Fungal Diversity
DOI 10.1007/s13225-015-0323-z
Towards a natural classification and backbone tree
for Pleosporaceae
Hiran A. Ariyawansa & Kasun M. Thambugala & Dimuthu S. Manamgoda &
Ruvishika Jayawardena & Erio Camporesi & Saranyaphat Boonmee &
Dhanushka N. Wanasinghe & R. Phookamsak & Singang Hongsanan &
Chonticha Singtripop & Ekachai Chukeatirote & Ji-Chuan Kang &
E. B. Gareth Jones & Kevin D. Hyde
Received: 2 January 2015 / Accepted: 8 February 2015
# School of Science 2015
Abstract The family Pleosporaceae includes numerous
saprobic, opportunistic human, and plant pathogenic taxa.
The classification of genera and species Pleosporaceae has
been a major challenge due to the lack of a clear understanding
of the importance of the morphological characters used to
distinguish taxa as well as the lack of reference strains. Recent
treatments concluded that Pleospora and some other genera in
Pleosporaceae are likely polyphyletic. In order to establish
the evolutionary relationships and to resolve the polyphyletic
nature of Pleospora and allied genera, we sequenced the 18S
nrDNA, 28S nrDNA, ITS, GAPDH, RPB2 and TEF1-alpha
gene regions of Pleosporaceae species and phylogenetically
analysed this data. Multigene phylogenies strongly support
the monophyletic nature of Pleosporaceae among the other
families in Pleosporales, and the acceptance of the genera
Alternaria, Bipolaris, Clathrospora, Comoclathris,
Curvularia, Dactuliophora, Decorospora, Diademosa,
Exserohilum, Extrawettsteinina, Gibbago,
Neocamarosporium, Paradendryphiella, Platysporoides,
Pleos pora , Poroce rc osp ora, P seud oyu con ia a nd
Pyrenophora. Austropleospora, Dendryphion, Edenia and
Macrospora are excluded from the family based on morphology coupled with molecular data. Two novel species,
Alternaria murispora in this paper and Comoclathris sedi
Electronic supplementary material The online version of this article
(doi:10.1007/s13225-015-0323-z) contains supplementary material,
which is available to authorized users.
H. A. Ariyawansa : K. D. Hyde (*)
Key Laboratory for Plant Diversity and Biogeography of East Asia,
Kunming Institute of Botany, Chinese Academy of Sciences,
Kunming 650201, People’s Republic of China
e-mail: kdhyde3@gmail.com
H. A. Ariyawansa : K. M. Thambugala : D. S. Manamgoda :
R. Jayawardena : E. Camporesi : S. Boonmee : D. N. Wanasinghe :
R. Phookamsak : S. Hongsanan : C. Singtripop : E. Chukeatirote :
K. D. Hyde
Institute of Excellence in Fungal Research, Mae Fah Luang
University, Chiang Rai 57100, Thailand
H. A. Ariyawansa : K. D. Hyde
World Agroforestry Centre, East Asia and Central,
Heilongtan, Kunming 650201, People’s Republic of China
H. A. Ariyawansa : J.<C. Kang
The Engineering and Research Center for Southwest
Bio-Pharmaceutical Resources of National Education Ministry of
China, Guizhou University, Guiyang 550025, Guizhou Province,
China
H. A. Ariyawansa : K. M. Thambugala : D. S. Manamgoda :
R. Jayawardena : E. Camporesi : S. Boonmee : D. N. Wanasinghe :
R. Phookamsak : S. Hongsanan : C. Singtripop : E. Chukeatirote :
K. D. Hyde
School of Science, Mae Fah Luang University, Chiang Rai 57100,
Thailand
E. Camporesi
A.M.B. Gruppo Micologico Forlivese BAntonio Cicognani^, Via
Roma 18, Forlì, Italy
E. B. G. Jones
Department of Botany and Microbiology, King Saudi University,
Riyadh, Saudi Arabia
Fungal Diversity
are introduced. The sexual morph of Alternaria alternata is
re-described and illustrated using modern concepts from fresh
collections. The paraphyletic nature of Pleospora is resolved
based on the available morpho-molecular data, but further
sampling with fresh collections, reference or ex-type strains
and molecular data are needed to obtain a natural classification
of genera and the family.
Keywords Backbone tree . Dothideomycetes . Pleosporales .
Plant pathogens . Saprobes
Introduction
Pleosporaceae is the largest family in the Pleosporales and is
representative of the order (Zhang et al. 2012; Hyde et al.
2013; Wijayawardene et al. 2014). The classification in the
Pleosporaceae has primarily been based on the Pleospora
type of centrum development (Dong et al. 1998) and asci that
are interspersed with pseudoparaphyses in the ascomata. These pseudoparaphyses originate above the hymenial layer and
grow downward among the asci to fuse at the base of the
locule (Wehmeyer 1975). Ascomata are perithecial, initially
immersed and become erumpent and are usually black and
sometimes hairy or setose. Asci are bitunicate and
fissitunicate, cylindrical, with an ocular chamber and the
hamathecium comprises cellular pseudoparaphyses are. Ascospores are usually brown and phragmosporous or
dictyosporous (Dong et al. 1998; Kirk et al. 2001). Species
are pathogenic or saprobic on wood and dead herbaceous
stems or leaves, as well as pathogenic in humans (Sivanesan
1984; Carter and Boudreaux 2004).
Historic overview of Pleosporaceae
The family Pleosporaceae was introduced by Nitschke (1869)
based on the immersed ascomata and presence of
pseudoparaphyses, and was classified in Sphaeriales. The
family was transferred to Pseudosphaeriaceae and later raised
to ordinal rank as the Pseudosphaeriales (Theissen and
Sydo w 191 8). Ple osp ora cea e, Ven turia cea e an d
Lophiostomataceae were assigned under Pleosporales by
Luttrell (1955), while treating Pseudosphaeriales as a synonym of Pleosporales. Luttrell (1973) included eight families
in Pleosporales including Pleosporaceae. Preliminary genera
added to this family were based on ascospore characteristics,
including shape, colour, septation, pigmentation and presence
or absence of mucilaginous sheaths (Luttrell 1955, 1973;
Wehmeyer 1961, 1975; Eriksson 1981; Sivanesan 1984; Barr
1987b; Abler 2003). Many of these characters were also found
in other families, such as Leptosphaeriaceae,
Melanommataceae, Phaeosphaeriaceae and Sporormiaceae.
This led for confusion in the intergeneric and familial classification (Luttrell 1955, 1973; Wehmeyer 1961, 1975; von Arx
and Müller 1975; Sivanesan 1984; Barr 1987a, b; Eriksson
and Hawksworth 1986, 1991). Barr (1987b) redefined the
Pleosporaceae to include Clathrospora (= Comoclathris),
Kirschsteiniothelia, Lewia and Pleospora and grouped
Cochliobolus, Pyrenophora and Setosphaeria into the family
Pyrenophoraceae. Berbee (1996) disagreed, suggesting that
all those genera belong to the Pleosporaceae. Kodsueb et al.
(2006) showed the intergeneric relationships and phylogenetic
perspectives of the family Pleosporaceae based on sequence
analyses of partial 28S rDNA and accepted 14 genera in
Pleosporaceae. Based on multi-gene phylogenetic analysis,
Zhang et al. (2009) concluded that some species from Lewia,
Cochliobolus, Pleospora, Pyrenophora and Setosphaeria resided in the Pleosporaceae. Lumbsch and Huhndorf (2010)
accepted 13 genera in Pleosporaceae but Zhang et al. (2012)
included only 10 genera by excluding Monascostroma,
Kriegeriella, and Zeuctomorpha (see notes below). Hyde
et al. (2013) included 23 genera in this family based on morphology coupled with molecular data. Multi-gene phylogenetic studies has shown that the familial placement of
Pleosporaceae with respect to other families in order
Pleosporales is valid (Lumbsch and Huhndorf 2010; Zhang
et al. 2012; Hyde et al. 2013). Major circumscription changes
of the genera in Pleosporaceae from 1961 to 2013 are given in
Table 1.
Asexual morphs of Pleosporaceae
The asexual morphs of Pleosporaceae can be coelomycetous
or hyphomycetous (Zhang et al. 2012; Hyde et al. 2013) and
some sexual genera in Pleosporaceae have been linked with
asexual morphs (Zhang et al. 2012; Hyde et al. 2013, 2014).
The type species of the family Pleospora is linked to
Stemphylium, which causes leaf disease (Sivanesan 1984).
Bipolaris was shown to be the asexual morph of
BCochliobolus^, and the cause of plant disease or infection
in human beings (Khan et al. 2000; Manamgoda et al.
2014). The nomenclatural conflict in this complex was resolved by giving priority to the more commonly used
established generic name Bipolaris (Manamgoda et al.
2012a, b; 2014). At the same time, Manamgoda et al.
(2012a, b) showed that Curvularia grouped with
BPseudocochliobolus^. The type species of Pleoseptum
(P. yuccaesedum A.W. Ramaley & M.E. Barr) has been linked
with Camarosporium yuccaesedum Fairm (Ramaley and Barr
1995) and Pyrenophora has the asexual morph Drechslera
(Farr et al. 1989). Ariyawansa et al. (2014c) proposed to
Major circumscription changes of the genera in Pleosporaceae
Wehmeyer
1961
Wehmeyer 1975
Eriksson and
Kirk et al. 2001
Hawksworth 1998
Kodsueb et al.
2006
Lumbsch and
Huhndorf 2010
Zhang et al.
2012
Hyde et al.
2013
Wijayawardene
et al. 2014
This study
Clathrospora
Platyspora
Pleospora
Pyrenophora
Allonecte
Catharinia
Clathrospora
Cochliobolus
Cucurbitaria
Didymopleella
Eudarluca
Fenestella
Gibberidea
Gilletiella
Herpotrichia
Herpotrichiella
Letendraea
Licopolia
Melanomma
Cochliobolus
Extrawettsteinina
Falciformispora
Kirschsteiniothelia
Kriegeriella
Leptosphaerulina
Lewia
Macrospora
?Macroventuria
Monascostroma
Platysporoides
Pleospora
Pseudoyuconia
Pyrenophora
Setosphaeria
Cochliobolus
Extrawettsteinina
Falciformispora
Kirschsteiniothelia
Kriegeriella
Leptosphaerulina
Lewia
Macrospora
Macroventuria
Platysporoides
Pleospora
Pseudoyuconia
Pyrenophora
Setosphaeria
Teratosphaeria
Cochliobolus
Decorospora
Extrawettsteinina
Falciformispora
Kriegeriella
Lewia
Macrospora
Monacostroma
Platysporoides
Pleospora
Pseudoyuconia
Pyrenophora
Setosphaeria
Zeuctomorpha
Cochliobolus
Crivellia
Decorospora
Extrawettsteinina
Kriegeriella
Lewia
Macrospora
Platysporoides
Pleospora
Pseudoyuconia
Pyrenophora
Setosphaeria
Zeuctomorpha
Cochliobolus
Crivellia
Extrawettsteinina
Lewia
Macrospora
Platysporoides
Pleospora
Pseudoyuconia
Pyrenophora
Setosphaeria
Alternaria
Alternariaster
Bipolaris
Brachycladium
Chalastospora
Clathrospora
Curvularia
Decorospora
Dendryphion
Edenia
Embellisia
Exserohilum
Extrawettsteinina
Macrospora
Marielliottia
Alternaria
Bipolaris
Clathrospora
Curvularia
Decorospora
Dendryphion
Edenia
Exserohilum
Extrawettsteinina
Marielliottia
Neocamarosporium
Paradendryphiella
Pleoseptum
Platysporoides
Pseudoyuconia
Melanopsamma
Metasphaeria
Mycomicrothelia
Ophiobolus
Otthia
Paralimyces
Platyspora
Wettsteinina
Zeuctomorpha
Tremateia
Wettsteinina
Zeuctomorpha
Platysporoides
Pleospora
Pseudoyuconia
Pyrenophora
Setosphaeria
Sinomyces
Teretispora
Pyrenophora
Stemphylium
Alternaria
Bipolaris
Clathrospora
Comoclathris
Curvularia
Dactuliophora
Decorospora
Diademosa
Exserohilum
Extrawettsteinina
Gibbago
Neocamarosporium
Paradendryphiella
Platysporoides
Pleospora/
Stemphylium
Porocercospora
Pseudoyuconia
Pyrenophora
Pleomassaria
Pleospora
Pteridiospora
Pyrenophora
Teichospora
Teratosphaeria
Thyridaria
Thyridium
Tomasellia
Trematosphaeria
Trichometasphaeria
Zignoella
Ulocladium
Fungal Diversity
Table 1
Fungal Diversity
conserve Pyrenophora over Drechslera by giving priority to
the oldest name. Based on the combined gene analysis of
GAPDH, RPB2 and TEF1, Woudenberg et al. (2013)
synonymised Allewia, Brachycladium, Chalastospora,
Chmelia, Crivellia, Embellisia, Lewia, Nimbya, Sinomyces,
Teretispora, Ulocladium, Undifilum and Ybotromyces under
Alternaria. In same study, Woudenberg et al. (2013) treated
the 24 internal clades in the Alternaria complex as sections,
which is a continuation of a recent proposal for the taxonomic
treatment of lineages in Alternaria. Furthermore,
Alternariaster, a genus formerly seen as part of Alternaria
was transferred to Leptosphaeriaceae based on molecular data. Recently, new asexual genera were introduced by different
researches based on both morphology and phylogeny. i.e.
Porocercospora was introduced as a new genus in
Pleosporaceae by Amaradasa et al. (2014) to accommodate
the buffalo grass false-smut pathogen, while Johnalcornia
was introduced to accommodate Bipolaris aberrans, which
clusters sister to the newly described Porocercospora (Tan
et al. 2014). Paradendryphiella was introduced by
Woudenberg et al. (2013) to accommodate two marine
Dendryphiella species (D. arenariae Nicot, D. salina (G.K.
Sutherl.) Pugh & Nicot) that did not group with the type species D. vinosa (Berk. & M.A. Curtis) Reisinger (Jones et al.
2008; Woudenberg et al. 2013).
Taxa in Alternaria, Bipolaris, Stemphylium and phoma like
species are more common asexual morphs in Pleosporaceae
and can be saprobic or parasitic on various hosts. Phoma betae
A.B... Frank is a notorious pathogen of sugar beet, which
causes zonate leaf spot. Alternaria porri (Ellis) Cif.,
Stemphylium solani G.F. Weber, S. botryosum Wallr and
S. vesicarium (Wallr.) E.G. Simmons can cause leaf blight of
garlic (Zhang et al. 2009). Phoma incompta Sacc & Martelli is
a pathogen on olive, and S. botryosum, the asexual morph of
Pleospora herbarum (Pers.) Rabenh, causes leaf disease of
olive trees (Malathrakis 1979). Some Curvularia species have
been reported as human pathogens, causing respiratory tract,
cutaneous, and corneal infections (Carter and Boudreaux
2004).
We have been studying the families of Dothideomycetes
based on morphology and molecular phylogeny, in order to
provide a natural classification of this large class. Hyde et al.
(2013) provide an account of all 105 families of
Dothideomycetes, while Wijayawardene et al. (2014) provided an outline of Dothideomycetes and suggestions on the on
the names to use where asexual and sexual genera were
linked. The largest order in Dothideomycetes is Pleosporales
and we also have provided a natural classification of this large
order (Ariyawansa et al. 2013a, b, c; Ariyawansa et al. 2014a,
b, c, d, e, f; Zhang et al. 2012). Zhang et al. (2012) provided an
account for 105 genera in Pleosporales, however, at that time
little verified molecular data were available. The aim of the
present study was to delineate the phylogenetic lineages
within Pleosporaceae, and to build a robust taxonomy to act
as a backbone tree for the family. Phylogenetic analysis were
conducted using sequence data from parts of the 18S nrDNA
(SSU), 28S nrDNA (LSU), the internal transcribed spacer
regions 1 and 2 and intervening 5.8S nrDNA (ITS) and
RNA polymerase second largest subunit (RPB2) gene regions
of extype, reference and putative strains of all available allied
genera in Pleosporaceae. We also focused on the phylogenetic lineages within Alternaria and allied genera, and to create a
stable taxonomy. Phylogenetic inferences were conducted on
sequence data of parts of the SSU, LSU, ITS, glyceraldehyde3-phosphate dehydrogenase (GAPDH), RPB2 and translation
elongation factor 1-alpha (TEF1) gene regions of extype and
reference strains of Alternaria species and all available allied
genera.
Material and methods
Specimen examination
Fresh material of species of Pleosporaceae were collected in
Thailand, Germany and Italy during 2011–2014. Specimens
were taken to the laboratory in Ziplock plastic bags. The samples were processed and examined following the method described in Ariyawansa et al. (2013a, b). Fresh and herbarium
material were examined under a Motic SMZ 168 dissecting
microscope to locate and isolate ascomata fruiting bodies.
Hand sections of the fruiting structures were mounted in water
for microscopic studies and photomicrography. The taxa were
examined using a Nikon ECLIPSE 80i compound microscope
and photographed with a Canon 450D digital camera fitted to
the microscope. Measurements were made with the Tarosoft
(R) Image Frame Work program and images used for figures
processed with Adobe Photoshop CS3 Extended version 10.0
software (Adobe Systems, USA). Isolations were made from
single ascospores, following a modified method of
Ariyawansa et al. (2013b) and Chomnunti et al. (2014). Contents of the sectioned fruiting body were transferred to a drop
of sterile water on a flame-sterilized slide. Drops of the spore
suspension were pipetted and spread on a Petri-dish containing 2 % water agar (WA) and incubated at 25 °C. Germinated
ascospores were transferred singly to MEA media (Alves et al.
