StudieS in Mycology 75: 1–36.
available online at www.studiesinmycology.org
redisposition of phoma-like anamorphs in Pleosporales
J. de Gruyter1–3*, J.H.C. Woudenberg1, M.M. Aveskamp1, G.J.M. Verkley1, J.Z. Groenewald1, and P.W. Crous1,3,4
1
CBS-KNAW Fungal Biodiversity Centre, P.O. Box 85167, 3508 AD Utrecht, The Netherlands; 2National Reference Centre, National Plant Protection Organization, P.O. Box 9102,
6700 HC Wageningen, The Netherlands; 3Wageningen University and Research Centre (WUR), Laboratory of Phytopathology, Droevendaalsesteeg 1, 6708 PB Wageningen,
The Netherlands; 4Microbiology, Department of Biology, Utrecht University, Padualaan 8, 3584 CH Utrecht, The Netherlands
*Correspondence: Hans de Gruyter, j.de.gruyter@minlnv.nl
Studies in Mycology
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.
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. Hard copy: June 2013.
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
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
Copyright CBS-KNAW Fungal Biodiversity Centre, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
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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),
2
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
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, Melanomma, Paraphaeosphaeria,
Pleomassaria,
Preussia,
Roussoella,
Splanchnonema,
Sporormiella, Thyridaria, Trematosphaeria and Westerdykella.
www.studiesinmycology.org
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 Bayesian
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,
3
Species name, final identification Former identification cBS no.
other no.
ItS
SSu
LSu
Act
tuB
cHS-1
Host, substrate
country
Pleurophoma sp.
CBS 131286 PD 83/367
JF740329
Kerria japonica (Rosaceae)
Netherlands
Pleurophoma sp.
CBS 131287 PD 83/831
JF740330
Fraxinus excelsior
(Oleaceae)
Netherlands
CBS 543.70
EU754130
Populus canadensis
(Salicaceae)
Netherlands
Pyrenochaeta sp.
CBS 350.82
JF740265
Picea abies (Pinaceae)
Germany
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)
USA
Halimione portulacoides
(Chenopodiaceae)
Netherlands
Aposphaeria corallinolutea sp. nov.
Aposphaeria populina
Chaetodiplodia sp.
Chaetodiplodia sp.
CBS 453.68
CBS 216.75
Chaetosphaeronema hispidulum
DAOM 226212
Cochliobolus sativus
Coniothyrium carteri comb. nov.
Coniothyrium dolichi comb. nov.
Coniothyrium glycines comb. nov.
Coniothyrium multiporum comb. nov.
JF740115
EU754045
EU754144
Anthyllis vulneraria
(Fabaceae)
Germany
DQ677995
DQ678045
(Poaceae)
Unknown
GQ387593
Quercus sp. Fagaceae)
Netherlands
GQ387594
Quercus robur (Fagaceae)
Germany
GQ387610
Dolichos biforus
(Fabaceae)
India
Phoma carteri
CBS 101633 PD 84/74
JF740180
Phoma carteri
CBS 105.91
JF740181
Pyrenochaeta dolichi
CBS 124143 IMI 217261
JF740182
Pyrenochaeta dolichi
CBS 124140 IMI 217262
JF740183
GQ387550
GQ387611
Dolichos biforus
(Fabaceae)
India
Phoma glycinicola
CBS 124455
JF740184
GQ387536
GQ387597
Glycine max (Fabaceae)
Zambia
Phoma glycinicola
CBS 124141 PG-1
JF740185
GQ387598
Glycine max (Fabaceae)
Zimbabwe
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
IMI 294986
GQ387533
CBS 400.71
AY720708
EU754054
EU754153
Chamaerops humilis
(Arecaceae)
Italy
Phoma septicidalis
CBS 188.71
JF740188
GQ387538
GQ387599
Air
Finland
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)
Netherlands
Didymella exigua
CBS 183.55
EU754056
EU754155
Rumex arifolius
(Polygonaceae)
France
Coniothyrium palmarum
Coniothyrium telephii comb. nov.
De Gruyter et al.
4
table 1. Isolates used in this study and their GenBank accession numbers. Name changes and newly generated sequences are indicated in bold.
table 1. (Continued).
www.studiesinmycology.org
Species name, final identification Former identification cBS no.
Didymella lycopersici, anam. Boeremia
lycopersici
Leptosphaeria conoidea
Leptosphaeria doliolum
LSu
JF740097
CBS 200.31
FJ427023
EU754088
Act
tuB
cHS-1
Host, substrate
country
GU237950
Lycopersicon esculentum
(Solanaceae)
Netherlands
GU371827
Elaeis guineensis
(Arecaceae)
Thailand
DQ678080
Juniperus nana
(Cupressaceae)
Switzerland
EU754187
Chenopodium album
(Chenopodiaceae)
Netherlands
Phoma heteromorphospora
CBS 448.68
Phoma heteromorphospora
CBS 115.96
PD 94/1576
JF740227
EU754188
Chenopodium album
(Chenopodiaceae)
Netherlands
Phoma dimorphospora
CBS 345.78
PD 76/1015
JF740203
GU238069
Chenopodium quinoa
(Chenopodiaceae)
Peru
Phoma dimorphospora
CBS 165.78
PD 77/884
JF740204
JF740098
JF740281
Chenopodium quinoa
(Chenopodiaceae)
Peru
Leptosphaeria conoidea,
anam. Phoma doliolum
CBS 616.75
ATCC 32813, IMI 199777,
PD 74/56
JF740201
JF740099
JF740279
Lunaria annua
(Brassicaceae)
Netherlands
Leptosphaeria conoidea,
anam. Phoma doliolum
CBS 125977
PD 82/888
JF740202
JF740280
Senecio sp. (Asteraceae)
Netherlands
Leptosphaeria doliolum
subsp. doliolum var.
doliolum, anam. Phoma
acuta subsp. acuta
CBS 505.75
PD 75/141
JF740205
Leptosphaeria doliolum
subsp. errabunda, anam.
