeceae) 

Pleospora typhicola, anam. 

Phoma typharum 

CBS 132.69 JF740105 JF740325 JF740124 Typha angustifolia 

(Typhaceae) 

Netherlands 

Netherlands 

India 

Greece 

Italy 

Netherlands 

CBS 602.72 JF740125 Typha sp. (Typhaceae) Netherlands 

Pleurophoma pleurospora Pleurophoma sp. CBS 116668 JF740326 Citysus scoparius 

(Fabaceae) 

Pleurophoma sp. CBS 130329 PD 82/371 JF740327 Lonicera sp. 

(Caprifoliaceae) 

Preussia funiculata CBS 659.74 GU296187 GU301864 Soil Senegal 

Netherlands 

Netherlands 

Pseudorobillarda phragmitis CBS 398.61 IMI 070678 EU754203 Phragmitis australis 

(Poaceae) 

Pyrenochaeta cava CBS 257.68 IMI 331911 JF740260 EU754100 EU754199 Wheat field soil Germany 

UK 

Pyrenochaeta lycopersici CBS 267.59 JF740261 GQ387551 GQ387612 Lycopersicon esculentum 

(Solanaceae) 

Pyrenochaeta nobilis CBS 407.76 EU930011 EU754107/ 

DQ898287 

EU754206 Laurus nobilis (Lauraceae) Italy 

Netherlands 

10




Pyrenochaetopsis leptospora CBS 101635 PD 71/1027 JF740262 GQ387566 GQ387627 Secale cereale (Poaceae) Europe 

Pyrenochaetopsis pratorum comb. nov. Phoma pratorum CBS 445.81 PDDCC 7049, PD 80/1254 JF740263 GU238136 Lolium perenne, leaf 

(Poaceae) 

CBS 286.93 PD 80/1252 JF740264 JF740331 Dactylis glomerata 

(Poaceae) 

Pyrenophora tritici-repentis OSC 100066 AY544716 AY544672 (Poaceae) Italy 

New 

Zealand 

New 

Zealand 

Roussoella hysterioides CBS 125434 HH 26988 AB524622 Sasa kurilensis (Poaceae) Japan 

Setomelanomma holmii CBS 110217 GQ387572 GQ387633 Picea pungens (Pinaceae) USA 

Setophoma terrestris CBS 335.29 GQ387526 GQ387587 Allium sativum (Alliaceae) USA 

Sporormiella minima CBS 524.50 DQ678003 DQ678056 Dung of goat Panama 

Stagonosporopsis cucurbitacearum CBS 133.96 GU238234 GU238181 Cucurbita sp. 

(Cucurbitaceae) 

New 

Zealand 

Subplenodomus apiicola comb. nov. Phoma apiicola CBS 285.72 JF740196 GU238040 Apium graveolens var. 

rapaceum (Umbelliferae) 

Subplenodomus drobnjacensis comb. 

nov. 

Phoma apiicola CBS 504.91 PD 78/1073 JF740197 JF740273 Apium graveolens 

(Umbelliferae) 

Phoma drobnjacensis CBS 269.92 PD 88/896 JF740211 JF740100 JF740285 Eustoma exaltatum 

(Gentianaceae) 

Phoma drobnjacensis CBS 270.92 PD 83/650 JF740212 JF740286 Gentiana makinoi ‘Royal 

Blue’ (Gentianaceae) 

Subplenodomus valerianae comb. nov. Phoma valerianae CBS 630.68 PD 68/141 JF740251 GU238150 Valeriana phu 

(Valerianaceae) 

Phoma valerianae CBS 499.91 PD 73/672 JF740252 JF740319 Valeriana officinalis 

(Valerianaceae) 

Subplenodomus violicola comb. nov. Phoma violicola CBS 306.68 FJ427054 GU238231 GU238156 Viola tricolor (Violaceae) Netherlands 

Germany 

Netherlands 

Netherlands 

Netherlands 

Netherlands 

Netherlands 

Phoma violicola CBS 100272 FJ427055 JF740322 Viola tricolor (Violaceae) New 

Zealand 

Thyridaria rubronotata CBS 419.85 GU301875 Acer pseudoplatanus 

(Aceraceae) 

Netherlands 

Trematosphaeria pertusa CBS 122368 FJ201990 Fraxinus excelsior 

(Oleaceae) 

France 

Westerdykella capitulum comb. nov. Phoma capitulum CBS 337.65 PD 91/1614, ATCC 16195, 

HACC 167, IMI 113693 

GU238054 Saline soil India 

Westerdykella minutispora comb. nov. Phoma minutispora CBS 509.91 PD 77/920 GU238108 Saline soil India 

Westerdykella ornata CBS 379.55 GU301880 Mangrove mud Mozambique 


11


CBS 246.79 Pleospora calvescens 

100 

CBS 432.77 Ascochyta obiones 

99 

CBS 206.80 Ascochyta hyalospora 

CBS 523.66 Pleospora betae, anam. Phoma betae 

Pleosporaceae (A) 

100 

100 DAOM 226212 Cocliobolus sativus 

100 CBS 191.86 Pleospora herbarum 

100 

OSC100066 Pyrenophora tritici-repentis 

CBS 132.69 Pleospora typhicola 

100 

64 

CBS 165.78 Phoma dimorphospora 

CBS 448.68 Phoma heteromorphospora 

56 

CBS 269.92 Phoma drobnjacensis 

100 

60 

CBS 275.63 Leptosphaeria maculans, anam. Phoma lingam 

CBS 119951 Leptosphaeria biglobosa 

Leptosphaeriaceae (B) 

57 CBS 551.93 Phoma tracheiphila 

CBS 539.63 Phoma vasinfecta 

CBS 122783 Plectophomella visci 

68 

99 

CBS 363.93 Cucurbitaria berberidis 

CBS 257.68 Pyrenochaeta cava 

CBS 407.76 Pyrenochaeta nobilis 

100 

Cucurbitariaceae (C) 

81 81 CBS 101635 Pyrenochaetopsis leptospora 

CBS 267.59 Pyrenochaeta lycopersici 

54 

89 CBS 389.80 Leptosphaeria slovacica 

100 CBS 505.75 Leptosphaeria doliolum subsp. doliolum Leptosphaeriaceae (D) 

CBS 616.75 Leptosphaeria conoidea 

88 

100 

CBS 216.75 Chaetosphaeronema hispidulum 

CBS 335.29 Setophoma terrestris 

CBS 110109 Phaeosphaeria nodorum 

CBS 138.96 Neosetophoma samarorum 

Phaeosphaeriaceae (E) 

99 80 

CBS 111.79 Paraphoma radicina 

CBS 110217 Setomelanomma holmii 

100 CBS 124455 Phoma glycinicola 

93 CBS 124140 Pyrenochaeta dolichi 

77 CBS 105.91 Phoma carteri 

Clade F 

CBS 101636 Phoma septicidalis 

CBS 400.71 Coniothyrium palmarum 

CBS 102202 Neophaeosphaeria filamentosa 

100 CBS 133.96 Stagonosporopsis bryoniae 

CBS 378.67 Didymella lycopersici 

100 

CBS 183.55 Didymella exigua 

100 

CBS 615.75 Phoma herbarum 

Didymellaceae (G) 

CBS 560.81 Phoma paspali 

100 CBS 122788 Paraconiothyrium minitans 

100 CBS 652.86 Paraphaeosphaeria michotii 

Montagnulaceae 

100 

CBS 797.95 Coniothyrium fuckelii 

97 

CBS 331.37 Neottiosporina paspali 

Massarinaceae 

CBS 252.60 Pyrenochaeta romeroi 

CBS 674.75 Pyrenochaeta mackinnonii 

CBS 524.50 Sporormiella minima 

Sporormiaceae 

100 

0.1 

Fig. 1. The phylogeny of Phoma lingam and Phoma betae, the type species of Phoma sections Plenodomus and Pilosa, based on the strict consensus tree from a Bayesian 

analysis of 48 LSU/SSU sequences. The Bayesian posterior probabilities are given at the nodes. The tree was rooted to Sporormiella minima (CBS 524.50). 

respectively. The tree (Fig. 1) was rooted to Sporormiella minima 

(CBS 524.50). The Bayesian analysis resulted in 6 5442 trees after 

3 272 000 generations, from which the burn-in was discarded and 

the consensus tree and posterior probabilities were calculated 

based on 56 028 trees (Fig. 1). 

The families that belong to Pleosporineae, represented by the 

species grouping in clades A–G, clustered in a strongly supported 

clade (99 % posterior probability). Clade A, representing those 

species classified in Pleosporaceae, was strongly supported (100 

%) and included two subclades. Pleospora betae (anam. Ph. 

betae), clustered with Pleospora calvescens (anam. Ascochyta 

caulina), A. obiones and A. hyalospora; all recorded as pathogens 

on Chenopodiaceae. The generic type species Pleospora 

herbarum, a plurivorous species, grouped with Cochliobolus 

sativus, Pyrenophora tritici-repentis and Pleospora typhicola 

(anam. Ph. typhina), all recorded from Poaceae. Clade B includes 

Leptosphaeria maculans (anam. Ph. lingam) and clustered with 

Leptosphaeria biglobosa. In clade B also other important plant 

pathogens of Phoma section Plenodomus can be found, such as Ph. 

tracheiphila, Ph. vasinfecta, Ph. drobnjacensis, and Plectophomella 

12


visci. Phoma heteromorphospora, type species of Phoma section 

Heterospora (Boerema et al. 1997) and Ph. dimorphospora also 

grouped in this Leptosphaeria clade, in congruence with previous 

findings (de Gruyter et al. 2009, Aveskamp et al. 2010). 

Leptosphaeria doliolum (anam. Ph. acuta), type species of 

the genus Leptosphaeria, is found in Clade D, clustering with 

L. conoidea and L. slovacica. Leptosphaeria doliolum and its 

relatives comprise a sister clade C with species classified in 

Cucurbitariaceae, including Cucurbitaria berberidis, the three 

Pyrenochaeta species, Py. cava, Py. lycopersici and Py. nobilis, 

and Pyrenochaetopsis leptospora. 

Phaeosphaeria nodorum and its relatives Neosetophoma 

samarorum, Setophoma terrestris, Chaetosphaeronema 

hispidulum, Paraphoma radicina and Setomelanomma holmii, 

represent Phaeosphaeriaceae in clade E as has previously been 

found (de Gruyter et al. 2009, 2010). 

A distinct clade F includes Ph. glycinicola, Ph. carteri, Ph. 

septicidalis, and the taxonomic confusing species Pyrenochaeta 

dolichi (Grondona et al. 1997). The position of Coniothyrium 

palmarum and Neophaeosphaeria filamentosa could not be 

clarified, but both species are also treated below in a phylogeny 

including close relatives based on ITS and LSU regions (Fig. 

2). Didymella exigua, type species of the genus Didymella, and 

Ph. herbarum represent Didymellaceae, and clustered in a wellsupported 

clade (G) in congruence with previous studies (de Gruyter 

et al. 2009, 2010, Aveskamp et al. 2010). The molecular phylogeny 

of species which group in this analysis outside of Pleosporineae in 

Montagnulaceae, Massarinaceae and Sporormiaceae were further 

analysed utilising LSU sequence data of a broader range of taxa 

(Fig. 5). 

Phoma section Plenodomus and close allies 

The aligned sequence matrix obtained for the LSU and ITS regions 

had a total length of 1 921 nucleotide characters, 1 332 and 589 

respectively. The combined dataset used in the analyses included 

87 taxa and contained 1921 characters with 298 and 118 unique site 

patterns for LSU and ITS respectively. The tree (Fig. 2) was rooted 

to Ph. herbarum (CBS 615.75), the representative isolate of the type 

species of Phoma (Boerema et al. 2004). The Bayesian analysis 

resulted in 100 002 trees after 5 000 000 generations, from which 

the burn-in was discarded and the consensus tree and posterior 

probabilities were calculated based on 90 930 trees (Fig. 2). 

The species currently classified in Leptosphaeria and Phoma 

section Plenodomus grouped in clades A and B representing 

Leptosphaeriaceae, including the type species Ph. lingam and 

Leptosphaeria doliolum, respectively. Isolates of the taxa that 

represent Cucurbitariaceae, Cucurbitaria berberidis and its related 

species Pyrenochaeta cava, Py. nobilis, Py. lycopersici and 

Pyrenochaetopsis leptospora, clustered in a distinct clade D only 

distantly related to Leptosphaeriaceae. This finding agrees with a 

recent study (de Gruyter et al. 2010). Phoma pratorum clustered 

with Pyrenochaetopsis leptospora. 

Leptosphaeria biglobosa grouped in a subclade A1 with Ph. 

wasabiae, the cause of black rot disease on Wasabia japonica 

(Brassicaceae) and Ph. pimpinellae, a necrotroph on Pimpinella 

anisum (Apiaceae). Leptosphaeria maculans, considered as closely 

related to the L. biglobosa complex, proved to be more distantly 

related in clade A1. In this subclade, other important pathogens can 

be found, such as Ph. tracheiphila, a quarantine organism on Citrus 

spp. (Rutaceae), Ph. vasinfecta, a pathogen on Chrysanthemum 

spp. (Asteraceae), L. lindquistii (anam. Ph. macdonaldii), a 

worldwide pathogen on Helianthus annuus (Asteraceae) and 

Ph. lupini, a seed borne pathogen known from Lupinus spp. 

(Fabaceae). Subclade A1 also comprises both varieties of Ph. 

enteroleuca, opportunistic pathogens on deciduous trees and 

shrubs, and the necrotrophic species L. agnita (anam. Ph. agnita), 

Ph. congesta (both recorded on Asteraceae), Ph. conferta (mainly 

on Brassicaceae), L. hendersoniae (on Salicaceae), L. fallaciosa, L. 

collinsoniae (mainly on Lamiaceae) and L. libanotis (on Apiaceae). 

Plectophomella visci, recorded from leaves of Viscum album 

(Viscaceae), also clustered in the Leptosphaeriaceae. The genus 

Plenodomus is re-introduced here to accommodate the species in 

subclade A1, which are allied to Ph. lingam. 

Subclade A2 comprises pathogenic species often causing leaf 

spots such as Ph. apiicola on Apium graveolens (Apiaceae), Ph. 

drobnjacensis (on Gentianaceae), Ph. violicola (on Violaceae) as 

well as the necrotrophic species Ph. valerianae, on Valeriana spp. 

(Valerianaceae). Phoma apiicola and Ph. valerianae were classified 

in Phoma section Phoma, and Ph. violicola was classified in Phoma 

sect. Peyronellaea; however, the relationship of these species in 

Leptosphaeriaceae is clearly demonstrated (Fig. 2), and therefore 

the species are transferred to the new genus Subplenodomus. 

These results are in congruence with a recent study where Ph. 

violicola, Ph. apiicola and Ph. valerianae clustered in a clade 

representing both Leptosphaeriaceae and Pleosporaceae 

(Aveskamp et al. 2010). 

Four Leptosphaeria species, L. macrospora (soil) and 

the necrotrophic species L. nitschkei (on Asteraceae), L. 

praetermissa, on Rubus idaeus (Rosaceae) and L. dryadis, 

on Dryas spp. (Rosaceae) grouped in a subclade A3 and are 

transferred here to a new genus Paraleptosphaeria. Phoma 

korfii also clustered in this subclade. The European species Ph. 

heteromorphospora, type species of Phoma section Heterospora, 

and the American counterpart Ph. dimorphospora, both pathogens 

on Chenopodiaceae, grouped in a distinct subclade A4. Phoma 

sect. Heterospora is raised to generic rank to accommodate both 

species in Leptosphaeriaceae. 

Clade B comprises necrotrophic species related to the type 

species L. doliolum (anam. Ph. acuta). The phylogeny of this 

species complex, and the closely related species Ph. veronicicola, 

Ph. macrocapsa and Ph. sydowii, is treated below. The necrotrophic 

species Ph. sclerotioides, L. conoidea (anam. Ph. doliolum), L. 

slovacica (anam. Ph. leonuri) and Ph. pedicularis also proved to 

be related. The species Ph. rubefaciens and Ph. etheridgei also 

belong to clade B, but these species, both recorded on trees, are 

more distantly related. 

The Phoma species in clades A and B are in majority currently 

described as anamorphs of the genus Leptosphaeria, or belong to 

Phoma section Plenodomus. These Phoma anamorphs are only 

distantly related to the type species Ph. herbarum and its relatives 

in Didymellaceae, and therefore these species described in section 

Plenodomus are excluded from the genus Phoma. Clade C is more 

distantly related to Leptosphaeriaceae and comprises species that 

are related to Coniothyrium palmarum in Coniothyriaceae. Two 

subclades are recognised in clade C: Ph. glycinicola, Py. dolichi 

and Ph. carteri group with the generic type species C. palmarum, 

whereas two isolates of Ph. septicidalis group with Ph. multipora. The 

teleomorph Neophaeosphaeria filamentosa clustered basal to this 

clade. Clade D includes the genera Cucurbitaria, Pyrenochaetopsis 

and Pyrenochaeta, which represent Cucurbitariaceae. This finding 

is in congruence with previous studies (de Gruyter et al. 2010). 


13

Fig 2 


A1 

A2 

A3 

A4 

53 

70 

51 

0.1 

100 CBS 121.89 Plenodomus agnitus comb. nov. Leptosphaeria agnita 

100 CBS 126584 Plenodomus agnitus comb. nov. Leptosphaeria agnita 

100 

CBS 414.62 Plenodomus fallaciosus comb. nov. Leptosphaeria fallaciosa 

99 

CBS 248.92 Plenodomus lupini comb. nov. Phoma lupini 

CBS 122783 Plenodomus visci comb. nov. Plectophomella visci 

83 

100 CBS 143.84 Plenodomus influorescens comb. nov. Phoma enteroleuca var. influorescens 

100 PD73.1382 Plenodomus influorescens comb. nov. Phoma enteroleuca var. influorescens 

100 CBS 381.67 Plenodomus lindquistii comb. nov. Leptosphaeria lindquistii 

CBS 386.80 Plenodomus lindquistii comb. nov. Leptosphaeria lindquistii 

CBS 120227 Plenodomus collinsoniae comb. nov. Leptosphaeria collinsoniae 

100 CBS 119951 Plenodomus biglobosus comb. nov. Leptosphaeria biglobosa 

CBS 127249 Plenodomus biglobosus comb. nov. Leptopshaeria biglobosa 

100 CBS 120119 Plenodomus wasabiae 

94 CBS 120120 Plenodomus wasabiae 

67 

CBS 101637 Plenodomus pimpinellae comb. nov. Leptosphaeria pimpinellae 

100 

CBS 539.63 Plenodomus chrysanthemi comb. nov. Phoma vasinfecta 

CBS 127250 Plenodomus tracheiphilus comb. nov. Phoma tracheiphila 

98 

CBS 551.93 Plenodomus tracheiphilus comb. nov. Phoma tracheiphila 

CBS 244.64 Plenodomus congestus comb. nov.Leptopshaeria congesta 

CBS 113795 Plenodomus libanotidis comb. nov. Leptosphaeria libanotis 

100 

100 CBS 260.94 Plenodomus lingam Leptosphaeria maculans, Phoma lingam 

CBS 275.63 Plenodomus lingam Leptosphaeria maculans, Phoma lingam 

94 

CBS 113702 Plenodomus hendersoniae comb. nov. Leptosphaeria hendersoniae 

CBS 139.78 Plenodomus hendersoniae comb. nov. Leptosphaeria hendersoniae 

CBS 375.64 Plenodomus confertus comb. nov. Leptosphaeria conferta 

100 CBS142.84 Plenodomus enteroleucus comb. nov. Phoma enteroleuca var. enteroleuca 

CBS 831.84 Plenodomus enteroleucus comb. nov. Phoma enteroleuca var. enteroleuca 

100 CBS 285.72 Subplenodomus apiicola comb. nov. Phoma apiicola 

CBS 504.91 Subplenodomus apiicola comb. nov. Phoma apiicola 

100 CBS 499.91 Subplenodomus valerianae comb. nov. Phoma valerianae 

CBS 630.68 Subplenodomus valerianae comb. nov. Phoma valerianae 

CBS 269.92 Subplenodomus drobnjacensis comb. nov. Phoma drobnjacensis 

CBS 270.92 Subplenodomus drobnjacensis comb. nov. Phoma drobnjacensis 

CBS 100272 Subplenodomus violicola comb. nov. Phoma violicola 

CBS 306.68 Subplenodomus violicola comb. nov. Phoma violicola 

CBS 114198 Paraleptosphaeria macrospora comb. nov. Leptosphaeria macrospora 

CBS 306.51 Paraleptosphaeria nitschkei comb. nov. Leptosphaeria nitschkei 

CBS 114591 Paraleptosphaeria praetermissa comb. nov. Leptosphaeria praetermissa 

CBS 643.86 Paraleptosphaeria dryadis comb. nov. Leptosphaeria dryadis 

CBS 101638 Paraleptosphaeria orobanches comb. nov. Phoma korfii 

100 CBS 115.96 Heterospora chenopodii comb. nov. Phoma heteromorphospora 

100 CBS 448.68 Heterospora chenopodii comb. nov. Phoma heteromorphospora 

CBS 165.78 Heterospora dimorphospora comb. nov. Phoma dimorphospora 

100 CBS 345.78 Heterospora dimorphospora comb. nov. Phoma dimorphospora 

91 CBS 145.84 Leptosphaeria veronicae comb. nov. Phoma veronicicola 

CBS 126583 Leptosphaeria veronicae comb. nov. Phoma veronicicola 

98 

CBS 505.75 Leptosphaeria doliolum 

82 CBS 125979 Leptosphaeria doliolum 


100 CBS 155.94 Leptosphaeria doliolum 

CBS 617.75 Leptosphaeria errabunda comb. nov. 

100 

CBS 125978 Leptosphaeria errabunda comb. nov. 