2006).
Herbarium specimens were obtained on loan from the
Swedish Museum of Natural History (S) and the New York
Botanical Garden (NY). Voucher specimens are deposited in
the herbarium of Mae Fah Luang University (MFLU), Chiang
Rai, Thailand, Kunming Institute of Botany (KIB) and New
Zealand Fungal Herbarium-Landcare Research (PDD), New
Zealand. Living cultures are deposited at the Mae Fah Luang
University Culture Collection (MFLUCC), International collection of microorganisms from plants (ICMP) and
Fungal Diversity
Queensland Plant Pathology Herbarium (BRIP), the latter under Material Transfer Agreement No. 4/2010 (MTA). Each
genus is listed along with a description of the type species,
except in cases where there is only a single species in the
genus.
DNA extraction, PCR amplification and sequencing
Single ascospore fungal isolates were grown on MEA or PDA
for 28 days at 25 °C in the dark. Genomic DNA was extracted
from the growing mycelium using the Biospin Fungus Genomic DNA Extraction Kit (BioFlux®) following the manufacturer’s protocol (Hangzhou, P.R. China). Otherwise DNA extracted directly from ascomata using a DNA extraction kit
(E.Z.N.A.® Forensic DNA kit, D3591- 01,Omega Bio-Tek)
following Telle and Thines (2008).
The amplification procedure was performed in a 50 μl reaction volume containing 5–10 ng DNA, 0.8 units Taq polymerase, 1X PCR buffer, 0.2 mM d’NTP, 0.3 μM of each
primer with 1.5 mM MgCl2 (Cai et al. 2009). The PCR reactions for amplification of the recently ratified universal fungal
barcode ITS1-5.8S-ITS2 of the nuclear ribosomal DNA operon (Schoch et al. 2009), were performed under standard conditions (White et al. 1990; Stielow et al. 2010). PCR conditions for amplifying the partial SSU and LSU r-DNA followed
the protocol of Phillips et al. (2008). Amplification of GAPD
H, RPB2 and TEF1 followed the protocol of Woudenberg
et al. (2013). The PCR products were observed on 1 % agarose
electrophoresis gels stained with ethidium bromide. Purification and sequencing of PCR products were carried at Shanghai
Sangon Biological Engineering Technology and Services Co.,
(China).
DNA sequence data was obtained from the internal transcribe spacer (ITS), small and large subunits of the nuclear
ribosomal RNA genes (SSU, LSU) and the protein coding
gene, GAPDH (TUB). Primer sets used for these genes were
as follows: ITS: ITS5/ITS4 SSU: NS1/NS4; LSU: LR0R/
LR5; RPB2–5F2/fRPB2–7cR (Liu et al. 1999; Sung et al.
2007). The GAPDH region with gpd1 and gpd2 (Berbee
et al. 1999) and the TEF1 gene with the primers EF1-728F
and EF1-986R (Carbone and Kohn 1999). Primer sequences
are available at the WASABI database at the AFTOL website
(aftol.org). Sequences are deposited at NCBI GenBank under
the accession numbers provided in Supplementary Table 1.
Alignments are deposited in TreeBASE.
Sequence alignment and phylogenetic analysis
Multiple sequence alignments were generated with MAFFT v.
6.864b (http://mafft.cbrc.jp/alignment/server/index.html).
The alignments were checked visually and improved
manually where necessary. Two different datasets were used
to estimate two phylogenies; a Pleosporineae family tree and
an Alternaria phylogeny. The first tree focuses on
phylogenetic placement of Pleosporaceae in sub order
Pleosporineae, the second one was generated to show the
placement of newly reported Alternaria species and their
sexual state. All introns and exons were aligned separately.
Regions containing many leading or trailing gaps were
removed from the ITS, SSU, LSU, GAPDH, TEF1 and
RPB2 alignments prior to tree building. The alignments
were checked visually and improved manually where
necessary. All sequences obtained from GenBank and used
by Ariyawansa et al. (2014a), Boonmee et al. (2014), Hyde
et al. (2013), Phookamsak et al. (2014) Schoch et al. (2009),
Suetrong et al. (2009), Verkley et al. (2014), and Zhang et al.
(2012) are listed in supplementary Table 1.
Maximum likelihood analyses including 1000 bootstrap
replicates were run using RAxML v. 7.2.6 (Stamatakis 2006;
Stamatakis et al. 2008). The online tool Findmodel (http://
www.hiv.lanl.gov/content/sequence/findmodel/findmodel.
html) was used to determine the best nucleotide substitution
model for each partition. The best scoring tree was selected
with a final likelihood value of −19492.551787. The resulting
replicates were plotted on to the best scoring tree obtained
previously. Maximum Likelihood bootstrap values (ML)
equal or greater than 50 % are given below or above each
node in black (Fig. 1).
The model of evolution was performed by using
MrModeltest 2.2 (Nylander 2004). Posterior probabilities
(PP) (Rannala and Yang 1996; Zhaxybayeva and Gogarten
2002) were determined by Markov Chain Monte Carlo sampling (MCMC) in MrBayes v. 3.0b4 (Huelsenbeck and
Ronquist 2001). Six simultaneous Markov chains were run
for 5,000,000 generations and trees were sampled every
100th generation and 50,000 trees were obtained. The first
10,000 trees, representing the burn-in phase of the analyses,
were discarded while the remaining trees were used for calculating posterior probabilities in the majority rule consensus
tree (critical value for the topological convergence diagnostic
set to 0.01) (Crous et al. 2006). Bayesian Posterior Probabilities (BYPP) equal or greater than 0.90 is given below or
above each node (Fig. 1).
In order to determine the species limits in Pleospora
herbarum species complex, Alternaria complex and the newly introduced Comoclathris sedi, we applied the criteria of
Genealogical Concordance Phylogenetic Species Recognition
(GCPSR) (Taylor et al. 2000; Dettman et al. 2003). Dettman
et al. (2003) emphasised that species should be recognised if
they satisfy one of two criteria: genealogical concordance or
genealogical non-discordance. Clades were genealogically
concordant if they were present in at least some of the gene
trees and genealogically non-discordant if they were strongly
supported (MP≥70 %; ML≥70 %) in a single gene and not
contradicted at or above this level of support in any other
single gene tree. This criterion prohibited poorly supported
Fungal Diversity
Fungal Diversity
RAxML tree based on a combined dataset of ITS, SSU, LSU and
RPB2 of 98 strains representing the Pleosporineae. Bootstrap support
values for maximum likelihood greater than 50 % (black) and bayesian
posterior probabilities greater than 0.90 (green) below and above the
nodes. Halojulella avicenniae is the out group taxon. The original
isolate numbers are noted after the species names. Ex-type culture
numbers are in bold. Newly generated strains in this study are indicated
in red. The type species of each genus is indicated in blue
Fig. 1
non-monophyly at one locus from undermining wellsupported monophyly at another locus. Phylogenetic trees
and data files were viewed in MEGA v. 5 (Tamura et al.
2011), TreeView v. 1.6.6 (Page 1996) and FigTree v. 1.4
(Rambaut and Drummond 2008).
Results and discussion
Phylogeny
Phylogeny of Pleosporineae
The final Pleosporineae alignment included 98 strains,
representing seven families, and consisted of 3170 characters
(SSU 941, LSU 863, ITS 493, RPB2 858). In the SSU alignment a large insertion at position 446 in the isolates
Chaetosphaeronema hispidulum (Corda) Moesz (CBS
216.75), Phoma fallens Sacc (Pleospora fallens) (CBS
161.78) and Ophiosphaerella herpotricha (Fr.) J. Walker
(CBS 620.86) was excluded from the phylogenetic analyses.
Four different alignments corresponding to each individual
gene, and a combined alignment of the four genes were
analysed. Comparison of the alignment properties and nucleotide substitution models are provided in Table 2. Results of
the partition-homogeneity test (P=0.107) indicate that the
ITS, LSU, SSU and RPB2 gene trees reflect the same underlying phylogeny. Therefore, these datasets were combined and
analyzed by using several tree-building programs, the
resulting trees compared and the best tree is presented in
Fig. 1. New sequences are deposited in GenBank (Table 1).
The combined ITS, LSU, SSU and RPB2 gene dataset
from seven families in the Pleosporales sub order
Pleosporaneae is shown in Fig 1. All trees (ML and BYPP)
were similar in topology and not significantly different (data
not shown). A best scoring RAxML tree is shown in Fig. 1,
with the value of −19492.551787. Phylogenetic trees obtained
Table 2
from Maximum Likelihood and Bayesian analysis yielded
trees with similar overall topology at subclass and family relationships in agreement with previous work based on Maximum Likelihood analysis (Schoch et al. 2009; Suetrong et al.
2009; Zhang et al. 2012; Ariyawansa et al. 2014a; Boonmee
et al. 2014; Hyde et al. 2013, Phookamsak et al. 2014, Verkley
et al. 2014; Wijayawardene et al. 2014). The support values
for the different phylogenetic methods vary, with the Bayesian
posterior probabilities being higher than the RAxML bootstrap support values.
In the multi-locus phylogeny inferred from the combined
dataset shown in Fig. 1, several well-supported sub-clades can
be identified in the family Pleosporaceae, which are
interpreted as appropriate for the delimitation of genera, i.e.
Alternaria, Bipolaris, Comoclathris, Curvularia,
Decorospora, Dendryphion, Exserohilum, Gibbago,
Johnalcornia, Neocamarosporium, Porocercospora,
Pleospora and Pyrenophora.
The Pleospora sensu stricto clade comprises of eleven
strains along with the type strain of Pleospora herbarum/
Stemphylium herbarum (CBS 191.86) including our strains
(MFLUCC 14–0261, MFLUCC 13–0344 and MFLUCC
13–0266). In order to determine the species limits of
Pleospora herbarum, a combine gene tree (Fig. 1.) was used.
Nodes that were supported (≥70 %) in the combined gene
phylogeny were recognised as taxa within Pleospora
herbarum sensu stricto. Thus the putative strains of
P. tomatonis E.G. Simmons (CBS 109844), P. sedicola E.G.
Simmons (CBS 109843), which clustered within Pleospora
herbarum sensu stricto are treated as strains of Pleospora
herbarum in this study. Further work may yet prove this is a
species complex.
A recently introduced genus, Paradendryphiella forms a
basal clade to the Pleospora sensu stricto clade including
the type strain of Paradendryphiella salina (CBS 302.84)
and Dendryphiella salina (CBS 142.60). Another poorlysupported clade forming the major part of the ingroup of the
tree comprises the putative strains of Gibbago trianthemae
E.G. Simmons (NFCCI 1886 and GT-VM). The Bipolaris
clade comprises four strains with B. oryzae (MFLUCC 10–
0714), B. cynodontis (Marignoni) Shoemaker (ICMP 6128),
B. melinidis Alcorn (BRIP 12898) and B. maydis (Y. Nisik. &
C. Miyake) Shoemaker (CBS 134.39, previously known as
Cochliobolus heterostrophus). Two monophyletic genera
Porocercospora seminalis (Ellis & Everh.) Amaradasa et al.,
Comparison of alignment properties of genes and nucleotide substitution models used in Pleosporineae phylogenetic analysis
Genes /loci
LSU
SSU
ITS
RPB2
Alignment strategy (MAFFT v6)
Nucleotide substitution models for Bayesian analysis
(determined by MrModeltest)
FFT-NS-Il
SYM+I+G
FFT-NS-I
GTR+I+G
FFT-NS-I+manual
GTR+I+G
FFT-NS-I+manual
GTR+I+G
Fungal Diversity
(CPC 21349 and CPC 21332) and Johnalcornia aberrans
(Alcorn) Y.P. Tan & R.G. Shivas (BRIP 16281) form wellsupported clades sister to the Curvularia, Exserohilum and
Bipolaris clades. Curvularia forms a relatively wellsupported clade within the family Pleosporaceae. This clade
contains three strains with C. lunata (Wakker) Boedijn (CBS
730.96), the type species of Curvularia, and the highly supported clades of the following species: C. ravenelii (M.A.
Curtis ex Berk.) Manamgoda et al. (BRIP 13165) and
C. heteropogonis Alcorn (CBS 284.91). Exserohilum forms
a well-supported clade sister to Curvularia containing the putative strains of Setosphaeria monoceras Alcorn (CBS
154.26), Exserohilum sp. (C950801) and Exserohilum sp.
(NK93).
The recently redefined genus Alternaria forms a wellsupported clade sister to Pyrenophora containing 18 strains
with A. alternata (Fr.) Keissl (CBS 916.96) the type species of
Alternaria, along with the three sexual strains of A. alternata
(MFLUCC 14–1184, MFLUCC 14–1185 and ICMP). Apart
from A. alternata strains, the clade consist of A. arborescens
E.G. Simmons (CBS 102605), A. solani (Ellis & G. Martin)
L.R. Jones & Grout (CBS 116651), A. macrospora Zimm.,
(CBS 117228), A. nobilis (Vize) E.G. Simmons (CBS
116490), Lewia infectoria (Fuckel) M.E. Barr & E.G.
Simmons (CBS 210.86), Lewia ethzedia E.G. Simmons
(CBS 197.86), Chalastospora gossypii (Jacz.) U. Braun &
Crous (CPC 15567), Chalastospora obclavata Crous & U.
Braun (CBS 124120), Embellisia abundans E.G. Simmons
(CBS 534.83), A. anigozanthi R.D. Raabe (CBS 121920),
Lewia eureka E.G. Simmons (DAOM 195275) and Crivellia
papaveracea (De Not.) Shoemaker & Inderb (CBS 116607).
Furthermore the type strain of Xenobotryosphaeria,
X. calamagrostidis Quaedvl et al., (CBS 303.71) also clustered within the Alternaria clade.
The Pyrenophora clade comprises five strains with
P. phaeocomes (Rebent.) Fr., (DAOM 222769), the type species of Pyrenophora, P. dictyoides A.R. Paul & Parbery
(DAOM 63666 and DAOM 75616), Drechslera dematioidea
( B u b á k & Wr ó b l . ) S c h a r i f ( C B S 1 0 8 9 6 3 ) a n d
P. chaetomioides Speg., (DAOM 208989). The type species
of Comoclathris, C. lanata Clem, was not available for study,
but the five verified strains of Comoclathris species form a
well-supported clade within the family Pleosporaceae. i. e. C.
compressa (Harkn.) Shoemaker & C.E. Babc (CBS 156.53
and CBS 157.53) and the novel species C. sedi (MFLUCC
13–0817, MFLUCC 13–0754, MFLUCC 13–0763 and
MFLUCC 13–0607) along with a putative strain of Pleospora
ambigua (Berl. & Bres.) Wehm (CBS 366.52) cluster in a
well-supported clade within the Pleosporaceae outside
Alternaria s. str. A putative strain of Pleospora ambigua
(CBS 366.52) also clustered with our new species
Comoclathris sedi and is thus treated as an additional strain
of C. sedi. Furthermore, Comoclathris incompta (Pleospora
incompta (Sacc. & Martelli) Gruyter & Verkley) (CBS
467.76) and Comoclathris typhicola (Pleospora typhicola
(Cooke) Sacc. 1875) (CBS 132.69) also forms two distinct
clades within the Comoclathris clade.
Another well supported clade (Fig. 1) was formed by
Decorospora, D. gaudefroyi (Pat.) Inderb et al., (pp4723)
the type species, along with two putative strains of Pleospora
sp. ATCC MYA-3203 and ATCC MYA-3202 basal to the
Comoclathris clade. The Neocamarosporium clade comprises
seven strains with Neocamarosporium goegapense Crous &
M. J. Wing f. ( C PC 2 367 6 T ) , t h e t y p e s p e c i e s of
Neocamarosporium: N. betae (CBS 109410, IMI 156653,
ICMP 10945) and N. calvescens (CBS 344.78, CBS 246.79
and CBS 432.77). The type species of Clathrospora,
C. elynae, forms a well-supported clade, located basal to the
Pleosporaceae (Fig. 1), outside the Alternaria complex.
The genus Edenia, with E. gomezpompae M.C. González
et al., as the type (C1cT, CBS 124106, JLCC34533 and
11G048) clusters outside Pleosporaceae and formed a distinct
clade in Phaeosphaeriaceae. During preliminary analysis (data not shown), the type strains of Austropleospora osteospermi
R.G. Shivas & L. Morin (BRIP 51628) and Dendryphion
europaeum Crous & R.K. Schumach (CPC 23231 and CPC
22943) clustered outside the suborder Pleosporineae, thus are
excluded from the final analysis (data not shown).
Pleospora fallens (Sacc.) Gruyter & Verkley (CBS 161.78)
treated here as Phoma fallens Sacc., forms a basal clade in
family Pleosporaceae. Confusion surrounding this species is
discussed in the latter part of this study.
Phylogeny of Alternaria
In order to define the phylogeny and taxonomy of Alternaria
and allied genera, 114 strains were included in the Alternaria
complex alignment. For the Alternaria tree, the gene boundaries were: 1–533 bp for ITS, 539–1080 bp for GPDH, 1087–
1941 bp for LSU, 1947–2811 bp for RPB2 and 2817–3080 bp
for TEF. All phylogenies, different phylogenetic methods and
gene regions or gene combinations used on this dataset (data
not shown, trees and alignments deposited in TreeBASE),
show a weak support at the deeper nodes of the tree. The only
well-supported node (Bayesian posterior probability of 1.0,
RAxML Maximum Likelihood support value of 99) in all
phylogenies separates Pyrenophora and the Pleospora/
Stemphylium clade from the Alternaria complex (Fig. 1).