Phoma acuta subsp.
errabunda
CBS 541.66
PD 66/221
Phoma acuta subsp. acuta
f.sp. phloxis
CBS 155.94
Phoma acuta subsp. acuta
f.sp. phloxis
GQ387576
JF740126
JF740144
JF740162
Urtica dioica (Urticaceae)
Netherlands
JF740206
JF740284
JF740127
JF740145
JF740163
Rudbeckia sp. (Asteraceae)
Netherlands
PD 77/80
JF740207
JF740282
JF740128
JF740146
JF740164
Phlox paniculata
(Polemoniaceae)
Netherlands
CBS 125979
PD 78/37
JF740208
JF740283
JF740129
JF740147
JF740165
Phlox paniculata
(Polemoniaceae)
Netherlands
Leptosphaeria doliolum
subsp. doliolum var.
doliolum, anam. Phoma
acuta subsp. acuta
CBS 504.75
PD 74/55
JF740209
JF740130
JF740148
JF740166
Urtica dioica (Urticaceae)
Netherlands
Leptosphaeria doliolum
subsp. doliolum var.
doliolum, anam. Phoma
acuta subsp. acuta
CBS 130000 PD 82/701
JF740210
JF740131
JF740149
JF740167
Urtica dioica (Urticaceae)
Netherlands
Leptosphaeria doliolum
subsp. errabunda, anam.
Phoma acuta subsp.
errabunda
CBS 617.75
JF740216
JF740132
JF740150
JF740168
Solidago sp. (hybrid)
(Asteraceae)
Netherlands
ATCC 32814, IMI 199775,
PD 74/201
GQ387515
JF740289
5
Phoma sections Plenodomus, Pilosa
Leptosphaeria errabunda comb. nov.
SSu
BCC 21118
Herpotrichia juniperi
Heterospora dimorphospora comb. nov.
ItS
CBS 378.67
Falcisormispora lignatilis
Heterospora chenopodii comb. nov.
other no.
Species name, final identification Former identification cBS no.
LSu
Act
tuB
cHS-1
Host, substrate
country
JF740290
JF740133
JF740151
JF740169
Delphinium sp.
(Ranunculaceae)
Netherlands
JF740218
JF740134
JF740152
JF740170
Aconitum sp.
(Ranunculaceae)
Netherlands
CBS 129998 PD 84/462
JF740219
JF740135
JF740153
JF740171
Gailardia (Asteraceae)
Netherlands
Leptosphaeria doliolum
subsp. errabunda, anam.
Phoma acuta subsp.
errabunda
CBS 129997 PD 78/631
JF740220
JF740136
JF740154
JF740172
Achillea millefolium
(Apiaceae)
Netherlands
Leptosphaeria etheridgei comb. nov.
Phoma etheridgei
CBS 125980
DAOM 216539, PD 95/1483
JF740221
JF740291
Populus tremuloides
(Salicaceae)
Canada
Leptosphaeria macrocapsa comb. nov.
Phoma macrocapsa
CBS 640.93
PD 78/139
JF740237
JF740304
Mercurialis perennis
(Euphorbiaceae)
Netherlands
Leptosphaeria pedicularis comb. nov.
Phoma pedicularis
CBS 126582
PD 77/710
JF740223
JF740293
Gentiana punctata
(Gentianaceae)
Switzerland
Phoma pedicularis
CBS 390.80
PD 77/711
JF740224
JF740294
Pedicularis sp.
(Scrophulariaceae)
Switzerland
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
Phoma sclerotioides
CBS 144.84
CECT 20025, PD 82/1061
JF740192
JF740269
Medicago sativa
(Fabaceae)
Canada
Phoma sclerotioides
CBS 148.84
PD 80/1242
JF740193
JF740270
Medicago sativa
(Fabaceae)
Canada
Leptosphaeria slovacica,
anam. Phoma leonuri
CBS 389.80
PD 79/171
JF740247
JF740315
Balota nigra (Lamiaceae)
Netherlands
Leptosphaeria slovacica,
anam. Phoma leonuri
CBS 125975
PD 77/1161
JF740248
JF740316
Balota nigra (Lamiaceae)
Netherlands
Phoma sydowii
CBS 385.80
PD 74/477
JF740244
JF740313
JF740139
JF740157
JF740175
Senecio jacobaea
(Asteraceae)
UK
Phoma sydowii
CBS 125976
PD 84/472
JF740245
JF740314
JF740140
JF740158
JF740176
Senecio jacobaea
(Asteraceae)
Netherlands
Leptosphaeria rubefaciens comb. nov.
Leptosphaeria sclerotioides comb. nov.
Leptosphaeria slovacica
Leptosphaeria sydowii comb. nov.
other no.
ItS
PD 74/61
JF740217
Leptosphaeria doliolum
subsp. errabunda, anam.
Phoma acuta subsp.
errabunda
CBS 125978
Leptosphaeria doliolum
subsp. errabunda, anam.
Phoma acuta subsp.
errabunda
CBS 129999 PD 78/569
Leptosphaeria doliolum
subsp. errabunda, anam.
Phoma acuta subsp.
errabunda
SSu
JF740101
JF740138
JF740137
JF740156
JF740155
JF740174
JF740173
De Gruyter et al.
6
table 1. (Continued).
table 1. (Continued).
www.studiesinmycology.org
Species name, final identification Former identification cBS no.
Leptosphaeria veronicae comb. nov.
ItS
LSu
Act
tuB
cHS-1
Host, substrate
country
JF740141
JF740159
JF740177
Papaver rhoeas
(Papaveraceae)
Switzerland
CBS 297.51
Phoma veronicicola
CBS 145.84
CECT 20059, PD 78/273
JF740254
JF740320
JF740142
JF740160
JF740178
Veronica chamaedryoides
(Scrophulariaceae)
Netherlands
Phoma veronicicola
CBS 126583
PD 74/227
JF740255
JF740321
JF740143
JF740161
JF740179
Veronica ‘Shirley Blue’
(Scrophulariaceae)
Netherlands
JF740246
AB521718
AB521735
Fagus sylvatica (Fagaceae)
UK
CBS 473.64
ETH 2945
GU296170
GU301840
Fagus sylvatica (Fagaceae)
Switzerland
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)
Unknown
GU301845
Wood
France
DQ678072
Fagus sp. (Fagaceae)
Belgium
GQ387516
GQ387577
Yucca rostrata (Agavaceae)
Mexico
GQ387517
GQ387578
Phlox paniculata
(Polemoniaceae)
Netherlands
EU754073
EU754172
Paspalum notatum
(Poaceae)
USA
GQ387613
Human, black grain
mycetoma
Venezuela
GQ387614
Human, mycetoma
Mexico
GU238075
Soil
Netherlands
GU237960
Rubus sp. (Rosaceae)
Denmark
EU754197
Malus sp. (Rosaceae)
USA
EU238093
Wisconsin tank
Netherlands
EU754200
Human, cutaneous lesions
USA
EU754173
Unknown
UK
CBS 122786 PD 99/1064-1
EU754174
Clematis sp.