CBS 640.93 Leptosphaeria macrocapsa comb. nov. Phoma macrocapsa 

62 100 

CBS 385.80 Leptosphaeria sydowii comb. nov. Phoma sydowii 

100 CBS 125976 Leptosphaeria sydowii comb. nov. Phoma sydowii 

100 CBS 144.84 Leptosphaeria sclerotioides comb. nov. Phoma sclerotioides 

92 CBS 148.84 Leptosphaeria sclerotioides comb. nov. Phoma sclerotioides 

98 CBS 616.75 Leptosphaeria conoidea 

100 

100 CBS 125977 Leptosphaeria conoidea 

100 CBS 389.80 Leptosphaeria slovacica 

CBS 125975 Leptosphaeria slovacica 

100 CBS 390.80 Leptosphaeria pedicularis comb. nov. Phoma pedicularis 

CBS 126582 Leptosphaeria pedicularis comb. nov. Phoma pedicularis 

100 CBS 223.77 Leptosphaeria rubefaciens comb. nov. Phoma rubefaciens 

CBS 387.80 Leptosphaeria rubefaciens comb. nov. Phoma rubefaciens 

CBS 125980 Leptosphaeria etheridgei comb. nov. Phoma etheridgei 

92 IMI 217261 Coniothyrium dolichi comb. nov. Pyrenochaeta dolichi 

100 IMI 217262 Coniothyrium dolichi comb. nov. Pyrenochaeta dolichi 

IMI294986 Coniothyrium glycines comb. nov. Phoma glycinicola 

100 IMI 124141 Coniothyrium glycines comb. nov. Phoma glycinicola 

100 IMI 101633 Coniothyrium carteri comb. nov. Phoma carteri 

CBS 105.91 Coniothyrium carteri comb. nov. Phoma carteri 

CBS 400.71 Coniothyrium palmarum 

Coniothyriaceae (C) 

97 CBS 188.71 Coniothyrium telephii comb. nov. Phoma septicidalis 

100 CBS 856.97 Coniothyrium telephii comb. nov. Phoma septicidalis 

CBS 101636 Coniothyrium telephii comb. nov. Phoma septicidalis 

100 CBS 353.65 Coniothyrium multiporum comb. nov. Phoma multipora 

CBS 501.91 Coniothyrium multiporum comb. nov. Phoma multipora 

CBS 102202 Neophaeosphaeria filamentosa 

CBS 267.59 Pyrenochaeta lycopersici 

100 CBS 286.93 Pyrenochaetopsis pratorum comb. nov. 

100 

CBS 445.81 Pyrenochaetopsis pratorum comb. nov. 

CBS 101635 Pyrenochaetopsis leptospora 

CBS 257.68 Pyrenochaeta cava 

Cucurbitariaceae (D) 

CBS 363.93 Cucurbitaria berberidis 

CBS 407.76 Pyrenochaeta nobilis 

CBS 615.75 Phoma herbarum 

Didymellaceae (E) 

56 

61 

66 

71 

100 

77 

90 

91 

100 

100 

100 

99 

100 

87 

100 

64 

64 

99 

96 

Fig. 2. The phylogeny of Phoma section Plenodomus and Leptosphaeria, based on the strict consensus tree from a Bayesian analysis of 87 LSU/ITS sequences. The Bayesian 

posterior probabilities are given at the nodes. The tree was rooted to Phoma herbarum (CBS 615.75). 

Leptosphaeriaceae (A) 

Leptosphaeriaceae (B) 

14


Phylogeny of the Leptosphaeria doliolum complex 

The aligned sequence matrix obtained for the ITS, ACT, TUB and 

CHS-1 regions had a total length of 1 345 nucleotide characters; 

ITS 522, ACT 240, TUB 332 and CHS-1 251, respectively. The 

combined dataset used in the analyses included 18 taxa and 

contained 1 345 characters with 98 unique site patterns. The 

tree (Fig. 3) was rooted to “Ph. pedicularis” (CBS 390.80). The 

Bayesian analysis resulted in 6 002 trees after 30 000 generations, 

from which the burn-in was discarded and the consensus tree and 

posterior probabilities were calculated based on 3 341 trees. 

The phylogenetic tree revealed two clades with high posterior 

probabilities, 98 and 99 % respectively, clade A with Ph. acuta 

subsp. errabunda and Ph. macrocapsa, and clade B with Ph. 

acuta subsp. acuta (anamorph of Leptosphaeria doliolum) and Ph. 

acuta subsp. acuta f. sp. phlogis. Phoma sydowii, a necrotroph on 

Asteraceae, Senecio spp. in particular, proved to be closely related 

to Ph. acuta subsp. errabunda. The isolate CBS 297.51 preserved 

as Ph. acuta is similar to Ph. sydowii, a synonym of L. sydowii, see 

below. Phoma veronicicola, as a necrotroph specifically occurring 

on Veronica spp. (Scrophulariaceae), also proved to be related to 

Leptosphaeria doliolum. 

Phylogeny of Phoma section Pilosa 

The aligned sequence matrix obtained for the ACT region had a 

total length of 252 nucleotide characters (20 taxa), and contained 

165 unique sites. The tree was rooted to Ph. lingam (CBS 147.24 

and CBS 260.94). The Bayesian analysis resulted in 34 802 trees 

after 174 000 generations, from which the burn-in was discarded, 

and the consensus tree and posterior probabilities were calculated 

based on 11 728 trees (Fig. 4). 

The phylogenetic tree representing the Pleosporaceae includes 

Ph. betae, type species of Phoma section Pilosa. This section is 

characterised by producing pycnidia that are covered by mycelial 

hairs. Phoma betae clearly groups with other pycnidial fungi 

pathogenic on Chenopodiaceae, including Ascochyta obiones, 

A. hyalospora and A. caulina and Chaetodiplodia sp. All species 

produce similar hairy pycnidia, but are classified in Ascochyta or 

Coniothyrium due to conidial septation, or brown pigmentation of 

conidia, respectively. 

A subclade comprises the cosmopolitan Pleospora herbarum 

and related species. The species involved are associated with 

various hosts or substrates. The most closely related Ph. incompta is 

a specific pathogen on Olea europea (Oleaceae). Phoma incompta 

was classified in Phoma section Sclerophomella because of its 

thick-walled pycnidia (de Gruyter & Noordeloos 1992, Boerema 

& de Gruyter 1998). The pycnidial characters of Ph. incompta, 

pycnidia covered with mycelial hairs and with an indistinct ostiole 

visible as a pallid spot (de Gruyter & Noordeloos 1992) however, 

agrees with those of Ph. betae and Ph. typhina. 

Phoma fallens proved to be closely related to Ph. glaucispora 

in keeping with the similar in vitro characters, especially the low 

growth-rate and the size and shape of its conidia (Boerema et 

al. 2004). Both species originate from southern Europe, and 

have been associated with spots on fruits and leaves of Olea 

europea, or leaf spots on Nerium oleander, respectively. An isolate 

preserved as Leptosphaeria clavata, CBS 259.51, proved to be 

closely related. The origin of the isolate, deposited by E. Müller, 

is unknown; however, it is likely that the isolate was obtained from 

Poaceae, Triticum vulgare or Dactylis glomerata (Müller 1950). 

Phoma flavigena, once isolated from water and also recorded 

from southern Europe, proved to be more distantly related in 

Pleosporaceae. 

Phylogeny of Phoma-like anamorphs excluded from 

the suborder Pleosporineae 

The aligned sequence matrix obtained for the LSU regions had 

a total length of 808 nucleotide characters, with 208 unique 

site patterns. The phylogenetic tree (Fig. 5) was rooted to 

Pseudorobillarda phragmitis (CBS 398.61). The Bayesian analysis 

resulted in 48 402 trees after 242 000 generations, from which 

the burn-in was discarded and the consensus tree and posterior 

probabilities were calculated based on 24 876 trees. 

Clade A includes the reference isolates of the teleomorph 

Paraphaeosphaeria and the anamorph Paraconiothyrium classified 

in Montagnulaceae. This teleomorph/anamorph relation agrees 

with previous molecular phylogenetic studies (Verkley et al. 2004, 

Damm et al. 2008, de Gruyter et al. 2009). Other Phoma-like 

species in this clade are Ph. lini, Plenodomus fusco-maculans, 

Pleurophoma pleurospora (CBS 101461) and Asteromella tilliae. 

Phoma lini, a saprobe frequently recorded on dead stems of 

Linum spp., was described in Phoma section Phoma (de Gruyter 

et al. 1993). Re-examination of the conidia revealed that they are 

hyaline and thin-walled; however, also darker, greenish to yellowish 

Coniothyrium-like conidia were observed. The conidiogenous cells 

are Phoma-like, doliiform to ampulliform. 

The isolate Asteromella tiliae (CBS 265.94) clearly represents a 

species of Paraconiothyrium, and therefore, the teleomorph name 

Didymosphaeria petrakiana, Didymosphaeriaceae, is probably 

incorrect. It was already mentioned by Butin & Kehr (1995) that 

“considering the taxonomical placement of the teleomorph, the 

authors were informed about forthcoming taxonomic changes”. 

The morphological characters of the isolate CBS 101461, 

considered as representing the generic type species Pleurophoma 

pleurospora, resembles Paraconiothyrium as was previously 

discussed (de Gruyter et al. 2009). The sterile ex-type strain of 

Plenodomus fusco-maculans, CBS 116.16, recorded from Malus 

sp., also grouped with the Paraconiothyrium isolates. 

Coniothyrium fuckelii clustered in the Paraphaeosphaeria/ 

Paraconiothyrium clade, in agreement with previous studies (Damm 

et al. 2008, Aveskamp et al. 2010), and therefore, the species is 

transferred to the genus Paraconiothyrium. Two Phoma-like species 

obtained from Citysus scoparius and Lonicera sp. respectively 

(CBS 116668 and CBS 130329), cluster near Montagnulaceae 

and Massarinaceae. The morphological characters of the species 

are typical for Pleurophoma pleurospora. The taxonomic position 

of both isolates at familial rank could not be determined. The 

morphology of Phoma flavescens proved to be most similar to that 

of Paraconiothyrium, it definitely does not belong to Phoma, and 

therefore the species is transferred to Paraconiothyrium. Sequence 

data of additional species clustering nearby are required to resolve 

the current classification of Ph. flavescens. None of the Phomalike 

anamorphs included in this study grouped in clade B, which 

represents Massarinaceae. 

Clade C includes the recently assigned ex-epitype strain 

of Trematosphaeria pertusa, isolate CBS 122368 (Zhang et 

al. 2008) and Falcisformispora lignatilis. Both T. perusa and F. 

lignatilis represent Trematosphaeriaceae (Suetrong et al. 2009). 

A second isolate preserved as Trematosphaeria pertusa, CBS 

400.97, proved to be only distantly related, and clustered in clade 


15


Fig 3 

CBS 617.75 Leptosphaeria errabunda comb. nov. 


99 




Clade A 

98 

CBS 640.93 Leptosphaeria macrocapsa comb. nov. 

99 

83 

CBS 297.51 Leptosphaeria sydowii comb. nov. 

CBS 125976 Leptosphaeria sydowii comb. nov. 

52 

92 

99 

CBS 385.80 Leptosphaeria sydowii comb. nov. 


CBS 130000 Leptosphaeria doliolum 

99 

97 

CBS 125979 Leptosphaeria doliolum ‘Phoma acuta subsp. acuta f. sp. phlogis’ 

CBS 155.94 Leptosphaeria doliolum ‘Phoma acuta subsp. acuta f. sp. phlogis’ 


Clade B 


99 

CBS 145.84 Leptosphaeria veronicae comb. nov. 

CBS 126583 Leptosphaeria veronicae comb. nov. 

Fig 4 

0.1 

CBS 390.80 Leptosphaeria pedicularis comb. nov. ‘Phoma pedicularis’ 

Fig. 3. The phylogeny of the Leptosphaeria doliolum complex, based on the strict consensus tree from a Bayesian analysis of 18 ITS/ACT/TUB/CHS-1 sequences. The Bayesian 

posterior probabilities are given at the nodes. The tree was rooted to Leptosphaeria pedicularis comb. nov. (CBS 390.80). 

99 CBS 109410 Pleospora betae 

CBS 523.66 Pleospora betae 

70 CBS 786.68 Pleospora halimiones nom. nov. ‘Ascochyta obiones’ 

CBS 432.77 Pleospora halimiones nom. nov. ‘Ascochyta obiones’ 

99 

100 

CBS 206.80 Pleospora chenopodii ‘Ascochyta hyalospora’ 

CBS 344.78 Pleospora chenopodii ‘Ascochyta hyalospora’ 

52 

CBS 246.79 Pleospora calvescens ‘Ascochyta caulina’ 

CBS 343.78 Pleospora calvescens ‘Ascochyta caulina’ 

CBS 453.68 Chaetodiplodia sp. 

75 99 CBS 467.76 Pleospora incompta comb. nov. ‘Phoma incompta’ 

69 

52 

CBS 526.82 Pleospora incompta comb. nov. ‘Phoma incompta’ 

CBS 191.86 Pleospora herbarum 

100 88 

100 CBS 132.69 Pleospora typhicola ‘Phoma typhina’ 

CBS 602.72 Pleospora typhicola ‘Phoma typhina’ 

100 CBS 161.78 Pleospora fallens comb. nov. ‘Phoma fallens’ 

CBS 284.70 Pleospora fallens comb. nov. ‘Phoma glaucispora’ 

79 

CBS 296.51 Pleospora angustis nom. nov. 

CBS 314.80 Pleospora flavigena comb. nov. ‘Phoma flavigena’ 

CBS 260.94 Plenodomus lingam ‘Leptosphaeria maculans’, Phoma lingam’ 

CBS 147.24 Plenodomus lingam ‘Leptosphaeria maculans’, Phoma lingam’ 

0.1 

Fig. 4. The phylogeny of Phoma-like anamorphs in the Pleosporaceae based on the strict consensus tree from a Bayesian analysis of 20 ACT sequences. The Bayesian 

posterior probabilities are given at the nodes. The tree was rooted to Plenodomus lingam (CBS 147.24, 260.94). 

16

Fig 5 


57 

97 

82 

CBS 101461 Paraconiothyrium maculicutis sp. nov. Pleurophoma pleurospora 

CBS 265.94 Paraconiothyrium tiliae comb. nov. Asteromella tiliae 

CBS 797.95 Paraconiothyrium fuckelii comb. nov. Coniothyrium fuckelii 

99 

97 

94 

CBS 116.16 Paraconiothyrium fusco-maculans comb. nov. Plenodomus fusco-maculans 

Montagnulaceae (A) 

CBS 253.92 Paraconiothyrium lini comb. nov. Phoma.lini 

CBS 122788 Paraconiothyrium minitans 

CBS 122786 Paraconiothyrium minitans 

CBS 652.86 Paraphaeosphaeria michotii 

55 94 

99 

CBS 473.64 Massarina eburnea 

H 3953 Massarina eburnea 

CBS 331.37 Neottiosporina paspali 

CBS 675.92 Byssothecium circinans 

Massarinaceae (B) 

83 

62 

91 

99 

99 

100 

CBS 116668 Pleurophoma pleurospora 

PD 82.371 Pleurophoma pleurospora 

CBS 178.93 Paraconiothyrium flavescens comb. nov. Phoma flavescens 

CBS 252.60 Medicopsis romeroi comb. nov. Pyrenochaeta romeroi 

CBS 122784 Medicopsis romeroi comb. nov. Pyrenochaeta romeroi 

Trematosphaeriaceae (C) 

BCC 21118 Falcisormispora lignatilis 

CBS 122368 Trematosphaeria pertusa 

CBS 221.37 Massaria platani 

Massariaceae 

CBS 419.85 Thyridaria rubronotata 

CBS 350.82 Aposphaeria populina 

99 

PD 84.221 Aposphaeria populina 

64 

92 

99 

98 

86 

CBS 543.70 Aposphaeria populina 

CBS 400.97 Melanomma pulvis-pyrius 

CBS 371.75 Melanomma pulvis-pyrius 

CBS 200.31 Herpotrichia juniperi 

PD 83.367 Aposphaeria corallinolutea sp. nov. 

Melanommataceae (D) 

PD 83.831 Aposphaeria corallinolutea sp. nov. 

CBS 283.53 Beverwykella pulmonaria 

73 

99 

99 

99 

92 

CBS 279.74 Pleomassaria siparia 

CBS 337.65 Westerdykella capitulum comb. nov. Phoma capitulum 

CBS 379.55 Westerdykella ornata 

Sporormiaceae (E) 

CBS 509.91 Westerdykella minutispora comb. nov. Phoma minutispora 

CBS 524.50 Sporormiella minima 

CBS 659.74 Preussia funiculata 

95 

CBS 125434 Roussoella hysterioides 

CBS 110022 Nigrograna mackinnonii comb. nov. Pyrenochaeta mackinnonii 

CBS 674.75 Nigrograna mackinnonii comb. nov. Pyrenochaeta mackinnonii 

Didymosphaeriaceae 

CBS 398.61 Pseudorobillarda phragmitis 

0.1 

Fig. 5. LSU The phylogeny of Phoma-like isolates excluded from the Pleosporineae, based on the strict consensus tree from a Bayesian analysis of 40 LSU sequences. The 

Bayesian posterior probabilities are given at the nodes. The tree was rooted to Pseudorobillarda phragmitis (CBS 398.61). 


17


D with Aposphaeria populina and Melanomma pulvis-pyrius in 

Melanommataceae. This isolate is considered as an incorrect 

identification (Mugambi & Huhndorf 2009), and we consider this 

sterile isolate as representative of Melanomma pulvis-pyrius. 

Clade C also comprises the human pathogen Pyrenochaeta 

romeroi. This species certainly does not belong to Pyrenochaeta 

(de Gruyter et al. 2010) and therefore, we describe the new 

genus Medicopsis in Trematosphaeriaceae to accommodate this 

species. 

A well-supported clade D represents the Melanommataceae 

and includes Melanomma pulvis-pyrius, Herpotrichia juniperi and 

Beverwijkella pulmonaria, in congruence with Zhang et al. (2009). 

There were four Phoma-like isolates present in the collections of CBS 

and PD, i.e. CBS 350.82, PD 83/367, PD 83/831 and PD 84/221, 

which could not be identified according to their morphological 

characters. The isolates were preserved as Pleurophoma spp. 

This study demonstrates that two strains represent Aposphaeria 

populina, whereas the other two strains represent the new species 

described here as Aposphaeria corallinolutea. Further studies 

in Melanommataceae are needed to clarify the phylogeny of 

Aposphaeria in Melanommataceae. 

Sporormiaceae (clade E) is represented by Sporormiella 

minima and Preussia funiculata. Phoma capitulum and Ph. 

minutispora, well-defined soil-borne fungi from Asia, group in this 

clade. Both species are related with the anamorph Westerdykella 

ornata, and therefore the species are transferred to Westerdykella 

in Sporormiaceae. 

Pyrenochaeta mackinnonii could not be assigned to familial 

rank. A blast search in GenBank with its LSU sequence suggested 

a relation with Versicolorisporum triseptum. However, the typical 

3-septate conidia of this anamorph are different. Neither could V. 

triseptum be assigned at familial rank in Pleosporales (Tanaka et 

al. 2009). We therefore introduce the new genus Nigrograna to 

accommodate Py. mackinnonii. 

TAXONOMY 

Leptosphaeriaceae M.E. Barr, Mycotaxon 29: 503. 1987. 

Heterospora (Boerema, Gruyter & Noordel.) Gruyter, Verkley 

& Crous, stat. nov. MycoBank MB564701. 

Basionym: Phoma sect. Heterospora Boerema, Gruyter & Noordel., 

Persoonia 16: 336. 1997. 

Type species: Heterospora chenopodii (Westend.) Gruyter, 

Aveskamp & Verkley, see below (= Phoma heteromorphospora Aa 

& Kesteren). 

Heterospora chenopodii (Westend.) Gruyter, Aveskamp & 

Verkley, comb. nov. MycoBank MB564702. 

Basionym: Phyllosticta chenopodii Westend., Bull. Acad. Roy. Sci. 

Belgique Ser. 2, 2: 567. 1857; not Phyllosticta chenopodii Sacc., 

Syll. Fung. 3: 55. 1884 = Phoma exigua Desm. var. exigua; not 

Plenodomus chenopodii (P. Karst. & Har.) Arx, Verh. Kon. Ned. 

Akad. Wetensch., Afd. Natuurk., Sect. 2. 51: 72. 1957 ≡ Phoma 

chenopodiicola Gruyter, Noordel. & Boerema, Persoonia 15: 395. 

1993; not Phoma chenopodii Pavgi & U.P. Singh, Mycopathol. 

Mycol. Appl. 30: 265. 1966. nom. illeg. = Phoma chenopodii S. 

Ahmad, Sydowia 2: 79. 1948. 

≡ Septoria westendorpii G. Winter, Hedwigia 26: 26. 1887. nom. nov.; not 

Phoma westendorpii Tosquinet, Westend., Bull. Acad. Roy. Sci. Belgique 

Ser. 2, 2: 564. 1857. 

≡ Phoma variospora Aa & Kesteren, Persoonia 10: 268. 1979. nom. nov., 

nom. illeg.; not Phoma variospora Shreem., Indian J. Mycol. Pl. Pathol. 8: 

221. 1979 (“1978”). 

≡ Phoma heteromorphospora Aa & Kesteren, Persoonia 10: 542. 1980. 

nom. nov. 

Specimens examined: Belgium, Beverloo, from leaves of Chenopodium suecicum 

(album) and Chenopodium urbicum (Chenopodiaceae), no date, G.D. Westendorp, 

Herb. Crypt. (Ed. Beyaert-Feys), No. 959. BR, holotype of Phyllosticta chenopodii 

Westend. ex herb. G.D. Westendorp. Netherlands, Baarn, from leaf spots in 

Chenopodium album, 3 Jul. 1968, H.A. van der Aa, epitype designated here CBS 

H-16386, culture ex-epitype CBS 448.68; Heelsum, from leaf spots in Chenopodium 

album, Sep. 1994, J. de Gruyter, CBS 115.96 = PD 94/1576. 

Notes: Van der Aa & van Kesteren (1979) provided a nom. nov. 

since the epithet “chenopodii” was occupied in Phoma. For more 

details of the taxonomy of the species see van der Aa & van 

Kesteren (1979). Although Leptosphaeria chenopodii-albi was 

described from leaves of Chenopodium album (Crane & Shearer 

1991) no cultures are available for comparison. 

Heterospora dimorphospora (Speg.) Gruyter, Aveskamp & 


Basionym: Phyllosticta dimorphospora Speg., Anales Mus. Nac. 

Buenos Aires 13: 334. 1910. 

≡ Phoma dimorphospora (Speg.) Aa & Kesteren, Persoonia 10: 269. 

1979. 

= Stagonospora chenopodii Peck, Rep. (Annual) New York State Mus. Nat. 