The overall topology of the concatenated genes analysis of
ITS, GPDH, LSU, RPB2 and TEF was in agreement to the
phylogenetic trees obtained from Maximum Likelihood and
Bayesian analysis yielded trees with previous work based on
Maximum Likelihood and Bayesian analysis of Woudenberg
et al. (2013) and Lawrence et al. (2013).
In the Alternaria clade, six monotypic lineages and 24
internal clades occur consistently in the individual and
Fungal Diversity
combined phylogenies, although positions vary between the
different gene regions or combinations used and are similar
with Woudenberg et al. (2013) and Lawrence et al. (2013).
The support values for the different phylogenetic methods
vary, with the Bayesian posterior probabilities being higher
than the RAxML bootstrap support values (Fig. 2). The recently introduced genus, Xenobotryosphaeria,
X. calamagrostidis (CBS 303.71) clustered within the
Alternaria complex in sect. Infectoriae, thus giving rise to
confusion (further discussed in the Taxonomy section of
Alternaria).
Seven alignments were analysed corresponding to single
gene analyses of ITS, SSU, LSU, GAPDH, TEF1 and RPB2
and combined alignments of the both Alternaria and
Pleosporineae. Comparison of the alignment properties and
nucleotide substitution models are provided in Tables 2 and 3.
=Chalastospora E.G. Simmons, CBS Diversity Ser.
(Utrecht) 6: 668 (2007).
=Chmelia Svob.-Pol., Biológia, Bratislava 21: 82 (1966).
=Crivellia Shoemaker & Inderb., in Inderbitzin, Shoemaker, O’Neill, Turgeon & Berbee, Can. J. Bot. 84(8): 1308
(2006).
=Embellisia E.G. Simmons, Mycologia 63(2): 380 (1971).
=Nimbya E.G. Simmons, Sydowia 41: 316 (1989).
=Sinomyces Yong Wang bis & X.G. Zhang, Fungal Biology 115(2): 192 (2009).
=Teretispora E.G. Simmons, CBS Diversity Ser. (Utrecht)
6: 674 (2007).
=Ulocladium Preuss, Linnaea 24: 111 (1851).
=Undifilum B.M. Pryor, Creamer, Shoemaker, McLainRomero & Hambl., Botany 87(2): 190 (2009).
=Ybotromyces Rulamort, Bull. Soc. bot. Centre-Ouest,
Nouv. sér. 17: 192 (1986).
Taxonomy
Type species: Alternaria alternata (Fr.) Keissl., Beih. bot.
Zbl., Abt. 2 29: 434 (1912).
Pleosporaceae Nitschke, Verh. naturh. Ver. preuss. Rheinl.
26: 74 (1869).
Facesoffungi number: FoF 00500
Pathogenic or saprobic on wood and dead herbaceous stems
or leaves or pathogen of humans. Sexual morph: Ascomata
perithecial, initially immersed and becoming erumpent to
nearly superficial, black, globose, subglobose or ovoid, sometimes hairy or setose, ostiolate. Ostiole papillate or apapillate,
sometimes with a pore-like ostiole, ostiolar canal filled with or
lacking periphyses. Peridium relatively thin, usually thick at
the sides, thinner at the base. Hamathecium of hyaline, septate, cellular pseudoparaphyses interspersed with asci. Asci 8spored, bitunicate, fissitunicate, cylindrical, with or without a
pedicel, with an ocular chamber. Ascospores uniseriate or
biseriate, partially overlapping, phragmosporous or muriform,
brown or pale brown, with or without mucilaginous sheath.
Asexual morph: coelomycetous or hyphomycetous, and the
conidiogenous cells can be phialidic, annellidic or sympodial
blastic.
Type: Pleospora Rabenh. ex Ces. & De Not., Comm. Soc.
crittog. Ital. 1(4): 217 (1863)
Bipolaris Shoemaker, Can. J. Bot. 37(5): 882 (1959).
= Cochliobolus Drechsler, Phytopathology 24: 973 (1934).
Type species: Bipolaris maydis (Y. Nisik. & C. Miyake) Shoemaker 1959.
Clathrospora Rabenh., Hedwigia 1(18): 116 (1857).
Type species: Clathrospora elynae Rabenh., Hedwigia 1: 116
(1857).
Comoclathris Clem., Gen. fung. (Minneapolis): 37, 173
(1909).
Type species: Comoclathris lanata Clem. [as ‘Comochlatris’],
Gen. fung. (Minneapolis): 1–227 (1909).
Curvularia Boedijn, Bull. Jard. bot. Buitenz, 3 Sér. 13(1): 123
(1933).
= Pseudocochliobolus Tsuda, Ueyama & Nishih.,
Mycologia 69(6): 1117 (1978) [1977].
Type species: Curvularia lunata (Wakker) Boedijn, Bull.
Jard. bot. Buitenz, 3 Sér. 13(1): 127 (1933).
Dactuliophora C.L. Leakey, Trans. Br. mycol. Soc. 47(3):
341 (1964).
Type species: Dactuliophora tarrii C.L. Leakey, Trans. Br.
mycol. Soc. 47(3): 343 (1964).
Genera accepted in Pleosporaceae
Alternaria Nees, Syst. Pilze (Würzburg): 72 (1816) [1816–17].
= Lewia M.E. Barr & E.G. Simmons, in Simmons,
Mycotaxon 25(1): 289 (1986).
= Allewia E.G. Simmons, Mycotaxon 38: 260 (1990).
=Brachycladium Corda, Icon. fung. (Prague) 2: 14 (1838).
Decorospora Inderb. et al., in Inderbitzin et al., Mycol. Progr.
1(4): 657 (2002).
Type species: Type species: Decorospora gaudefroyi (Pat.)
Inderb. et al., in Inderbitzin et al., Mycol. Progr. 94(4): 657 (2002).
≡ Pleospora gaudefroyi Pat., Tab. analyt. Fung. (Paris)(7):
40 (no. 602) (1886).
Fungal Diversity
Fungal Diversity
RAxML tree based on a combined dataset of ITS, LSU GAPDH,
RPB2 and TEF1 of 114 strains representing the Alternaria-complex.
Bootstrap support values for maximum likelihood greater than 50 %
and Bayesian posterior probabilities greater than 0.90 (green) are
indicated below or above the nodes. Pleospora herbarum is the out
group taxon. The original isolate numbers are noted after the species
names. Newly generated strains in this study are indicated in red
Fig. 2
Diademosa Shoemaker & C.E. Babc., Can. J. Bot. 70(8):
1641 (1992).
Type species: Diademosa californiana (M.E. Barr) Shoemaker & C.E. Babc. [as ‘californianum’], Can. J. Bot. 70(8): 1641
(1992).
Basionym: Graphyllium californianum M.E. Barr, Mem.
N. Y. bot. Gdn 62: 40 (1990).
Exserohilum K.J. Leonard & Suggs, Mycologia 66(2): 289
(1974).
= Setosphaeria K.J. Leonard & Suggs, Mycologia 66(2):
294 (1974).
= Luttrellia Khokhr. & Gornostaĭ, in Gornostaĭ in
Azbukina et al. (Eds), Vodorosli, Griby i Mkhi Dal’nego
Vostoka [Algae, Fungi and Mosses of the Soviet Far-East]
(Vladivostok): 80 (1978).
Type species: Exserohilum turcicum (Pass.) K.J. Leonard &
Suggs, Mycologia 66(2): 291 (1974).
Extrawettsteinina M.E. Barr, Contr. Univ. Mich. Herb. 9(8):
538 (1972).
Type species: Extrawettsteinina minuta M.E. Barr, Contr.
Univ. Mich. Herb. 9(8): 538 (1972).
Gibbago E.G. Simmons, Mycotaxon 27: 108 (1986).
Type species: Gibbago trianthemae E.G. Simmons,
Mycotaxon 27: 108 (1986).
Neocamarosporium Crous & M.J. Wingf., Persoonia, Mol.
Phyl. Evol. Fungi 32: 273 (2014).
Type species: Neocamarosporium goegapense Crous &
M.J. Wingf., Persoonia, Mol. Phyl. Evol. Fungi 32:
273 (2014).
Paradendryphiella Woudenberg & Crous, Stud. Mycol.
75(1): 207 (2013).
Type species: Paradendryphiella salina (G.K. Sutherl.)
Woudenberg & Crous, Stud. Mycol. 75(1): 207 (2013).
Table 3
Basionym: Cercospora salina G.K. Sutherl., New Phytol.
15: 43 (1916).
Platysporoides (Wehm.) Shoemaker & C.E. Babc., Can. J.
Bot. 70(8): 1648 (1992).
Type species: Platysporoides chartarum (Fuckel) Shoemaker
& C.E. Babc., Can. J. Bot. 70(8): 1650 (1992).
Basionym: Pleospora chartarum Fuckel, Jb. nassau. Ver.
Naturk. 23–24: 133 (1870) [1869–70].
Pleospora Rabenh. ex Ces. & De Not., Comm. Soc. crittog.
Ital. 1(4): 217 (1863).
Type species: Pleospora herbarum (Pers.) Rabenh.,
Klotzschii Herb. Viv. Mycol.: no. 547 (1854).
Basionym: Sphaeria herbarum Pers., Syn. meth. fung.
(Göttingen) 1: 78 (1801).
Porocercospora Amaradasa et al., Mycologia 106(1): 81 (2014).
Type species: Porocercospora seminalis (Ellis & Everh.)
Amaradasa et al., Mycologia 106(1): 81 (2013).
Basionym: Cercospora seminalis Ellis & Everh., J. Mycol.
4(1): 4 (1888).
Pseudoyuconia Lar.N. Vassiljeva, Nov. sist. Niz. Rast. 20: 71
(1983).
Type species: Pseudoyuconia thalictri (G. Winter) Lar.N.
Vassiljeva [as ‘thalicti’], Nov. sist. Niz. Rast. 20: 71 (1983).
Basionym: Leptosphaeria thalictri G. Winter, Hedwigia
10: 40 (1872)
Pyrenophora Fr., Summa veg. Scand., Section Post. (Stockholm): 397 (1849).
= Drechslera S. Ito, Proc. Imp. Acad. Japan 6: 355 (1930).
= Marielliottia Shoemaker, Can. J. Bot. 76(9): 1559 (1999)
[1998].
Type species: Pyrenophora phaeocomes (Rebent.) Fr., Summa
veg. Scand., Section Post. (Stockholm): 397 (1849).
Basionym: Sphaeria phaeocomes Rebent., Prodr. fl.
neomarch. (Berolini): 338 (1804).
Excluded genera
Austropleospora R.G. Shivas & L. Morin et al., Fungal Diversity 40(1): 70 (2010).
Comparison of alignment properties of genes and nucleotide substitution models used in Alternaria complex phylogenetic analysis
Genes /loci
LSU
ITS
GAPDH,
TEF1
RPB2
Alignment strategy (MAFFT v6)
Nucleotide substitution models for Bayesian
analysis (determined by MrModeltest)
FFT-NS-Il
GTR+I+G
FFT-NS-I+manual
GTR+I+G
FFT-NS-I+manual
GTR+I+G
FFT-NS-I+manual
GTR+I+G
FFT-NS-I+manual
SYM+I+G
Fungal Diversity
Type species: Austropleospora osteospermi R.G. Shivas & L.
Morin, in Morin, Shivas, Piper & Tan, Fungal Diversity 40(1):
70 (2010).
Dendryphion Wallr., Fl. crypt. Germ. (Norimbergae) 2: 300
(1833).
Type species: Dendryphion comosum Wallr., Fl. crypt. Germ.
(Norimbergae) 2: 300 (1833).
Edenia M.C. González et al., in González et al., Mycotaxon
101: 254 (2007).
Type species: Edenia gomezpompae M.C. González et al., in
González et al., Mycotaxon 101: 254 (2007).
Kriegeriella Höhn., Annls mycol. 16(1/2): 39 (1918).
Type species: Kriegeriella mirabilis Höhn., Annls mycol.
16(1/2): 39 (1918).
Macrospora Fuckel, Jb. nassau. Ver. Naturk. 23–24: 139
(1870) [1869–70].
= Nimbya E.G. Simmons, Sydowia 41: 316 (1989).
Type species: Macrospora scirpicola (DC.) Fuckel, Jb. nassau. Ver. Naturk. 23–24: 139 (1870) [1869–70].
Basionym: Sphaeria scirpicola DC., in Lamarck & de Candolle, Fl. franç., Edn 3 (Paris) 2: 300 (1805).
Monascostroma Höhn., Annls mycol. 16(1/2): 160 (1918).
Type species: Monascostroma innumerosum (Desm.) Höhn.
[as ‘innumerosa’], Annls mycol. 16(1/2): 160 (1918).
Zeuctomorpha Sivan et al., Bitunicate Ascomycetes and their
Anamorphs (Vaduz): 572 (1984).
Type species: Zeuctomorpha arecae Sivan et al., in
Sivanesan, Bitunicate Ascomycetes and their Anamorphs (Vaduz): 572 (1984).
Herein we accepted 18 genera in family Pleosporaceae
based on morphology coupled with molecular data and exclude 7 genera, which have previously been classified under
the family (sensu Hyde et al. 2013; Wijewardana et al. 2014).
Treatment of genera in Pleosporaceae
Alternaria Nees, Syst. Pilze (Würzburg): 72 (1816) [1816–
17]
Facesoffungi number: FoF 00501
Pathogenic or saprobic on wood and dead herbaceous stems
or leaves. Sexual morph: Ascomata small, solitary to clustered, erumpent to (nearly) superficial at maturity, globose to
ovoid, dark brown, smooth, apically papillate, ostiolate.
Ostiole papilla short, blunt. Peridium relatively thin.
Hamathecium of cellular pseudoparaphyses. Asci (4–6–) 8spored, bitunicate, fissitunicate, cylindrical to cylindro-
clavate, straight or somewhat curved, with a short, furcate
pedicel and minute ocular chamber. Ascospores ellipsoid to
fusoid, muriform, slightly constricted at septa, yellow-brown,
without guttules, smooth-walled. Asexual morph: Stroma
rarely formed, setae and hyphopodia absent. Conidiophores
macronematous, mononematous, simple or irregularly and
loosely branched, pale brown or brown, solitary or in fascicles. Conidiogenous cells integrated, terminal becoming intercalary, polytretic, sympodial, or sometimes monotretic,
cicatrized. Conidia catenate or solitary, dry, ovoid, obovoid,
cylindrical, narrowly ellipsoid or obclavate, beaked or nonbeaked, pale or medium olivaceous-brown to brown, smooth
or verrucose, with transverse and with or without oblique or
longitudinal septa (Woudenberg et al. 2013).
Type species: Alternaria alternata (Fr.) Keissl., Beih. bot.
Zbl., Abt. 2 29: 434 (1912), Figs. 3 and 4
Basionym: Torula alternata Fr., Syst. mycol. (Lundae)
3(2): 500 (1832)
Pathogen or saprobe on wood and dead herbaceous stems or
leaves. Sexual morph: Ascomata 170–20×190–240 μm (x=
190×440 μm, n=10), small, solitary to clustered, erumpent to
(nearly) superficial at maturity, globose to ovoid, dark brown,
smooth, apically papillate, ostiolate. Ostiole papilla short,
blunt. Peridium 38–50 μm (x=45 μm, n=10) wide, thin, comprising two cell types, outer layer composed of small heavily
pigmented, thick-walled cells of textura angularis, inner layer
composed of lightly pigmented or hyaline, thin-walled cells of
textura angularis. Hamathecium of 2–2.5 μm (x=2.5 μm, n=
10) broad, long, cellular pseudoparaphyses. Asci 170–190×
23–30 μm (x= 180 × 25 μm, n = 20), (4–6–) 8-spored,
bitunicate, fissitunicate, cylindrical to cylindro-clavate,
straight or somewhat curved, with a short, furcate pedicel
and minute ocular chamber. Ascospores 37–43×13–14 μm
(x=37×13.5 μm, n=40), ellipsoid to fusoid, muriform, slightly constricted at septa, with 3–7 transverse septa, 1–2 series of
longitudinal septa through the two original central segments,
end cells without septa, or with 1 longitudinal or oblique septum, or with a Y-shaped pair of septa, brown, without guttules,
smooth-walled. Asexual morph: Conidiophores 40–50×3–
6 μm solitary to clustered, simple or branched, straight or
flexuous, sometimes geniculate, pale, olivaceous or golden
brown, smooth. Conidia 20–63×9–18 μm, mostly branched,
obclavate, obpyriform, ovoid or ellipsoidal, often with a short
conical or cylindrical beak, pale to mid golden brown, up to 8
transverse and usually several longitudinal or oblique septa,
smooth or verruculose (Ellis 1971).
Material examined: ITALY, Monte Cervarola, Sestola, on
the dead stem, 26 March 2012, E. Camporesi IT181-1 MFLU
14–0755, ICMP, living culture (MFLUCC 14–1184). ITALY,
Monte Cervarola, Sestola, on the stem of Achillea sp.
(Asteraceae) 15 July 2013, E. Camporesi IT 181–2 MFLU
Fungal Diversity
Fig. 3 Alternaria alternata (MFLU 14–0755) a–b. Ascomata on host substrate. c. Section of ascoma (TS). d. Close up of peridium. e.