(Ranunculaceae)
Netherlands
EU754139
Tilia platyphyllos (Tiliaceae)
Austria
GU301828
Dryas octopetala
(Rosaceae)
Switzerland
CBS 371.75
Melanomma pulvis-pyrius
DQ678020
CBS 400.97
Neophaeosphaeria filamentosa
CBS 102202 BPI 802755
Neosetophoma samarorum
CBS 138.96
Neottiosporina paspali
CBS 331.37
JF740259
PD 82/653
Pyrenochaeta mackinnonii
CBS 674.75
FMC 270
Pyrenochaeta mackinnonii
CBS 110022
Paraconiothyrium flavescens comb. nov.
Phoma flavescens
CBS 178.93
Paraconiothyrium fuckelii comb. nov.
Coniothyrium fuckelii
CBS 797.95
Paraconiothyrium fusco-maculans comb.
nov.
Plenodomus fuscomaculans
CBS 116.16
Paraconiothyrium lini comb. nov.
Phoma lini
CBS 253.92
Paraconiothyrium maculicutis sp. nov.
Pleurophoma pleurospora
CBS 101461 IMI 320754, UTHSC 87-144
GQ387552
PD 82/1062
GU238204
PD 70/998
EU754074
CBS 122788 PD 07/03486739
Paraconiothyrium minitans
Paraconiothyrium tiliae comb. nov.
Asteromella tiliae
CBS 265.94
Paraleptosphaeria dryadis comb. nov.
Leptosphaeria dryadis
CBS 643.86
JF740213
7
Phoma sections Plenodomus, Pilosa
H 3953, HHUF 26621, JCM
14422
Massarina eburnea
Nigrogana mackinnonii comb. nov.
SSu
Phoma sydowii
Massarina eburnea
Medicopsis romeroi comb. nov.
other no.
Species name, final identification Former identification cBS no.
other no.
ItS
Host, substrate
country
UPSC 2686
JF740238
JF740305
Rumex domesticus
(Chenopodiaceae)
Norway
CBS 306.51
JF740239
JF740308
Cirsium spinosissimum
(Asteraceae)
Switzerland
Phoma korfii
CBS 101638 PD 97/12070
JF400230
JF740299
Epifagus virginiana
(Orobanchaceae)
USA
Leptosphaeria praetermissa
CBS 114591
JF740241
JF740310
Rubus idaeus (Rosaceae)
Sweden
Paraleptosphaeria macrospora comb.
nov.
Phoma macrospora
CBS 114198
Paraleptosphaeria nitschkei comb. nov.
Leptosphaeria nitschkei
Paraleptosphaeria orobanches comb.
nov.
Paraleptosphaeria praetermissa comb.
nov.
SSu
LSu
Act
tuB
cHS-1
Paraphaeosphaeria michoti
CBS 652.86
ETH 9483
GQ387520
GQ387581
Typha latifolia (Typhaceae)
Switzerland
Paraphoma radicina
CBS 111.79
IMI 386094, PD 76/437
EU754092
EU754191
Malus sylvestris
(Rosaceae)
Netherlands
Phaeosphaeria nodorum
CBS 110109
EU754076
EU754175
Lolium perenne
(Gramineae)
Denmark
Phoma herbarum
CBS 615.75
EU754087
EU754186
Rosa multiflora (Rosaceae)
Netherlands
Phoma paspali
CBS 560.81
PD 92/1569
GU238227
G238124
Paspalum dilatum
(Poaceae)
New
Zealand
Leptosphaeria agnita,
anam. Phoma agnita
CBS 121.89
PD 82/903
JF740194
JF740271
Eupatorium cannabinum
(Asteraceae)
Netherlands
Leptosphaeria agnita,
anam. Phoma agnita
CBS 126584
PD 82/561
JF740195
JF740272
Eupatorium cannabinum
(Asteraceae)
Netherlands
Leptosphaeria biglobosa
CBS 119951
JF740198
JF740274
Brassica rapa
(Brassicaceae)
Netherlands
CBS 127249 DAOM 229269
JF740199
JF740275
Brassica juncea
(Brassicaceae)
France
GU238151
Chrysanthemum sp.
(Asteraceae)
Greece
Plenodomus agnitus comb. nov.
Plenodomus biglobosus comb. nov.
FJ427022
JF740102
Plenodomus chrysanthemi comb. nov.
Phoma vasinfecta,
synanam. Phialophora
chrysanthemi
CBS 539.63
JF740253
Plenodomus collinsoniae comb. nov.
Leptosphaeria collinsoniae
CBS 120227 JCM 13073, MAFF 239583
JF740200
JF740276
Vitis coignetiae (Vitaceae)
Japan
Plenodomus confertus comb. nov.
Leptosphaeria conferta,
anam. Phoma conferta
CBS 375.64
AF439459
JF740277
Anacyclus radiatus
(Asteraceae)
Spain
Plenodomus congestus comb. nov.
Leptosphaeria congesta,
anam. Phoma congesta
CBS 244.64
AF439460
JF740278
Erigeron canadensis
(Asteraceae)
Spain
Plenodomus enteroleucus comb. nov.
Phoma enteroleuca var.
enteroleuca
CBS 142.84
JF740214
JF740287
Catalpa bignonioides
(Bignoniaceae)
Netherlands
Phoma enteroleuca var.
enteroleuca
CBS 831.84
JF740215
JF740288
Triticum aestivum
(Poaceae)
Germany
Leptosphaeria fallaciosa
CBS 414.62
JF740222
JF740292
Satureia montana
(Lamiaceae)
France
Plenodomus fallaciosus comb. nov.
PD 81/654, CECT20063
ETH 2961
GU238230
De Gruyter et al.
8
table 1. (Continued).
table 1. (Continued).
www.studiesinmycology.org
Species name, final identification Former identification cBS no.
Plenodomus hendersoniae comb. nov.
Plenodomus influorescens comb. nov.
Phoma intricans
CBS 113702
Phoma intricans
CBS 139.78
Phoma enteroleuca var.
influorescens
CBS 143.84
Phoma enteroleuca var.
influorescens
other no.