Hist. 40: 60. 1887. (sometimes erroneously listed as Stag. chenopodii “House”). 

Specimens examined: Argentina, La Plata, from leaves of Chenopodium hircinum 

(Chenopodiaceae), 13 Oct. 1906, C. Spegazzini, Colect. micol. Museo Inst. 

Spegazzini, No. 11.353, LPS, holotype of Phyllosticta dimorphospora Speg. 

Lima, from stem of Chenopodium quinoa, 1977, L.J. Turkensteen, CBS 165.78 = 

PD 77/884. Peru, from lesions in stems of Chenopodium quinoa, 1976, V. Otazu, 

epitype designated here CBS H-16203, culture ex-epitype CBS 345.78 = PD 

76/1015. 

Note: For more details of the taxonomy of the species see van der 

Aa & van Kesteren (1979). 

Leptosphaeria Ces. & De Not., Comment. Soc. Crittog. Ital. 

1: 234. 1863. 

= Leptophoma Höhn., Sitzungsber. Kaiserl. Akad. Wiss., Math.-Naturwiss. Cl., 

Abt. 1. 124: 73. 1915. 

Type species: Leptosphaeria doliolum (Pers. : Fr.) Ces. & De Not., 

see below. 

Note: For full synonymy, including the species listed below, see 

Crane & Shearer (1991) and Boerema et al. (2004). 

Leptosphaeria conoidea (De Not.) Sacc., Fungi Venet. Nov. 

Vel. Crit. Ser. 2: 314. 1875. 

Basionym: Leptosphaeria doliolum var. conoidea De Not., Mycoth. 

Veneti, No. 76. 1873. 

= Leptosphaeria doliolum subsp. pinguicula Sacc., Michelia 2: 598. 1882. 

= Phoma acuta subsp. amplior Sacc. & Roum., Rev. Mycol. 6: 30. 1884. 

≡ Phoma hoehnelii subsp. amplior (Sacc. & Roum.) Boerema & Kesteren, 

Trans. Brit. Mycol. Soc. 67: 299. 1976. 

= Phoma doliolum P. Karst., Meddel. Soc. Fauna Fl. Fenn. 16: 9. 1888. 

= Plenodomus microsporus Berl., Bull. Soc. Mycol. France 5: 55. 1889. 

Specimens examined: Netherlands, Zaltbommel, from dead stem of Lunaria annua 

(Brassicaceae), Jan. 1974, G.H. Boerema, CBS 616.75 = ATCC 32813 = IMI 199777 

= PD 74/56; Montfoort, Senecio sp. (Asteraceae), 1982, CBS 125977 = PD 82/888. 

18


Leptosphaeria doliolum (Pers. : Fr.) Ces. & de Not., 

Comment. Soc. Crittog. Ital. 1: 234. 1863. 

Basionym: Sphaeria doliolum Pers. : Fr., Icon. Desc. Fung. Min. 

Cognit. (Leipzig) 2: 39. 1800. 

= Sphaeria acuta Hoffm. : Fr, Veg. cryptog. 1: 22. 1787. Syst. Mycol. 2: 507. 

1823. 

≡ Phoma acuta (Hoffm. : Fr.) Fuckel, Jahrb. Nassauischen Vereins 

Naturk. 23–24: 125. 1870 (as “acutum”). 

≡ Leptophoma acuta (Hoffm. : Fr.) Höhn., Sitzungsber. Kaiserl. Akad. 

Wiss., Math.-Naturwiss. Cl., Abt. 1. 124: 73. 1915. 

≡ Plenodomus acutus (Hoffm. : Fr.) Bubák, Ann. Mycol. 13: 29. 1915. 

[as “(Fuckel)”]. 

= Phoma phlogis Roum., Rev. Mycol. 6: 160. 1884. 

= Phoma hoehnelii var. urticae Boerema & Kesteren, Trans. Brit. Mycol. Soc. 

67: 299. 1976. 

Specimens examined: Netherlands, from stem of Rudbeckia sp. (Asteraceae), 

Sep. 1966, M.M.J. Dorenbosch, CBS 541.66 = PD 66/221; from stem of Urtica 

dioica (Urticaceae), 1974, G.H. Boerema, CBS 504.75 = PD 74/55; Rhenen, from 

Urtica dioica, Feb. 1975, G.H. Boerema, CBS 505.75 = PD 75/141; Wageningen, 

from stem of Phlox paniculata (Polemoniaceae), 1977, G.H. Boerema, CBS 155.94 

= PD 77/80; from stem of Phlox paniculata, 1978, G.H. Boerema, CBS 125979 = PD 

78/37; from stem of Urtica dioica, 1982, G.H. Boerema, CBS 130000 = PD 82/701. 

Notes: Isolate CBS 541.66 was preserved as Phoma acuta subsp. 

errabunda (teleom. Leptosphaeria errabunda, see below); however, 

the isolate clustered with L. doliolum. Both isolates CBS 155.94 

and CBS 125979 were considered as forma specialis “phlogis” 

(Boerema et al. 1994) of the anamorph Ph. acuta subsp. acuta. 

The subspecies acuta was created by the differentiation of Phoma 

acuta subsp amplior Sacc. & Roum., but the latter is a synonym of 

Ph. doliolum, reclassified here as L. conoidea, see above. Sphaeria 

acuta Hoffm. was applied as basionym for different anamorphs an 

a teleomorph of various species of Leptosphaeria leading to a 

confusing nomenclature. The epitet has been unambiguously tied 

to Ph. acuta by Boerema & Gams (1995). 

Leptosphaeria errabunda (Desm.) Gruyter, Aveskamp & 


Basionym: Phoma errabunda Desm., Ann. Sci. Nat., Bot. Ser. 3, 

11: 282. 1849. 

≡ Phoma acuta subsp. errabunda (Desm.) Boerema, Gruyter & Kesteren, 

Persoonia 15: 465. 1994. 

= Leptophoma doliolum Höhn., Sitzungsber. Kaiserl. Akad. Wiss., Math.- 

Naturwiss. Cl., Abt. 1. 124: 75. 1915; not Phoma doliolum P. Karst. = 

Leptosphaeria conoidea (De Not.) Sacc., see above. 

≡ Plenodomus doliolum (Höhn.) Höhn., Ber. Deutsch. Bot. Ges. 36: 139. 

1918. 

≡ Phoma hoehnelii Kesteren, Netherlands J. Pl. Pathol. 78: 116. 1972. 

nom. nov. 

= Leptosphaeria doliolum subsp. errabunda Boerema, Gruyter & Kesteren, 

Persoonia 15: 466. 1994. 

Specimens examined: Netherlands, Leeuwarden, from stem of Delphinium sp. 

(Ranunculaceae), 1974, CBS 125978 = PD 74/61; Ferwerderadeel, from Solidago 

sp., hybrid (Asteraceae), Mar. 1974, G.H. Boerema, CBS 617.75 = ATCC 32814 = IMI 

199775 = PD 74/201; from stem of Aconitum sp. (Ranunculaceae), CBS 129999 = PD 

78/569; from stem of Achillea millefolium (Asteraceae), CBS 129997 = PD 78/631; 

from Gailardia sp. (Asteraceae), 1984, G.H. Boerema, CBS 129998 = PD 84/462. 

Notes: The isolate CBS 617.75 = ATTC 32814 was deposited as 

the anamorph Ph. hoehnelii var. hoehnelii, but interpreted as L. 

doliolum subsp. conoidea (Dong et al. 1998). The isolate clustered 

with L. errabunda in this study. 

Leptosphaeria etheridgei (L.J. Hutchison & Y. Hirats.) 

Gruyter, Aveskamp & Verkley, comb. nov. MycoBank 

MB564712. 

Basionym: Phoma etheridgei L.J. Hutchison & Y. Hirats., Canad. J. 

Bot. 72: 1425. 1994. 

Specimen examined: Canada, Alberta, from bark of gall, on trunck of Populus 

tremuloides (Salicaceae), Jul. 1989, P. Crane, holotype DAOM 216539, culture exholotype 

DAOM 216539 = CBS 125980 = PD 95/1483. 

Leptosphaeria macrocapsa (Trail) Gruyter, Aveskamp & 


Basionym: Phoma macrocapsa Trail, Scott. Naturalist (Perth) 8: 

327. 1886. 

≡ Plenodomus macrocapsa (Trail) H. Ruppr., Sydowia 13: 20. 1959. 

Specimen examined: Netherlands, from stem of Mercurialis perennis 

(Euphorbiaceae), 1978, G.H. Boerema, CBS 640.93 = PD 78/139. 

Leptosphaeria pedicularis (Fuckel) Gruyter, Aveskamp & 


Basionym: Phoma pedicularis Fuckel, Reisen Nordpolarmeer 

3: 318. 1874 (as “pedicularidis”); not Phoma pedicularis Wehm., 

Mycologia 38: 319. 1946 (= Phoma herbicola Wehm). 

= Sphaeronaema gentianae Moesz, Bot Közlem. 14: 152. 1915 (as 

“Sphaeronema”). 

≡ Plenodomus gentianae (Moesz) Petr., Ann. Mycol. 23: 54. 1925. 

Specimens examined: Switzerland, Kanton Graubünden, Albulapass, from dead 

stem of Pedicularis sp. (Scrophulariaceae), 1977, CBS 390.80 = PD 77/711 = ATCC 

42535 = IMI 248430; Zürich, from Gentiana punctata (Gentianaceae), 1977, CBS 

126582 = PD 77/710. 

Leptosphaeria rubefaciens (Togliani) Gruyter, Aveskamp & 


Basionym: Phoma rubefaciens Togliani, Ann. Sper. Agr. II, 7: 1626. 

1953. 

Specimens examined: Switzerland, Zürich, Albis, from twig of Quercus sp. 

(Fagaceae), Aug. 1976, W. Gams, CBS 223.77. Netherlands, Oploo, from wood of 

Tilia (×) europaea (Tiliaceae), 1978, G.H. Boerema, CBS 387.80 = ATCC 42533 = 

IMI 248432 = PD 78/809. 

Leptosphaeria sclerotioides (Sacc.) Gruyter, Aveskamp & 


Basionym: Phoma sclerotioides Sacc., Fungi Herb. Bruxelles 21. 

1892; Syll. Fung. 11: 492. 1895. 

= Plenodomus sclerotioides Preuss, Klotzsch. Herb. Vivum Mycol. Sistems 

Fungorum German., No. 1281. 1849. nom. nud. (no description). 

= Plenodomus meliloti Mark.-Let., Bolezni Rast. 16: 195. 1927. 

Specimens examined: Canada, British Columbia, from Medicago sativa 

(Fabaceae), 1980, J. Drew Smith, CBS 148.84 = PD 80/1242; Alberta, from root 

of Medicago sativa, Mar. 1984, G.H. Boerema, CBS 144.84 = CECT 20025 = 

PD 82/1061. 

Note: Seven varieties of this species have been recognised 

(Wunsch et al. 2011) in a phylogenetic analysis using 10 loci. 

Leptosphaeria slovacica Picb., Sborn. Vysoké Skoly. 

Zemed. v Brno 7: 7. 1927. 

= Phoma leonuri Letendre, Revue Mycol. 6: 229. 1884. 

≡ Plenodomus leonuri (Letendre) Moesz & Smarods in Moesz, Magyar 

Bot. Lapok 31: 38. 1932. 

Specimens examined: Netherlands, from dead stem of Ballota nigra (Lamiaceae), 

1977, CBS 125975 = PD 77/1161; Arnhem, from dead stem of Ballota nigra, 1979, 

G.H. Boerema, CBS 389.80 = PD 79/171. 


19


Leptosphaeria sydowii (Boerema, Kesteren & Loer.) 


MB564717. 

Basionym: Phoma sydowii Boerema, Kesteren & Loer., Trans. Brit. 

Mycol. Soc. 77: 71. 1981. nom. nov. 

= Sphaeronaema senecionis Syd. & P. Syd., Ann. Mycol. 3: 185. 1905; not 

Phoma senecionis P. Syd., Beibl. Hedwigia 38: 136. 1899. 

≡ Plenodomus senecionis (Syd. & P. Syd.) Bubák, Ann. Mycol. 13: 29. 

1915. 

≡ Plenodomus senecionis (Syd. & P. Syd.) Petr., Ann. Mycol. 19: 192. 

1921. Isonym. 

= Plenodomus rostratus Petr., Ann. Mycol. 21: 199. 1923; not Phoma rostrata 

O’Gara, Mycologia 7: 41. 1915; not Leptosphaeria rostrata M.L. Far & H.T. 

Horner, Nova Hedwidgia 15: 250. 1968. 

Specimens examined: Switzerland, Kt. Zürich, Zollikon, from Papaver rhoeas 

(Papaveraceae), Oct. 1949, E. Müller, CBS 297.51. Netherlands, from Senecio 

jacobaea (Asteraceae), G.H. Boerema, 1984, CBS 125976 = PD 84/472. UK, 

Scotland, Isle of Lewis, Hebrides, from dead stem of Senecio jacobaea, 1974, 

R.W.G. Dennis, CBS 385.80 = PD 74/477. 

Notes: Leptosphaeria senecionis (Fuckel) G. Winter was suggested 

as the possible teleomorph (Boerema et al. 2004). Because the 

teleomorph connection has not been proven, however, we did not 

include it as a synonym that would have priority as the correct 

name. The isolate CBS 297.51 was originally identified as L. 

doliolum var. doliolum. 

Leptosphaeria veronicae (Hollós) Gruyter, Aveskamp & 


Basionym: Sphaeronaema veronicae Hollós, Ann. Hist.-Nat. Mus. 

Natl. Hung. 4: 341. 1906. 

≡ Phoma veronicicola Boerema & Loer., Trans. Brit. Mycol. Soc. 84: 297. 

1985. nom. nov.; not Phoma veronicae Roum., Revue Mycol. 6: 160. 1884. 

Specimens examined: Netherlands, from stem of Veronica “Shirley Blue” 

(Scrophulariaceae), 1974, CBS 126583 = PD 74/227; Huis ter Heide, from dead 

stem of Veronica chamaedryoides, Mar. 1978, H.A. van Kesteren, neotype CBS 

H-7632, culture ex-neotype CBS 145.84 = CECT 20059 = PD 78/273. 

Paraleptosphaeria Gruyter, Verkley & Crous, gen. nov. 

MycoBank MB564720. 

Pseudothecia immersed, subglobose, solitary or aggregated, 

thick-walled, pseudoparenchymatous to scleroplectenchymatous, 

ostiolate, unilocular. Asci bitunicate, broadly ellipsoidal, 8-spored, 

interascal filaments pseudoparaphyses, Ascospores biseriate, 

broadly fusiform, transversally 3–5-septate, hyaline to yellowbrownish. 

Conidiomata pycnidial, globose to subglobose, 

scleroplectenchymatous, with papillate pore, unilocular. 

Conidiogenous cells phialidic, ampulliform to doliiform. Conidia 

hyaline, aseptate, oblong to ellipsoidal. Sclerotia sometimes 

produced. 

Type species: Paraleptosphaeria nitschkei (Rehm ex G. Winter) 

Gruyter, Aveskamp & Verkley (see below). 

Notes: Munk (1957) recognised Leptosphaeria section Para- 

Leptosphaeria, an invalid taxon, as a heterogenous group. The 

section was differentiated from Eu-Leptosphaeria, which included 

the generic type species L. doliolum. Leptosphaeria nitschkei was 

considered a typical representative of section Eu-Leptosphaeria 

(Müller & von Arx 1950). However, this molecular phylogeny 

demonstrates that L. nitschkei is only distantly related to L. doliolum. 

We introduce Paraleptosphaeria to accomodate L. nitschkei and its 

relatives. These necrotrophic species are morphologically closely 

allied to Leptosphaeria. The former classification of Leptosphaeria 

in sections Eu-Leptosphaeria and Para-Leptosphaeria cannot be 

upheld from a evolutionary point of view, as two other species 

attributed to section Eu-Leptosphaeria, namely L. agnita and L. 

maculans (Munk 1957), were found to group in Plenodomus. 

Paraleptosphaeria dryadis (Johanson) Gruyter, Aveskamp 

& Verkley, comb. nov. MycoBank MB564721. 

Basionym: Melanomma dryadis Johanson, Hedwigia 29: 160. 

1890. 

≡ Leptosphaeria dryadophila Huhndorf, Bull. Illinois Nat. Hist. Surv. 34: 

484 (1992). nom. illeg. via nom. superfl. 

= Leptosphaeria dryadis Rostr., Bot. Tidsskr. 25: 305. 1903. 

Specimen examined: Switzerland, Kt. Ticino, Leventina, Alpe Campolungo, from 

Dryas octopetala (Rosaceae), 24 July 1980, A. Leuchtmann, CBS 643.86. 

Note: An explanation of the nomenclature of Leptosphaeria dryadis 

has been provided by Chen et al. (2002). 

Paraleptosphaeria macrospora (Thüm.) Gruyter, Aveskamp 


Basionym: Leptosphaeria macrospora Thüm. Mycotheca Univ. 

1359. 1879. nom. nov. 

≡ Metasphaeria macrospora (Fuckel) Sacc., Syll. Fung. 2: 158. 1883. 

Replaced synonym: Pleospora macrospora Fuckel, Jahrb. 

Nassauischen Vereins Naturk. 23–24: 138. 1870. nom. illeg., 

Art. 53.1.; not Pleospora macrospora (De Not.) Ces. & De Not., 

Comment. Soc. Crittog. Ital. 1: 218. 1863. 

Specimen examined: Norway, Troms, Tromsöya, from Rumex domesticus 

(Polygonaceae), 20 Aug. 1988, K. & L. Holm, CBS 114198 = UPSC 2686. 

Paraleptosphaeria nitschkei (Rehm ex G. Winter) Gruyter, 

Aveskamp & Verkley, comb. nov. MycoBank MB564723. 

Basionym: Leptosphaeria nitschkei Rehm ex G. Winter, 

Ascomyceten, Fascicle 1, No. 15. 1870. nom. nud.; Flora, Jena 

und Regensburg 55: 510. 1872. 

Specimens examined: Austria, Ötscher in Niederösterreich, c. 4500’, from Cacalia 

sp. (= Adenostyles sp, Asteraceae), June 1869, Lojka, holotype of Leptosphaeria 

nitschkei Rehm Ascomyceten 15b, S. Switzerland, Kt. Graubünden, Lü, from 

Cirsium spinosissimum (Asteraceae), 16 July 1948, E. Müller, epitype designated 

here CBS H-20822, culture ex-epitype CBS 306.51. 

Note: The name Leptosphaeria nitschkei was considered a nom. 

nud. by Crane and Shearer (1991) who cited Art. 32.1 but gave no 

further explanation. In Flora, Jena und Regensburg 55: 510. 1872 

Rehm refers to additional notes by G. Winter that include a Latin 

description. Therefore, we consider this name as valid, following 

Müller (1950) who provided a detailed description in vivo. 

Paraleptosphaeria orobanches (Schweinitz : Fr.) Gruyter, 


Basionym: Sclerotium orobanches Schweinitz, Schriften Naturf. 

Ges. Leipzig 1: 57. 1822 : Fr., Syst. Mycol. 2: 257. 1822. 

= Phoma korfii Boerema & Gruyter, Persoonia 17: 275. 1999. 

Specimen examined: USA, Ringwood Swamp, Lloyd-Cornell, from stem of Epifagus 

virginiana (Orobanchaceae), 13 Sep. 1995, T. Uturriaga, R.P. Korf, P. Mullin, 

holotype of Sclerotium orobanches Schweinitz, CUP 63537, culture ex-holotype 

CBS 101638 = PD 97/12070. 

20


Note: A Phoma synanamorph of Sclerotium orobanches was 

reported by Yáňez-Morales et al. (1998) and described as Phoma 

korfi (Boerema & Gruyter 1999). 

Paraleptosphaeria praetermissa (P. Karst.) Gruyter, 


Basionym: Sphaeria praetermissa P. Karst., Bidrag Kannedom 

Finlands Natur Folk 23: 89. 1873. 

≡ Leptosphaeria praetermissa (P. Karst.) Sacc., Syll. Fung. 2: 26. 1883. 

Specimen examined: Sweden, Dalarna, Folkärna, from Rubus idaeus (Rosaceae), 

21 Mar. 1993, K. & L. Holm, CBS 114591. 

Plenodomus Preuss, Linnaea 24: 145. 1851. 

≡ Phoma sect. Plenodomus (Preuss) Boerema, Kesteren & Loer., Trans. 

Brit. Mycol. Soc. 77: 61. 1981. 

= Diploplenodomus Diedicke, Ann. Mycol. 10: 140. 1912. 

= Plectophomella Moesz, Magyar Bot. Lapok 21: 13. 1922. 

= Apocytospora Höhn., Mitt. Bot. Lab. TH Wien 1: 43. 1924. 

= Deuterophoma Petri, Boll. R. Staz. Patalog. Veget. Roma 9: 396. 1929. 

Type species: Plenodomus rabenhorstii Preuss, Linnaea 24: 145. 

1851 (dubious synonym, see below) = Plenodomus lingam (Tode : 

Fr.) Höhn., see below. 

Note: For full synonymy of the anamorph names of the species 

listed below, see Boerema et al. (1994). For additional synonyms 

of the teleomorph names of the species below that have been 

recorded on Asteraceous hosts, see Khashnobish et al. (1995). 

Plenodomus agnitus (Desm.) Gruyter, Aveskamp & Verkley, 

comb. nov. MycoBank MB564726. 

Basionym: Sphaeria agnita Desm., Ann. Sci. Nat., Bot. Ser. 3, 16: 

313. 1851. 

≡ Leptosphaeria agnita (Desm.) Ces. & De Not., Comm. Soc. Crittog. Ital. 

1: 236. 1863. 

= Plenodomus chondrillae Died, Ann. Mycol.. 9: 140. 1911; Krypt.-fl. 

Brandenburg 9: 236. 1912. 

= Phoma agnita Gonz. Frag., Mem. Real Acad. Ci. Barcelona 15: 6. 1920. 

Specimens examined: Netherlands, from stem of Eupatorium cannabinum 

(Asteraceae), 1982, W.M. Loerakker, CBS 126584 = PD 82/561; from stem of 

Eupatorium cannabinum, 1982, W.M. Loerakker, CBS 121.89 = PD 82/903. 

Plenodomus biglobosus (Shoemaker & H. Brun) Gruyter, 


Basionym: Leptosphaeria biglobosa Shoemaker & H. Brun, Canad. 