Pseudoparaphyses. f. Asci with 8-spores. g–j. Muriform, brown ascospores. Scale bar: c=80 μm, d=30 μm, e–f=20 μm, g–j=5 μm
14–0756 living culture (MFLUCC 14–1185). ITALY, Monte
Cervarola, Sestola, on the stem of Portulaca sp.
(Portulacaceae), 5 September 2012, E. Camporesi IT 466
(MFLU 14–0757); (KIB, PDD).
Notes: Alternaria was introduced by Nees (1816) and is a
ubiquitous genus that includes saprobic, endophytic and pathogenic species associated with a wide variety of substrates
(Woudenberg et al. 2013). Recent studies based on DNA data
as well as literature reviews revealed multiple paraphyletic
genera within the Alternaria complex, and Alternaria species
clades that do not always correlate to species-groups based on
morphological characteristics (Woudenberg et al. 2013).
Based on the combined gene analysis of GAPDH, RPB2
and TEF1, Woudenberg et al. (2013) concluded that the
Alternaria clade contains 24 internal clades and six
monotypic lineages, the grouping of which are recognised as
Alternaria and thus synonymised Allewia, Brachycladium,
Chalastospora, Chmelia, Crivellia, Embellisia, Lewia,
Nimbya, Sinomyces, Teretispora, Ulocladium, Undifilum and
Ybotromyces under Alternaria senso stricto Furthermore,
Woudenberg et al. (2013) treated 24 internal clades in the
Fungal Diversity
Fig. 4 Alternaria alternata
(Redrawn from Alternaria
alternata in Ellis (1971), Fig
330.). Scale bars: 650 μm
Alternaria complex as sections. Our phylogeny based on analysis of ITS, GAPDH, LSU, RPB2 and TEF1 sequence data
(Fig. 2) produced similar results as Woudenberg et al. (2013).
We therefore follow their recent proposal for the taxonomic
treatment of lineages in Alternaria.
In the present study we collected the sexual morph of
Alternaria alternata from Italy in 2013 and 2014. Cultures
did not produce the asexual morph but phylogenetically our
collections (MFLUCC 14–1185, MFLUCC 14–1184) are
100 % similar with the type strain of Alternaria alternata
(CBS 916.96). Therefore here we have given the description
of the sexual morph of Alternaria alternata based on our
collections.
We carried out a separate phylogenetic study for the
Alternaria section to show the placement of some newly introduce species in this study and some sexual reports of
Alternaria (Fig. 2). During our phylogenetic analysis we observed that the type strain of Xenobotryosphaeria
calamagrostidis (CBS 303.71) forms a clade within the
Alternaria section Infectoriae and clustered with Alternaria
infectoria.
The sexual morph of Xenobotryosphaeria calamagrostidis
is characterised by globose, smooth, superficial ascomata, a
peridium with 3–4 layers of brown textura angularis,
bitunicate, clavate, short pedicellate, 6–8-spored, asci and
multiseriate, hyaline, granular, broadly ellipsoid, aseptate
ascospores. This is a clear morphological deviation from the
sexual morph of Alternaria infectoria. Xenobotryosphaeria is
typical of genera in the Botryosphaeriales, but is phylogenetically distinct (Crous et al. 2006; Phillips et al. 2008; Liu et al.
2012). It also resembles species of Muyocopron
(Muyocopronaceae), but the latter genus differs in that it has
circular, flattened ascomata, as well as prominent
pseudoparaphyses, which are absent in Xenobotryosphaeria.
Therefore the confusion surrounding Xenobotryosphaeria remains unresolved.
During the study we found a novel species of Alternaria,
A. murispora from Germany and this is described below.
Alternaria murispora Ariyawansa & K.D. Hyde, sp. nov.,
Fig. 5
Index Fungorum number: IF 550952, Facesoffungi
number: FoF 00502
Etymology: Named after its muriform ascospores.
Holotype: MFLU 14–0758
Saprobic on dead stem. Sexual morph: Ascomata 110–160×
100–200 μm (x= 130 × 150 μm, n = 10), small, scattered,
erumpent to (nearly) superficial at maturity, globose or spheroid, dark brown, smooth, apically papillate, ostiolate. Ostiole
papilla short and blunt. Peridium 7–15 μm (x=12 μm, n=10),
thin, comprising two cell types, outer layer composed of small
Fungal Diversity
Fig. 5 Alternaria murispora a, b. Ascomata immersed in host substrate. c. Section of ascoma (TS). d. Close up of peridium. e Cellular
pseudoparaphyses. f–i. Asci with 8-spores. j–m. Brown ascospores. Scale bars: c–e=50 μm, f–i=30 j–m=10 μm
heavily pigmented, thick-walled cells of textura angularis,
inner layer composed of lightly pigmented or hyaline, thinwalled cells of textura angularis. Hamathecium of 1–2.5 μm
(x= 1.8 μm, n = 20), cellular, septate, pseudoparaphyses
branching and anastomosing between and above asci. Asci
75–106×11–16 μm (x=92×14 μm, n=20), (4–6–) 8-spored,
bitunicate, fissitunicate, cylindrical to subcylindrical, straight
or somewhat curved, with a short, furcate pedicel and minute
ocular chamber. Ascospores 16–18×5–7 μm (x=17×6 μm,
n=40), overlapping seriate, ellipsoid to fusoid, muriform, initially 3-transseptate become 4–5 at maturity, constricted at the
central septum, pale brown when immature, dark brown at
maturity, without guttules, smooth-walled, with a 1–2.5 μm
wide sheath. Asexual morph: undetermined.
Material examined: GERMANY, Frankfurt, on dead stem,
28 November 2013, A. D Ariyawansa (MFLU 14–0758,
holotype).
Notes: Single spore isolation was not successful and therefore DNA was extracted directly from the fruiting body of the
fungus. Therefore no living culture is available.
Notes: The novel species Alternaria murispora is introduced here based on both morphology and phylogeny.
Alternaria murispora fits well with the general concept of
Alternaria in having superficial, globose to ovoid ascomata
with short, blunt ostiole, cellular pseudoparaphyses, cylindrical asci with a short, furcate pedicel and ellipsoid to fusoid,
brown ascospores which are slightly constricted at the septa.
Morphologically A. murispora is closely related to
A. conjuncta E.G. Simmons (= Lewia scrophulariae (Desm.)
M.E. Barr & E.G. Simmons), but differs in having comparatively smaller asci (75–106 μm versus 120–140 μm), smaller
ascospores (16–18 μm versus 23–25 μm) and in the number
of septa at maturity (3–5 μm versus 6–7 μm). This is also
supported phylogenetically. i.e. A. conjuncta forms a sister
Fungal Diversity
clade to A. californica E.G. Simmons & S.T. Koike, while
A. murispora forms a sister clade with A. triticina Prasada &
Prabhu, in section Infectoriae (Fig. 2).
Bipolaris Shoemaker, Can. J. Bot. 37(5): 882 (1959)
= Cochliobolus Drechsler, Phytopathology 24: 973 (1934)
Facesoffungi number: FoF 00503
Pathogenic or saprobic on wood and dead herbaceous stems
or leaves. Sexual morph Ascomata brown or black, immersed, erumpent, partially embedded or superficial, free or
on flat stroma, mostly globose to ellipsoidal, sometimes flaskshaped or flattened on hard substrata, smooth or covered with
vegetative filaments. Ostiole arising centrally, papillate or
with a neck. Peridium comprising pseudoparenchymatous
cells of equal thickness or slightly thickened at apex.
Hamathecium dense septate, filiform, branched
pseudoparaphyses. Asci 2–8 spored, bitunicate, fissitunicate,
clavate, cylindrical-clavate or broadly fusoid, straight or
slightly curved, thin-walled, often becoming more or less
distended prior to dehiscence, short pedicellate, rounded at
apex. Ascospores fasciculate, filiform or flageliform, hyaline
or sometimes pale yellow or pale brown at maturity, septate,
helically coiled within ascus, degree of ascospore coiling
moderate to very strongly coiled, sometimes with free ends,
often with a thin mucilaginous sheath (modified from
Manamgoda et al. 2012). Asexual morph Conidiophores pale
to dark brown, single, branched, sometimes arranged in small
groups, straight to flexuous or geniculate with smooth or verrucose conidiogenous node. Conidia mostly curved, canoeshaped, fusoid or obclavate, rarely straight, 3–14
pseudoseptate (usually more than 6), hyaline, pale or dark
brown, reddish brown or pale to deep olivaceous, germinating
by production of one or two germination tubes by polar cells.
Hilum often slightly protruding or truncate sometimes inconspicuous. Septum ontogeny first septum median to sub median, second septum delimits basal cell and third delimits distal
cell.
Type species: Bipolaris maydis (Y. Nisik. & C. Miyake) Shoemaker, Canad. J. Bot. 33: 882 (1959), Fig. 6.
Basionym: Helminthosporium maydis Y. Nisik. & C.
Miyake, Journal of Plant Protection, Tokyo13: 20 (1926).
≡ Drechslera maydis (Y. Nisik. & C. Miyake) Subram.&
B.L. Jain, Curr. Sci. 35: 354 (1966)
= Helminthosporium maydis Brond., Ill. Iconogr.
Microscop.Cryptog. France 15. 1856–1857 (as
‘Helmisporium’), nom. rej.prop. (Rossman, Manamgoda &
Hyde, 2013b)
= Ophiobolus heterostrophus Drechsler in J. Agric. Res.
31: 701. 1925, nom. rej.prop (Rossman et al., 2013b)
= Cochliobolus heterostrophus (Drechsler) Drechsler,
Phytopathology 24: 973 (1934).
Facesoffungi number: FoF 00504
Pathogenic or saprobic on wood and dead herbaceous stems
or leaves. Sexual morph of Bipolaris luttrellii on Sach’s agar
medium: Ascomata 260–370 μm (x=316 μm, n=10) diam.,
superficial or slightly immersed, black, sub globose to ellipsoidal. Ostiole, subconical to campanulate, ostiolar beak and
upper part of ascomata covered by densely arranged setae.
Pseudoparaphyses filiform, hyaline, septate. Asci 140–205×
18–26 μm (x=178×22 μm, n=20), 1–8-spored, bitunicate,
fissitunicate, hyaline, subcylindrical, short pedicellate.
Ascospores 180–285×6–8 μm (x=235×7 μm, n=20), filiform, hyaline, tapered slightly towards apex and base, tightly
coiled inside ascus, sometimes slightly coiled to straight at
upper most part, (7–)8(−12)-distoseptate. Asexual morph of
Bipolaris maydis on PDA: Conidiophores 105–470×5–7 μm
(x=286 μm, n=20), usually arising singly or in small groups,
simple or rarely branched, septate, straight or flexuous, geniculate at upper part,olivaceous brown. Conidiogenous nodes
dark brown, distinct. Occasionally secondary sporulation observed. Conidia 66–102×14–18 μm (x=94×16 μm, n=40)
μm, pale to mid dark brown, smooth, slightly curved, fusiform, distoseptate. Hilum distinct, 3–5 μm wide, germination
tubes arising from both ends of conidia.
Material examined: AUSTRALIA, on Dactyloctenium
aegyptium, 3 June1985, J.L. Alcorn, (BRIP 14791, holotype
of dried culture of Cochliobolus luttrellii) and USA, North
Carolina, isolated from Zea mays L?, Olin Yoder C5, resulting
from six crosses, culture sporulating on Zea mays (BPI
892696, neotype of Bipolaris maydis).
Notes: Bipolaris was introduced by Shoemaker (1959) and
it is considered an important plant pathogen associated with
over 60 host genera (Sivanesan 1987; Manamgoda et al. 2011;
Agrios 2005; Hyde et al. 2014). Cochliobolus Drechsler
(1934) is the sexual stage of Bipolaris. There was no clear
morphological boundary between the asexual genera
Bipolaris and Curvularia, and some species show intermediate morphology which thus caused confusion for many plant
pathologists and mycologists for their correct identification
(Sivanesan 1987; Manamgoda et al. 2011; Hyde et al. 2014).
Based on combined gene analysis of rDNA ITS (internal transcribed spacer), LSU, GPDH and EF1-α, Manamgoda et al.
(2012) resolved the taxonomic confusion in Bipolaris and
Curvularia complex. Multilocus phylogeny showed that
Bipolaris and Curvularia form two well supported clades in
previous studies (Manamgoda et al. 2011, 2012) as well as in
the present study. Further, the nomenclatural conflict in this
complex was resolved giving priority to the more commonly
used established generic names Bipolaris and Curvularia thus
Cochliobolus was synonymised under Bipolaris. Recently
Manamgoda et al. (2014) revised the genus Bipolaris based
Fungal Diversity
Fig. 6 a-h: Bipolaris luttrellii (holotype of dried culture of Cochliobolus
luttrellii, j-n), Bipolaris maydis (neotype of Bipolaris maydis, a–h) a.
Ascomata on host substrate. b. Section of the ascomata. c. Close up of
the peridium d-e. Subcylindrical asci with a short pedicle note: helical
arrangement of the ascospores j. Conidiophore. k-n. Conidia. Scale bars:
b=100 μm, c=10 μm, d–g=60 μm, h–j=30 μm
on DNA sequence data derived from living cultures of fresh
isolates, available ex-type cultures from worldwide collections and observation of type and additional specimens. They
accepted 47 species in the genus Bipolaris and clarify the
taxonomy, host associations, geographic distributions and
species’ synonymies while epi- or neotypes were designated
for Bipolaris cynodontis (Marignoni) Shoemaker, B. oryzae
(Breda de Haan) Shoemaker, B. victoriae (F. Meehan & H.C.
Murphy) Shoemaker, B. yamadae (Y. Nisik.) Shoemaker and
B. zeicola (G.L. Stout) Shoemaker.
Fungal Diversity
In our phylogeny, Bipolaris forms a robust clade sister to
Curvularia and Porocercospora. Therefore we accept
Bipolaris as a well established genus in Pleosporaceae based
on both morphology and phylogeny.
Clathrospora Rabenh., Hedwigia 1(18): 116 (1857)
Facesoffungi number: FoF 00505
Saprobic on wood and stems. Sexual morph: Ascomata
semi-immersed, scattered on putrid host stems and foliage, brown to blackish brown, subglobose or nearly globose, with a central sunken ostiole open via a circular lid,
asci and pseudoparaphyses forming at the base of the
peridium. Peridium composed of 3–5 layers of brown,
relatively thick-walled cells of textura angularis, inner
cells flattened, thin-walled and lighter. Hamathecium
composed of dense, hyaline, filiform, pseudoparaphyses
which are longer than the asci. Asci 8-spored, bitunicate,
fissitunicate, thick-walled, cylindrical to clavate, with a
short pedicle and shallow ocular chamber. Ascospores
biseriate, fusiform, muriform, 7-transseptate, two or many
rows of longitudinal septa, applanate, constricted only at
the central septum, dark brown to brown, surrounded by a
thin, hyaline mucilaginous sheath. Asexual morph:
Alternaria like (Zhang et al. 2012).
Type species: Clathrospora elynae Rabenh., Hedwigia 1: 116
(1857), Fig. 7.
≡ Clathrospora elymae Rabenh. (1857).
≡ Pleospora elynae (Rabenh.) Ces. & De Not.,
Commentario della Società Crittogamologica Italiana 1 (4):
218 (1863).
Facesoffungi number: FoF 00506
Saprobic on wood and stems. Sexual morph: Ascomata
220–140×145–175 μm (x=170×150 μm, n=10), semi-immersed, scattered on the putrid host stems and foliage,
subglobose or nearly globose, brown to blackish brown,
with a central sunken ostiole open via a circular lid.
Peridium 20–55 μm (x=38, n=20), composed of 3–5 layers
of brown, relatively thick-walled cells of textura angularis,
inner cells flattened, thin-walled and lighter. Hamathecium
composed of dense, 2–3 μm diam (x=2, n=20), hyaline,
filiform, pseudoparaphyses, longer than the asci. Asci
160–230 × 24–48 μm (x= 190 × 35 μm, n = 20), 8-spored,
bitunicate, fissitunicate, thick-walled, cylindrical to clavate,
with a short pedicle and minute ocular chamber. Ascospores
40–65×18–27 μm (x=53×23 μm, n=40), biseriate, fusiform, muriform, 7-transseptate, two or many rows of longitudinal septa, constricted only at the central septum,
applanate, dark brown to brown, surrounded by a thin, hyaline mucilaginous sheath. Asexual morph: Alternaria-like
(Zhang et al. 2012).
Material examined: SWITZERLAND, on the stem of Carex
curvula, September 1898, Winter (BPI 627748, isotype).
Notes: Shoemaker and Babcock (1992) assigned
Clathrospora to Diademaceae and included an additional nine
species and provided a key to the genus based on the number
of septa and length of ascospores. Clathrospora was characterized by ascomata circular lid-like opening and applanate,
muriform ascospores. Currently, 50 Clathrospora species are
listed in the genus in Index Fungorum (2015). Molecular studies based on combine gene analysis showed that two putative
strains of Clathrospora, C. elynae (CBS 196.54) and
C. diplospora (IMI 68086) clustered in Pleosporaceae
(Ariyawansa et al. 2014a). We obtained similar results in the
phylogenetic tree produced from combined ITS, nrLSU,
nrSSU and RPB2 sequence analysis (Fig. 1). Clathrospora
elynae, the type of Clathrospora, formed a separate clade with
low bootstrap support (58 %) within Pleosporaceae. Based on
the phylogenetic result together with the morphological characters (slightly papillate ostiole with applanate, muriform ascospores and Alternaria- like asexual morph) we refer
Clathrospora to Pleosporaceae.