ItS
UPSC 1843
JF740225
JF740226
PD 78/883, CECT 20064
SSu
LSu
Act
tuB
cHS-1
Host, substrate
country
JF740295
Salix cinerea (Salicaceae)
Sweden
JF740296
Pyrus malus (Rosaceae)
Netherlands
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
CBS 386.80
PD 77/336
JF400232
JF740301
Helianthus annuus
(Asteraceae)
former
Yugoslavia
Leptosphaeria lindquistii,
anam. Phoma macdonaldii
CBS 381.67
JF400233
JF740302
Helianthus annuus
(Asteraceae)
Canada
Leptosphaeria maculans,
anam. Phoma lingam
CBS 275.63
MUCL 9901, UPSC 1025
JF400234
JF740306
Brassica sp. (Brassicaceae)
UK
Leptosphaeria maculans,
anam. Phoma lingam
CBS 260.94
PD 78/989
JF400235
JF740116
Brassica oleracea
(Brassicaceae)
Netherlands
Leptosphaeria maculans,
anam. Phoma lingam
CBS 147.24
JF740117
Unknown
Unknown
Plenodomus lupini comb. nov.
Phoma lupini
CBS 248.92
Plenodomus pimpinellae comb. nov.
Leptosphaeria pimpinellae,
anam. Phoma pimpinellae
Plenodomus tracheiphilus comb. nov.
Plenodomus lingam
PD 79/141
JF740103
JF740307
JF740303
Lupinus mutabilis
(Fabaceae)
Peru
CBS 101637 PD 92/41
JF740240
JF740309
Pimpinella anisum
(Apiaceae)
Israel
Phoma tracheiphila
CBS 551.93
PD 81/782
JF740249
JF740317
Citrus limonium (Rutaceae)
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
EU754195
Viscum album (Viscaceae)
France
Plenodomus wasabiae
Phoma wasabiae
CBS 120119
FAU 559
JF740257
JF740323
Wasabia japonica
(Brassicaceae)
Taiwan
Phoma wasabiae
CBS 120120 FAU 561
JF740258
JF740324
Wasabia japonica
(Brassicaceae)
Taiwan
AY004341
Betula verrucosa
(Betulaceae)
Netherlands
EU754096
CBS 279.74
Pleomassaria siparia
Pleospora angustis nom. nov.
Leptosphaeria clavata
CBS 296.51
Pleospora betae
Pleospora betae, anam.
Phoma betae
CBS 523.66
Pleospora betae, anam.
Phoma betae
CBS 109410 PD 77/113
Pleospora calvescens,
anam. Ascochyta caulina
CBS 246.79
Pleospora calvescens
JF740256
JF740104
PD 66/270, IHEM 3915
PD 77/655
EU754080
EU754032
JF740122
Unknown
Switzerland
EU754179
JF740118
Beta vulgaris
(Chenopodiaceae)
Netherlands
EU754178
JF740119
Beta vulgaris
(Chenopodiaceae)
Netherlands
EU754131
JF740120
Atriplex hastata
(Chenopodiaceae)
Germany
9
Phoma sections Plenodomus, Pilosa
JF740236
Species name, final identification Former identification cBS no.
Host, substrate
country
JF740121
Atriplex hastata
(Chenopodiaceae)
Netherlands
JF740109
Chenopodium quinoa
(Chenopodiaceae)
Bolivia
PD 68/682
JF740110
Atriplex hastata
(Chenopodiaceae)
Netherlands
CBS 161.78
LEV 1131
JF740106
Olea europaea
(Oleaeceae)
New
Zealand
Phoma glaucispora
CBS 284.70
PD 97/2400
JF740107
Nerium oleander
(Apocynaceae)
Italy
Pleospora flavigena comb. nov.
Phoma flavigena
CBS 314.80
PD 91/1613
JF740108
Water
Romania
Pleospora halimiones nom. nov.
Ascochyta obiones
CBS 432.77
IMI 282137
JF740113
Halimione portulacoides
(Chenopodiaceae)
Netherlands
Ascochyta obiones
CBS 786.68
JF740114
Halimione portulacoides
(Chenopodiaceae)
Netherlands
JF740123
Medicago sativa
(Fabaceae)
India
Pleospora chenopodii
Pleospora fallens comb. nov.
CBS 343.78
Ascochyta hyalospora
CBS 206.80
PD 74/1022
Pleospora calvescens,
anam. Ascochyta caulina
CBS 344.78
Phoma fallens
CBS 191.86
Pleospora herbarum
Pleospora incompta comb. nov.
Pleospora typhicola
Pleurophoma pleurospora
other no.
Pleospora calvescens,
anam. Ascochyta caulina
ItS
SSu
JF740095
JF740096
IMI 276975
GU238232
LSu
JF740266
JF740267
GU238160
Act
tuB
cHS-1
Phoma incompta
CBS 467.76
JF740111
Olea europaea
(Oleaeceae)
Greece
Phoma incompta
CBS 526.82
JF740112
Olea europaea
(Oleaeceae)
Italy
Pleospora typhicola, anam.
Phoma typharum
CBS 132.69
JF740124
Typha angustifolia
(Typhaceae)
Netherlands
Pleospora typhicola, anam.
Phoma typharum
CBS 602.72
JF740125
Typha sp. (Typhaceae)
Netherlands
Pleurophoma sp.
CBS 116668
JF740326
Citysus scoparius
(Fabaceae)
Netherlands
Pleurophoma sp.
CBS 130329 PD 82/371
JF740327
Lonicera sp.
(Caprifoliaceae)
Netherlands
GU301864
Soil
Senegal
EU754203
Phragmitis australis
(Poaceae)
UK
JF740105
Preussia funiculata
CBS 659.74
Pseudorobillarda phragmitis
CBS 398.61
IMI 070678
GU296187
Pyrenochaeta cava
CBS 257.68
IMI 331911
Pyrenochaeta lycopersici
JF740325
JF740260
EU754100
EU754199
Wheat field soil
Germany
CBS 267.59
JF740261
GQ387551
GQ387612
Lycopersicon esculentum
(Solanaceae)
Netherlands
Pyrenochaeta nobilis
CBS 407.76
EU930011
EU754107/
DQ898287
EU754206
Laurus nobilis (Lauraceae)
Italy
Pyrenochaetopsis leptospora
CBS 101635
JF740262
GQ387566
GQ387627
Secale cereale (Poaceae)
Europe
PD 71/1027
De Gruyter et al.