J. Bot. 79: 413. 2001. 

Specimens examined: France, Le Rheu, from stem of Brassica juncea 

(Brassicaceae), CBS 127249 = DAOM 229269. Netherlands, from Brassica rapa 

(Brassicaceae), 2006, R. Veenstra, CBS 119951. 

Notes: Leptosphaeria biglobosa was originally described as a less 

virulent segregate of L. maculans (Shoemaker & Brun 2001). The 

species, also indicated as Tox 0 isolates, has been described from 

cultivated Brassica species as the cause of upper stem lesions 

and considered as less damaging than L. maculans (West et al. 

2002). However, in Poland L. biglobosa is the predominant cause 

of these symptoms (Jedryczka et al. 1999, Huang et al. 2005). The 

current species concept of L. biglobosa is broadly defined with six 

distinct subclades recognised by multilocus phylogenetic analyses 

of ITS, β-tubulin and actin sequences (Mendes-Pereira et al. 2003, 

Vincenot et al. 2008). These subclades are named after the host or 

geographic origin of the isolates involved. It has been suggested 

that the clades represent distinct subspecies formed over time by 

reproductive isolation (Mendes-Pereira et al. 2003). Alignments of 

the ITS sequences of Ph. wasbiae, Ph. pimpinellae and L. biglobosa 

isolates were compared with those of the representative strains 

of the L. biglobosa subclades obtained from GenBank, and both 

Ph. wasbiae and Ph. pimpinellae grouped in this species complex 

(unpubl. data). Both species are maintained here, awaiting a 

redescription of the taxa representing all clades in the L. biglobosa 

complex. 

Plenodomus chrysanthemi (Zachos, Constantinou & 

Panag.) Gruyter, Aveskamp & Verkley, comb. nov. MycoBank 

MB564728. 

Basionym: Cephalosporium chrysanthemi Zachos, Constantinou & 

Panag., Ann. Inst. Phytopath. Benaki, N.S. 55. 1960. 

≡ Phialophora chrysanthemi (Zachos, Constantinou & Panag.) W. Gams, 

Cephalosporium-artige Schimmelpilze (Stuttgart): 207. 1971. 

= Phoma vasinfecta Boerema, Gruyter & Kesteren, Persoonia 15: 484. 1994. 

Specimen examined: Greece, from Chrysanthemum sp. (Asteraceae), Apr. 1963, 

D.G. Zachos, holotype CBS H-7576, culture ex-holotype CBS 539.63. 

Note: The species was also described as Phoma tracheiphila f. sp. 

chrysanthemi (Baker et al. 1985). 

Plenodomus collinsoniae (Dearn. & House) Gruyter, 


Basionym: Leptosphaeria collinsoniae Dearn. & House, Bull. New 

York State Mus. Nat. Hist. 233–234: 36. 1921. 

Specimen examined: Japan, Osawa river, Komukai, Miyagi, from Vitis coignetiae 

(Vitaceae), 27 Sep. 2003, Y. Takahashi, CBS 120227 = JCM 13073 = MAFF 239583. 

Plenodomus confertus (Niessl ex Sacc.) Gruyter, Aveskamp 


Basionym: Leptosphaeria conferta Niessl ex Sacc., Syll. Fung. 2: 

20. 1883. 

= Phoma conferta P. Syd. ex Died., Krypt.-fl. Brandenburg 9: 142. 1912. 

Specimen examined: Spain, Cais do Tejo, from dead stem of Anacyclus radiatus 

(Asteraceae), Mar. 1961, M.T. Lucas, CBS 375.64. 

Plenodomus congestus (M.T. Lucas) Gruyter, Aveskamp & 


Basionym: Leptosphaeria congesta M.T. Lucas, Trans. Brit. Mycol. 

Soc. 46: 362. 1963. 

= Phoma congesta Boerema, Gruyter & Kesteren, Persoonia 15: 461. 1994. 

Specimen examined: Spain, Póvoa de Santa Iria, Estremadura, from stem 

of Erigeron canadensis (Asteraceae), Mar. 1961, M.T. Lucas, holotype of 

Leptosphaeria congesta M.T. Lucas, dried culture LISE 1638, culture ex-holotype 

CBS 244.64. 

Plenodomus enteroleucus (Sacc.) Gruyter, Aveskamp & 


Basionym: Phoma enteroleuca Sacc. var. enteroleuca, Michelia 1: 

358. 1878. 

Specimens examined: France, Alencon, from Pyrus communis (Rosaceae), 1878, 

C. C. Gillet, holotype of Phoma enteroleuca var. enteroleuca, Herb. Sacc. ’19’, 

PAD. Germany, Monheim, from leaf spots of Triticum aestivum (Poaceae), 15 Aug. 

1984, M. Hossfeld, CBS H-3684, culture CBS 831.84. Netherlands, Bennekom, 

from discoloured wood of Catalpa bignonioides (Bignoniaceae), 1981, G.H. 

Boerema, epitype designated here CBS H-16209, culture ex-epitype CBS 142.84 

= PD 81/654 = CECT 20063. 


21


Plenodomus fallaciosus (Berl.) Gruyter, Aveskamp & 


Basionym: Leptosphaeria fallaciosa Berl., Bull. Soc. Mycol. France. 

5: 43. 1889. 

Specimen examined: France, Var, Ste. Baume, from Satureia montana (Lamiaceae), 

July 1951, E. Müller, CBS 414.62 = ETH 2961. 

Plenodomus hendersoniae (Fuckel) Gruyter, Aveskamp & 


Basionym: Cucurbitaria hendersoniae Fuckel, Symb. Myc. p. 172. 

1870. 

≡ Melanomma hendersoniae (Fuckel) Sacc., Syll. Fung. 2: 109. 1883. 

≡ Chiajaea hendersoniae (Fuckel) Höhn., Sitzungsber. Kaiserl. Akad. 

Wiss., Math.-Naturwiss. Cl., Abt. 1. 129: 152. 1920. 

≡ Leptosphaeria hendersoniae (Fuckel) L. Holm, Symb. Bot. Upsal. 14: 

26. 1957. 

= Phoma intricans M.B. Schwarz, Meded. Phytopath. Lab. Willie Commelin 

Scholten 8: 44. 1922. 

Specimens examined: Sweden, Uppland, Jerusalem, from Salix cinerea 

(Salicaceae), 10 Apr. 1986, K. & L. Holm, CBS 113702 = UPSC 1843. Netherlands, 

Wilhelminadorp, from bark of Pyrus malus (Rosaceae), June 1977, H.A.Th. van der 

Scheer, CBS 139.78. 

Plenodomus influorescens (Boerema & Loer.) Gruyter, 


Basionym: Phoma enteroleuca var. influorescens Boerema & Loer., 


Specimens examined: Netherlands, from Lilium sp. (Liliaceae), 1973, G.H. 

Boerema, PD 73/1382; Emmeloord, from Fraxinus excelsior (Oleaceae), 1978, J.D. 

Janse, holotype of Phoma enteroleuca var. influorescens, CBS H-16208, culture ex 

holotype CBS 143.84 = PD 78/883 = CECT 20064. 

Note: The isolate PD 73/1382 is no longer available for study. 

Plenodomus libanotidis (Fuckel) Gruyter, Aveskamp & 


Basionym: Pleospora libanotidis Fuckel, Jahrb. Nassauischen 

Vereins Naturk. 27–28: 24. 1873 as “libanotis”. 

≡ Leptosphaeria libanotidis (Fuckel) Sacc., Syll. Fung. 2: 16. 1883 as 

“libanotis”. 

= Phoma sanguinolenta Rostr., Tidsskr. Landokon. 5(7): 384. 1888; not 

Phoma sanguinolenta Grove, J. Bot. 23: 164. 1885. 

≡ Phoma rostrupii Sacc., Syll. Fung. 11: 490. 1895. nom. nov. 

Specimen examined: Sweden, Uppland, Gröna strand, from Seseli libanotis 

(Apiaceae), 19 May 1987, K. & L. Holm, CBS 113795 = UPSC 2219. 

Plenodomus lindquistii (Frezzi) Gruyter, Aveskamp & 


Basionym: Leptosphaeria lindquistii Frezzi, Revista Invest. 

Agropec., Sér. 5, 5: 79. 1968. 

= Phoma macdonaldii Boerema, Persoonia 6: 20. 1970. 

Specimens examined: Canada, from Helianthus annuus (Asteraceae), 1967, W.C. 

McDonald, CBS 381.67. Former Yugoslavia, from stem of Helianthus annuus, 

1977, A. Maric, CBS 386.80 = PD 77/336. 

Note: Strain CBS 381.67 is ex-holotype of Phoma macdonaldii 

Boerema, pycnidial state of Leptosphaeria lindquistii Frezzi 

(Boerema 1970). 

Plenodomus lingam (Tode : Fr.) Höhn., Sitzungsber. Kaiserl. 

Akad. Wiss., Math.-Naturwiss. Cl., Abt. 1. 120: 463. 1911. 

Basionym: Sphaeria lingam Tode : Fr., Fungi mecklenb. 2: 51. 

1791. : Fr., Syst. Mycol. 2: 507. 1823. 

≡ Phoma lingam (Tode : Fr.) Desm., Ann. Sci. Nat., Bot. Ser. 3, 11: 281. 

1849. 

= Sphaeria maculans Desm., Ann. Sci. Nat., Bot. Ser. 3, 6: 77. 1846. nom. 

illeg. 

≡ Leptosphaeria maculans (Desm.) Ces. & De Not., Comment. Soc. 

Crittog. Ital. 1: 235. 1863. 

= Plenodomus rabenhorstii Preuss, Linnaea 24: 145. 1851. nom. dub. 

Specimens examined: Netherlands, near Goes, from Brassica oleracea 

(Brassicaceae), 1978, M.M.J. Dorenbosch, CBS 260.94 = PD 78/989. Origin 

unknown, Mar. 1924, A. Weber, CBS 147.24. UK, from Brassica sp. (Brassicaceae), 

1963, B.C. Sutton, CBS 275.63 = MUCL 9901= UPSC 1025. 

Notes: The combination Plen. lingam as published by van Höhnel 

(1911) was preferred over Plen. rabenhorstii Preuss (1851) by 

Boerema & van Kesteren (1964) because the type material of 

Plen. rabenhorstii had been lost during the Second World War. 

Therefore, Plen. rabenhorstii is indicated here as a nomen 

dubium. Leptosphaeria maculans causes a serious stem base 

canker (blackleg) on cultivated Brassica spp. (Brassicaceae) in 

Europe, Australia and North America (West et al. 2001, Fitt et al. 

2006). 

Plenodomus lupini (Ellis & Everh.) Gruyter, Aveskamp & 


Basionym: Phoma lupini Ellis & Everh., Bull. Washburn Lab. Nat. 

Hist. 1: 6. 1884. 

≡ Asteromella lupini (Ellis & Everh.) Petr., Sydowia 9: 495. 1955; not 

Phoma lupini N.F. Buchw., Møller, Fungi Faeröes 2: 153. 1958. nom. illeg. 

Specimen examined: Peru, Andes region, from stem lesion of Lupinus mutabilis 

(Fabaceae), May 1992, J. de Gruyter, CBS 248.92 = PD 79/141. 

Plenodomus pimpinellae (Lowen & Sivan.) Gruyter, 


Basionym: Leptosphaeria pimpinellae Lowen & Sivan., Mycotaxon 

35: 205. 1989. 

= Phoma pimpinellae Boerema & Gruyter, Persoonia 17: 278. 1999. 

Specimen examined: Israel, Mt Carmel near Kibbutz Oren, from dead stems of 

Pimpinella anisum (Apiaceae), 9 Dec. 1987, R. Rowen, 523-88 NY, holotype of 

Leptosphaeria pimpinellae Lowen & Sivan, culture ex-holotype CBS 101637 = PD 

92/41. 

Plenodomus tracheiphilus (Petri) Gruyter, Aveskamp & 


Basionym: Deuterophoma tracheiphila Petri, Boll. Staz. Patol. Veg. 

Roma 9: 396. 1929. 

≡ Bakerophoma tracheiphila (Petri) Cif., Ist. Bot. Reale Univ. Reale Lab. 

Crittog. Pavia Atti Ser. 5, 5: 307. 1946. 

≡ Phoma tracheiphila (Petri) L.A. Kantsch. & Gikaschvili, Trudy Inst. 

Zasch. Rast. Tibilisi 5: 20. 1948. 

Specimens examined: Israel, from Citrus limonium (Rutaceae), Oct. 1993, J. de 

Gruyter, CBS 551.93 = PD 81/782. Italy, from Citrus sp. (Rutaceae), CBS 127250 

= PD 09/04597141. 

Note: The species produces a Phialophora-like synanamorph. 

Plenodomus visci (Moesz) Gruyter, Aveskamp & Verkley, 


Basionym: Plectophomella visci Moesz, Magyar Bot. Lapok 21: 13. 

1922. 

= Apocytospora visci Höhn., Mitt. Bot. Lab. TH Wien 1: 43. 1924. 

22


Specimen examined: Hungary, Tata-Tóváros, from leaves of Viscum album 

(Viscaceae), 22 Oct. 1911, G. von Moesz, BP, holotype of Plectophomella visci 

Moesz. France, from Viscum album, 1974, epitype designated here CBS H-20823, 

culture ex-epitype CBS 122783 = PD 74/1021. 

Notes: Plectophomella visci is the type species of the genus 

Plectophomella. This genus was accepted by Sutton (1980) based 

on the eustromatic conidiomata; branched, septate conidiophores, 

phialidic conidiogenesis and small, hyaline conidia. However, the 

phylogenetic analyses clearly demonstrated the placement of 

Plectophomella grouping in the Plenodomus clade and therefore it 

is treated as a synonym. 

Plenodomus wasabiae (Yokogi) J.F. White & P.V. Reddy, 

Canad. J. Bot. 76: 1920. 1999 (1998). 

Basionym: Phoma wasabiae Yokogi, Ann. Phytopathol. Soc. Japan 

2: 549. 1933. 

Specimens examined: Taiwan, from Wasabia japonica (syn. Eutrema wasabi) 

(Brassicaceae), A. Rossman, CBS 120119 = FAU 559; from Wasabia japonica, A. 

Rossman, CBS 120120 = FAU 561. 

Subplenodomus Gruyter, Verkley & Crous, gen. nov. 


Etymology: Although the genus resembles Plenodomus in the 

production of thick-walled pycnidia, the pycnidial cell wall of 

Subplenodomus often remains pseudoparenchymatous, similar to 

the pycnidial wall of species of Phoma. 

Conidiomata pycnidial, globose to papillate, or with an elongated neck, 

solitary or aggregated, thin-walled pseudoparenchymatous, or thickwalled 

scleroplectenchymatous, ostiolate, unilocular. Conidiogenous 

cells phialidic, ampulliform to doliiform. Conidia hyaline, aseptate, 

ellipsoid to cylindrical. Chlamydospores sometimes produced, 

olivaceous, unicellular in chains, or multicellular, dictyosporousbotryoid 

or forming pseudosclerotioid structures. 

Type species: Subplenodomus violicola (P. Syd.) Gruyter, 

Aveskamp & Verkley (see below) 

Subplenodomus apiicola (Kleb.) Gruyter, Aveskamp & 


Basionym: Phoma apiicola Kleb., Z. Pflanzenkrankh. 20: 22. 1910. 

Specimens examined: Germany, from tuber of Apium graveolens var. rapaceum 

(Apiaceae), Feb. 1972, Diercks, culture CBS 285.72. Netherlands, from stem base 

of Apium graveolens, 1978, J. de Gruyter, CBS 504.91 = PD 78/1073. 

Subplenodomus drobnjacensis (Bubák) Gruyter, 


Basionym: Phoma drobnjacensis Bubák, Bot. Közlem. 14: 63. 1915 

= Pyrenochaeta gentianae Chevassut, Bull. Soc. Mycol. France. 81: 36. 

1965. 

Specimens examined: Netherlands, from stem base of Gentiana makinoi “Royal 

Blue” (Gentianaceae), 1983, M.M.J. Dorenbosch, CBS 270.92 = PD 83/650; 

Naaldwijk, from red-brown root of Eustoma exaltatum (Gentianaceae), 1988, M.M.J. 

Dorenbosch, CBS 269.92 = PD 88/896. 

Subplenodomus valerianae (Henn.) Gruyter, Aveskamp & 


Basionym: Phoma valerianae Henn., Nyt Mag. Naturvidensk. 42: 

29. 1904. 

= Phyllosticta valerianae-tripteris f. minor Unamuno, Mem. Real Soc. Esp. 

Hist. Nat. 15: 348. 1929. 

Specimens examined: Netherlands, Arnhem, from dead stem of Valeriana phu 

(Valerianaceae), Sep. 1968, G.H. Boerema, CBS 630.68 = PD 68/141; Elburg, from 

stem base of Valeriana officinalis, 1973, M.M.J. Dorenbosch, culture CBS 499.91 

= PD 73/672. 

Subplenodomus violicola (P. Syd.) Gruyter, Aveskamp & 


Basionym: Phoma violicola P. Syd., Beibl. Hedwigia 38: 137. 1899. 

= Phyllosticta violae f. violae-hirtae Allesch. Rabenh.-Fl., Ed. 2, Pilze 6: 156. 

1898. 

= Phoma violae-tricoloris Died., Ann. Mycol. 2: 179. 1904. 

= Phyllosticta violae f. violae-sylvaticae Gonz. Frag., Trab. Mus. Nac. Ci. Nat., 

Ser. Bot. 7: 35. 1914. 

Specimens examined: Netherlands, Baarn, from leaf spot in Viola tricolor, 10 Mar. 

1968, H.A. van der Aa, CBS 306.68. New Zealand, Auckland, Henderson, from leaf 

spot in Viola tricolor (Violaceae), 1997, J. Jury, CBS 100272. 

Coniothyriaceae W.B. Cooke. Revista Biol. (Lisbon) 12: 

289. 1983. 

Coniothyrium carteri (Gruyter & Boerema) Verkley & 

Gruyter, comb. nov. MycoBank MB564775. 

Basionym: Phoma carteri Gruyter & Boerema, Persoonia 17(4): 

547. 2002 (“2001”). nom. nov. 

Replaced synonym: Pyrenochaeta minuta J.C. Carter, Bull. 

Illinois Nat. Hist. Surv. 21: 214. 1941; not Phoma minuta Wehm., 

Mycologia 38: 318. 1946, nor Phoma minuta Alcalde, Anales Inst. 

Bot. Cavanilles 10: 235. 1952; not Coniothyrium minutum (Berl.) 

O. Kuntze, Revis. Gen. Pl. 3: 459. 1898 = Phoma cava, syn. of 

Pyrenochaeta cava; not Coniothyrium minutum (Died) Petr. & Syd., 

Feddes Repert. Spec. Nov. Regni Veg. Beih. 42: 349. 1927. 

Specimens examined: Germany, isolated from Quercus robur (Fagaceae), 1991, 

CBS 105.91. Netherlands, from shoot of Quercus sp. (Fagaceae), 1984, M.M.J. 

Dorenbosch, CBS 101633 = PD 84/74. 

Coniothyrium dolichi (Mohanty) Verkley & Gruyter, comb. 

nov. MycoBank MB564776. 

Basionym: Pyrenochaeta dolichi Mohanty, Indian Phytopathol. 11: 

85. 1958. 

Specimen examined: India, Nani Tal, Sarichuan, from leafspot of Dolichos biflorus 

(Fabaceae), 20 Oct. 1955, N.N. Mohandy, CBS 124140 = IMI 217262, CBS 124143 

= IMI 217261. 

Notes: A synanamorph was noted and described as a Coniosporium 

state based on the dark brown to black, dictyosporous conidia 

(Mohanty 1958). This synanamorph was considered later as 

Monodictys-like (Grodona et al. 1997). 

Coniothyrium glycines (R.B. Stewart) Verkley & Gruyter, 


Basionym: Pyrenochaeta glycines R.B. Stewart, Mycologia 49: 

115. 1957. 

≡ Phoma glycinicola Gruyter & Boerema, Persoonia 17: 554. 2002 

(“2001”). nom. nov. nom. inval.; not Phoma glycines Sawada, Special. 

Publ. Coll. Agric., Natl. Taiwan Univ. 8: 129. 1959. nom. inval. = Phoma 

glycines Sawada ex J.K. Bai & G.Z. Lu, Fl. Fungorum Sin. 15: 33. 2003. 

Specimens examined: Zambia, on Mt. Makulu, from leaf of Glycine max (Fabaceae), 

Mar. 1985, J.M. Waller, CBS 124455 = IMI 294986. Zimbabwe, from a leaf of 

Glycine max (Fabaceae), 2001, C. Lavy, CBS 124141 = PG1. 


23


Coniothyrium multiporum (V.H. Pawar, P.N. Mathur 

& Thirum.) Verkley & Gruyter, comb. nov. MycoBank 

MB564778. 

Basionym: Phoma multipora V.H. Pawar, P.N. Mathur & Thirum., 


≡ Phoma multipora V.H. Pawar & Thirum., Nova Hedwigia 12: 501. 1966. 

nom. nud. 

Specimens examined: Egypt, CBS 501.91 = PD 83/888. India, Bombay, Bandra, 

from saline soil, 15 Jan. 1958, M.J. Thirumalachar, Isotype CBS H-16492, culture 

ex-isotype CBS 353.65 = ATCC 16207 = HACC 164 = IMI 113689. 

Coniothyrium palmarum Corda, Icon. Fungorum. (Corda) 

4: 38. 1840. 

≡ Clisosporium palmarum (Corda) Kuntze, Revis. Gen. Pl. 3: 458. 1898. 

≡ Microdiplodia palmarum (Corda) Died., Ann. Mycol. 11: 47. 1913. 

Specimens examined: Italy, Sardegna, near Dorgali, from a dead petiole of 

Chamaerops humilis (Arecaceae), Aug. 1970, W. Gams, CBS H-10891–10893, 

culture CBS 400.71. 

Coniothyrium telephii (Allesch.) Verkley & Gruyter, comb. 


Basionym: Pyrenochaeta telephii Allesch., Ber. bayer. bot. Ges. 4: 

33. 1896. 

≡ Phoma septicidalis Boerema, Versl. Meded. Plantenziektenk. Dienst 

Wageningen 153 (Jaarb. 1978): 20. 1979. nom. nov.; not Phoma telephii 

(Vestergr.) Kesteren, Netherlands J. Pl. Pathol. 78: 117. 1972. 