Comoclathris Clem., Gen. fung. (Minneapolis): 37, 173
(1909)
= Platyspora Wehm., World Monograph of the Genus
Pleospora and its Segregates: 254 (1961).
Facesoffungi number: FoF 00507
Saprobic on dead wood or stems. Sexual morph: Ascomata
semi-immersed to superficial, scattered or aggregated,
subglobose or nearly globose, brown to blackish brown coriaceous, ascomata opening via a large circular aperture or lid.
Peridium comprising 3–4 layers of brown, relatively thickwalled cells of textura angularis. Hamathecium composed
of dense, hyaline, filiform, septate pseudoparaphyses. Asci
8-spored, bitunicate, fissitunicate, cylindrical to cylindroclavate with an ocular chamber. Ascospores uniseriate or partially overlapping, fusiform, muriform, applanate, brown to
reddish-brown, surrounded by a thick, hyaline, mucilaginous
sheath. Asexual morph: Alternaria-like (Zhang et al. 2012).
Ty p e s p e c i e s : C o m o c l a t h r i s l a n a t a C l e m . [ a s
‘Comochlatris’], Gen. fung. (Minneapolis): 1–227 (1909),
Fig. 8.
Facesoffungi number: FoF 00508
Saprobic on dead stem. Sexual morph: Ascomata 175–245×
142–197 μm (x=205×159 μm, n=20), scattered or aggregated on the host stem, subglobose or nearly globose, superficial,
coriaceous, covered with a pale membrane, brown to blackish
brown, with a central ostiole. Peridium 10–22 μm wide, comprising 3–4 layers of brown, relatively thick-walled cells of
textura angularis, inner cells flattened, thin-walled and lighter.
Fungal Diversity
Fig. 7 Clathrospora elynae (isotype) a. Herbarium material. b. Close up
of ascomata. c. Section of the ascoma d. Close up of the peridium. e.
Hyaline, filiform, pseudoparaphyses. f–h. Cylindrical to clavate asci with
a short pedicle and ocular chamber. i-k. Dark brown to brown muriform
ascospores surrounded by a thin, hyaline mucilaginous sheath. Scale
bars: b=100 μm, c=10 μm, d–g=60 μm, h–j=30 μm
Fungal Diversity
Fig. 8 Comoclathris lanata (holotype). a. Herbarium material showing
habit of fungus on host stem. b. Erumpent ascomata. c. Section through
ascoma. Note the arrangement of asci and external setae. d.
Section showing peridial cells of ascoma. e. Hyaline, filform,
pseudoparaphyses. f. Asci with short knob-like pedicels and shallow
ocular chamber. g–i. Muriform, applanate ascospores with a thick
sheath. Scale Bars: b =200 μm, c=40 μm, d–e=10 μm, f =20 μm, g–
i=10 μm
Hamathecium composed of dense 2–4 μm wide, septate, hyaline, filiform, pseudoparaphyses longer than the asci. Asci
108–149 × 20–30 μm (x= 24 × 125 μm, n = 20), 8-spored,
bitunicate, fissitunicate, thick-walled, cylindrical to cylindroclavate, with a short knob-like pedicel, and indistinct shallow
ocular chamber. Ascospores 20–32×8–13 μm (x=12×28 μm,
n=20 ), 1–2 overlapping seriate, fusiform, muriform, with 4–
5-transverse septa and 1–2-longitudinal septa, not constricted
at the septa, applanate, brown to reddish-brown, surrounded
by a distinct, hyaline, mucilaginous 3–8 μm wide sheath.
Asexual morph: undetermined.
Material examined: USA, Colorado, on stem of Leptotaenia
multifida Nutt (Umbelliferae), 8 July 1907, F.E. & E.S. Clements (COLO 62872, holotype).
Notes: Comoclathris, typified by Comoclathris lanata, was
introduced by Clements (1909). The genus is characterized by
ascomata with circular lid-like openings and applanate
reddish-brown to dark reddish-brown, muriform ascospores,
with single longitudinal septa (Ariyawansa et al. 2014a).
Zhang et al. (2012) tentatively placed Comoclathris in the
Pleosporaceae based on Alternaria-like asexual morphs and
Fungal Diversity
this was followed by Woudenberg et al. (2013). Comoclathris
shares common characters with Pleospora herbarum, the type
of Pleospora, in having cylindrical to cylindro-clavate asci
with an ocular chamber and muriform, brown or pale brown
ascospores, with or without sheath. Comoclathris and
Pleospora differ in the opening of ascomata (opening via a
large circular aperture or lid versus opening by a central pore
and applanate ascospores). Comoclathris and Pleoseptum
share similar characters in having globose, black, ascomata
and cylindrical to cylindro-clavate asci with muriform, yellowish to dark brown ascospores. Comoclathris differs from
Pleoseptum in having superficial ascomata with circular lidlike openings composed of comparatively thin peridium and
applanate and fusiform ascospores surrounded by a distinct
hyaline, mucilaginous thick sheath (Ariyawansa et al.
2014a). In Pleoseptum ascomata are immersed, usually with
a papillate apex, with a relatively broad peridium and ovoid to
fusoid ascospores (Ariyawansa et al. 2014a). Comoclathris
was considered to differ from Clathrospora as in the latter
genus species have two or more rows of longitudinal septa
as compared with a single row in Comoclathris (Ariyawansa
et al. 2014a). Shoemaker and Babcock (1992) provided a key
to 21 species of Comoclathris. Presently 32 epithets are listed
for Comoclathris in Index Fungorum (2015).
Molecular data for Comoclathris lanata, the type species of
Comoclathris, is not available. Two putative strains of
Comoclathris compressa (CBS 157.53 and CBS 156.53)
however, cluster together in a well-supported clade within
the family Pleosporaceae (Ariyawansa et al. 2014a). Based
on the phylogenetic result, coupled with the morphological
characters (Alternaria-like asexual morph), we agree with
Ariyawansa et al. (2014a), Zhang et al. (2012) and
Woudenberg et al. (2013) in placing Comoclathris in
Pleosporaceae, but re-collection of the type species and
epitypification or a reference specimen (Ariyawansa et al.
2014a) with molecular data is essential to establish the correct
placement of the genus.
During this study we have proposed several new combinations in order to resolve the paraphyletic nature of
Pleospora and to make a stable taxonomy for the family
Pleosporaceae. Pleospora incompta (Sacc. & Martelli)
Gruyter & Verkley) (CBS 467.76) was introduced by de
Gruyter et al. (2013) in order to accommodate Phoma
incompta Sacc. & Martelli. In the same study Pleospora
typhicola Cook was proposed by de Gruyter et al. (2013)
to accommodate Phoma typhina (Sacc. & Malbr.) van der
Aa & Vanev in the family Pleosporaceae but during our
study Pleospora typhicola (Cooke) Sacc. 1875) (CBS
132.69) and Pleospora incompta (CBS 467.76) form separate clades within the genus Comoclathris (Fig. 1). Therefore we provide two new combinations namely
Comoclathris incompta and C. typhicola to accommodate
Pleospora incompta and P. typhicola in Comoclathris.
Comoclathris incompta (Sacc. & Martelli) Ariyawansa & K.
D. Hyde, comb. nov.,
Fungorum number: IF 550953
Basionym: Phoma incompta Sacc. & Martelli, Syll. Fung.
10: 146.1892.
≡ Pleospora incompta (Sacc & Martelli) Gruyter &
Verkley, in Gruyter et al., Stud. Mycol. 75: 25 (2012).
Comoclathris typhicola (Cooke) Ariyawansa & K. D. Hyde,
comb. nov.,
Fungorum number: IF 550954
Basionym: Sphaeria typhicola Cooke, Grevillea 5: 121.
1877.
≡ Pleospora typhicola (Cooke) Sacc., Reliq. Libert 2: no.
152 (1875)
≡ 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.
We also collected a novel species of Comoclathris, C. sedi
from Italy and this is described below.
Type species: Comoclathris sedi Wanasinghe, Ariyawansa,
E. Camporesi & K.D. Hyde, sp. nov., Fig. 9.
Index Fungorum number: IF 550955, Facesoffungi
number: FoF 00509
Etymology: The specific epithet sedi is based on the host
genus from which the fungus was isolated.
Holotype: MFLU 14 0758
Saprobic on dead stem. Sexual morph: Ascomata 200–250×
290–350 μm (x=230×320 μm, n=10), scattered or aggregated on the host stem, subglobose or nearly globose, superficial,
coriaceous, brown to blackish brown with a blunt ostiole.
Peridium 20–38 μm (x=25 μm, n=10) wide, comprising 3–
4 layers of brown, relatively thick-walled cells of textura
angularis, inner cells flattened, thin-walled and lighter.
Hamathecium composed of dense 1.5–2.5 μm (x=2 μm, n=
10) wide, septate, hyaline, filiform, pseudoparaphyses longer
than the asci. Asci 80–110×16–18 μm (x=98×18 μm, n=20),
8-spored, bitunicate, fissitunicate, cylindrical to cylindro-clavate, with a short knob-like pedicel, and indistinct shallow
ocular chamber. Ascospores 19–20 × 8–10 μm (x= 20 ×
19 μm, n=40 ), 1–2 overlapping seriate, fusiform, muriform,
Fungal Diversity
Fig. 9 Comoclathris sedi (holotype). a. Habit of the fungus on host
stem. b. Superficial ascomata. c. Thick section through ascoma. d.
Dark brown setae. e. Hyaline pseudoparaphyses. f-h. Asci with shallow
ocular chamber. i-k. Muriform ascospores with thick sheath. Scale Bars:
b=200 μm, c=40 μm, f–h=10 μm, i–k=20 μm
with 4–5-transverse septa and 1–2-longitudinal septa, not constricted at the septa, brown to reddish brown, surrounded by a
distinct, hyaline, mucilaginous 5–9 μm wide sheath. Asexual
morph: undetermined.
(MFLUCC 13 0817, BRIP); ITALY, Monte Cervarola,
Sestola, on dead branch of Clematis vitalba (Ranunculaceae),
21 July 2012, E. Camporesi IT 589, (KIB, paratype) – extype
living culture (ICMP); ITALY, Almazzago, on dead stem of
Rosa sp. (Rosaceae), 8 August 2013, N. Camporesi IT 1408
(MFLU14-0760) – living culture (MFLUCC 13 0763).
Notes: The novel taxon, Comoclathris sedi was initially
isolated on dead stem of Sedum sp. But later the same species
Material examined: ITALY, Stavel, Ortignano-Raggiolo, on
stem of Sedum sp. (Crassulaceae), 6 June 2012, E. Camporesi
IT 416, (MFLU 14 0759, holotype) – extype living culture
Fungal Diversity
was isolated from Clematis vitalba, Rosa sp. and Digitalis sp.
and they are morphologically and phylogenetically identical.
Comoclathris sedi shows similarities with Comoclathris, in
having globose, black, ascomata with setae and cylindrical
to cylindro-clavate asci having knob-like pedicel with
muriform, yellowish to dark brown applanate ascospores.
Our new species differs from other species of Comoclathris
in having superficial, 200–250×290–350 μm ascomata ()
with distinct ostioles, , a peridium comprising a single layer
of brown, relatively thick-walled cells of textura angularis,
and fusiform to ellipsoidal, muriform, brown to reddish brown
ascospores with 2–5-transverse septa and 1–4-longitudinal
septa. The phylogenetic analysis of combined ITS, LSU,
SSU nrDNA and RPB2 sequence data provides strong evidence that Comoclathris sedi belongs in Pleosporaceae and
forms a robust clade within the genus Comoclathris sister to
Comoclathris compressa (CBS 157.53 and CBS 156.53) with
relatively high bootstrap support (Fig. 1); thus a new species is
proposed.
Curvularia Boedijn, Bull. Jard. bot. Buitenz, 3 Sér. 13(1): 123
(1933)
= Pseudocochliobolus Tsuda, Ueyama & Nishih.,
Mycologia 69(6): 1117 (1978) [1977]
= Curvusporium Corbetta [as ‘Curvosporium’], Riso 12(3):
28, 30 (1963)
= Malustela Bat. & J.A. Lima, Publções Inst. Micol. Recife
263: 5 (1960)
Facesoffungi number: FoF 00510
Pathogenic or saprobic on wood and dead herbaceous stems
or leaves or humans. Sexual morph: Ascomata superficial,
globose to ellipsoidal, dark brown to black, free or frequently
developing from columnar stromata or flat stromata, with a
well defined ostiolar neck. Peridium coriaceous, carbonaceous, pseudoparenchymatous. Hamathecium comprising filiform, septate and sometimes branched pseudoparaphyses,.
Asci 1–8-spored, bitunicate, fissitunicate, cylindrical to cylindrical clavate, pedicel short and minute ocular chamber.
Ascospores fasciculate, usually parallel, loosely coiled or
highly coiled at the extremities of the ascus, filamentous, filiform to flageliform and somewhat tapered at the extremities,
3–20 septate, hyaline or somewhat pigmented at maturity.
Asexual morph: Conidiophores branched or unbranched,
straight to flexuous, septate, smooth to verruculose, often geniculate sometimes nodulose. Conidiogenous cells polytretic,
integrated, sometime when mature becoming intercalary, sympodial, cylindrical with smooth to verrucose conidiogenous
nodes. Conidia straight oblong, ellipsoidal, clavate, fusiform,
subcylindrical or lunate, rounded at the ends or sometimes
tapering slightly towards the base, pale brown, medium reddish brown to dark brown, 3–10 distoseptate (usually 3–5),
conidial wall smooth to verrucose. Hilum protuberant in some
species. Stromata formed in some species (obtained from
Manamgoda et al. 2012).
Type species: Curvularia lunata (Wakker) Boedijn, Bull.
Jard. bot. Buitenz, 3 Sér. 13(1): 127 (1933), Fig. 10.
≡ Acrothecium lunatum Wakker, De ziekten van het
suikerriet op Java, die niet door dieren veroorzaakt worden:
196 (1898).
= Helminthosporium curvulum Sacc., Atti della Accademia
Scientifica Veneto-Trentino-Istriana 10: 89 (1916).
= Helmisporium curvulum Sacc. (1916).
Facesoffungi number: FoF 00511
Pathogenic or saprobic on wood and dead herbaceous stems
or leaves. Sexual morph not observed in culture. Asexual
morph: Conidiophores 39–430×4–9 μm, branched or unbranched, straight to flexuous, septate, smooth to verruculose,
often geniculate sometimes nodulose. Conidiogenous cells 4–
20×3–13 μm, polytretic, integrated, sometime when mature
becoming intercalary, sympodial, cylindrical, cicatrized or
sometimes swollen. Conidiogenous nodes smooth to verrucose. Conidia 21–31×9–13 μm, straight oblong, ellipsoidal,
clavate, fusiform, sub-cylindrical or lunate, rounded at the
ends or sometimes tapering slightly towards the base, pale
brown, medium reddish brown to dark brown, 3–10
distoseptate (usually 3–5), conidial wall smooth to verrucose.
Material examined: THAILAND, Chiang Rai, on stem of
Zea mays, 8 July 2012, D.S Manamgoda (MFLU 10–0555)
– extype living culture (MFLUCC 12–0181).
Notes: Curvularia was introduced by Boedijin (1933) and
it is an important pathogen in humans and plants (Sivanesan
1987; Agrios 2005, Manamgoda et al. 2012, 2014). Bipolaris
and Curvularia share many morphological similarities
(Sivanesan 1987). There are some Bipolaris species with
short, straight conidia showing intermediate conidial characters between these two genera (Manamgoda et al. 2012). Confusion surrounding Bipolaris and Curvularia was resolved by
Manamgoda et al. (2012) based on combined genes of ITS,
GPDH, LSU, EF1-α sequence data thus Manamgoda et al.
(2012) revealed that the traditionally circumscribed Bipolaris
and Curvularia cannot be combined into a single monophyletic genus (Goh et al. 1998; Shimizu et al. 1998; Berbee et al.
1999). A similar phylogeny was found in the present study,
where Bipolaris and Curvularia form two well-supported
clades in the family Pleosporaceae, thus we accept
Curvularia as a separate genus in Pleosporaceae.
Dactuliophora C.L. Leakey, Trans. Br. mycol. Soc. 47(3):
341 (1964).
Facesoffungi number: FoF 00512
Fungal Diversity
Fig. 10 Curvularia lunata (MFLU 10–0555). a, b. Fungus on host stem. c, d. Conidiophores. e-i. Conidia. j. Atypical bifurcate conidium. Scale Bars:
c–j =10 μm
Parasitic on leaves. Mycelium generally immersed, hyaline
and diffuse in the leaf tissues, aggregated irregularly in the
epidermal or deeper leaf tissues into plectenchymic masses
from which sclerotiophores and sclerotia develop,
‘sclerotiophore’ remains after the disjunction of the sclerotium
as a superficial or occasionally more or less immersed structure, being the external continuation of the immersed aggregation. Hyphae at the circumference are dematiaceous and
relatively large while those in the centre of the ring so formed
are generally paler and relatively smaller. Sclerotia maybe
glaborus, hispidulous, hispid or sparsely setose, spherical to
subspherical, ellipsoidal, pyriform or rostrate, wholly or partly
composed of dematiaceous cells on the outside, hyaline and
undifferentiated, separates from sclerotiophore by the fracture
of many thin-walled cells joining the base of the mature sclerotium to the centre of the sclerotiophore. Note: No other
structures have been observed other than sclerotia (Leakey
1964).