10
table 1. (Continued).
table 1. (Continued).
www.studiesinmycology.org
Species name, final identification Former identification cBS no.
Pyrenochaetopsis pratorum comb. nov.
Phoma pratorum
other no.
ItS
CBS 445.81
PDDCC 7049, PD 80/1254
JF740263
CBS 286.93
PD 80/1252
JF740264
AY544716
OSC 100066
Pyrenophora tritici-repentis
SSu
LSu
Act
tuB
cHS-1
Host, substrate
country
GU238136
Lolium perenne, leaf
(Poaceae)
New
Zealand
JF740331
Dactylis glomerata
(Poaceae)
New
Zealand
AY544672
(Poaceae)
Italy
AB524622
Sasa kurilensis (Poaceae)
Japan
Roussoella hysterioides
CBS 125434
Setomelanomma holmii
CBS 110217
GQ387572
GQ387633
Picea pungens (Pinaceae)
USA
Setophoma terrestris
CBS 335.29
GQ387526
GQ387587
Allium sativum (Alliaceae)
USA
Splanchnonema platani
CBS 221.37
DQ678013
DQ678065
Platanus occidentalis
(Platanaceae)
USA
Sporormiella minima
CBS 524.50
DQ678003
DQ678056
Dung of goat
Panama
Stagonosporopsis cucurbitacearum
CBS 133.96
GU238234
GU238181
Cucurbita sp.
(Cucurbitaceae)
New
Zealand
JF740196
GU238040
Apium graveolens var.
rapaceum (Umbelliferae)
Germany
JF740273
Apium graveolens
(Umbelliferae)
Netherlands
JF740285
Eustoma exaltatum
(Gentianaceae)
Netherlands
Subplenodomus apiicola comb. nov.
Subplenodomus drobnjacensis comb.
nov.
Subplenodomus valerianae comb. nov.
Subplenodomus violicola comb. nov.
HH 26988
CBS 285.72
Phoma apiicola
CBS 504.91
PD 78/1073
JF740197
Phoma drobnjacensis
CBS 269.92
PD 88/896
JF740211
Phoma drobnjacensis
CBS 270.92
PD 83/650
JF740212
JF740286
Gentiana makinoi ‘Royal
Blue’ (Gentianaceae)
Netherlands
Phoma valerianae
CBS 630.68
PD 68/141
JF740251
GU238150
Valeriana phu
(Valerianaceae)
Netherlands
Phoma valerianae
CBS 499.91
PD 73/672
JF740252
JF740319
Valeriana officinalis
(Valerianaceae)
Netherlands
Phoma violicola
CBS 306.68
FJ427054
GU238156
Viola tricolor (Violaceae)
Netherlands
Phoma violicola
CBS 100272
FJ427055
JF740322
Viola tricolor (Violaceae)
New
Zealand
JF740100
GU238231
Thyridaria rubronotata
CBS 419.85
GU301875
Acer pseudoplatanus
(Aceraceae)
Netherlands
Trematosphaeria pertusa
CBS 122368
FJ201990
Fraxinus excelsior
(Oleaceae)
France
11
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
GU301880
Mangrove mud
Mozambique
Westerdykella ornata
CBS 379.55
Phoma sections Plenodomus, Pilosa
Phoma apiicola
De Gruyter et al.
CBS 246.79 Pleospora calvescens
CBS 432.77 Ascochyta obiones
99
CBS 206.80 Ascochyta hyalospora
Pleosporaceae (A)
CBS 523.66 Pleospora betae, anam. Phoma betae
100
DAOM 226212 Cocliobolus sativus
100
100
CBS 191.86 Pleospora herbarum
100
OSC100066 Pyrenophora tritici-repentis
CBS 132.69 Pleospora typhicola
CBS 165.78 Phoma dimorphospora
100
64
CBS 448.68 Phoma heteromorphospora
56
CBS 269.92 Phoma drobnjacensis
60
CBS 275.63 Leptosphaeria maculans, anam. Phoma lingam
100
Leptosphaeriaceae (B)
CBS 119951 Leptosphaeria biglobosa
57
CBS 551.93 Phoma tracheiphila
CBS 539.63 Phoma vasinfecta
CBS 122783 Plectophomella visci
CBS 363.93 Cucurbitaria berberidis
68
CBS 257.68 Pyrenochaeta cava
99
CBS
407.76
Pyrenochaeta nobilis
Cucurbitariaceae (C)
100
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
CBS 216.75 Chaetosphaeronema hispidulum
CBS 335.29 Setophoma terrestris
88
CBS 110109 Phaeosphaeria nodorum
Phaeosphaeriaceae (E)
100
CBS 138.96 Neosetophoma samarorum
CBS 111.79 Paraphoma radicina
99
80
CBS 110217 Setomelanomma holmii
CBS 124455 Phoma glycinicola
100
93
CBS 124140 Pyrenochaeta dolichi
Clade F
CBS 105.91 Phoma carteri
77
CBS 101636 Phoma septicidalis
CBS 400.71 Coniothyrium palmarum
CBS 102202 Neophaeosphaeria filamentosa
CBS 133.96 Stagonosporopsis bryoniae
100
100
CBS 378.67 Didymella lycopersici
100
CBS 183.55 Didymella exigua
Didymellaceae (G)
100
CBS 615.75 Phoma herbarum
CBS 560.81 Phoma paspali
CBS 122788 Paraconiothyrium minitans
100
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.
12
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
Phoma sections Plenodomus, Pilosa
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
www.studiesinmycology.org
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
De Gruyter et al.