Specimens examined: Finland, Helsinki, Asko Kahanpää, obtained from air, 

Jan. 1971, CBS H-16567, culture CBS 188.71; Oulu, from mineral wool between 

walls, Dec. 1996, K. Poldmaa, CBS 856.97. Zimbabwe, from leaf of Glycine max 

(Fabaceae), CBS 101636 = PD 86/1186. 

Cucurbitariaceae G. Winter, Rabenh, Krypt.-Fl., Ed 2, 308. 

1885. 

Neophaeosphaeria filamentosa (Ellis & Everh.) Câmara, 

M.E. Palm & A.W. Ramaley, Mycol. Res. 107: 519. 2003. 

Basionym: Leptosphaeria filamentosa Ellis & Everh., J. Mycol. 4: 

76. 1888. 

≡ Paraphaeosphaeria filamentosa (Ellis & Everh.) M.E. Barr, Mycotaxon 

43: 392. 1992. 

Specimen examined: Mexico, from Yucca rostrata (Asparagaceae), Stevens, CBS 

102202 = BPI 802755. 

Pyrenochaetopsis pratorum (P.R. Johnst. & Boerema) 


MB564780. 

Basionym: Phoma pratorum P.R. Johnst. & Boerema, New Zealand 

J. Bot. 19: 395. 1981. 

Specimens examined: New Zealand, Rakura, near Hamilton, from a leaf of Lolium 

perenne (Poaceae), 1980, P.R. Johnston, isotype CBS H-7625, CBS H-7626, 

culture CBS 445.81 = PDDCC 7049 = PD 80/1254; Dactylis glomerata (Poaceae), 

1980, CBS 286.93 = PD 80/1252. 

Pleosporaceae Nitschke, Verh. Naturhist. Vereines Preuss. 

Rheinl. 26: 74. 1869. 

Pleospora angustis Gruyter & Verkley, nom. nov. MycoBank 

MB564781. 

≡ Leptosphaeria clavata A.L. Guyot, Revue Mycol. (Paris) 11: 62. 1946. 

≡ Massariosphaeria clavata (A.L. Guyot) Shoemaker & C.E. Babc., 

Canad. J. Bot. 67: 1582.1989; not Pleospora clavata Gucevič (”as 

clavatis”), Novosti Sist. Nizsh. Rast. 7: 168. 1970. 

Specimen examined: Switzerland, 1951, E. Müller, CBS 296.51. 

Notes: The origin of the isolate deposited by E. Müller is unknown; 

however, it is likely that the isolate was obtained from Poaceae, 

Triticum vulgare or Dactylis glomerata (Müller 1950). Pleospora 

clavata Gucevič was obtained from Lonicera alseuosmoides and 

refers to a different species. 

Pleospora betae (Berl.) Nevod., Grib. ross. Exs., No. 247. 

1915. 

Basionym: Pyrenophora echinella var. betae Berl. Nuovo Giorn. 

Bot. Ital. 20: 208. 1888. 

= Pleospora betae Björl., Bot. Not. 1944: 218. 1944. (later homonym). nom. 

illeg. 

≡ Pleospora bjoerlingii Byford, Trans. Brit. Mycol. Soc. 46: 614. 1963. 

nom. nov. 

= Phoma betae A.B. Frank, Z. Rúbenzucker-Ind. 42: 904, tab. 20. 1892. 

= Phyllosticta betae Oudem., Ned. Kruidk. Arch. Ser. 2, 2: 181. 1877. 

= Gloeosporium betae Dearn. & E.T. Barthol., Mycologia 9: 356. 1917. 

Specimens examined: Netherlands, Wageningen, from Beta vulgaris 

(Chenopodiaceae), Sep. 1966, M.M.J. Dorenbosch, CBS H-16156, culture CBS 

523.66 = IHEM 3915 = PD 66/270; from Beta vulgaris, 1977, G.H. Boerema, CBS 

109410 = PD 77/113. 

Note: The name Phoma betae A.B. Frank has been conserved 

against Phyllosticta tabifica and any combination based on that 

name (Shoemaker & Redhead 1999). 

Pleospora calvescens (Fr.) Tul. & C. Tul., Selecta Fung. 

Carpol. (Paris) 2: 266. 1863. 

Basionym: Sphaeria calvescens Fr., Ann. Sci. Nat., Bot. Ser. 2, 19: 

353. 1843. 

≡ Leptosphaeria calvescens (Fr.) Sacc., Syll. fung. 2: 24. 1883. 

≡ Pyrenophora calvescens (Fr.) Sacc., Syll. fung. 2: 279. 1883. 

= Chaetodiplodia caulina P. Karst., Hedwigia 23: 62. 1884. 

≡ Ascochyta caulina (P. Karst.) v.d. Aa & Kesteren, Persoonia 10: 271. 

1979. 

= Microdiplodia henningsii Staritz, Hedwigia 53: 163. 1913. 

Specimens examined: Germany, Munkmarsch, from leaf spots in Atriplex hastata 

(Chenopodiaceae), 20 July 1977, G.H. Boerema, CBS H-8980, culture CBS 246.79 

= PD 77/655. Netherlands, Texel, from dead stem of Atriplex hastata, June 1978, 

H.A. van der Aa, CBS H-8976, culture CBS 343.78. 

Note: For additional synonyms see Boerema et al. (1993). 

Pleospora chenopodii Ellis & Kellerman, J. Mycol. 4: 26. 

1888. 

= Diplodia hyalospora Cooke & Ellis, Grevillea 7: 5. 1878; not Pleospora 

hyalospora Ellis & Everh., Proc. Acad. Nat. Sci. Philadelphia. 42: 238. 1890. 

≡ Ascochyta hyalospora (Cooke & Ellis) Boerema, S.B. Mathur & Neerg., 

Netherlands J. Pl. Pathol. 83: 156. 1977. 

= Diplodina ellisii Sacc., Syll Fung. 3: 417. 1884 

Specimens examined: Bolivia, isolated from Chenopodium quinoa 

(Chenopodiaceae), 1974, S.B. Mathur, CBS H-9051, CBS H-9052, culture 

CBS 206.80 = PD 74/1022. Netherlands, Zoutelande, from Atriplex hastata 

(Chenopodiaceae), Aug. 1968, H.A. van Kesteren, CBS 344.78 = PD 68/682. 

Note: Isolate CBS 344.78 was originally identified as Ascochyta 

caulina but was identical to Pleospora chenopodii in the present 

study. 

24


Pleospora fallens (Sacc.) Gruyter & Verkley, comb. nov. 


Basionym: Phoma fallens Sacc., Syll. Fung. 10: 146. 1892. 

= Phyllosticta glaucispora Delacr., Bull. Soc. Mycol. France 9: 266. 1893. 

≡ Phoma glaucispora (Delacr.) Noordel. & Boerema, Versl. Meded. 

Plantenziektenk. Dienst Wageningen 166 (Jaarb. 1987): 108. 1989 

(“1988”). 

= Phyllosticta oleandri Gutner, Trudy Bot. Inst. Akad. Nauk S.S.S.R., Ser. 2, 

Sporov. Rast. 1: 306. 1933. 

Specimens examined: Italy, Capri, Villa Jovis, from a leaf spot of Nerium oleander 

(Apogynaceae), CBS H-16639, culture CBS 284.70 = PD 97/2400. New Zealand, 

Levin, from leaf spot of Olea europaea (Oleaceae), 1978, G.F. Laundon, CBS 

161.78 = LEV 1131. 

Pleospora flavigena (Constantinou & Aa) Gruyter & Verkley, 


Basionym: Phoma flavigena Constantinou & Aa, Trans. Brit. Mycol. 

Soc. 79: 343. 1982. 

Specimen examined: Romania, Bucuresti, isolated from water, 1980, K. Fodor, CBS 

H-1418, holotype of Phoma flavigena Constantinou & Aa, culture ex-holotype CBS 

314.80 = PD 91/1613. 

Pleospora halimiones Gruyter & Verkley, nom. nov. 


≡ Diplodina obiones Jaap (as “obionis”), Verh. Bot. Vereins Prov. 

Brandenburg 47: 96. 1905; not Pleopora obiones P. Crouan & H. Crouan, 

Fl. Finistère: 22. 1867. 

≡ Ascochytula obiones (Jaap) Died., Ann. Mycol. 10: 141. 1912. 

≡ Ascochyta obiones (Jaap) P.K. Buchanan, Mycol. Pap. 156: 28 1987. 

= Coniothyrium obiones Jaap (as “obionis”), Schriften Naturwiss. Vereins 

Schleswig-Holstein 14: 29. 1907. 

Specimens examined: Netherlands, Texel, from leaf spots in Halimione 

portulacoides (Chenopodiaceae), 27 Oct. 1968, H.A. van der Aa, CBS H-9127, CBS 

H-9129, culture CBS 786.68; Texel, De Cocksdorp, from dead stems of Halimione 

portulacoides, 6 July 1977, H.A. van der Aa, CBS H-9126, CBS H-9125, culture CBS 

432.77 = IMI 282137. 

Notes: Isolate CBS 453.68 preserved as Chaetodiplodia sp. and 

also isolated from dying stems and leaf sheaths of Halimione 

portulacoides on Texel, is not the same as Pleo. halimiones and is 

probably a different species. 

Pleospora herbarum (Pers.) Rabenh., Bot. Zeitung (Berlin) 

15: 428. 1857; Klotzschii Herb. Viv. Mycol. 2: no. 547 (1854.) 

Basionym: Sphaeria herbarum Pers., Syn. Meth. Fung. 1: 78. 1801. 

= Stemphylium herbarum E.G. Simmons, Sydowia 38: 291. 1986 (1985). 

Specimen examined: India, Uttar Pradesh, from a leaf of Medicago sativa 

(Fabaceae), 1986 (isolated in 1983), E.G. Simmons, CBS 191.86 = IMI 276975. 

Note: This isolate is the ex-type culture of Stemphylium herbarum. 

Pleospora incompta (Sacc. & Martelli) Gruyter & Verkley, 


Basionym: Phoma incompta Sacc. & Martelli, Syll. Fung. 10: 146. 

1892. 

Specimens examined: Greece, Crete, from branch of Olea europaea (Oleaceae), 

1976, N. Malathrakis, CBS H-16394, culture CBS 467.76. Italy, from branch of Olea 

europaea, Mar. 1982, CBS H-16392, culture CBS 526.82. 

Pleospora typhicola (Cooke) Sacc., Syll. Fung. 2: 264. 

1883. 

Basionym: Sphaeria typhicola Cooke, Grevillea 5: 121. 1877. 

≡ Clathrospora typhicola (Cooke) Höhn., Ann. Mycol. 16: 88. 1918. 

≡ Pyrenophora typhicola (Cooke) E. Müll., Sydowia 5: 256. 1951. 

≡ Macrospora typhicola (Cooke) Shoemaker & C.E. Babc., Canad. J. Bot. 

70: 1644. 1992. 

= Phyllosticta typhina Sacc. & Malbr., Sacc., Michelia 2: 88. 1880. 

≡ Phoma typhina (Sacc. & Malbr.) Aa, van der Aa & Vanev, A revision of 

the species described in Phyllosticta: 468. 2002. 

= Phoma typharum Sacc., Syll. Fung. 3: 163. 1884. 

Specimens examined: Netherlands, Texel, from dead leaves of Typha angustifolia 

(Typhaceae), 1969, W. Gams, CBS H-16597, culture CBS 132.69; Staverden, from 

leaf spots of Typha sp., 24 June 1972, G.S. de Hoog, CBS H-16598, culture CBS 

602.72. 

Phoma-like anamorphs excluded from the suborder 

Pleosporineae 

Montagnulaceae M.E. Barr, Mycotaxon 77: 194. 2001. 

Paraconiothyrium Verkley, Stud. Mycol. 50: 327. 2004. 

Type species: Paraconiothyrium estuarinum Verkley & M. da Silva, 

Stud. Mycol. 50: 327. 2004. 

Paraconiothyrium flavescens (Gruyter, Noordel. & 

Boerema) Verkley & Gruyter, comb. nov. MycoBank 

MB564786. 

Basionym: Phoma flavescens Gruyter, Noordel. & Boerema, 

Persoonia 15(3): 375. 1993. 

Specimen examined: Netherlands, Nagele, from soil, rhizosphere of Solanum 

tuberosum (Solanaceae), CBS 178.93 = PD 82/1062. 

Paraconiothyrium fuckelii (Sacc.) Verkley & Gruyter, 


Basionym: Coniothyrium fuckelii Sacc., Nuovo Giorn. Bot. Ital. 8: 

200. 1876; Michelia 1: 207. 1878 

≡ Clisosporium fuckelii (Sacc.) Kuntze, Revis. Gen. Pl. 3: 458. 1898. 

≡ Microsphaeropsis fuckelii (Sacc.) Boerema, 2003, Persoonia 18: 160. 

2003. 

Specimen examined: Denmark, Geelskov, from a dead stem of Rubus sp. 

(Rosaceae), 1995, A.M. Dahl-Jensen, CBS 797.95. 

Notes: Coniothyrium fuckelii var. sporulosum has been redisposed 

as Paraconiothyrium sporulosum (Verkley et al. 2004) and it is 

clearly different from Paraconiothyrium fuckelii (Damm et al. 2008). 

Paraconiothyrium fusco-maculans (Sacc.) Verkley & 

Gruyter, comb. nov. MycoBank MB564788. 

Basionym: Phoma fusco-maculans Sacc., Michelia 2: 275. 1881 

≡ Plenodomus fusco-maculans (Sacc.) Coons, J. Agric. Res. 5: 714. 1916. 

Specimens examined: Italy, Selva, from decorticated wood of Malus pumila 

(Rosaceae), Oct. 1880, PAD, holotype of Phoma fusco-maculans Sacc. USA, from 

wood of Malus sp. (Rosaceae), July 1916, G.H. Coons, epitype designated here 

CBS H-20825, culture ex-epitype CBS 116.16. 

Notes: Plenodomus fusco-maculans was discussed by Boerema & 

Loerakker (1985) and de Gruyter et al. (2010). The holotype of the 

basionym Aposphaeria fusco-maculans was studied and considered 

to be Aposphaeria pulviscula (Boerema et al. 1996). However, the 

description of A. fusco-maculans given by Boerema et al. (1996) 

fits the generic concept of Paraconiothyrium, in congruence with 

the molecular phylogeny of the culture CBS 116.16. 


25


Fig. 6. Paraconiothyrium maculicutis sp. nov. CBS 101461. A–B. Fourteen day old cultures on OA (A) and MA (B). C–D. Pycnidia. E. Phoma-like conidiogenous cells. F–G. 

Conidia, initially hyaline to pale olivaceous (F), then becoming olivaceous (G). Scale bars: C–D = 20 μm; E = 10 μm; F–G = 5 μm. 

Paraconiothyrium lini (Pass.) Verkley & Gruyter, comb. 


Basionym: Phoma lini Pass., Diagn. Funghi Nuovi 4, No. 81. 1890. 

Specimen examined: Netherlands, from Wisconsin tank, 1970, CBS 253.92 = PD 

70/998. 

Paraconiothyrium maculicutis Verkley & Gruyter, sp. nov. 

MycoBank MB564796. Fig. 6. 

Etymology: Latin, cutis = skin; maculae = spots. 

Pycnidia in vitro 50–125 μm diam, globose to subglobose, 

glabrous or with mycelial outgrowth, scattered, non-ostiolate 

or ostiolate, pycnidial wall made up of 5–7 layers of cells. 

Conidiogenous cells 1.5–3 × 0.5–2.5 μm, indeterminate or 

ampulliform to filiform in a later state, up to 10 μm in length. 

Conidia 1.5–2.5 × 0.5–1.5 μm, ellipsoidal, initially hyaline, then 

discolouring to olivaceous. 

Description in vitro: Colonies on OA 50–52 mm diam after 7 d, 

margin entire; colony olivaceous buff to greenish olivaceous/grey 

olivaceous, with greenish olivaceous to pale olivaceous grey, 

finely floccose to woolly aerial mycelium; reverse smoke-grey to 

greenish olivaceous, with olivaceous patches. Colonies on MEA 

43–44 mm diam after 7 d, margin entire; colony pale olivaceous 

grey to greenish olivaceous, with isabelline to cinnamon at centre, 

with compact pale olivaceous grey, finely floccose to woolly aerial 

mycelium; reverse buff to honey, isabelline to olivaceous near 

margin. Pycnidia globose to subglobose, olivaceous to brick, finally 

olivaceous black, scattered, mainly on the agar, 50–125 μm diam, 

glabrous or with mycelial outgrowth, non-ostiolate or ostiolate, 

pycnidial wall made up of 5–7 layers of cells. Conidiogenous cells 

1.5–3 × 0.5–2.5 μm, ampulliform to filiform in a later state, up to 10 

μm in length. Conidia 1.5–2.5 × 0.5–1.5 μm, av. 1 × 2 μm, length/ 

width ratio = 1.5–3.2, av. 2.2, ellipsoidal, initially hyaline, then 

discolouring to olivaceous. Chlamydospores absent. NaOH spot 

test: negative. Crystals absent. 

Specimen examined: USA, Texas; San Antonio, Fort Sam Houston, from human, 

cutaneous lesions, 1989, D.P. Dooley, holotype CBS H-20824, culture ex-holotype 

CBS 101461 = IMI 320754 = UTHSC 87-144. 

Notes: Isolate CBS 101461 was identified as Pleurophoma 

pleurospora (Dooly et al. 1989). However, in vitro data and the 

molecular phylogeny demonstrate that this isolate does not belong 

to Pleurophoma pleurospora, see below, and therefore is described 

as a new species in the genus Paraconiothyrium. 

Paraconiothyrium minitans (W.A. Campb.) Verkley, Stud. 

Mycol. 50: 332. 2004. 

Basionym: Coniothyrium minitans W.A. Campb., Mycologia 39: 

191. 1947. 

Specimens examined: Netherlands, Boskoop, from stem of Clematis sp. 

(Ranunculaceae), 1999, J. de Gruyter, CBS 122786 = PD 99/1064-1. UK, CBS 

122788 = PD 07/03486739. 

Paraconiothyrium tiliae (F. Rudolphi) Verkley & Gruyter, 


Basionym: Asteroma tiliae F. Rudolphi, Linnaea 4: 514. 1829. 

26


Fig. 7. Pleurophoma pleurospora. CBS 130329. A–B. Fourteen day old cultures on OA (A) and MA (B). C. Pycnidia. D–H. Conidiogenous cells, septate conidiophores with 

acropleurogenous conidiogenesis (D–G) or Phoma-like (H). I. Conidia. Scale bars: C = 50 μm; D–G, I = 10 μm; H = 5 μm. 

≡ Asteromella tiliae (F. Rudolphi) Butin & Kehr, Mycol. Res. 99: 1193. 

1995. nom. inval., Art. 33.4. 

Specimen examined: Austria, Amlach, from a leaf of Tilia platyphyllos (Tiliaceae), 

10 Sep. 1993, H. Butin, neotype IMI 362854, lectotype designated here CBS 

H-20826, culture ex-lectotype CBS 265.94. 

Pleurophoma pleurospora (Sacc.) Höhn., Sitzungsber. 

Kaiserl. Akad. Wiss., Math.-Naturwiss. Cl., Abt. 1. 123: 117. 

1914. Fig. 7. 

Basionym: Dendrophoma pleurospora Sacc., Michelia 2: 97. 1880. 

Description in vitro: Colonies on OA 14–18 mm diam after 7 d 

(18–28 mm after 14 d), margin entire to undulate; colony greenish 

olivaceous/olivaceous to rosy-buff and sepia, with white, felty 

aerial mycelium; reverse olivaceous grey to greenish olivaceous/ 

olivaceous. Colonies on MEA 11–16 mm diam after 7 d (19–29 

mm after 14 d), colony margin undulate; colony pale olivaceous 

grey/ olivaceous grey to dark mouse-grey with rosy-buff tinges, 

with white, floccose, compact aerial mycelium, reverse umber/ 

brown olivaceous to olivaceous/olivaceous black. Pycnidia globose 

to subglobose,olivaceous to olivaceous black, abundant, scattered, 

mainly on the agar, 30–120 μm diam, solitary or aggregated, 

covered by mycelial outgrowths or setae-like hyphae, up to 50 μm, 

non-papillated, without or with ostiole, walls made up of 2–5 layers 

of cells, outer layer(s) pigmented; conidial exudate not observed. 

Conidiogenous cells of two types; ampulliform to doliiform, 4–6.5 × 

2–5.5 μm, or filiform, septate, branched, acropleurogenous, up to 

60 μm long. Conidia 3.5–5.5 × 1.5–2.5 μm, av. 4.5 × 2 μm, length/ 

width ratio = 1.5–3, av. 2.1, cylindrical to oblong, without or with 

some minute, polar orientated guttules. Chlamydospores absent. 

NaOH spot test: a weak reddish discolouring may occur on MA, not 

specific. Crystals absent. 

Specimens examined: France, Perpignan, from leaf of Laurus nobilis (Lauraceae), 

PAD, holotype of Dendrophoma pleurospora Sacc. Netherlands, from wood of 

Lonicera sp. (Caprifoliaceae), lectotype designated here CBS H-20626, culture 

ex-lectotype CBS 130329 = PD 82/371; Molenhoek, Heumense Schans, from twig 

lesions of Cytisus scoparius (Fabaceae), 23 Aug. 2004, G. Verkley & M. Starink, 

CBS 116668. 

Notes: A specimen derived from isolate CBS 130329 is assigned 

here as lectotype of Pleurophoma pleurospora, the type species of 

the genus (von Höhnel 1914). The species is known from branches 

and bare wood of trees and shrubs (Sutton 1980, Boerema et al. 

1996) and the isolate from Cytisus scoparius demonstrates that the 

species also may occur on green twigs. The isolates showed two 

types of conidiogenesis characteristic for the genus Pleurophoma; 

Phoma-like, ampulliform to doliiform conidiogenous cells, as well as 

Pyrenochaeta-like branched, filiform, septate, acropleurogenous. 

As a result, species of the genus Pleurophoma can easily be 

confused with taxa classified in the genera Phoma, Paraphoma, 

Pyrenochaeta and Pyrenochaetopsis. 


27


Paraphaeosphaeria michotii (Westend.) O.E. Erikss., Arkiv 

før Botanik 6: 406. 1967. 