Type species: Dactuliophora tarrii C.L. Leakey, Trans. Br.
mycol. Soc. 47(3): 343 (1964), Fig. 11.
Facesoffungi number: FoF 00513
Fungal Diversity
Fig. 11 Dactuliophora tarrii
(Redrawn from the holotype in
Leakey 1964, Figs. 1 and 2).
Scale bars: = 50 μm
Parasitic on leaves. Leaf spots broadly zonate on upper leaf
surface, rosette-like altering white and brown bands, lower surface densely covered with sclerotiophores and sclerotia, zonation less distinct. Mycelium immersed, hyaline, generally diffused throughout the tissues in the leaf spot and aggregated in
plectenchymic masses beneath the sclerotiophores.
Sclerotiophores consist of a fully erumpent ring of
dematiaceous hyphae continuous with a subepidermal
plectenchymic mycelial mass. Sclerotia 35–135 μm diam.,
scattered, specially on the paler parts of the leaf spot, generally
hypophyllous but occasionally amphigenous, spherical to
subspherical, bearing 0–10 (usually 3–7) scattered setae or
sometimes glabrous, dematiaceous external cells (8–12 μm
diam.), internal cells hyaline, undifferentiated. Sclerotial setae
3–4 μm diam., when present more or less cylindrical but sometimes slightly sinuate, slightly attenuated distally to a blunt tip,
dematiaceous or hyaline, 0–13 septate (Leakey 1964).
Notes: Dactuliophora was introduced by Leakey (1964) to
accommodate four species of Dactuliophora, namely
D. elongata C.L. Leakey, D. glycines C.L. Leakey,
D. harrisiae C.L. Leakey and D. tarrii C.L. Leakey and typified with D. tarrii (Leakey 1964). The genus is characterised
on the basis of the production of a cup-like ‘sclerotiophores’,
which are parasitic on several economically important crops
(Leakey 1964). Wijayawardene et al. (2014) placed
Dactuliophora in Pleosporaceae thus we follow this classification. Currently five Dactuliophora species are listed in Index Fungorum (2015) including D. anuae S.M. Singh., and no
molecular data is available for any of these species. Therefore
fresh collections of the type species of the genus are needed so
that molecular data can be obtained to verify the natural taxonomic affinities of this genus.
Decorospora Inderb et al., in Inderbitzin et al., Mycol. Progr.
1(4): 657 (2002).
Facesoffungi number: FoF 00514
Saprobic on dead wood and maritime plants in marine habitats. Sexual morph: Ascomata subglobose to ellipsoidal,
immersed, ostiolate, epapillate or short papillate, carbonaceous, black. Peridium composed of thick-walled cells with
large lumina, forming a textura angularis in longitudinal section. Hamathecium composed of septate, branch
pseudoparaphyses. Asci 8-spored, bitunicate, fissitunicate,
clavate, short pedicellate, thick-walled, with a clear ocular
chamber. Ascospores biseriate, ellipsoidal, muriform, brown,
covered by a gelatinous sheath that is slightly constricted
around the centre and drawn out at each apex into 2 or rarely
3 subconical extensions. Asexual morph: undetermined.
Decorospora gaudefroyi (Pat.) Inderb. et al., in Inderbitzin
et al., Mycol. Progr. 1(4): 657 (2002), Fig. 12
Basionym: Pleospora gaudefroyi Pat., Tabl. analyt. Fung.
France (Paris) 10: 40 (no. 602) (1886)
= Pleospora salsolae Fuckel var. schoberiae (Sacc.) Sacc.,
Syll. fung. (Abellini) 2: 248 (1883)
≡ Pleospora schoberiae (Sacc.) Berl., Icon. fung.
(Abellini) 2: 23 (1895)
= Pleospora lignicola J. Webster & M. T. Lucas, Trans. Br.
mycol. Soc. 44(3): 431 (1961)
= Pleospora salicorniae Jaap, Verh. Bot. Ver. Prov. Brandenburg 49, 16. 1907 (non Pleospora salicorniae P. A. Dang.
1888)
= Pleospora herbarum var. salicorniae Jaap, Annls mycol.
14(1/2): 17 (1916).
Facesoffungi number: FoF 00515
Saprobic on dead wood in marine habitats. Sexual morph:
Ascomata perithecioid, small, 200–250 μm in diam., globose
to subglobose, immersed to erumpent, becoming superficial
on the substrate at maturity, carbonaceous, black, solitary or
several clustered, short ostiolate, papillate. Ostiole central,
short, brown to black. Peridium thin, composed of thickwalled cells, forming a textura angularis in both surface view
and longitudinal section, up to 12 μm. Hamathecium composed of 2 μm wide, hypha-like, cellular, septate, branched
pseudoparaphyses anastomosing above the asci. Asci
80–125 × 15–25 μm (x= 104 × 18 μm, n = 10), 8-spored,
bitunicate, fissitunicate, clavate, short pedicellate, thick-
Fungal Diversity
Fungal Diversity
Decorospora gaudefroyi (holotype). a. Ascomata on the host
surface. b. Peridium cells from surface view. c. Section of ascoma. d.
Peridium. e–f. Branched pseudoparaphyses. g. Young asci amongst
pseudoparaphyses. h. Tip of the ascus. Note the ocular chamber. i.
Pedicellate ascus base. j–l. Ascospores. k, l Ascospores stained in
Indian ink. Note the sheath. Scale bars: a=300 μm, b=10 μm, c=
50 μm, d−e=10 μm, f=5 μm, g=30 μm, h−l=10 μm
Fig. 12
walled with large lumina, apically rounded with an ocular
chamber. Ascospores 20–25×7–10 μm (x=21.2×9 μm, n=
10), biseriate, ellipsoidal, muriform, 6–7 transverse septa
and 2–3 longitudinal septa in each segment, slightly constricted at the centre transverse septum, brown, covered by a gelatinous thick sheath with 2 or rarely 3 extensions. Asexual
morph: undetermined.
Material examined: France, Marais de la Pointe de Touquet,
Etaples, on Suaeda maritima (L.) Dumort, 15 Aug. 1879,
O. Hariot (PC 0084490, holotype)
Notes: Decorospora gaudefroyi (Pat.) Inderbn et al., is a
marine ascomycete firstly described as Pleospora gaudefroyi
Pat. from northern France in 1886 and described by Inderbn
et al. (2002). Yusoff et al. (1994) described the ascospores of
P. gaudefroyi at the ultrastructural level without the unfolded
sheath extensions. Based on analysis of partial SSU and ITS
ribosomal DNA gene data transferred P. gaudefroyi to a new
genus Decorospora, as it did not group in Pleospora. Morphological characters of D. gaudefroyi include black,
erumpent ascomata becoming superficial on the substrate at
maturity, septate and branched pseudoparaphyses,
fissitunicate, clavate asci with thick wall, as well as yellowbrown ascospores with seven transverse and 1–3 longitudinal
septa with a gelatinous sheath with a tripartite outer boundary.
The characteristic sheath makes these species easy to distinguish from any other marine ascomycetes. D. gaudefroyi was
considered to be a synonym of Pleospora herbarum (Fr.)
Rabh. which is the type species of Pleospora (Yusoff et al.
1994). However, the presence of the ascospore sheath with its
apical extensions and its marine habitat distinguish it from
other marine Pleospora species (Inderbn et al. (2002).
Decorospora is placed in Pleosporaceae, which is well-supported by partial SSU and ITS ribosomal DNA
sequences.
During our phylogenetic analysis, the type strain of
Decorospora gaudefroyi (pp4723) forms a robust clade basel
to Comoclathris. Furthermore the putative strains of
Pleospora sp. ATCC MYA-3203 and Pleospora sp. ATCC
MYA-3202 are included. Thus we renamed them as
Decorospora sp. in this study.
Diademosa Shoemaker & C.E. Babc., Can. J. Bot. 70(8):
1641 (1992).
Facesoffungi number: FoF 00516
Saprobic on stems and wood. Sexual morph: Ascomata immersed, initially erumpent becoming superficial, scattered,
depressed-globose, some flattened at the base, opening a
disc-like lid of brown prismatic cells with setae. Peridium
composed of brown pseudoparenchyma cells of textura
angularis. Hamathecium comprising numerous, septate, hyaline, cellular pseudoparaphyses. Asci 8-spored, bitunicate,
fissitunicate, clavate with short narrow pedicel and minute
ocular chamber. Ascospores biseriate, partially overlapping,
fusiform, straight, frequently circular in section but narrowing
to one end, with transverse and vertical septa, pale brown to
dark brown, smooth walled. Asexual morph: undetermined.
Type species: Diademosa californiana (M.E. Barr) Shoemaker & C.E. Babc. [as ‘californianum’], Can. J. Bot. 70(8): 1641
(1992), Fig. 13
Basionym: Graphyllium californianum M.E. Barr, Mem.
N. Y. bot. Gdn 62: 40 (1990).
Facesoffungi number: FoF 00517
Saprobic on stem and wood. Sexual morph: Ascomata 200–
365×240–425 μm (x=315×275 μm, n=10), immersed, initially erumpent becoming superficial, scattered, depressedglobose, some flattened at the base, opening a disc-like lid
of brown prismatic cells with setae. Peridium 24–59 μm (x=
32 μm, n=20), composed brown pseudoparenchyma cells of
textura angularis. Hamathecium of dense, 2–3 μm diam. (x=
2 μm, n = 20), numerous, septate, hyaline, cellular
pseudoparaphyses. Asci 140–175 × 24–28 μm (x= 150 ×
26 μm, n=20), 8-spored, bitunicate, fissitunicate, clavate with
short narrow pedicel and minute ocular chamber. Ascospores
50–65×26–32 μm (x=57×29 μm, n=40), biseriate or discontinuously arranged, partially overlapping, fusiform, straight,
cylindrical, frequently circular in end view with transverse
and vertical septa, muriform, constricted at first septum, pale
brown to dark brown, smooth walled with narrow sheath.
Asexual morph: undetermined.
Material examined: USA, Bump-Cold Boiling Lake Trail,
Lassen Volcanic National Park, Shasta, California, on branch
of Wyethia Nutt (Asteraceae), 12 July 1966, W.B. Cooke &
D.L. Hawksworth. (NY, holotype).
Notes: Diademosa was established by Shoemaker and
Babcock (1992) and typified by D. californiana, based on
the ascomatal opening via a circular lid and ascospores
being frequently circular in end view. Diademosa
californiana was initially introduced as Graphyllium
californianum by Barr (1990) and referred to Hysteriaceae
based on the pore or slit like opening. Re-examination of the
type specimens by Shoemaker and Babcock (1992) concluded
that Diademosa opened by a flat lid similar with Diadema and
assigned it in to Diademaceae. The lid is hard to observe in
sections unless they are mounted directly in lactic acid
Fungal Diversity
Fig. 13 Diademosa californiana (holotype) a, b. Ascomata on host
substrate.c. Section of ascoma. d. Section of peridium. e. Septate,
hyaline, cellular pseudoparaphyses f. Light to dark brown setae. g, h.
Ascus with minute pedicel bearing 8 irregularly arranged ascospores. ik. Ascospores. l. Ascospore stained in Indian ink and showing narrow
sheath. Scale bars: d–e=200μm, f–h=10μm, g–h=50μm, i–l=10μm
because excessive swelling occurs in water (Shoemaker and
Babcock 1992). Diademosa differs from Comoclathris in having cylindrical ascospores which are frequently circular in
section, but narrow of one end, as compared with the flattened
ascospores of Comoclathris. Ariyawansa et al. (2014a) placed
Diademosa in Pleosporaceae based on its similarities with
other genera in this family, but confirmation of the phylogenetic status of this genus depends on recollecting the fungus
and epitypification with molecular sequences.
Vostoka [Algae, Fungi and Mosses of the Soviet Far-East]
(Vladivostok): 80 (1978)
Facesoffungi number: FoF 00518
Exserohilum K.J. Leonard & Suggs, Mycologia 66(2): 289
(1974)
= Setosphaeria K.J. Leonard & Suggs, Mycologia 66(2):
294 (1974).
= Luttrellia Khokhr. & Gornostaĭ, in Gornostaĭ in
Azbukina et al. (Eds), Vodorosli, Griby i Mkhi Dal’nego
Saprobic or pathogenic on leaves or wood. Sexual morph:
Ascomata superficial, globose or ellipsoid with or without an
ostiole. Ostiole surrounded by simple, short, rigid, brown
hairs, similar hairs scattered over surface of upper part of
ascomata. Peridium coriaceous-carbonaceous,
pseudoparenchymous. Pseudoparaphyses filamentous. Asci
1–8-spored, bitunicate, cylindrical to cylindrical-clavate.
Ascospores hyaline, fusoid, trans-septate, surrounded by a
mucilaginous sheath. Asexual morph: Conidiophores cylindrical, simple, olivaceous brown, upwardly geniculate.
Conidia porogenous, acrogenous and pseudopleurogenous,
subclylindrical to fusoid or broad obclavate-rostrate,
Fungal Diversity
pseudoseptate, olivaceous to brown, hilum protruding from
basal cell, germinating by bipolar germ tubes.
Exserohilum turcicum (Pass.) K.J. Leonard & Suggs,
Mycologia 66(2): 291 (1974), Fig. 14
≡ Helminthosporium turcicum Pass., Boln Comiz. Agr.
Parmense: 3 (1876).
≡ Bipolaris turcica (Pass.) Shoemaker, Canadian Journal of
Botany 37 (5): 884 (1959).
≡ Drechslera turcica (Pass.) Subram. & B.L. Jain, Current
Science 35 (14): 355 (1966).
≡ Luttrellia turcica (Pass.) Khokhr., Vodorosli, Griby i
Mkhi Dal’nego Vostoka: 81 (1978).
= Helminthosporium inconspicuum Cooke & Ellis,
Grevillea 6 (39): 88 (1878).
= Trichometasphaeria turcica Luttr., Phytopathology
48(5): 282 (1958).
Facesoffungi number: FoF 00519
Pathogenic on living leaves of Zea mays. Sexual morph:
undetermined. Asexual morph: Conidiophores (111-)147–
164(−215)×7.3–8.5(−11) μm (x=158×8.2 μm, n=20) erect
singly or in groups of 2–5, long, straight or flexuous, cylindrical,
unbranched, or branched below, 2–5-septate, grayish brown,
inner wall layers of conidiogenous cell are continuous with
the conidial wall, pale to medium brown. Conidia (41-)112–
127×(17-)22–23(−29) μm (x=99×23 μm, n=20), ellipsoid-fusiform, widest at the middle and tapering toward ends, long,
straight or slightly curved, 2–8-distoseptate, some euseptate,
grayish brown, thick-walled, smooth, scars thickened.
Material examined: ZAMBIA, Katete, on living leaves of
Zea mays (Poaceae), 9 February 1957, A. Angus, NY (Ex
herbarium: IMI 69726).
Notes: The genus Exserohilum was proposed by Leonard
and Suggs (1974) to accommodate species which were previously classified in Bipolaris, in which the conidial hilum was
distinctly protuberant. In the same study, Leonard and Suggs
(1974) introduced a new genus Setosphaeria to place the sexual morphs of Exserohilum, which is segregated from
Keissleriella on the basis of lack of a clypeus, lysigenous
development of the ostiole, occurrence of setae on the perithecial wall, the absence of periphyses in the ostiole and the
hyphomycetous conidial states. As a genus can now only
have one name, Wijayawardene et al. (2014) proposed to conserved Exserohilum over Setosphaeria on the basis having of
more epithets (Index Fungorum 2015), and is more widely use
in the literature.
Fig. 14 Exserohilum turcicum (holotype). a. Herbarium material. b. Conidiophores and conidia on leaf surface of Zea mays. c. Conidiophores. d.
Conidiophore and conidium. e. I Immature and mature conidia. Scale bars: b=200 μm, c, d, h, i=50 μm, e–g=20 μm
Fungal Diversity
During our phylogenetic study putative strains of
Exserohilum monoceras (= Setosphaeria monoceras CBS
154.26), Exserohilum sp. (C950801) and Exserohilum sp.
(NK93) formed a well-supported clade sister to Curvularia.
Therefore based on phylogeny and morphology we accept
Exserohilum as a separate genus in the family Pleosporaceae.
Extrawettsteinina M.E. Barr, Contr. Univ. Mich. Herb. 9(8):
538 (1972).
Facesoffungi number: FoF 00520
Saprobic on dead leaves and wood. Sexual morph: Ascomata
scattered, erumpent to superficial, globose or conical, dark
brown to black, carbonaceous. Papilla black, with a porelike ostioles, ostiolar canal filled with periphyses. Peridium a
single layer composed of compressed, thick-walled cells, fusing with the host at the outside. Hamathecium of dense broad,
cellular pseudoparaphyses. Asci 8-spored, bitunicate,
fissitunicate, cylindrical, apex rounded with and a minute ocular chamber with short pedicel. Ascospores uniseriate, slightly overlapping, ellipsoid, obovate-clavate, 3-septate, slightly
to not constricted at the septum, wall smooth. Asexual
morph: undetermined.
Extrawettsteinina minuta M.E. Barr, Contr. Univ. Mich.
Herb. 9(8): 538 (1972), Fig. 15
≡ Kriegeriella minuta (M.E. Barr) Arx & E. Müll., Stud.