CBS 121.89 Plenodomus agnitus comb. nov. Leptosphaeria agnita
CBS 126584 Plenodomus agnitus comb. nov. Leptosphaeria agnita
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
100
CBS
143.84
Plenodomus influorescens comb. nov. Phoma enteroleuca var. influorescens
83
PD73.1382 Plenodomus influorescens comb. nov. Phoma enteroleuca var. influorescens
100
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
A1
CBS 101637 Plenodomus pimpinellae comb. nov. Leptosphaeria pimpinellae
67
56
CBS 539.63 Plenodomus chrysanthemi comb. nov. Phoma vasinfecta
100
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
CBS
113702
Plenodomus hendersoniae comb. nov. Leptosphaeria hendersoniae
94
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
61
CBS 504.91 Subplenodomus apiicola comb. nov. Phoma apiicola
100 CBS 499.91 Subplenodomus valerianae comb. nov. Phoma valerianae
66
CBS 630.68 Subplenodomus valerianae comb. nov. Phoma valerianae
A2 71
100 CBS 269.92 Subplenodomus drobnjacensis comb. nov. Phoma drobnjacensis
77
CBS 270.92 Subplenodomus drobnjacensis comb. nov. Phoma drobnjacensis
CBS 100272 Subplenodomus violicola comb. nov. Phoma violicola
100 CBS 306.68 Subplenodomus violicola comb. nov. Phoma violicola
CBS 114198 Paraleptosphaeria macrospora comb. nov. Leptosphaeria macrospora
100
100
CBS 306.51 Paraleptosphaeria nitschkei comb. nov. Leptosphaeria nitschkei
CBS
114591 Paraleptosphaeria praetermissa comb. nov. Leptosphaeria praetermissa
99
A3
100 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
A4
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
70
82 CBS 125979 Leptosphaeria doliolum
CBS 541.66 Leptosphaeria doliolum
CBS 155.94 Leptosphaeria doliolum
100
CBS 617.75 Leptosphaeria errabunda comb. nov.
100
CBS 125978 Leptosphaeria errabunda comb. nov.
CBS 640.93 Leptosphaeria macrocapsa comb. nov. Phoma macrocapsa
100
62
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 CBS 125977 Leptosphaeria conoidea
100
100 CBS 389.80 Leptosphaeria slovacica
CBS 125975 Leptosphaeria slovacica
87
100 CBS 390.80 Leptosphaeria pedicularis comb. nov. Phoma pedicularis
CBS 126582 Leptosphaeria pedicularis comb. nov. Phoma pedicularis
53
100 CBS 223.77 Leptosphaeria rubefaciens comb. nov. Phoma rubefaciens
90
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
64
100 IMI 124141 Coniothyrium glycines comb. nov. Phoma glycinicola
64 100 IMI 101633 Coniothyrium carteri comb. nov. Phoma carteri
CBS 105.91 Coniothyrium carteri comb. nov. Phoma carteri
Coniothyriaceae (C)
CBS 400.71 Coniothyrium palmarum
100
97 CBS 188.71 Coniothyrium telephii comb. nov. Phoma septicidalis
100
CBS 856.97 Coniothyrium telephii comb. nov. Phoma septicidalis
96
51
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
Cucurbitariaceae (D)
91 99
CBS 257.68 Pyrenochaeta cava
CBS 363.93 Cucurbitaria berberidis
CBS 407.76 Pyrenochaeta nobilis
Didymellaceae (E)
CBS 615.75 Phoma herbarum
100
100
100
Leptosphaeriaceae (A)
Leptosphaeriaceae (B)
0.1
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).
14
Phoma sections Plenodomus, Pilosa
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 growthrate 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
www.studiesinmycology.org
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
D with Aposphaeria populina and Melanomma pulvis-pyrius in
15
De Gruyter et al.
CBS 617.75 Leptosphaeria errabunda comb. nov.
CBS 129998 Leptosphaeria errabunda comb. nov.
CBS 129997 Leptosphaeria errabunda comb. nov.
99
CBS 129999 Leptosphaeria errabunda comb. nov.
CBS 125978 Leptosphaeria errabunda comb. nov.
Clade A
98
CBS 640.93 Leptosphaeria macrocapsa comb. nov.
CBS 297.51 Leptosphaeria sydowii comb. nov.
83
99
CBS 125976 Leptosphaeria sydowii comb. nov.
CBS 385.80 Leptosphaeria sydowii comb. nov.
52
CBS 504.75 Leptosphaeria doliolum
99
92
97
CBS 130000 Leptosphaeria doliolum
CBS 125979 Leptosphaeria doliolum ‘Phoma acuta subsp. acuta f. sp. phlogis’
Clade B
CBS 155.94 Leptosphaeria doliolum ‘Phoma acuta subsp. acuta f. sp. phlogis’
99
CBS 505.75 Leptosphaeria doliolum
CBS 541.66 Leptosphaeria doliolum
CBS 145.84 Leptosphaeria veronicae comb. nov.
99
CBS 126583 Leptosphaeria veronicae comb. nov.
CBS 390.80 Leptosphaeria pedicularis comb. nov. ‘Phoma pedicularis’
0.1
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).
CBS 109410 Pleospora betae
99
CBS 523.66 Pleospora betae
70
99
CBS 786.68 Pleospora halimiones nom. nov. ‘Ascochyta obiones’
CBS 432.77 Pleospora halimiones nom. nov. ‘Ascochyta obiones’
CBS 206.80 Pleospora chenopodii ‘Ascochyta hyalospora’
100
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.
99
75
69
CBS 467.76 Pleospora incompta comb. nov. ‘Phoma incompta’
CBS 526.82 Pleospora incompta comb. nov. ‘Phoma incompta’
CBS 191.86 Pleospora herbarum
52
100
100
CBS 132.69 Pleospora typhicola ‘Phoma typhina’
CBS 602.72 Pleospora typhicola ‘Phoma typhina’
88
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, CBS 260.94).
16
Phoma sections Plenodomus, Pilosa
CBS 101461 Paraconiothyrium maculicutis sp. nov. ‘Pleurophoma pleurospora’
82
CBS 265.94 Paraconiothyrium tiliae comb. nov. ‘Asteromella tiliae’
57
CBS 797.95 Paraconiothyrium fuckelii comb. nov. ‘Coniothyrium fuckelii’
97
94
CBS 116.16 Paraconiothyrium fusco-maculans comb. nov. ‘Plenodomus fusco-maculans’
Montagnulaceae (A)
CBS 253.92 Paraconiothyrium lini comb. nov. ‘Phoma.lini’
97
CBS 122788 Paraconiothyrium minitans
99
CBS 122786 Paraconiothyrium minitans
CBS 652.86 Paraphaeosphaeria michotii
99
CBS 473.64 Massarina eburnea
H 3953 Massarina eburnea
55 94
Massarinaceae (B)
CBS 331.37 Neottiosporina paspali
CBS 675.92 Byssothecium circinans
CBS 116668 Pleurophoma pleurospora
99
PD 82.371 Pleurophoma pleurospora
CBS 178.93 Paraconiothyrium flavescens comb. nov. ‘Phoma flavescens’
62
CBS 252.60 Medicopsis romeroi comb. nov. ‘Pyrenochaeta romeroi’
100
CBS 122784 Medicopsis romeroi comb. nov. ‘Pyrenochaeta romeroi’
91
83
Trematosphaeriaceae (C)
BCC 21118 Falcisormispora lignatilis
99
CBS 122368 Trematosphaeria pertusa
Lentitheciaceae?