Basionym: Sphaeria michotii Westend., Bull. Acad. Roy. Sci. 

Belgique Ser. 2, 7: 87. 1859. 

Specimen examined: Switzerland, Kt. Obwalden, from Typha latifolia (Typhaceae), 

18 May 1980, A. Leuchtmann, CBS 652.86 = ETH 9483. 

Massarinaceae Munk, Friesia 5: 305. 1956. 

Byssothecium circinans Fuckel, Bot. Zeitung (Berlin) 19: 

251. 1861. 

≡ Leptosphaeria circinans (Fuckel) Sacc., Syll. Fung. 2: 88. 1883. 

≡ Passeriniella circinans (Fuckel) Sacc., Syll. Fung. 11: . 1895. 

≡ Trematosphaeria circinans (Fuckel) G. Winter, Rabenh. Krypt.-Fl., ed 

1(2): 277. 1887. 

≡ Heptameria circinans (Fuckel) Cooke, Grevillea 18: 30. 1889. 

= Melanomma vindelicorum Rehm, Ber. Nat. Ver. Augsburg: 116. 1881. 

≡ Trematosphaeria vindelicorum (Rehm) Sacc., Syll. Fung. 2: 122. 1883. 

Specimen examined: USA, South Dakota, from rotten crown of Medicago sativa 

(Fabaceae), G. Semeniuk, CBS 675.92 = ATCC 52767 = ATCC 52678 = IMI 266220. 

Massarina eburnea (Tul. & C. Tul.) Sacc., Syll. Fung. 2: 153. 

1883. 

Basionym: Massaria eburnea Tul. & C. Tul., Select. Fung. Carpol. 

(Paris) 2: 239. 1863. 

Specimens examined: Switzerland, Zürich, from Fagus sylvatica (Fagaceae), S.K. 

Bose, CBS 473.64 = ETH 2945. UK, Wales, isolated from dead branch of Fagus 

sylvatica, HHUF 26621, JCM 14422 = H3953. 

Neottiosporina paspali (G.F. Atk.) B. Sutton & Alcorn, 

Austral. J. Bot. 22: 519. 1974. 

Basionym: Stagonospora paspali G.F. Atk., Bull. Cornell Univ. 

(Science) 3: 33. 1897. 

Specimen examined: USA, Florida, from Paspalum notatum (Poaceae), Oct. 1937, 

R.K. Voorhees, CBS 331.37. 

Trematosphaeriaceae Suetrong et al. Cryptogamie Mycol. 

32: 347. 2011. 

Falciformispora lignatilis K.D. Hyde, Mycol. Res. 96: 27. 

1992. 

Specimen examined: Thailand, Pinruan Ban Bang, from Elaeis guineensis 

(Arecaceae), BCC 21118. 

Medicopsis Gruyter, Verkley & Crous, gen. nov. MycoBank 

MB564791. 

Etymology: refers to Medi- medica, Latin, -opsis, refers to, Greek. 

The description of the type species as the cause of a mycetoma 

suggest this is a human pathogen. However, the mycetoma 

described was secondary to a wound produced by a thorn of Palito 

blanco tree, and the species was found later on Hordeum vulgare. 

Pycnidia solitary or confluent, on upper surface of the agar, globose 

to pyriform with elongated neck, setose, ostiolate, olivaceous 

to olivaceous-black, the wall with pseudoparenchymatal cells. 

Conidiogenous cells hyaline, phialidic, ampulliform to doliiform, to 

elongated. Conidia sub-hyaline to yellowish, ellipsoid, aseptate, 

catenulate. 

Type species: Medicopsis romeroi (Borelli) Gruyter, Verkley & 

Crous (see below). 

Medicopsis romeroi (Borelli) Gruyter, Verkley & Crous, 


Basionym: Pyrenochaeta romeroi Borelli, Dermatol. Venez. 1: 326. 

1959. 

Specimens examined: Venezuela, from human, maduromycosis, no date, D. Borelli, 

UAMH 2892, holotype of Pyrenochaeta romeroi Borelli, culture ex-holotype CBS 

252.60 = ATCC 13735 = FMC 151 = UAMH 10841. Country unknown, from Hordeum 

vulgare (Poaceae), 1984, M.M.J. Dorenbosch, CBS 122784 = PD 84/1022. 

Notes: The species was described as a human pathogen of 

tropical origin, and it may cause suppurative subcutaneous 

or deep nonmycetomatous infections, or a subcutaneous 

phaeohyphomycotic cyst (Badali et al. 2010). However, the species 

also occurs in plant material. 

Trematosphaeria pertusa (Pers.) Fuckel, Jahrb. 

Nassauischen Vereins Naturk 23–24: 161. 1870. 

Basionym: Sphaeria pertusa Pers., Syn. Meth. Fung. 1: 83. 1801. 

Specimen examined: France, Deux Sèvres, from bark of a dead stump of Fraxinus 

excelsior (Oleaceae), 25 Apr. 2004, Jacques Fournier, epitype IFRD 2002, culture 

ex-epitype CBS 122368. 

Note: The epitype IFRD 2002 was designated by Zhang et al. 

(2008). 

Massariaceae Nitschke. Verh. Naturhist. Vereines Preuss. 

Rheinl. 26: 73. 1869. 

Massaria platani Ces., Fungi Eur. Exsicc. Klotzsch. Herb. 

Vivi Mycol. no. 323. 1861. 

Specimen examined: USA, from Platanus occidentalis (Platanaceae), Jan. 1937, 

C.L. Shear, CBS 221.37. 

Melanommataceae G. Winter, Rabenh. Krypt.-Fl., ed 1(2): 

220 (1885) [as “Melanommeae”] 

Aposphaeria corallinolutea Gruyter, Aveskamp & Verkley, 

sp. nov. MycoBank MB564798. Fig. 8. 

Etymology: The name refers to the coral coloured colony on OA, 

and the luteous exudate diffusing into the agar medium. 

Pycnidia in vitro 65–215 μm diam, solitary or aggregated to 

confluent, globose to subglobose, ostiolate or non-ostiolate. 

Conidiogenous cells 7–9 × 2–4 μm, ampuliform to filiform. Conidia 

3–5 × 1–2 μm, ellipsoidal to allantoid, eguttulate or with some small, 

polar guttules. 

Description in vitro: Colonies on OA 13–15 mm diam after 14 d, 

margin entire to somewhat lobated; colony vinaceous to brick, 

with white at centre, ochraceous near margin due to a diffusible 

pigment, with white, felty or poorly developed aerial mycelium; 

reverse cinnamon to brick. Colonies on MEA 15–20 mm diam after 

14 d, margin entire to somewhat lobated; colony white with dull 

green and grey olivaceous sectors and primrose tinges, with white, 

felty aerial mycelium; reverse sepia to brown olivaceous, greenish 

28


Fig. 8. Aposphaeria corallinolutea sp. nov. CBS 131287. A–B. Fourteen day old cultures on OA (A) and MA (B). C–D. Pycnidia. E–H. Conidiogenous cells. I. Conidia. Scale 

bars: C = 50 μm; D = 20 μm; E–I = 10 μm. 

grey at centre, white near margin. Pycnidia globose to subglobose, 

olivaceous to brick, then olivaceous black, solitary or aggregated, 

65–215 μm diam, non-setose or with short setae-like outgrowths 

up to 25 μm long, with or without distinct ostiole, pycnidial wall 

consisting of 3–5 layers of cells. Conidiogenous cells 7–9 × 2–4 

μm, ampulliform to filiform. Conidia 3–5 × 1–2 μm, av. 4 × 1.5 

μm, length/width ratio is 1.7–3.3, av. = 2.5, ellipsoidal to allantoid, 

eguttulate or with some small, polar guttules. Chlamydospores 

absent, NaOH test negative. Crystals produced in the agar, small, 

orange coloured. 

Specimens examined: Netherlands, from wood of Fraxinus excelsior (Oleaceae), 

1983, M.M.J. Dorenbosch, holotype CBS H-20625, culture ex-holotype CBS 

131287 = PD 83/831; from wood of Kerria japonica (Rosaceae), 1983, M.M.J. 

Dorenbosch, CBS 131286 = PD 83/367. 

Aposphaeria populina Died., Krypt.-Fl. Brandenburg 9: 

206. 1912 (vol. dated “1915”). Fig. 9. 

Description in vitro: Colonies on OA 21–24 mm diam after 7 d 

(32–37 mm diam after 14 d), margin entire to undulate; colony 

grey olivaceous/olivaceous to pale luteous/luteous, with white to 

pale olivaceous grey, finely felty to woolly aerial mycelium; reverse 

luteous to orange, greenish olivaceous to olivaceous or grey 

olivaceous/olivaceous grey to iron-grey, a rosy-buff discolouring 

near margin may occur. Colonies on MEA 16–20 mm diam after 7 d 

(30–37 mm diam after 14 d), margin entire to undulate; colony pale 

olivaceous grey with rosy-vinaceous tinges to peach or olivaceous 

grey, with white, woolly aerial mycelium; reverse saffron to pale 

olivaceous/olivaceous grey, sometimes with dark vinaceous tinges, 

rosy-buff near margin. Pycnidia globose to subglobose, olivaceous 

to olivaceous black, scattered, 55–305 μm diam, glabrous or with 

mycelial outgrowths, non-ostiolate or ostiolate, pycnidial wall 

composed of up to 10 layers of cells. Conidiogenous cells 5–11.5 

× 1.5–3 μm, ampulliform to filiform. Conidia hyaline, subglobose to 

ellipsoidal, with 1–3 minute guttules, 1–2 × 1–1.5 μm, av. 1.5 × 1 

μm, length/width ratio is 1.0–2.0, av. = 1.4. Chlamydospores and 

crystals absent, NaOH test negative. 

Specimens examined: Germany, Triglitz, from twigs of Populus canadensis 

(Salicaceae), Mar. 1904. O. Jaap, B, holotype; from branch scars of Picea abies, 

(Pinaceae), Feb. 1982, H. von Aufess, CBS 350.82. Netherlands, Valkenswaard, 

from fallen twig of Populus canadensis (Salicaceae), 23 Mar. 1970, H.A. van der 

Aa, epitype designated here CBS H-9336, culture ex lectotype CBS 543.70; from 

wood of Cornus mas (Cornaceae), 1984, M.M.J. Dorenbosch, CBS 130330 = PD 

84/221. 

Beverwykella pulmonaria (Beverw.) Tubaki, Trans. Mycol. 

Soc. Japan 16: 139. 1975. 

Basionym: Papulaspora pulmonaria Beverw., Antonie van 

Leeuwenhoek 20: 11. 1954. 

Specimen examined: Netherlands, Baarn, from submerged leaf in rain water barrel 

of Fagus sylvatica (Fagaceae), Apr. 1953, A.L. van Beverwijk, culture CBS 283.53 

= ATCC 32983 = IFO 6800. 

Herpotrichia juniperi (Duby) Petr., Ann. Mycol. 23: 43. 

1925. 

Basionym: Sphaeria juniperi Duby, Klotzsch. Herb. Vivum Mycol. 

Sistems Fungorum German., no. 1833. 1854. 

Specimen examined: Switzerland, Andermatt, from Juniperus nana (Cupressaceae), 

Nov. 1931, E. Gäumann, CBS 200.31. 


29


Fig. 9. Aposphaeria populina. CBS 543.70. A–B. Fourteen day old cultures on OA (A) and MA (B). C. Pycnidium with mycelial outgrowths (CBS 130330). D–E. Conidiogenous 

cells. F. Conidia. Scale bars: C = 20 μm; D–E = 10 μm; F = 5 μm. 

Melanomma pulvis-pyrius (Pers.) Fuckel, Jahrb. 

Nassauischen Vereins Naturk. 23–24: 160. 1870. 

Basionym: Sphaeria pulvis-pyrius Pers., Syn. Meth. Fung. 1: 86. 

1801. 

Specimens examined: Belgium, from wood of Fagus sp. (Fagaceae), CBS 400.97. 

France, Vosges, Bot. Garden Le Chitelet, from unidentified decaying wood, CBS 

371.75. 

Notes: Phoma-like anamorphs have been reported by Chesters 

(1938) and Sivanesan (1984), but no anamorphic stage was 

observed in IFRDCC 2044, CBS 109.77 or CBS 371.75 after 

culturing 3 mo on PDA (Zhang et al. 2008). CBS 400.97 was 

preserved as Trematosphaeria pertusa. 

Pleomassaria siparia (Berk. & Broome) Sacc., Syll. Fung. 

2: 239. 1883. 

Basionym: Sphaeria siparia Berk. & Broome, Ann. Mag. Nat. Hist. 

Ser. 2(9): 321. 1852. 

Specimen examined: Netherlands, Uden, from dead branch of Betula verrucosa 

(Betulaceae), 8 Dec. 1973, W.M. Loerakker, CBS H-258, CBS H-260, culture CBS 

279.74. 

Sporormiaceae Munk, Dansk Bot. Ark. 17(1): 450. 1957. 

(nom. inval., art. 36.1.) 

Preussia funiculata (Preuss) Fuckel, Jahrb. Nassauischen 

Vereins Naturk. 23–24: 91. 1870 (1869–70). 

Basionym: Perisporium funiculatum Preuss, Linnaea 24(1): 143. 

1851. 

Specimen examined: Senegal, from soil, CBS 659.74. 

Sporormiella minima (Auersw.) S.I. Ahmed & Cain, Canad. 

J. Bot. 50: 449. 1972. 

Basionym: Sporormia minima Auersw., Hedwigia 7: 66. 1868. 

Specimen examined: Panama, from dung of goat, CBS 524.50. 

Westerdykella Stolk, Trans. Brit. Mycol. Soc. 38: 422. 1955. 

Type species: Westerdykella ornata Stolk, see below. 

Westerdykella capitulum (V.H. Pawar, P.N. Mathur & 

Thirum) Gruyter, Aveskamp & Verkley, comb. nov. MycoBank 

MB564801. 

Basionym: Phoma capitulum V.H. Pawar, P.N. Mathur & Thirum., 


≡ Phoma capitulum V.H. Pawar & Thirum., Nova Hedwigia 12: 502. 1966 

(as ”capitula”). nom. nud., nom. inval. 

= Phoma ostiolata V.H. Pawar, P.N. Mathur & Thirum., Trans. Brit. Mycol. Soc. 

50: 262. 1967, var. ostiolata. 

≡ Phoma ostiolata V.H. Pawar & Thirum., Nova Hedwigia 12: 502. 1966. 

nom. nud., nom. inval. 

= Phoma ostiolata var. brunnea V.H. Pawar, P.N. Mathur & Thirum., Trans. Brit. 

Mycol. Soc. 50: 263. 1967. 

≡ Phoma ostiolata var. brunnea V.H. Pawar & Thirum., Nova Hedwigia 

12: 502. 1966. nom. nud. nom. inval. 

Specimen examined: India, Bandra, Bombay, from saline soil, 15 Jan. 1958, M.J. 

Thirumalachar, Isotype CBS H-7602, culture ex-isotype CBS 337.65 = ATCC 16195 

= HACC 167 = IMI 113693 = PD 91/1614. 

30


Westerdykella minutispora (P.N. Mathur ex Gruyter 

& Noordel.) Gruyter, Aveskamp & Verkley, comb. nov. 


Basionym: Phoma minutispora P.N. Mathur ex Gruyter & Noordel., 

Persoonia 15: 75. 1992 (as “collection name” originally also referred 

to Thirumalachar; = depositor). 

Replaced synonym: Phoma oryzae Cooke & Massee, Grevillea 

16: 15. 1887; not Phoma oryzae Catt., Arch. Triennale Bot. Crittog. 

Pavia 2–3: 118. 1879. nom. illeg. 

≡ Phyllosticta oryzae (Cooke & Massee) I. Miyake. J. Coll. Agric. Imp. 

Univ. Tokyo 2: 252. 1910. nom. illeg. 

Specimen examined: India, from saline soil, 1977, M.J. Thirumalachar, CBS 

H-5941, culture CBS 509.91 = PD 77/920. 

Westerdykella ornata Stolk, Trans. Brit. Mycol. Soc. 38: 

422. 1955. 

Specimen examined: Mozambique, from mangrove mud, CBS 379.55. 

Didymosphaeriaceae Munk, Dansk Bot. Ark. 15(2): 128. 

1953. 

Roussoella hysterioides (Ces.) Höhn., Sitzungsber. 

Kaiserl. Akad. Wiss., Math.-Naturwiss. Cl., Abt. 1. 128: 563. 

1919. 

Basionym: Dothidea hysterioides Ces., Atti Accad. Sci. Fis. 8: 24. 

1879. 

Specimen examined: Japan, Aomori, Shimokita Yagen, from culms of Sasa 

kurilensis (Poaceae), Y. Ooki, culture CBS 125434 = HH 26988. 

Family incertae sedis 

Nigrograna Gruyter, Verkley & Crous, gen. nov. MycoBank 

MB564794. 

Etymology: refers to Nigro-, black, Latin, -grana, grains, Latin. The 

description refers to the black grains produced by the type species. 

Pycnidia solitary or rarely confluent, on upper surface or submerged 

in agar, globose to subglobose or pyriform, with dark brown, 

septate mycelial outgrowths, with papillate ostioles, olivaceous 

to olivaceous-black, the wall with pseudoparenchymatous cells. 

Conidiogenous cells hyaline, phialidic, discrete. Conidia subhyaline, 

brown in mass, aseptate, ellipsoidal. 

Type species: Nigrograna mackinnonii (Borelli) Gruyter, Verkley & 

Crous (see below). 

Nigrograna mackinnonii (Borelli) Gruyter, Verkley & Crous, 


Basionym: Pyrenochaeta mackinnonii Borelli, Castellania 4: 230. 

1976. 

Specimens examined: Mexico, from a mycetoma of a human, Feb. 2002, R. Arenas, 

CBS 110022; Venezuela, from a black grain mycetoma of human, Aug. 1975, D. 

Borelli, holotype FMC 270, culture ex-holotype CBS 674.75. 

Thyridaria rubronotata (Berk. & Broome) Sacc., Syll. Fung. 

2: 141. 1883. 

Basionym: Melogramma rubronotatum Berk. & Broome, Ann. Mag. 

Nat. Hist. Ser. 3(3): 20. 1859. 

Specimen examined: Netherlands, Zuidelijk Flevoland, from a dead branch of Acer 

pseudoplatanus (Aceraceae), 13 Apr. 1985, N. Ernste, CBS H-18824, culture CBS 

419.85. 

DISCUSSION 

The genus Phoma has been shown to be highly polyphyletic 

and Phoma is now restricted to taxa in the Didymellaceae (de 

Gruyter et al. 2009, Aveskamp et al. 2010). Phoma anamorphs 

and Phoma-like species in Coniothyriaceae, Leptosphaeriaceae, 

Melanommataceae, Montagnulaceae, Pleosporaceae, 

Sporormiaceae and Trematosphaeriaceae are redisposed here as 

a result of this and previous studies. 

The delimitation of Leptosphaeriaceae in Pleosporineae 

from Cucurbitariaceae, Didymellaceae, Phaeosphaeriaceae and 

Pleosporaceae agrees with recent studies of Phoma-like species 

in Pleosporales (de Gruyter et al. 2009, Aveskamp et al. 2010, de 

Gruyter et al. 2010). Cucurbitariaceae is recognised as the fifth 

family in Pleosporineae in addition to the four families accepted by 

Zhang et al. (2009), which are Didymellaceae, Leptosphaeriaceae 

Phaeosphaeriaceae and Pleosporaceae. 

The genera Leptosphaeria, Paraleptosphaeria, 

Plenodomus, Subplenodomus and Heterospora 

Plenodomus lingam and L. doliolum, the type species of 

Plenodomus and Leptosphaeria respectively, were found to be 

distant genetically, which agrees with findings of previous molecular 

phylogenetic studies (Jasalavic et al. 1995, Morales et al. 1995, 

Dong et al. 1998, Câmara et al. 2002, Eriksson & Hawksworth 

2003, Wunsch & Bergstrom 2011). In our study the generic type 

species grouped in sister clades, which represent Leptosphaeria 

and Plenodomus. Species of Leptosphaeria produce dark brown, 

3-septate ascospores, which have been considered the primitive 

state with more recently evolved species producing ascospores that 

are paler in colour, longer and narrower, and more than 3-septate 

(Wehmeyer 1946). This hypothesis is supported by the results 

obtained in our study. Paraleptosphaeria is distinct but seems to 

be most closely related to Leptosphaeria producing 3(–5)-septate, 

yellow/brown or hyaline ascospores. Both genera include only 

necrotrophic species. Plenodomus and Subplenodomus include 

necrotrophs and plant pathogens. Ascospores in Plenodomus are 

3–7-septate, whereas in Subplenodomus no sexual state has thus 

far been recorded. The scleroplectenchymatous pycnidial cell wall 

is typical for Plenodomus, whereas in Subplenodomus the pycnidial 

cell wall is pseudoparenchymatous. Heterospora is closely allied 

to Subplenodomus and no sexual state has been recorded for 

this genus either. The distinctive characterisitics of the genera 

Heterospora, Leptosphaeria, Paraleptosphaeria, Plenodomus 

and Subplenodomus are summerised in Table 2. A blast search 

in GenBank using ITS sequences of five selected species of the 

Leptosphaeriaceae, namely L. doliolum, L. etheridgei, Plen. lingam, 

H. dimorphospora and Subplen. drobnjacensis, did not reveal close 

matches to other teleomorphic or anamorphic genera. 

Plectophomella visci grouped in Plenodomus in this study 

and in the Leptosphaeriaceae in a previous molecular phylogeny 

of Phoma and allied anamorph genera (de Gruyter et al. 2009). 

Plectophomella visci is the type species of Plectophomella (Moesz 


31


Table 2. Characteristics of ascospores, mitosporic state and pathogenicity of Leptosphaeria, Paraleptosphaeria, Plenodomus and 

Subplenodomus in vivo. 