Mycol. 9: 88 (1975)
Facesoffungi number: FoF 00521
Saprobic on dead leaves of Juniperus communis. Sexual
morph: Ascomata 100–150 × 210–350 μm (x= 120 ×
280 μm, n=10), scattered, erumpent to superficial, globose
or conical, dark brown to black, carbonaceous. Papilla black,
with a pore-like ostioles, ostiolar canal filled with periphyses.
Peridium 10–25 μm (x=21 μm, n=10), a single layer composed of compressed, thick-walled cells, fusing with the host
at the outside. Hamathecium of dense 1–1.5 μm (x=1.25 μm,
n=20), broad, cellular pseudoparaphyses. Asci 75–90×6–
8 μm (x=80×7 μm, n=10), 8-spored, bitunicate, fissitunicate,
cylindrical, apex rounded with and a minute ocular chamber
with short pedicel. Ascospores 10–12×4–5 μm (x=11.5×
4 μm, n=10), uniseriate, slightly overlapping, ellipsoid, obovate-clavate, 3-septate, slightly to not constricted at the septum, wall smooth. Asexual morph: undetermined.
Material examined: USA, Vermont, Stow Pinnacle trail, on
dead leaves of Juniperus communis L (Cupressaceae), 11 July
1964, H.E and M.E Bigelow (NY, 4278).
Notes: During our study we observed the holotype of
Extrawettsteinina minuta (NY, 01103003) collected by H. E.
Bigelow (19 August 1957) from Juniperus communis but we
could not find any ascomata in the type material. Therefore we
examined another collection of E. minuta (NY, 4278), which
was collected from same host and by the same collector as
present in Fig. 15.
Extrawettsteinina was introduced by Barr (1972) and typified by E. minuta. The genus was introduced to accommodate
three species, i.e. E. minuta, E. pinastri M.E. Barr and
E. mediterranea (E. Müll.) M.E. Barr, which are saprobic on
the dead leaves of gymnosperms and angiosperms, in North
America and Europe (Barr 1972). Subsequently, a fourth species was introduced, viz. E. andromedae (Auersw.) M.E. Barr
(based on Wettsteinina andromedae) by Barr (1987a).
Extrawettsteinina is characterized by superficial, conical
ascomata, containing a few saccate bitunicate asci and ellipsoidal, obovate-clavate, septate, smooth and hyaline ascospores which turn dull brown at maturity (Barr 1972). The
diagnostic character of Extrawettsteinina is its conic
ascomata, which are superficial on the substrate, and radiating
arrangement of wall cells, which makes it distinguishable
from comparable genera such as Stomatogene and
Wettsteinina. Morphologically, Extrawettsteinina is comparable with Kriegeriella. In particularly, E. pinastri could not be
distinguished from K. transiens or K. mirabilis. Thus,
K. transiens, including Extrawettsteinina pinastri, was treated
as synonyms of K. mirabilis, and was included in the section
of Kriegeriella (von Arx and Müller 1975; Barr 1975). The
other section of Kriegeriella, Extrawettsteinina, includes two
previous Kriegeriella species, i.e. K. minuta and
K. mediterranea. Barr (1987b) introduced a family,
Kriegeriellaceae (Dothideales) to accommodate Kriegeriella
and Extrawettsteinina. This proposal has rarely been
followed.
In this study we tentatively refer Extrawettsteinina in
Pleosporaceae following Zhang et al. (2012) and Lumbsch
and Huhndorf (2010). Currently no molecular data is available
for this genus therefore generic type needs to be collected,
isolated and sequenced to show the natural classification of
this genus.
Gibbago E.G. Simmons, Mycotaxon 27: 108 (1986)
Facesoffungi number: FoF 00522
Saprobic or pathogenic on leaves and wood. Sexual morph:
undetermined. Asexual morph: Conidiophores solitary or 2–
4 loosely fasciculate, erect, rarely distantly branched, simple
with a single apical conidiogenous locus, then often proliferating by means of a secondary conidiophore that arises immediately below the apical cell of the existing conidiophore,
septate, slightly pigmented. Conidia initially solitary, ellipsoid, beakless, pigmented, becoming transversely and longitudinal septate, with apical cells swelling slightly and producing secondary conidia similar to the initial ones (Simmons
1986).
Fungal Diversity
Fig. 15 Extrawettsteinina minuta (NY, 4278). a, b. Herbarium material.
c. Fruiting bodies on the host. d, e. Sections of ascomata. f. Cellular
pseudoparaphyses. g. Ascus with short pedicel. h-k. Ellipsoid, obovate-
clavate, 3-septate ascospores. Scale bars: d=80 μm, e=50 μm, f=10 μm,
g=30 μm, h–k=5 μm
Type species: Gibbago trianthemae E.G. Simmons,
Mycotaxon 27: 108 (1986), Fig. 16
Facesoffungi number: FoF 00523
on its similarities with Alternaria, Embellisia,
Ulocladium and Stemphylium and this was followed by
Wijayawardene et al. (2014). During our phylogenetic
analysis, the putative strains of Gibbago trianthemae
GT-VM and NFCCI 1886 forms a distinct clade basel
to Paradendryphiella and Pleospora clades with low
boostrap support. Therefore based on morphology
coupled with molecular data, we tentatively refer
Gibbago in Pleosporaceae, but this needs to be confirmed with further sequences of the monotypic genus
and type species G. trianthemae.
Notes: Gibbago was introduced by Simmons (1986)
and typified by G. trianthemae. The genus is
characterised by erect, simple with a single apical
conidiogenous locus, septate, slightly pigmented, having
solitary conidiophores with ellipsoid, beakless conidia
and producing secondary conidia. Gibbago was
assigned to Pleosporaceae by Simmons (1986) based
Fungal Diversity
Fig. 16 Gibbago trianthemae
(Redrawn from the holotype in
Simmons 1986, Fig. 1). Scale
bar: = 50 μm
Johnalcornia Y.P. Tan & R.G. Shivas, in Tan et al., Australas.
Pl. Path.: 10.1007/s13313-014-0315-6 (2014).
Facesoffungi number: FoF 00524
Type species: Johnalcornia aberrans (Alcorn) Y.P. Tan &
R.G. Shivas, in Tan et al., Australas. Pl. Path.: 10.1007/
s13313-014-0315-6 (2014), Fig. 17
≡ Bipolaris aberrans Alcorn, Mycotaxon 39: 364 (1990)
≡ Cochliobolus aberrans Alcorn, Mycotaxon 39: 362
(1990)
Facesoffungi number: FoF 00525
Parasitic on leaves. Sexual morph: Ascomata globose with a
short neck. Asci cylindrical to fusoid, straight or curved.
Ascospores hyaline, filiform, tightly coiled. Asexual morph:
Conidiophores solitary or in fascicles, simple or branched,
straight to flexuous, apically geniculate, cylindrical, smooth.
Conidiogenous cells integrated, cylindrical, proliferating
sympodially, smooth to verruculose, with thickened and conspicuous scars. Conidia solitary, straight to curved, smooth,
hilum inconspicuous, distoseptate, germinating from both polar cells, first conidial septum median or submedian, second
conidial septum delimiting the apical cell, third septum basal
(Description obtained from Tan et al. 2014).
Notes: Johnalcornia was introduced by Tan et al. (2014) to
accommodate Bipolaris aberrans and is monotypic. Combined gene analysis of EF-1α, GAPDH, ITS and LSU has
shown that J. aberrans forms a sister clade to the newly described genus Porocercospora. Johnalcornia differs from
Porocercospora in having distinctive conidia-like
Fungal Diversity
Fig. 17 Johnalcornia aberrans
(Redrawn from the holotype in
Tan et al. (2014), Fig. 2). Scale
bars: 10 μm
chlamydospores as well as comparatively thick-walled, geniculate conidiophores, with conidiogenous cells that have conspicuous scars. Johnalcornia further differs from related genera by forming the second conidial septum in the apical cell. In
our phylogenetic analysis we also obtain the similar results
where the type strain of J. aberrans forms a distinct clade
sister to Porocercospora thus we accept Johnalcornia as a
separate genus in the family Pleosporaceae.
Marielliottia Shoemaker, Can. J. Bot. 76(9): 1559 (1999)
[1998].
Facesoffungi number: FoF 00526
Pathogenic on cereals, grasses and seeds or saprobic on wood
and leaves. Sexual morph: undetermined. Asexual morph:
Conidiophores solitary to clustered, erect, septate, straight,
brown, closely geniculate from sympodial growth or distantly nodose, apex truncate or inflated, forming conidia
through a terminal pore, with flat, non-protruding scars,
smooth or sometimes slightly roughened. Conidiogenous
cells tretic, integrated, terminal, sympodially proliferating,
cylindrical or swollen, cicatrized or not cicatrized. One species additionally produces massive, subulate
macroconidiophores radiating from a pseudoparenchymatic
base. Conidia porogenous, first septum formed near base,
transversely 3-septate, exceptionally 2-septate when immature in one species and sometimes up to 5-septate in one
species, brown, straight, ovoid or obovoid; scar pore-like,
basal, dark brown, not protruding, set off by a hyaline zone,
atrium type (Alcorn 1983) or flat in a slightly papillate base.
Germ tubes usually two from the basal cell, rarely one from
the apex, but never from the central cells (Description obtained from Shoemaker 1998).
Type species: Marielliottia biseptata (Sacc. & Roum.) Shoemaker, Can. J. Bot. 76(9): 1560 (1999) [1998], Fig. 18
≡ Helminthosporium biseptatum Sacc. & Roum., Revue
Mycol. (Toulouse): 56 (1881).
≡ Brachysporium biseptatum (Sacc. & Roum.) Sacc.,
Sylloge Fungorum 4: 428 (1886).
≡ Drechslera biseptata (Sacc. & Roum.) M.J. Richardson
& E.M. Fraser, Transactions of the British Mycological Society 51 (1): 148 (1968).
≡ Pyrenophora biseptata (Sacc. & Roum.) Crous, CBS
Biodiversity Series 13: 257 (2013)
= Helminthosporium biforme E.W. Mason & S. Hughes,
Transactions of the British Mycological Society 30: 114
(1948).
= Drechslera biforme (E.W. Mason & S. Hughes) Subram.
& B.L. Jain (1966).
Facesoffungi number: FoF 00527
Saprobic on stems. Sexual morph: undetermined. Asexual
morph: Conidiophores two types. Macronematous
conidiophores from stroma, single to many in a cluster, erect
to divergent, long, broad at base, gradually narrowed towards
apex, closely septate at intervals of 10–18 mm, dark chocolate
brown; sporogenous area gnarled, frequently bent, with numerous dark brown scars, apex truncate, bearing an Bear^ of
conidia, 100–800×10–15 μm (at base), 9–11 μm wide near
middle, 6–8 μm wide near apex. Conidiogenous cells tretic,
integrated, terminal, sympodially proliferating, cylindrical,
cicatrized, stromata shallow ellipsoidal to depressed globose,
100–250×100–150 μm. Micronematous conidiophores arising from hyaline to pale yellow hyphae (in culture), slender at
base, sparingly septate, short, erect, sporogenous area gnarled,
densely scarred, dark chocolate brown, usually simple, rarely
Fungal Diversity
Fig. 18 Marielliottia biseptata
(Redrawn from Drechslera
biseptata in Ellis 1971, Fig 288.).
Scale bar: 100 μm
branched, 15–105×5– 8 μm. Conidiogenous cells tretic, integrated, terminal, sympodially proliferating, cylindrical,
cicatrized. Conidia obovoid, 3-septate, uppermost septum inconspicuous or lacking in some, rarely constricted at septa,
thick walled, light to dark olive brown except lighter basal
cell, narrowed to base, bearing a broad black nonprotruding
scar 3–4.5 μm wide, (21–)32–45(−49)×(9–)11–13(−17) μm.
Germination by one or two (three) oblique tubes near scar,
rarely one hypha from apex and very rarely through the scar
(Shoemaker 1998).
Notes: Marielliottia was introduced by Shoemaker (1998)
to accommodate three species formerly treated in Drechslera
namely D. biseptata (Sacc. & Roum.) M.J. Richardson &
E.M. Fraser, D. dematioidea (Bubák & Wróbl.) Scharif and
D. triseptata (Drechsler) Subram. & B.L. Jain. The genus is
typified with Marielliottia biseptata. The species differ from
Drechslera in having mostly three-septate, obovoid to ovoid
conidia that germinate primarily from the basal cell, occasionally from the apical cell, and not from the central cells (Shoemaker 1998). These fungi are occasional parasites of cereals
and grasses, are sometimes seed borne, and may infect nongrass plants (Shoemaker 1998).
Currently no sequence data is available for Marielliottia
species in GenBank. Therefore, fresh collections of the type
Fungal Diversity
species of the genus are needed so that molecular data can be
obtained to verify the natural taxonomic affinities of this genus. Based on the morphological similarities with Drechslera,
we tentatively classified Marielliottia in Pleosporaceae.
Neocamarosporium Crous & M.J. Wingf., Persoonia 32: 273
(2014).
Facesoffungi number: FoF 00528
Pathogenic or saprobic on dying leaves and stems. Sexual
morph: undetermined. Asexual morph: Conidiomata brown
to black, immersed, becoming erumpent, globose with papillate apex and central ostiole; wall of 3–6 layers of brown
textura angularis. Conidiophores reduced to conidiogenous
cells. Conidiogenous cells lining the inner layer of conidioma,
separate, hyaline, smooth, ampulliform; proliferating several
times percurrently near apex, or at the same level, giving rise
to prominent periclinal thickening. Conidia solitary, initially
hyaline, aseptate, thick-walled, developing a central septum
and then becoming muriformly septate, shape variable from
globose to obovoid to ellipsoid, golden brown, finely roughened, thick-walled (Description obtained from Crous et al.
2014).
Type species: Neocamarosporium goegapense Crous & M.J.
Wingf., Persoonia, Mol. Phyl. Evol. Fungi 32: 273 (2014),
Fig. 19
Facesoffungi number: FoF 0529
Pathogenic on dying leaves of Mesembryanthemum sp.
(Aizoaceae). Sexual morph: undetermined. Asexual morph:
Conidiomata 300 μm diam, brown to black, immersed, becoming erumpent, globose with papillate apex and central
ostiole. Conidiomata wall consists of 3–6 layers of brown
textura angularis. Conidiophores reduced to conidiogenous
cells. Conidiogenous cells 7–9×5–6 μm, lining the inner layer
Fig. 19 Neocamarosporium
goegapense (Redrawn from the
holotype of Neocamarosporium
goegapense in Crous et al. 2014).
Scale bars: 10 μm
of conidioma, separate, hyaline, smooth, ampulliform, proliferating several times percurrently near apex, or at the same
level, giving rise to prominent periclinal thickening. Conidia
(15–) 20–22(−24)×15–17(−19) μm, solitary, initially hyaline,
aseptate, thick-walled, developing a central septum and then
becoming muriformly septate, shape variable from globose to
obovoid to ellipsoid, golden brown, finely roughened, thickwalled. (Description obtained from Crous et al. 2014)
Notes: Neocamarosporium was introduced by Crous et al.
(2014) and typified with Neocamarosporium goegapense.
Neocamarosporium is morphologically similar to
Camarosporellum, though the latter appears to have holoblastic conidiogenesis. Based on a megablast search of NCBIs
GenBank nucleotide database of LSU and ITS sequences,
Crous et al. (2014) referred Neocamarosporium to
Pleosporaceae. Crous et al. (2014) also suggested that phylogenetically, Neocamarosporium is allied to a clade containing
taxa accommodated in Phoma, Chaetosphaeronema and
Pleospora, but morphologically quite distinct. In our phylogenetic analysis we observed that the type strain of
Neocamarosporium goegapense (CBS 138008) forms a robust clade sister to the Clathrospora and Decorospora clades.
Furthermore, putative strains of Phoma betae (CBS 109410),
Pleospora betae (IMI 156653) and Pleospora bjoerlingii
(ICMP10945 reside within the Neocamarosporium clade. de
Gruyter et al. (2013) reported Phoma betae (CBS 109410) as
the asexual state of Pleospora betae (IMI 156653), thus they
synonymised Phoma betae under Pleospora betae. Furthermore based on the morphological and DNA data they
synonymised Pleospora bjoerlingii under Pleospora betae.
Therefore in this study we introduce Neocamarosporium
betae to accommodate these species in genus
Neocamarosporium.
Neocamarosporium betae (Berl.) Ariyawansa & K. D.
Hyde, comb. nov.,
Fungal Diversity
Index Fungorum number: IF 550956
Basionym: Pyrenophora echinella var. betae Berl., Nuovo
Giorn.Bot. Ital. 20: 208. 1888.
≡ Pleospora betae (Berl.) Nevod., Grib. ross. Exs., No.
247. 1915.
= Pleospora betae Björl., Bot. Not. 1944: 218. 1944. (later
homonym), nom.
illeg.
≡ Pleospora bjoerlingii Byford, Trans. Brit. Mycol. Soc.
46: 614. 1963.
= 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.
During our study the putative strains of Pleospora
chenopodii (CBS 344.78), P. calvescens (CBS 246.79) and
P. halimiones (CBS 432.77) also forms a separate clade basel
to the type species of Neocamarosporium, N. goegapense. de
Gruyter et al. (2013) treated these species as separate species
but our phylogeny clearly show that these three species can be
treated a one species thus we propose Neocamarosporium
calvescens to accommodate Pleospora chenopodii,
P. calvescens and P. halimiones.
Neocamarosporium calvescens (Fr. ex Desm.) Ariyawansa &
K.D. Hyde, comb. nov.,
Index Fungorum number: IF 550957
Basionym: Sphaeria calvescens Fr. ex Desm., Ann. Sci.