CBS 221.37 Splanchnonema platani
CBS 419.85 Thyridaria rubronotata
CBS 350.82 Aposphaeria populina
99
98
PD 84.221 Aposphaeria populina
CBS 543.70 Aposphaeria populina
CBS 400.97 Melanomma pulvis-pyrius
86
CBS 371.75 Melanomma pulvis-pyrius
Melanommataceae (D)
CBS 200.31 Herpotrichia juniperi
92
64
99
PD 83.367 Aposphaeria corallinolutea sp. nov.
PD 83.831 Aposphaeria corallinolutea sp. nov.
CBS 283.53 Beverwykella pulmonaria
CBS 279.74 Pleomassaria siparia
73
99
99
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’
99
92
CBS 524.50 Sporormiella minima
CBS 659.74 Preussia funiculata
CBS 125434 Roussoella hysterioides
95
Didymosphaeriaceae
CBS 110022 Nigrograna mackinnonii comb. nov. ‘Pyrenochaeta mackinnonii’
CBS 674.75 Nigrograna mackinnonii comb. nov. ‘Pyrenochaeta mackinnonii’
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).
www.studiesinmycology.org
17
De Gruyter et al.
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.
≡ 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 &
Verkley, comb. nov. MycoBank MB564703.
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.
Type species: Heterospora chenopodii (Westend.) Gruyter,
Aveskamp & Verkley, see below (= Phoma heteromorphospora Aa
& Kesteren).
Note: For full synonymy, including the species listed below, see
Crane & Shearer (1991) and Boerema et al. (2004).
Heterospora chenopodii (Westend.) Gruyter, Aveskamp &
Verkley, comb. nov. MycoBank MB564702.
Leptosphaeria conoidea (De Not.) Sacc., Fungi Venet. Nov.
Vel. Crit. Ser. 2: 314. 1875.
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.
18
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.
Phoma sections Plenodomus, Pilosa
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).
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 &
Verkley, comb. nov. MycoBank MB564713.
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 &
Verkley, comb. nov. MycoBank MB564714.
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 &
Verkley, comb. nov. MycoBank MB564715.
Basionym: Phoma rubefaciens Togliani, Ann. Sper. Agr. II, 7: 1626.
1953.
Leptosphaeria errabunda (Desm.) Gruyter, Aveskamp &
Verkley, comb. nov. MycoBank MB564704.
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.
≡ 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.
Leptosphaeria sclerotioides (Sacc.) Gruyter, Aveskamp &
Verkley, comb. nov. MycoBank MB564716.
Basionym: Phoma errabunda Desm., Ann. Sci. Nat., Bot. Ser. 3,
11: 282. 1849.
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.
www.studiesinmycology.org
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
De Gruyter et al.
Leptosphaeria sydowii (Boerema, Kesteren & Loer.)
Gruyter, Aveskamp & Verkley, comb. nov. MycoBank
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 &
Verkley, comb. nov. MycoBank MB564718.
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 ParaLeptosphaeria, 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.
20
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
& Verkley, comb. nov. MycoBank MB564722.
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,
Aveskamp & Verkley, comb. nov. MycoBank MB564724.
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.
Phoma sections Plenodomus, Pilosa
Note: A Phoma synanamorph of Sclerotium orobanches was
reported by Yáňez-Morales et al. (1998) and described as Phoma
korfii (Boerema & Gruyter 1999).
Paraleptosphaeria praetermissa (P. Karst.) Gruyter,
Aveskamp & Verkley, comb. nov. MycoBank MB564725.
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,
Aveskamp & Verkley, comb. nov. MycoBank MB564727.
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 Tox0 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
www.studiesinmycology.org
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,
Aveskamp & Verkley, comb. nov. MycoBank MB564729.
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
& Verkley, comb. nov. MycoBank MB564730.
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 &
Verkley, comb. nov. MycoBank MB564731.
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 &
Verkley, comb. nov. MycoBank MB564753.
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
De Gruyter et al.
Plenodomus fallaciosus (Berl.) Gruyter, Aveskamp &
Verkley, comb. nov. MycoBank MB564732.
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 &
Verkley, comb. nov. MycoBank MB564754.
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.
Basionym: Cucurbitaria hendersoniae Fuckel, Symb. Myc. p. 172.
1870.
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.
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.
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 influorescens (Boerema & Loer.) Gruyter,
Aveskamp & Verkley, comb. nov. MycoBank MB564755.
Plenodomus lupini (Ellis & Everh.) Gruyter, Aveskamp &
Verkley, comb. nov. MycoBank MB564758.
≡ 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.
Basionym: Phoma enteroleuca var. influorescens Boerema & Loer.,
Trans. Brit. Mycol. Soc. 84: 290. 1985.
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 &
Verkley, comb. nov. MycoBank MB564756.
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 &
Verkley, comb. nov. MycoBank MB564757.
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.
22
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,
Aveskamp & Verkley, comb. nov. MycoBank MB564759.
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 &
Verkley, comb. nov. MycoBank MB564760.
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,
comb. nov. MycoBank MB564761.
Basionym: Plectophomella visci Moesz, Magyar Bot. Lapok 21: 13.
1922.
= Apocytospora visci Höhn., Mitt. Bot. Lab. TH Wien 1: 43. 1924.
Phoma sections Plenodomus, Pilosa
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.
MycoBank MB564769.
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.
= 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 &
Verkley, comb. nov. MycoBank MB564774.
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].
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.
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.
Type species: Subplenodomus violicola (P. Syd.) Gruyter,
Aveskamp & Verkley (see below)
Coniothyrium dolichi (Mohanty) Verkley & Gruyter, comb.
nov. MycoBank MB564776.
Subplenodomus apiicola (Kleb.) Gruyter, Aveskamp &
Verkley, comb. nov. MycoBank MB564770.
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,
Aveskamp & Verkley, comb. nov. MycoBank MB564771.
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 &
Verkley, comb. nov. MycoBank MB564772.
Basionym: Phoma valerianae Henn., Nyt Mag. Naturvidensk. 42:
29. 1904.
www.studiesinmycology.org
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,
comb. nov. MycoBank MB564777.