Genus Ascospores Mitosporic state Pathogenicity 

Leptosphaeria Ascospores 3-septate, (dark) brown Mitosporic state common, pycnidial cell wall usually directly 

scleroplectenchymatous, conidia mostly aseptate 

Paraleptosphaeria 

Plenodomus 

Ascospores 3–5-septate, hyaline to 

yellow/brown 

Ascospores 3–7-septate, pale yellow 

to brown 

Mitosporic state rare, pycnidial cell wall directly scleroplectenchymatous, 

conidia aseptate 

Mitosporic state common, pycnidial cell wall initially 

pseudoparenchymatous, later scleroplectenchymatous, conidia aseptate 

Subplenodomus No known sexual state Mitosporic state common, pycnidial cell wall mainly 

pseudoparenchymatous, conidia aseptate 

Heterospora No known sexual state Mitosporic state common, pycnidial cell wall pseudoparenchymatous, 

conidia of two types: small aseptate and large septate 

Necrotrophic 

Necrotrophic 

Necrotrophic or plant pathogenic 

Necrotrophic or plant pathogenic 

Plant pathogenic 

1922) and three additional species have been described in 

the genus. Two species were described from the bark of Ulmus 

spp., viz. Plectophomella ulmi (basionym Dothiorella ulmi) and 

Plectophomella concentrica (Redfern & Sutton 1981). Dothiorella 

ulmi is considered the appropriate name for Plectophomella ulmi 

(Crous et al. 2004). A third species, Plectophomella nypae, was 

described from Nypa fruticans (Arecaceae) (Hyde & Sutton 1992). 

As a result of the transfer of the type species Plectophomella visci 

to Plenodomus, the taxonomy of both Plectophomella concentrica 

and P. nypae needs to be reconsidered based on the outcome of a 

molecular study. 

Plenodomus chrysanthemi could not be differentiated from 

Plen. tracheiphilus based on comparison of their LSU and ITS 

sequences. Plenodomus vasinfecta was proposed by Boerema et 

al. (1994) for the species originally described as Phoma tracheiphila 

f. sp. chrysanthemi (Baker et al. 1985). Because these are part 

of the Plenodomus clade the name Plenodomus chrysanthemi is 

proposed with P. tracheiphila f. sp. chrysanthemi and P. vasinfecta 

as synonyms. Plenodomus chrysanthemi and Plen. tracheiphilus 

are host specific (Chrysanthemum and Citrus, respectively) and the 

scleroplectenchymatous conidiomatal wall of Plen. tracheiphilus 

differentiates this species from Plen. chrysanthemi, where only 

a parenchymatous wall has been observed (Boerema et al. 

1994). The results of this molecular study and the production of a 

Phialophora synanamorph by both species demonstrate the close 

relationship of both taxa. 

Plenodomus enteroleucus and Plen. influorescens have a 

similar ecological niche as opportunistic pathogens on woody 

plants in Europe. Both taxa were formerly described as varieties 

of Ph. enteroleuca, vars. enteroleuca and influorescens, and could 

be differentiated only by the fluorescence of var. enteroleuca under 

black light. However, the molecular phylogeny demonstrates the 

two varieties are only distantly related and they are raised from 

varietal status to species rank. The close relation of Plen. wasabiae 

with Plen. biglobosus agrees with the results of a previous study on 

the production of Phomalignin A and other yellow pigments, as well 

as ITS sequence analyses (Pedras et al. 1995). 

Subplenodomus apiicola, Subplen. drobnjacensis, Subplen. 

valerianae and Subplen. violicola all produce pycnidia with 

an elongated neck, resembling Plenodomus. The pycnidial 

wall remains usually pseudoparenchymatous. Pycnidia with 

a scleroplectenchymatous wall are only observed in Subplen. 

drobnjacensis. Subplenodomus apiicolus, Subplen. drobnjacensis 

and Subplen. valerianae produce relatively small conidia, up to 

4.5 × 2 μm (de Gruyter & Noordeloos 1992) in congruence with 

many of the Plenodomus species described; however, in contrast 

Subplen. violicola produces relatively large conidia, up to 11 × 3 μm 

(Boerema 1993). 

The grouping of species of Phoma section Plenodomus based 

on the host being either herbaceous plants or wood of trees and 

shrubs (Boerema 1982, Boerema et al. 1994) is not supported 

by the molecular phylogeny. The grouping of the species into two 

categories based on the production of pseudoparenchymatous 

pycnidia that become scleroplectenchymatous pycnidia (type I), 

versus always scleroplectenchymatous pycnidia (type 2) (Boerema 

et al. 1981), is partly supported by the molecular phylogeny. 

In the Leptosphaeria clade most species directly develop 

scleroplectenchymatous pycnidia, whereas in the Plenodomus 

clade the pycnidia generally are pseudoparenchymatous and 

become scleroplectenchymatous. 

Heterospora is established for two species of Phoma sect. 

Heterospora that cluster in the Leptosphaeriaceae, viz H. 

chenopodii and H. dimorphospora. All other species of Phoma sect. 

Heterospora are in the Didymellaceae (Aveskamp et al. 2010). 

The Leptosphaeria doliolum species complex 

The taxonomy of the generic type species Leptosphaeria doliolum 

and Phoma anamorphs is complex with a number of subspecies 

and varieties described in literature. Leptosphaeria doliolum subsp. 

doliolum and L. doliolum subsp. errabunda are morphologically 

very similar, as well as the anamorphs Ph. acuta subsp. errabunda 

and Ph. acuta subsp. acuta. It has been suggested that both 

taxa represent originally American and European counterparts 

(Boerema et al. 1994). Both subspecies of L. doliolum proved to be 

closely related in a phylogenetic analysis utilising LSU and ITS. A 

detailed multilocus phylogenetic study including the ITS, ACT, TUB 

and CHS genes, however, demonstrated that both subspecies 

could be clearly differentiated, and represent two subclades in 

the L. doliolum complex. All species allied with L. doliolum and L. 

errabunda are necrotrophic species. Surprisingly, L. macrocapsa 

grouped with the L. errabunda isolates. Leptosphaeria macrocapsa 

is described as a host-specialised necrotroph on Mercurialis 

perennis (Euphorbiaceae) in Europe (Boerema et al. 1994). The 

species is characterised by large pycnidia (Grove, 1935), with a 

conspicuously broad, long cylindrical neck (Boerema et al. 1994). 

This is different to the sharply delimited papilla or neck of variable 

length of the pycnidia of L. errabunda. Leptosphaeria sydowii, a 

necrotroph on Senecio spp. in particular (Asteraceae), proved to 

be closely related to L. errabunda. It can be concluded that the 

Leptosphaeria doliolum complex includes several necrotrophic 

species, with adapted host specificity. 

32


The genus Coniothyrium 

Coniothyrium palmarum is the type species of the genus 

Coniothyrium. Coniothyrium is characterised by ostiolate pycnidial 

conidiomata, annellidic conidiogenous cells, the absence of 

conidiophores, and brown, thick-walled, 0- or 1-septate, verrucose 

conidia. Coniothyrium is similar morphologically to some species 

in the genus Microsphaeropsis. However, Microsphaeropsis is 

characterised by the production of phialidic conidiogenous cells 

with periclinal thickening, and thin-walled, pale greenish brown 

conidia. 

Coniothyrium, Microsphaeropsis and Paraconiothyrium clearly 

grouped in different clades in a study of the partial SSU nrDNA 

(Verkley et al. 2004). In a subsequent study utilising SSU and LSU 

sequences, the generic type species Microsphaeropsis olivacea 

grouped in Didymellaceae, whereas Coniothyrium palmarum 

clustered with the genus Leptosphaeria in Leptosphaeriaceae 

(de Gruyter et al. 2009). In the present study C. palmarum 

and its relatives grouped in a distinct clade, which represents 

Coniothyriaceae. Phoma carteri, Ph. glycinicola, Ph. septicidalis 

and Pyrenochaeta dolichi grouped in this clade and are transferred 

to the genus Coniothyrium. The inclusion of these species with 

setose pycnidia and conidiogenesis with elongated conidiophores 

expands the morphological circumscription of Coniothyrium. 

Species with those characters are also found in other genera 

treated in this paper in the Cucurbitariaceae, Didymellaceae, 

Phaeosphaeriaceae, Leptosphaeriaceae, Montagnulaceae and 

Sporormiaceae, indicating convergent evolution. 

The Coniothyrium species included here are plurivorous or soilborne, 

such as C. palmarum, C. septicidalis and C. multiporum, or 

are associated with a specific host such as C. carteri on Quercus 

spp. (Fagaceae), C. glycinicola on Glycine max (Fabaceae) and 

C. dolichii on Dolichos biflorus (Fabaceae). The species also are 

diverse geographically. 

Coniothyrium palmarum was frequently found associated 

with leaf spots on Phoenix dactylifera (Arecaceae) in India and 

Cyprus (Sutton 1980). The C. palmarum isolates regularly used 

in phylogenetic studies are CBS 758.73, from leaf spots on 

Phoenix dactylifera in Israel, and CBS 400.71, from a dead petiole 

of Chaemeropsis humulis (Arecaceae) in Italy. The subtropical 

distribution of these species is similar to that of the most closely 

allied C. dolichi and C. glycinicola. Coniothyrium multiporum, 

recorded from marine soil, also is found in warm regions. 

Coniothyium carteri, in contrast, is reported from North America 

and Europe. 

Coniothyrium dolichi produces setose pycnidia with hyaline 

conidia (Mohanty 1958). The conidiogenesis was studied in 

detail later. Phoma-like ampulliform conidiogenous cells as well 

as conidiogenous cells on filiform, septate conidiophores were 

found in the same pycnidia leading to confusion regarding the 

classification of this species in Phoma or Pyrenochaeta (Grodona 

et al. 1997). This study clearly supports the classification in 

Coniothyrium. Coniothyrium glycinicola was originally placed 

in the genus Pyrenochaeta as Py. glycines due to its setose 

pycnidia (Stewart 1957). The conidiogenesis and hyaline 

conidia are Phoma-like and therefore, it was reclassified as Ph. 

glycinicola in Phoma sect. Paraphoma (de Gruyter & Boerema 

2002). However, in the original description it was noted that the 

conidia were greenish-yellow in mass (Stewart 1957), resembling 

Microsphaeropsis or Coniothyrium-like conidia. This study clearly 

supports the classification in Coniothyrium. Coniothyrium carteri 

produces setose pycnidia with hyaline conidia and therefore, the 

species was classified in Phoma section Paraphoma (de Gruyter & 

Boerema 2002). In spite of this similarity, C. carteri was determined 

to be only distantly related to the generic type species Paraphoma 

radicina (de Gruyter et al. 2010). Coniothyrium multiporum was 

described in Phoma section Phoma; however, it proved to be 

unrelated to Phoma in Didymellaceae (Aveskamp et al. 2010). The 

conidiogenesis may comprise elongated conidiophores (Pawar et 

al. 1967). Two isolates originally described as Ph. septicidalis are 

placed here in Coniothyrium telephii. Other strains deposited as 

Ph. septicidalis proved to be Pyrenochaeta unguis-hominis (de 

Gruyter et al. 2010). 

The anamorph of the genus Neophaeosphaeria was described 

as Coniothyrium-like, producing pigmented, aseptate conidia 

from holoblastic, percurrently proliferating conidiogenous cells 

with conspicuous annellations (Câmara et al. 2003). Although 

Neophaeosphaeria is related to Coniothyrium based on the 

molecular data, Neophaeosphaeria probably belongs to a separate 

phylogenetic clade. The grouping of N. filamentosa with the 

Coniothyrium species included in this study was poorly supported 

and N. filamentosa proved to be more distantly related in previous 

molecular phylogenetic studies (Verkley et al. 2004, Damm et al. 

2008, de Gruyter et al. 2010). 

Both anamorph genera Cyclothyrium and Cytoplea were 

considered to be related to Coniothyrium and Microsphaeropsis 

(Sutton 1980) based on morphological similarities. Cyclothyrium 

also resembles Paraconiothyrium but produces conidiogenous cells 

that are more elongated than in most species of Paraconiothyrium 

and the conidia are almost truncate at the base, or at least 

they are much less rounded at the base than the conidia of 

Paraconiothyrium (Verkley et al. 2004). The generic type species 

Cyclothyrium juglandis, the anamorph of Thyridaria rubronotata, 

proved to be related to Roussoella hysterioides, teleomorph 

of Cytoplea (Verkley et al. 2004). Based on present results R. 

hysterioides could not be assigned to familial rank. The clustering 

of this species in Massariaceae (Zhang et al. 2009) could not be 

confirmed. Moreover, Roussoella probably is not a monophyletic 

genus (Tanaka et al. 2009). Thyridaria rubronotata, the teleomorph 

of Cyclothyrium juglandis, proved to be related to Massariosphaeria 

phaeospora but was not assigned to familial rank (Schoch et al. 

2009). 

Coniothyrium-like anamorphs also have been linked to 

Mycosphaerella in the past. However, these species were 

subsequently accommodated in Colletogloeopsis (Cortinas et al. 

2006), Readeriella/Kirramyces (Crous et al. 2007) and are now 

known to be species of Teratosphaeria (Crous et al. 2009b). 

The genus Pleospora 

Pleospora is a large genus in Pleosporaceae, Pleosporales, and 

includes important pathogens that occur on both monocotyledons 

and dicotyledons. Anamorphs of Pleospora s. lat. have been 

described in various genera of coelomycetes and hyphomycetes as 

summarised by Zhang et al. (2009, 2012). A delimitation of Pleospora 

into two sections, Pyrenophora and Eu-Pleospora was made based 

on the size of fruiting bodies and ascospore septation and colour 

(Munk 1957). The genus Pyrenophora (Drechslera anamorphs) 

is recognised at the generic rank. However, Pleospora remains 

heterogenous (Wehmeyer 1961, Berbee 1996) and molecular 

phylogenetic studies demonstrated that Pleospora is polyphyletic in 

Pleosporaceae (Kodsueb et al. 2006, Wang et al. 2007, Inderbitzin 


33


et al. 2009). Taxa with a Stemphylium anamorph such as Pleospora 

sedicola and Pleo. tomatonis, as well as Pleo. halophola with no 

known anamorph, are closely related to Cochliobolus, whereas 

Pleo. herbarum and Pleo. ambigua were more distantly related 

in the Pleosporaceae (Kodsueb et al. 2006, Wang et al. 2007). A 

phylogenetic study of the genus Massariosphaeria demonstrated 

the polyphyly in the genera Pleospora, Kirschsteiniothelia, 

Massarina, Melanomma, Trematosphaeria and Massariosphaeria 

in the Loculoascomycetes (Wang et al. 2007) and the paraphyletic 

character of the genus Cochliobolus was demonstrated (Kodsueb 

et al. 2006, Mugambi & Huhndorf 2009). These findings support 

the previous speculation by several authors that ascomatal and 

ascospore morphologies have undergone convergent evolution 

among Pleosporales (Wang et al. 2007). 

Pleospora betae groups ambiguously in Pleosporaceae (Dong 

et al. 1998). SSU nrDNA sequence data supported the affinity of 

P. betae to Leptosphaeriaceae. Partial LSU nrDNA data supported 

the affinity of P. betae to Pleosporaceae (Dong et al. 1998), but 

bootstrap support values in that study were low. In a multigene 

phylogenetic study Pleo. betae was found as being basal to 

Pleosporaceae (Zhang et al. 2009). Our results demonstrate the 

sister group relationship of Pleo. betae and its relatives to the 

generic type species Pleo. herbarum. 

Pleospora betae has been often confused with Pleo. 

calvescens as was discussed by Boerema et al. (1987). 

Both species are pathogens of Chenopodiaceae and are 

morphologically rather similar and therefore, a phylogenetic 

relation of both species was inferred (Boerema 1984). In addition 

Ascochyta hyalospora, originally found on the American continent 

on Chenopodiaceae, also was supposed to be closely related. 

Our results demonstrate that Pleo. betae and Pleo. calvescens 

could be recognised at species rank and confirmed that A. 

hyalospora is related supporting our transfer to Pleospora as 

Pleo. chenopodii. The delimitation of both halophytic species 

Pleo. chenopodii and Pleo. calvescens needs further study; both 

species could not be clearly differentiated based on the ACT 

sequences alone. Additional studies are underway to elucidate 

these species boundaries, in which also the recently described 

halophyte, Ascochyta manawaorae (Verkley et al. 2010), will 

be included. Pleospora fallens and Pleo. incompta, formerly 

described in Phoma sect. Phoma and producing mainly glabrous 

pycnidia, grouped in the Pleo. herbarum clade. Pleospora 

typhicola, producing pilose pycnidia, also grouped in this clade. 

Phoma-like species excluded from the Pleosporineae 

The genus Paraconiothyrium was introduced by Verkley et al. 

(2004) as the anamorph of Paraphaeosphaeria. The morphological 

characters of Paraconiothyrium are variable. The conidiomata can 

be eustromatic to pycnidial, the phialidic conidiogenous cells are 

discrete or integrated, and the thin-walled conidia are aseptate 

or septate, smooth-walled or minutely warted, and hyaline to 

brown in a later stage (Verkley et al. 2004). The morphological 

characters of Ph. lini and Asteromella tilliae, redisposed here in 

Paraconiothyrium, fit this description. 

Paraconiothyrium fuckelii is a serious plant pathogen of 

Rosaceae (Horst & Cloyd 2007), but it also is recorded as an 

opportunistic human pathogen as summarised by de Hoog et 

al. (2000). The teleomorph is currently known as Leptosphaeria 

coniothyrium, but this is not likely considering the phylogeny of 

Leptosphaeriaceae in Pleosporales (Fig 1). The species was also 

described as Melanomma coniothyrium (Holm 1957); however, 

Melanomma is more distantly related in Melanommataceae. 

Neottiosporina paspali proved to be related to Paraconiothyrium. 

However, this species is characterised by conidia with an 

apical appendage (Sutton 1980) and resembles members of 

Massarinaceae. Pyrenochaeta romeroi is redescribed in the new 

genus Medicopsis, and its taxonomic position is most close to 

Trematosphaeriaceae. 

Aposphaeria corallinolutea could be recognised as a 

new species in Melanommataceae. Phoma capitulum and 

Ph. minutispora (Phoma section Phoma) clustered in the 

Sporormiaceae, most closely related to the holotype isolate of 

Westerdykella ornata. Other Phoma-like anamorphs have been 

recorded in Sporormiaceae, such as anamorphs of Sporormia 

aemulans (≡ Preussia aemulans) and Westerdykella dispersa 

(≡ Pycnidiophora dispersa) (von Arx & Storm 1967). The in vitro 

characters of W. capitulum and W. oryzae agree with the in vitro 

characters of Phoma-like anamorphs in the Sporormiaceae 

summarised by Boerema et al. (2004). The conidia produced are 

small, mostly 2–3 × 1–2 μm, arising from undifferentiated cells, but 

sometimes also elongated conidiogenous cells are observed. The 

colonies, often with a pink-yellow-red discolouration on OA, usually 

produce little aerial mycelium, whereas pycnidia are often produced 

in abundance. No matching sequences were found in a blast search 

in GenBank using the partial LSU sequences of W. capitulum and 

W. minutispora. Westerdykella minutispora from India was most 

similar to a sequence of Westerdykella nigra, isolate CBS 416.72, 

obtained from soil in Pakistan, and W. capitulum was most similar 

to a sequence of W. dispersa, isolate CBS 297.56, obtained from 

a seedling of Phlox drummondii, USA. These blast results support 

the redisposition of both species in the genus Westerdykella. 

ACKNOWLEDGEMENTS 

This project, ”Strengthening the Plant Health Infrastructure”, was supported by The 

Dutch Ministry of Economic Affairs, Agriculture and Innovation. We thank Mrs Trix 

Merkx and Mrs Karin Rosendahl-Peters for providing the strains from the culture 

collection of CBS and PD respectively and for their assistance in the deposit of 

strains. Mrs Arien van Iperen kindly helped us with the deposit of herbarium material. 

Thanks are due to Marjan Vermaas for her assistance in preparing the photoplates. 

We are indebted to Machiel E. Noordeloos and the reviewers for critical reading of 

the manuscript. 

REFERENCES 

Aa HA van der, Kesteren HA van (1979). Some pycnidial fungi occurring on Atriplex 

and Chenopodium. Persoonia 10: 267–276. 

Arx JA von, Storm PK (1967). Über einige aus dem erdboden isolierte, zu Sporormia, 

Preussia und Westerdykella gehörende Ascomyceten. Persoonia 4: 407–415. 

Aveskamp MM, Gruyter J de, Crous PW (2008). Biology and recent developments in 

the systematics of Phoma, a complex genus of major quarantine significance. 

Fungal Diversity 31: 1–18. 

Aveskamp MM, Gruyter J de, Woudenberg JHC, Verkley GJM, Crous PW (2010). 

Highlights of the Didymellaceae: A polyphasic approach to characterise Phoma 

and related pleosporalean genera. Studies in Mycology 65: 1–60. 

Aveskamp MM, Verkley GJM, Gruyter J de, Murace MA, Perelló A, Woudenberg 

JHC, Groenewald JZ, Crous PW (2009). DNA phylogeny reveals polyphyly of 

Phoma section Peyronellaea and multiple taxonomic novelties. Mycologia 101: 

359–378. 

Badali H, Chander J, Gulati N, Attri A, Chopra R, Najafzadeh MJ, Chhabra S, Meis 

JFGM, Hoog GS de (2010). Subcutaneous phaeohyphomycotic cyst caused by 

Pyrenochaeta romeroi. Medical Biology 48: 763–768. 

Baker KF, Davis-Clark LH, Wilhelm S, Snyder WC (1985). An aggressive vascularinhabiting 

Phoma (Phoma tracheiphila f.sp. chrysanthemi nov. f. sp.) weakly 

pathogenic to Chrysanthemum. Canadian Journal of Botany 63: 1730–1735. 

34


Berbee ML (1996). Loculoascomycete origins and evolution of filamentous 

Ascomycete morphology based on 18s rRNA gene sequence data. Molecular 

Biology and Evollution 13: 462–470. 

Boerema GH (1970). Additional notes on Phoma herbarum. Persoonia 6: 15–48. 

Boerema GH (1982). Phoma-soorten van de sectie Plenodomus. Verslagen en 

Mededelingen Plantenziektenkundige Dienst Wageningen 158 (Jaarboek 

1981): 28–30. 

Boerema GH (1984). Mycologisch-taxonomisch onderzoek. Ascochyta’s met 

lichtbruine conidien die pathogen zijn voor Chenopodiaceae. Verslagen en 

Mededelingen Plantenziektenkundige Dienst Wageningen 162 (Jaarboek 

1983): 31–34. 

Boerema GH (1993). Contributions towards a monograph of Phoma (Coelomycetes) 

– II. Section Peyronellaea. Persoonia 15: 197–221. 