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
De Gruyter et al.
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.,
Trans. Brit. Mycol. Soc. 50: 260. 1967.
≡ 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.
nov. MycoBank MB564779.
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.
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.
Basionym: Leptosphaeria filamentosa Ellis & Everh., J. Mycol. 4:
76. 1888.
≡ 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.
Specimen examined: Mexico, from Yucca rostrata (Asparagaceae), Stevens, CBS
102202 = BPI 802755.
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.
Neophaeosphaeria filamentosa (Ellis & Everh.) Câmara,
M.E. Palm & A.W. Ramaley, Mycol. Res. 107: 519. 2003.
≡ Paraphaeosphaeria filamentosa (Ellis & Everh.) M.E. Barr, Mycotaxon
43: 392. 1992.
Pyrenochaetopsis pratorum (P.R. Johnst. & Boerema)
Gruyter, Aveskamp & Verkley, comb. nov. MycoBank
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.,
24
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.
Phoma sections Plenodomus, Pilosa
Pleospora fallens (Sacc.) Gruyter & Verkley, comb. nov.
MycoBank MB564782.
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,
comb. nov. MycoBank MB564783.
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.
MycoBank MB564784.
≡ 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,
comb. nov. MycoBank MB564785.
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.
www.studiesinmycology.org
≡ 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.) 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,
comb. nov. MycoBank MB564787.
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
De Gruyter et al.
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.
nov. MycoBank MB564789.
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
26
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,
comb. nov. MycoBank MB564790.
Basionym: Asteroma tiliae F. Rudolphi, Linnaea 4: 514. 1829.
Phoma sections Plenodomus, Pilosa
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.
www.studiesinmycology.org
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
De Gruyter et al.
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: 326. 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.
Type species: Medicopsis romeroi (Borelli) Gruyter, Verkley &
Crous (see below).
Medicopsis romeroi (Borelli) Gruyter, Verkley & Crous,
comb. nov. MycoBank MB564792.
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,
Nassauischen Vereins Naturk 23–24: 161. 1870.
Jahrb.
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).
Lentitheciaceae Yin. Zhang, C.L. Schoch, J. Fourn., Crous
& K.D. Hyde, Stud. Mycol. 64: 93. 2009.
Splanchnonema platani (Ces.) M.E. Barr, Mycotaxon 15:
364. 1982.
Specimen examined: uSA, Florida, from Paspalum notatum (Poaceae), Oct. 1937,
R.K. Voorhees, CBS 331.37.
Basionym: Sphaeria (Massaria) platani Ces., in Rabenhorst,
Klotzschii Herb. Viv. Mycol.: no. 1842. 1854.
Trematosphaeriaceae Suetrong et al. Cryptogamie Mycol.
32: 347. 2011.
Specimen examined: uSA, from Platanus occidentalis (Platanaceae), Jan. 1937,
C.L. Shear, CBS 221.37.
Falciformispora lignatilis K.D. Hyde, Mycol. Res. 96: 27.
1992.
Note: This taxon was shown by Zhang et al. (2012) to cluster basal
to the Lentitheciaceae.
Specimen examined: thailand, Pinruan Ban Bang, from Elaeis guineensis
(Arecaceae), BCC 21118.
Melanommataceae G. Winter, Rabenh. Krypt.-Fl., ed 1(2):
220 (1885) [as “Melanommeae”]
Medicopsis Gruyter, Verkley & Crous, gen. nov. MycoBank
MB564791.
Aposphaeria corallinolutea Gruyter, Aveskamp & Verkley,
sp. nov. MycoBank MB564798. Fig. 8.
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.
Etymology: The name refers to the coral coloured colony on OA,
and the luteous exudate diffusing into the agar medium.
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.
28
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
Phoma sections Plenodomus, Pilosa
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.
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 grey at centre, white
near margin. Pycnidia globose to subglobose, olivaceous to brick,
then olivaceous black, solitary or aggregated, 65–215 μm diam, nonsetose 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
www.studiesinmycology.org
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.
29
De Gruyter et al.
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.
Specimen examined: Switzerland, Andermatt, from Juniperus nana (Cupressaceae),
Nov. 1931, E. Gäumann, CBS 200.31.
Melanomma pulvis-pyrius (Pers.) Fuckel,
Nassauischen Vereins Naturk. 23–24: 160. 1870.
Jahrb.
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.
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.
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.
Westerdykella Stolk, Trans. Brit. Mycol. Soc. 38: 422. 1955.
Pleomassaria siparia (Berk. & Broome) Sacc., Syll. Fung.
2: 239. 1883.
Westerdykella capitulum (V.H. Pawar, P.N. Mathur &
Thirum) Gruyter, Aveskamp & Verkley, comb. nov. MycoBank
MB564801.
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.
30
Type species: Westerdykella ornata Stolk, see below.
Basionym: Phoma capitulum V.H. Pawar, P.N. Mathur & Thirum.,
Trans. Brit. Mycol. Soc. 50: 261. 1967.
≡ 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.
Phoma sections Plenodomus, Pilosa
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.
Westerdykella minutispora (P.N. Mathur ex Gruyter
& Noordel.) Gruyter, Aveskamp & Verkley, comb. nov.
MycoBank MB564793.
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,
comb. nov. MycoBank MB564795.
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.
www.studiesinmycology.org
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
31
De Gruyter et al.
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
Necrotrophic
Paraleptosphaeria
Ascospores 3–5-septate, hyaline to
yellow/brown
Mitosporic state rare, pycnidial cell wall directly scleroplectenchymatous, Necrotrophic
conidia aseptate
Plenodomus
Ascospores 3–7-septate, pale yellow
to brown
Mitosporic state common, pycnidial cell wall initially
Necrotrophic or plant pathogenic
pseudoparenchymatous, later scleroplectenchymatous, conidia aseptate
Subplenodomus
No known sexual state
Mitosporic state common, pycnidial cell wall mainly
pseudoparenchymatous, conidia aseptate
Necrotrophic or plant pathogenic
Heterospora
No known sexual state
Mitosporic state common, pycnidial cell wall pseudoparenchymatous,
conidia of two types: small aseptate and large septate
Plant pathogenic
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
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
32
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
Phoma sections Plenodomus, Pilosa
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.
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
www.studiesinmycology.org
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
33
De Gruyter et al.
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
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.
34
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.
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