Boerema GH (1997). Contributions towards a monograph of Phoma (Coelomycetes) 

– V. Subdivision of the genus in sections. Mycotaxon 64: 321–333. 

Boerema GH and Coworkers (1993). Check-list for scientific names of common 

parasitic fungi. Libri botanici; Vol. 10. IHW-Verlag, Eching. 

Boerema GH, Gams W (1995). What is Sphaeria acuta Hoffm. : Fr.? Mycotaxon 

53: 355–360. 

Boerema GH, Gruyter J de (1998). Contributions towards a monograph of Phoma 

(Coelomycetes) – VII. Section Sclerophomella: Taxa with thick-walled 

pseudoparenchymatous pycnidia. Persoonia 17: 81–95. 

Boerema GH, Gruyter J de (1999). Contributions towards a monograph of Phoma 

(Coelomycetes) – III-Supplement: Additional species of section Plenodomus. 

Persoonia 17: 273–280. 

Boerema GH, Gruyter J de, Kesteren HA van (1994). Contributions towards a 

monograph of Phoma (Coelomycetes) – III-1. Section Plenodomus: Taxa often 

with a Leptosphaeria teleomorph. Persoonia 15: 431–487. 

Boerema GH, Gruyter J de, Noordeloos ME (1997). Contributions towards a 

monograph of Phoma (Coelomycetes) – IV. Section Heterospora: Taxa 

with large sized conidial dimorphs, in vivo sometimes as Stagonosporopsis 

synanamorphs. Persoonia 16: 335–371. 

Boerema GH, Gruyter J de, Noordeloos ME, Hamers MEC (2004). Phoma 

identification manual. Differentiation of specific and infra-specific taxa in 

culture. CABI Publishing, Wallingford, UK. 

Boerema GH, Kesteren HA van (1964). The nomenclature of two fungi parasitizing 

Brassica. Persoonia 3: 17–28. 

Boerema GH, Kesteren HA van, Loerakker WM (1981). Notes on Phoma. 

Transactions of the British Mycological Society 77: 61–74. 

Boerema GH, Loerakker WM (1985). Notes on Phoma 2. Transactions of the British 

Mycological Society 84: 289–302. 

Boerema GH, Loerakker WM, Hamers MEC (1987). Check-list for scientific names 

of common parasitic fungi. Supplement Series 2a (additions and corrections): 

Fungi on field crops: beet and potato; caraway, flax and oilseed poppy. 

Netherlands Journal of Plant Pathology 93, Suppl. 1: 1–20. 

Boerema GH, Loerakker WM, Hamers MEC (1996). Contributions towards a 

monograph of Phoma (Coelomycetes) – III-2. Misapplications of the type 

species-name and the generic synonyms of section Plenodomus (Excluded 

species). Persoonia 16: 141–190. 

Butin H, Kehr R (1995). Leaf blotch of lime associated with Asteromella tiliae comb. 

nov. and the latter’s connection to Didymosphaeria petrakiana. Mycological 

Research 99: 1191–1194. 

Câmara MPS, Palm ME, Berkum P van, O’Neill NR (2002). Molecular phylogeny of 

Leptosphaeria and Phaeosphaeria. Mycologia 94: 630–640. 

Câmara MPS, Ramaley AW, Castlebury LA, Palm ME (2003). Neophaeosphaeria 

and Phaeosphaeriopsis, segregates of Paraphaeosphaeria. Mycological 

Research 107: 516–522. 

Carbone I, Kohn LM (1999). A method for designing primer sets for speciation 

studies in filamentous ascomycetes. Mycologia 91: 553–556. 

Chaverri P, Salgado C, Hirooka Y, Rossman AY, Samuels GJ (2011). Delimitation of 

Neonectria and Cylindrocarpon (Nectriaceae, Hypocreales, Ascomycota) and 

related genera with Cylindrocarpon-like anamorphs. Studies in Mycology 68: 

57–78. 

Chen C-Y, David JC, Hsieh WH (2002). Leptosphaeria dryadis. IMI Descriptions of 

Fungi and Bacteria 154, Sheet 1533. CABI Bioscience, Egham, Surrey, UK. 

Chesters CGC (1938). Studies on British pyrenomycetes II. A comparative study of 

Melanomma pulvis-pyrius (Pers.) Fuckel, Melanomma fuscidulum Sacc. and 

Thyridaria rubro-notata (B. & Br.) Sacc. Transactions of the British Mycological 

Society 22: 116–150. 

Cortinas MN, Burgess T, Dell B, Xu DP, Crous PW, Wingfield BD, Wingfield MJ 

(2006). First record of Colletogloeopsis zuluense comb. nov., causing a stem 

canker of Eucalyptus in China. Mycological Research 110: 229–236. 

Crane, JL, Shearer CA (1991). A nomenclator of Leptosphaeria V. Cesati & G. de 

Notaris (Mycota-Ascomycotina-Loculoascomycetes). Illinois Natural history 

Survey Bulletin 34: 195–355. 

Crous PW, Braun U, Groenewald JZ (2007). Mycosphaerella is polyphyletic. Studies 

in Mycology 58: 1–32. 

Crous PW, Gams W, Stalpers JA, Robert V, Stegehuis G (2004). MycoBank: an 

online initiative to launch mycology into the 21st century. Studies in Mycology 

50: 19–22. 

Crous PW, Schoch CL, Hyde KD, Wood AR, Gueidan C (2009a). Phylogenetic 

lineages in the Capnodiales. Studies in Mycology 64: 17–47. 

Crous PW, Slippers B, Wingfield MJ, Rheeder J, Marasas WOF, Phillips AJL, Alves 

A, Burgess T, Barber P, Groenewald JZ (2006). Phylogenetic lineages in the 

Botryosphaeriaceae. Studies in Mycology 55: 235–253. 

Crous PW, Summerell BA, Carnegie AJ, Wingfield MJ, Hunter GC, Burgess TI, 

Andijc V, Barber PA, Groenewald JZ (2009b). Unravelling Mycosphaerella: do 

you believe in genera? Persoonia 23: 99–118. 

Crous PW, Verkley GJM, Groenewald JZ, Samson RA (2009c). Fungal Biodiversity. 

CBS Laboratory Manual Series. Centraalbureau voor Schimmelcultures, 

Utrecht, Netherlands. 

Damm U, Verkley GJM, Crous PW, Fourie PH, Haegi A, Riccioni L (2008). Novel 

Paraconiothyrium species on stone fruit trees and other woody hosts. 

Persoonia 20: 9–17. 

Dong J, Chen W, Crane JL (1998). Phylogenetic studies of the Leptosphaeriaceae, 

Pleosporaceae and some other Loculoascomycetes based on nuclear 

ribosomal DNA sequences. Mycological Research 102: 151–156. 

Dooly DP, Beckius ML, Jeffery BS, McAllister CK, Radentz WH, Feldman AR, 

Rinaldi MG, Bailey SR, Keeling JH (1989). Phaeohyphomycotic cutaneous 

disease caused by Pleurophoma in a cardiac transplant patient. The Journal of 

Infectious Diseases 159: 503–507. 

Eriksson OE, Hawksworth DL (2003). Saccharicola, a new genus for two 

Leptosphaeria species on sugar cane. Mycologia 95: 426–433. 

Fitt BDL, Brun H, Barbetti MJ, Rimmer SR (2006). World-wide importance of phoma 

stem canker (Leptosphaeria maculans and L. biglobosa) on oilseed rape 

(Brassica napus). European Journal of Plant Pathology 114: 3–15. 

Gräfenhan T, Schroers H-J, Nirenberg HI, Seifert KA (2011). An overview of the 

taxonomy, phylogeny, and typification of nectriaceous fungi in Cosmospora, 

Acremonium, Fusarium, Stilbella, and Volutella. Studies in Mycology 68: 79–113. 

Grondona I, Monte E, Garcia-Acha I, Sutton B (1997). Pyrenochaeta dolichi: an 

example of a confusing species. Mycological Research 101: 1404–1408. 

Grove WB (1935). British stem- and leaf-fungi (Coelomycetes) Vol. 1 

Sphaeropsidales. Cambridge, UK. Cambridge University Press. 

Gruyter J de, Aveskamp MM, Woudenberg JHC, Verkley GJM, Groenewald JZ, 

Crous PW (2009). Molecular phylogeny of Phoma and allied anamorph genera: 

towards a reclassification of the Phoma complex. Mycological Research 113: 

508–519. 

Gruyter J de, Boerema GH (2002). Contributions towards a monograph of Phoma 

(Coelomycetes) – VIII. Section Paraphoma: Taxa with setose pycnidia. 

Persoonia 17 (“2001”): 541–561. 

Gruyter J de, Boerema GH, Aa HA van der (2002). Contributions towards a 

monograph of Phoma (Coelomycetes) – VI-2. Section Phyllostictoides: Outline 

of its taxa. Persoonia 18: 1–53. 

Gruyter J de, Noordeloos ME (1992). Contributions towards a monograph of Phoma 

(Coelomycetes) – I-1. Section Phoma: Taxa with very small conidia in vitro. 

Persoonia 15: 71–92. 

Gruyter J de, Noordeloos ME, Boerema GH (1993). Contributions towards a 

monograph of Phoma (Coelomycetes) – I-2. Section Phoma: Additional taxa 

with very small conidia and taxa with conidia up to 7 mm long. Persoonia 15: 

369–400. 

Gruyter J de, Noordeloos ME, Boerema GH (1998). Contributions towards a 

monograph of Phoma (Coelomycetes) – I-3. Section Phoma: Taxa with conidia 

longer than 7 mm. Persoonia 16: 471–490. 

Gruyter J de, Woudenberg JHC, Aveskamp MM, Verkley GJM, Groenewald JZ, 

Crous PW (2010). Systematic reappraisal of species in Phoma section 

Paraphoma, Pyrenochaeta and Pleurophoma. Mycologia 102: 1066–1081. 

Hawksworth DL, Crous PW, Redhead SA, Reynolds DR, Samson RA,, et al. (2011). 

The Amsterdam declaration on fungal nomenclature. IMA Fungus 2: 105–112. 

Höhnel F von (1911). Fragmente zur Mykologie XIII (713): Über Leptosphaeria 

maculans (Desm.) und Sphaeria lingam Tode. Sitzungsberichte der Kaiserlichen 

Akademie der Wissenschaften in Wien. (Mathematisch-naturwissenschaftliche 

Klasse (Abteilung I) 120: 458–463. 

Höhnel F von (1914). Fragmente zur Mykologie XVI. (XVI. Mitteilung, Nr. 813 

bis 875). Sitzungsberichte der Kaiserlichen Akademie der Wissenschaften 

in Wien. (Mathematisch-naturwissenschaftliche Klasse (Abteilung I) 123: 

49–155. 

Holm L (1957). Études taxonomiques sur les Pléosporacées. Symbolae Botanicae 

Upsalienses 14: 5–188. 

Hoog GS de, Gerrits van den Ende AHG (1998). Molecular diagnostics of clinical 

strains of filamentous Basidiomycetes. Mycoses 41: 183–189. 

Hoog GS de, Guarro J, Gené J, Figueras MJ (2000). Atlas of Clinical Fungi. 2nd 

edition. Centraalbureau voor Schimmelcultures, Utrecht, Netherlands. 

Horst RK, Cloyd R (2007). Compendium of rose diseases and pests. 2nd edition. 

APS press, MN, USA. 


35


Huang YJ, Fitt BDL, Jedryczka M, Dakowska S, West JS, Gladders P, Steed JM, 

Li ZQ (2005). Patterns of ascospore release in relation to phoma stem canker 

epidemiology in England (Leptosphaeria maculans) and Poland (L. biglobosa). 

European Journal of Plant Pathology 111: 263–277. 

Huelsenbeck JP, Ronquist F (2001). MRBAYES: Bayesian inference of phylogenetic 

trees. Bioinformatics 17: 754–755. 

Hyde KD, Sutton BC (1992). Nypaella frondicola gen. et sp. nov., Plectophomella 

nypae sp. nov. and Pleurophomopsis nypae sp. nov. (Coelomycetes) from 

intertidal fronds of Nypa fruticans. Mycological Research 96: 210–214. 

Inderbitzin P, Mehta YR, Berbee ML (2009). Pleospora species with Stemphylium 

anamorphs: a four locus phylogeny resolves new lineages yet does not 

distinguish among species in the Pleospora herbarum clade. Mycologia 101: 

329–339. 

Jasalavic CA, Morales VM, Pelscher LE, Seguin-Swartz G (1995). Comparison 

of nuclear ribosomal DNA sequences from Alternaria species pathogenic to 

crucifers. Mycological Research 99: 604–614. 

Jedryczka M, Fitt BDL, Kachlicki P, Lewartowska E, Balesdent MH, Rouxel T (1999). 

Comparison between Polish and United Kingdom populations of Leptosphaeria 

maculans, cause of stem canker of winter oilseed rape. Zeitschrift für 

Pflanzenkrankheiten und Pflanzenschutz 106: 608–617. 

Khashnobish A, Shearer CA, Crane JL (1995). Reexamination of species of 

Leptosphaeria on asteraceous hosts. Mycotaxon 54: 91–106. 

Kodsueb R, Dhanasekaran V, Aptroot A, Lumyong S, McKenzie EHC, et al. (2006). 

The family Pleosporaceae: intergeneric relationships and phylogenetic 

perspectives based on sequence analyses of partial 28SrDNA. Mycologia 98: 

571–583. 

Lombard L, Crous PW, Wingfield BD, Wingfield MJ (2010a). Species concepts in 

Calonectria (Cylindrocladium). Studies in Mycology 66: 1–13. 

Lombard L, Crous PW, Wingfield BD, Wingfield MJ (2010b). Multigene phylogeny and 

mating tests reveal three cryptic species related to Calonectria pauciramosa. 

Studies in Mycology 66: 15–30. 

Lombard L, Crous PW, Wingfield BD, Wingfield MJ (2010c). Phylogeny and 

systematics of the genus Calonectria. Studies in Mycology 66: 31–69. 

Mendes-Pereira E, Balesdent MH, Brun H, Rouxel T (2003). Molecular phylogeny 

of the Leptosphaeria maculans - L. biglobosa species complex. Mycological 

Research 107: 1287–1304. 

Moesz G (1922). Mycologiai közlemények V. (Mykologische Mitteilungen V.). 

Magyar Botanikai Lapok 21: 5–16. 

Mohanty NN (1958). An undescribed species of Pyrenochaeta on Dolichos biflorus 

Linn. Indian Phytopathology 8: 85–87. 

Morales VM, Jasalavich CA, Pelcher LE, Petrie GA, Taylor JL (1995). Phylogenetic 

relationship among several Leptosphaeria species based on their ribosomal 

DNA sequences. Mycological Research 99: 593–603. 

Mugambi GK, Huhndorf SM (2009). Molecular phylogenetics of Pleosporales: 

Melanommataceae and Lophiostomataceae re-circumscribed 

(Pleosporomycetidae, Dothideomycetes, Ascomycota). Studies in Mycology 

64: 103–121. 

Müller E (1950). Die schweizerischen arten der gattung Leptosphaeria und 

ihrer verwandten. Sydowia, Annales Mycologici editi in Notitiam Scientiae 

Mycologicae Universalis. Horn, Austria 4: 185–319. 

Müller E, Arx JA von (1950). Einige aspecte zur systematik pseudosphärialer 

Ascomyceten. Berichte der Schweizerichen Botanischen Gesellschaft 21: 

329–397. 

Munk A (1957). Danish Pyrenomycetes: a preliminary flora. Dansk botanisk Arkiv 17, 

Dansk Botanisk Forening. Ejnar Munksgaard, Copenhagen, Denmark. 

Page RDM (1996). Treeview: An application to display phylogenetic trees on 

personal computers. Computer Applications in the Biosciences 12: 357–358. 

Pawar, VH, Mathur, PN, Thirumalachar MJ (1967). Species of Phoma isolated 

from marine soils in India. Transactions of the British Mycological Society 50: 

259–265. 

Pedras MSC, Taylor JL, Morales VM (1995). Phomaligin A and other yellow pigments 

in Phoma lingam and P. wasabiae. Phytochemistry 38: 1215–1222. 

Phillips AJL, Alves A, Pennycook SR, Johnston PR, Ramaley A, et al. (2008). 

Resolving the phylogenetic and taxonomic status of dark-spored teleomorph 

genera in the Botryosphaeriaceae. Persoonia 21: 29–55. 

Redfern DB, Sutton BC (1981). Canker and dieback of Ulmus glabra caused by 

Plectophomella concentrica, and its relationship to P. ulmi. Transactions of the 

British Mycological Society 77: 381–390. 

Rehner SA, Samuels GJ (1994). Taxonomy and phylogeny of Gliocladium analysed 

from nuclear large subunit ribosomal DNA sequences. Mycological Research 

98: 625–634. 

Schoch CL, Crous PW, Groenewald JZ, Boehm EWA, Burgess TI (2009). A classwide 

phylogenetic assessment of Dothideomycetes. Studies in Mycology 64: 

1–15. 

Schroers H-J, Gräfenhan T, Nirenberg HI, Seifert KA (2011). A revision of 

Cyanonectria and Geejayessia gen. nov., and related species with Fusariumlike 

anamorphs. Studies in Mycology 68: 115–138. 

Shoemaker RA, Brun H (2001). The teleomorph of the weakly aggressive segregate 

of Leptosphaeria maculans. Canadian Journal of Botany 79: 412–419. 

Shoemaker RA, Redhead SA (1999). Proposals to conserve the names of four 

species of fungi (Phoma betae, Helminthosporium avenae, Pyrenophora 

avenae and Pleospora tritici-repentis) against competing earlier synonyms. 

Taxon 48: 381–384. 

Sivanesan A (1984). The Bitunicate Ascomycetes and their Anamorphs. J. Cramer, 

Vaduz, Liechtenstein. 

Stewart RB (1957). An undescribed species of Pyrenochaeta on soybean. Mycologia 

49: 115–117. 

Suetrong S, Schoch CL, Spatafora JW, Kohlmeyer J, Volkmann-Kohlmeyer B 

(2009). Molecular systematics of the marine Dothideomycetes. Studies in 

Mycology 64: 155–173. 

Sutton BC (1980). The Coelomycetes. Fungi Imperfecti with Pycnidia, Acervuli and 

Stromata. CMI, Kew, UK. 

Swofford DL (2003). PAUP*: Phylogenetic analysis using parsimony (*and other 

methods), version 4.0b10. Sinauer Associates, Sunderland, Massachusetts. 

Tanaka K, Hirayama K, Yonezawa H, Hatakeyama S, Harada Y, Sano T, 

Shirouzu T, Hosoya T (2009). Molecular taxonomy of bambusicolous fungi: 

Tetraplosphaeriaceae, a new pleosporalean family with Tetraploa-like 

anamorphs. Studies in Mycology 64: 175–209. 

Verkley GJM, Silva M da, Wicklow DT, Crous PW (2004). Paraconiothyrium, a new 

genus to accommodate the mycoparasite Coniothyrium minitans, anamorphs of 

Paraphaeosphaeria, and four new species. Studies in Mycology 50: 323–335. 

Verkley GJM, Woudenberg JHC, Gruyter J de (2010). Ascochyta manawaorae 

Verkley, Woudenberg & de Gruyter, sp. nov. Persoonia 24: 128–129. 

Vilgalys R, Hester M (1990). Rapid genetic identification and mapping of 

enzymatically amplified ribosomal DNA from several Cryptococcus species. 

Journal of Bacteriology 172: 4238–4246. 

Vincenot L, Balesdent MH, Li H, Barbetti MJ, Sivasithamparam K, Gout L, Rouxel T 

(2008). Occurrence of a new subclade of Leptosphaeria biglobosa in Western 

Australia. Phytopathology 98: 321–329. 

Wang HK, Aptroot A, Crous PW, Hyde KD, Jeewon R (2007). The polyphyletic 

nature of Pleosporales: an example from Massariosphaeria based on rDNA 

and RBP2 gene phylogenies. Mycological Research 111: 1268–1276. 

Wehmeyer LE (1946). Studies on some fungi of northwestern Wyoming. III. 

Pleospora and Leptosphaeria. Lloydia. A quarterly Journal of Biological 

Science, Manasha 9: 203–240. 

Wehmeyer LE (1961). A world monograph of the genus Pleospora and its 

segregates. Univ. Michigan Press, USA. 

West JS, Balesdent MH, Rouxel T, Narcy JP, Huang YJ (2002). Colonization of 

winter oilseed rape tissues by A/Tox+ and B/Tox0 Leptosphaeria maculans 

(phoma stem canker) in France and England. Plant Pathology 51: 311–321. 

West JS, Kharbanda PD, Barbetti MJ, Fitt BDL (2001). Epidemiology and 

management of Leptosphaeria maculans (phoma stem canker) on oilseed rape 

in Australia, Canada and Europe. Plant Pathology 50: 10–27. 

White TJ, Bruns T, Lee S, Taylor J (1990). Amplification and direct sequencing of 

fungal ribosomal RNA genes for phylogenetics. In: PCR protocols: a guide to 

methods and applications. Innis MA, Gelfand DH, Sninsky JJ, White TJ, eds. 

Academic press, San Diego, CA, USA: 315–322. 

Woudenberg JHC, Aveskamp MM, Gruyter J de, Spiers AG, Crous PW (2009). 

Multiple Didymella teleomorphs are linked to the Phoma clematidina 

morphotype. Persoonia 22: 56–62. 

Wunsch MJ, Bergstrom GC (2011). Genetic and morphological evidence that 

Phoma sclerotioides, causal agent of brown root rot of alfalfa, is composed of a 

species complex. Phytopathology 101: 594–610. 

Yáňez-Morales M de J, Korf RP, Babcock JF (1998). Fungi on Epifagus 

(Orobanchaceae) – I. On Sclerotium orobanches and its Phoma synanamorph. 

Mycotaxon 67: 275–286. 

Zhang Y, Crous PW, Schoch CL, Hyde KD (2012). Pleosporales. Fungal Diversity 

53: 1–221. 

Zhang Y, Fournier J, Pointing SB, Hyde KD (2008). Are Melanomma pulvis-pyrius 

and Trematosphaeria pertusa congeneric? Fungal Diversity 33: 47–60. 

Zhang Y, Schoch CL, Fournier J, Crous PW, Gruyter J de (2009). Multi-locus 

phylogeny of the Pleosporales: a taxonomic, ecological and evolutionary 

reevaluation. Studies in Mycology 64: 85–102. 

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