Nat., Bot., 2e Sér., 19: 353. 1843.
≡ Pleospora calvescens (Fr. ex Desm.) Tul. & C. Tul.,
Selecta Fung. Carpol. 2: 266. 1863.
≡ Pyrenophora calvescens (Fr. ex Desm.) Sacc., Syll.
Fung. 2: 279. 1883.
≡ Chaetoplea calvescens (Fr. ex Desm.) Clem., Gen.
Fung., 2nd Ed.: 275. 1931.
≡ Leptosphaeria calvescens (Fr. ex Desm.) Crivelli, Ueber
Heterog. Ascomycet. Pleospora: 177. 1983.
= Phloeospora chenopodii Ellis & Kellerm., J. Mycol. 4:
26. 1888, [as BPhleospora^].
≡ Pleospora chenopodii (Ellis & Kellerm.) Gruyter & Redhead, Index Fung. 205: 1. 2014.
= Pleospora halimiones Gruyter & Verkley, Stud. Mycol.
75: 25. 2012.
Paradendryphiella Woudenberg & Crous, Stud. Mycol.
75(1): 207 (2013)
Facesoffungi number: FoF 00530
Saprobic or parasitic in terrestrial and marine habitats. Sexual
morph: undetermined. Asexual morph: Conidiophores
simple or branched, septate or not, straight or flexuous, often
nodose with conspicuous, subhyaline, brown pigmentation at
the apical region; at times reduced to conidiogenous cells.
Conidiogenous cells terminal or lateral, with denticles aggregated at apex, with prominent conidial scars, thickened but not
darkened; sometimes proliferating with a new head or a short,
inconspicuous sympodial rachis. Conidia produced
holoblastically, on narrow denticle, smooth, cylindrical to
obclavate, straight or slightly flexuous, 1–7 transverse septa,
pale to medium brown, often with dark septa (often constricted), and a darkened zone of pigmentation at the apex, and at
the hilum, which is thickened, and somewhat protruding, with
a minute marginal frill (Description obtained from
Woudenberg et al. 2013).
Type species: Paradendryphiella salina (G.K. Sutherl.)
Woudenberg & Crous, Stud. Mycol. 75(1): 207 (2013),
Fig. 20
≡ Cercospora salina G.K. Sutherl., New Phytol. 15: 43
(1916).
≡ Dendryphiella salina (G.K. Sutherl.) Pugh & Nicot,
Transactions of the British Mycological Society 47 (2): 266
(1964).
≡ Scolecobasidium salinum (G.K. Sutherl.) M.B. Ellis,
More dematiaceous Hyphomycetes: 192 (1976).
Facesoffungi number: FoF 0531
Saprobic or parasitic in marine habitats. Sexual morph: undetermined. Asexual morph: Conidiophores 30–40 × 2–
4 μm, simple or branched, septate or not, straight or flexuous,
often nodose with conspicuous, brown pigmentation at the
apical region; at times reduced to conidiogenous cells.
Conidiogenous cells terminal or lateral, with denticles aggregated at apex, with prominent conidial scars, thickened but not
darkened; sometimes proliferating with a new head or a short,
inconspicuous sympodial rachis. Conidia 16–65×5–9 μm,
produced holoblastically, on narrow denticle, smooth-walled,
cylindrical to obclavate, straight or slightly flexuous, 1–7
transverse septa, pale to medium brown, often with dark septa
(often constricted), and a darkened zone of pigmentation at the
apex, and at the hilum, which is thickened, and somewhat
protruding, with a minute marginal frill (Description of
Scolecobasidium salinum obtained from Ellis 1976).
Notes: Paradendryphiella was introduced by Woudenberg
et al. (2013) to accommodate two species Dendryphiella
arenariae and D. salina (= Cercospora salina).
Paradendryphiella is characterised by simple or branched conidiophores with a new head or a short, inconspicuous sympodial rachis conidiogenous cells and holoblastically produced cylindrical to obclavate, straight or slightly flexuous,
1–7 transverse septa, pale to medium brown conidia. The
phylogenetic trees show that Paradendryphiella nested within
the family Pleosporaceae and clustered sister to Pleospora
Fungal Diversity
Fig. 20 Paradendryphiella
salina (Redrawn from
Scolecobasidium salinum in Ellis
1976, Fig 138c.). Scale bar: 5 μm
senso stricto clade in Jones et al. (2008), Woudenberg et al.
(2013) and as well as in this study. Therefore we accept
Paradendryphiella as separate genus in the family
Pleosporaceae.
Platysporoides (Wehm.) Shoemaker & C.E. Babc., Can. J.
Bot. 70(8): 1648 (1992).
Facesoffungi number: FoF 00532
Saprobic in terrestrial habitats. Sexual morph: Ascomata
small, scattered, immersed, semi-immersed to nearly superficial, globose, subglobose, black, smooth; apex with a protruding papilla and pore-like ostiole, without periphyses. Peridium
thin, composed of a few layers of textura angularis.
Hamathecium of numerous, cellular pseudoparaphyses, anastomosing above the asci, septate. Asci bitunicate, fissitunicate,
cylindrical to cylindro-clavate, with a short, furcate pedicel
and minute ocular chamber. Ascospores broadly ellipsoid, reddish brown, muriform. Asexual morph: undetermined
Type species: Platysporoides chartarum (Fuckel) Shoemaker
& C.E. Babc., Can. J. Bot. 70(8): 1650 (1992), Fig. 21
Basionym: Pleospora chartarum Fuckel, Jb. nassau. Ver.
Naturk. 23–24: 133 (1870) [1869–70]
≡ Clathrospora chartarum (Fuckel) O.E. Erikss., Arkiv før
Botanik 6 (4–5): 352 (1967)
Facesoffungi number: FoF 00533
Saprobic in terrestrial habitats. Sexual morph: Ascomata
150–230 μm × 180–260 μm, scattered, immersed, semiimmersed to rarely superficial, globose, subglobose, black,
smooth; apex with a protruding papilla, ostiolate. Ostiole comprises apex with a protruding papilla and without periphyses.
Peridium 8–22 μm wide, composed of 2–4 layers of brown
cells of textura angularis, cells 5–9 μm diam., cell wall 1–
2.5 μm thick. Hamathecium of 2–3 μm broad, long cellular
pseudoparaphyses, anastomosing above the asci, septate. Asci
110–140×12.5–16.5 μm (x=121:5×14:7 mm, n=10), (6-)8spored, bitunicate, fissitunicate, cylindrical to cylindro-clavate, with a short, furcate pedicel, 8–17 μm long,and minute
ocular chamber. Ascospores 20–26× 8–11 μm (x=23:7 ×
9 μm, n=10), obliquely uniseriate and partially overlapping,
flattened, broadly ellipsoid in front view, reddish brown,
muriform, 3 transverse septa, one longitudinal septum in each
central cell, 1 oblique septum in each end cell, constricted at
all septa, granulate, with a 2–3 μm wide mucilaginous sheath.
Asexual morph: undetermined.
Material examined: GERMANY, Rhineland-Palatinate, on
dead leaves, 1894, Herbier Barby-Boissier (HUH 00290381,
holotype).
Notes: Platysporoides was introduced as a subgenus of
Pleospora by Wehmeyer (1961) and was typified by
Pleospora chartarum. Shoemaker and Babcock (1992) raised
Platysporoides to generic rank and placed it in the
Pleosporaceae based on its Bapplanodictyospore^ and Bterete
pored beak of the ascomata^. Currently, eleven species are
included in this genus (Shoemaker and Babcock 1992). Another comparable pleosporalean family is Diademaceae,
which is distinguished from Platysporoides by its ascoma
opening as Ban intraepidermal discoid lid^ (Shoemaker and
Babcock 1992). Aigialus grandis is another pleosporalean
fungus with flattened and muriform ascospores as well as
papilla and ostioles, which belongs to Aigialaceae, a phylogenetically well supported marine family (Suetrong et al.
2009). Thus, it is highly likely that flattened and muriform
ascospores have evolved on separate occasions and not taxonomically informative.
Currently no molecular data is available for this genus and
therefore it is essential to epitypify the generic type with fresh
collections. Therefore molecular data can be used to resolve
the correct natural classification of Platysporoides. We tentatively place Platysporoides in Pleosporaceae based on its
Fungal Diversity
Fungal Diversity
Platysporoides chartarum (holotype). a, b. Specimen and
herbarium material. c. Ascomata on substrate. d. Section through
ascoma. e. Upper walled cells of ascoma. f. Peridium. g. Ascus when
immature in Melzer’s reagent. h. Ascus when immature in Cotton Blue
reagent. i. Ascus at maturity. j. Hamathecium in Cotton Blue reagent. k.
Ocular chamber in Melzer’s reagent. l. Ascospores at maturity. m, n.
Ascospore when immature with Sheath. o. Ascospore when immature
in Cotton Blue reagent. p. Ascospore at maturity with sheath in India
Ink. Scale bars: d=40 μm, e–f=10 μm, g–i=100 μm, j=10 μm, k–p=
5 μm
Fig. 21
similarities with the other genera of this family, such as, in
having globose to subglobose ascomata with pore-like ostiole,
a peridium composed of a few cell layers of textura angularis,
cellular pseudoparaphyses, fissitunicate, cylindrical to
cylindro-clavate asci with broadly ellipsoid, reddish brown,
muriform ascospores.
Pleospora Rabenh. ex Ces. & De Not., Comm. Soc. crittog.
Ital. 1(4): 217 (1863)
= Stemphylium Wallr., Fl. crypt. Germ. (Norimbergae) 2:
300 (1833)
= Cleistotheca Zukal, Österreichische Botanische
Zeitschrift 53 (5): 163 (1893)
= Cleistothecopsis Stevens & True, Illinois Agric. Exper.
Stat. Bull. Nr. 220: 507 (1919).
Facesoffungi number: FoF 00534
Habitat terrestrial or aquatic, saprobic or parasitic on stems
and leaves. Sexual morph: Ascomata small to medium-sized,
immersed, erumpent to superficial, base not easy to remove
from the substrate, broadly to narrowly oblong and flattened,
ostiolate. Ostiole papillate, black, smooth, ostiolar canal filled
with hyaline cells. Peridium thin, usually with two layers,
thick at the sides and thinner at the base, outer layer composed
of heavily pigmented, thick-walled cells of textura angularis,
inner layer composed of hyaline thin-walled cells of textura
angularis, coriaceous. Hamathecium of cellular, hyaline, septate, broad, dense pseudoparaphyses. Asci 8-spored,
bitunicate, fissitunicate, cylindrical to clavate, with furcate
pedicel and minute ocular chamber. Ascospores uniseriate or
partially overlapping, mostly ellipsoidal, muriform, brown or
pale brown, with or without sheath. Asexual morph:
Stemphylium. Mycelium immersed or nearly superficial,
brown. Stroma sometimes present. Conidiophores
macronematous, mononematous, scattered or caespitose, unbranched or rarely loosely branched, straight or flexuous, usually nodose with a number of vesicular swellings, pale to mid
brown or olivaceous brown, smooth or in part verruculose.
Conidiogenous cells monoblastic, integrated, terminal,
percurrent, at first clavate or subsphearical with the wall at
the apex thin. Conidia solitary, dry, acrogenous, oblong,
rounded at the ends, ellipsoidal, obclavate or subsphearical,
pale to mid dark or olivaceous brown, smooth, verrucose or
echinulate, muriform, often constricted at one or more of the
septa, cicatrized at the base. Colonies effuse, grey, brown,
olivaceous brown or black, velvety or cottony (Ellis 1971).
Type species: Pleospora herbarum (Pers.) Rabenh.,
Klotzschii Herb. Viv. Mycol. 2: no. 547 (1854), Fig. 22
≡ Sphaeria herbarum Pers., Syn. meth. fung. (Göttingen)
1: 78 (1801)
Facesoffungi number: FoF 00535
Habitat terrestrial, marine, saprobic or parasitic on stems and
leaves. Sexual morph: Ascomata 110–240×250–460 μm
small to medium-sized, immersed, erumpent to superficial,
base not easy to remove from the substrate, broadly to narrowly oblong and flattened, ostiolate. Ostiole papillate, black,
smooth, ostiolar canal filled with hyaline cells. Peridium 30–
50 μm (x=42 μm, n=10) thin, usually with two layers, thick
at the sides and thinner at the base, outer layer of heavily
pigmented thick-walled cells of textura angularis, inner layer
composed of hyaline thin-walled cells of textura angularis,
coriaceous. Hamathecium of 2–3 μm (x=2.5 μm, n=10) cellular, hyaline, septate, broad, dense pseudoparaphyses. Asci
100–225 × 25–30 μm (x= 142 × 28 μm, n = 20), 8-spored,
bitunicate, fissitunicate, cylindrical to clavate, with furcate
pedicel and minute ocular chamber. Ascospores 25–40×12–
15 μm (x=33×14 μm, n=40), uniseriate or partially overlapping, mostly ellipsoidal, muriform, brown or pale brown, with
or without sheath. Asexual morph: Stemphylium.
Conidiophores 4–6 μm (x=5 μm, n=10) macronematous,
mononematous, scattered or caespitose, unbranched or rarely
loosely branched, straight or flexuous, usually nodose with a
number of vesicular swellings, pale to mid brown or olivaceous brown, smooth or in part verruculose. Conidiogenous
cells monoblastic, integrated, terminal, percurrent, at first clavate or subsphearical with the wall at the apex thin. Conidia
16–22 × 11–18 μm (x= 21 × 16 μm, n = 20) solitary, dry,
acrogenous, oblong, rounded at the ends, ellipsoidal,
obclavate or subspherical, pale to mid dark or olivaceousbrown, smooth, verrucose or echinulate, muriform, often constricted at one or more of the septa, cicatrized at the base.
Colonies effuse, grey, brown, olivaceous-brown or black, velvety or cottony.
Material examined: ITALY, Forlì-Cesena, Montevescovo,
on dead stem of Brassica nigra (Brassicaceae), 01 January
2012, E. Camporesi IT 82 (MFLU 12–2216) – living culture
(MFLUCC 14–0261); ITALY, Tessello - Cesena, on dead
stem of Cirsium sp (Asteraceae), 16 December 2013, E.
Camporesi IT 956 (MFLU 14–0762) – living culture
(MFLUCC 13–0344); ITALY, Massera - Predappio, on dead
fruits of Lunaria rediviva (Brassicaceae), 16 December 2013,
E. Camporesi IT 958 (MFLU 14–0763) – living culture
(MFLUCC 13–0266, ICMP)
Fungal Diversity
Fig. 22 Pleospora herbarum (MFLU12-2216). a. Ascomata on host
substrate. b. Close up of ascoma .c . Section of ascoma. d. Close up of
the peridium. e. Cellular, hyaline, septate, broad pseudoparaphyses. f–h.
Asci with short, broad pedicel bearing 8 ascospores i–l. Mature and
immature ascospores with mucilaginous sheath. Stemphylium sp. m–p.
Conidiophores with pale to mid dark or olivaceous brown, smooth
conidia. Scale bars: c=100 μm, d=50 μm, e=20 μm, f–g=60 μm, i–
k=10 μm, l–o=10 μm
Notes: Pleospora was originally described by Rabenhorst
(1857) and is typified by Pleospora herbarum (Pers.)
Rabenh., (Kirk et al. 2008). Initially the genus was included in Sphaeriales and later Pseudosphaeriales and
Pleosporales, respectively (Wehmeyer 1961). All species
belonging to Pleospora have muriform ascospores
(Wehmeyer 1961, 1975). Pseudoparaphyses arrangement
(downward growing) within the ascomata of BPleospora-
type^ development (Luttrell 1951) is considered to be the
main feature for the genus. Various authors had included
and excluded different species in Pleospora at various
times. Barr (1981) placed Curreya in Pleospora however,
von Arx and van der Aa (1983) treated it as separate
genus, because of its Coniothyrium asexual morph. Petrak
(1952) transferred Graphyllium to Pleospora, and noted
that the elongate ascomata and closely grouped rows of
Fungal Diversity
small ascomata are not sufficient to recognize the genus.
Barr (1987b, 1990b) supported this proposal. Due to the
heterogenous nature of Pleospora several subgenera have
been included. i.e. Teichosporoides contains species of
Pleospora with immersed ascomata, and Pleosphaeria
with superficial and setose ascomata (Wehmeyer 1961).
Pleospora species have a wide host range, especially on
monocotyledons as well as dicotyledonous plants
(Wehmeyer 1975).
There are numerous species of Dothideomycetes that have
muriform ascospores and have at one time or another been
placed in Pleospora (Wehmeyer 1961). Pleospora is however,
rather distinctive and should be confined to species with characters that are similar to Pleospora herbarum. The ascomata
in P. herbarum are immersed and usually become erumpent,
the peridium is unusual in having three strata, a thin inner
layer of thin-walled, hyaline to light brown flattened cells, a
relatively wide central layer of thin-walled, hyaline to light
brown angular cells, and an outer very thin black amorphous
cells, which gives the blackened colour to the ascomata. The
asci are broadly clavate and have a distinctive, squarish, wide,
ocular chamber and ascospores are muriform with at least
three longitudinal septa per transverse row, and surrounded
by a mucilaginous sheath. The asexual morph is Stemphyllium
(Zhang et al. 2012; Hyde et al. 2013). For this reason several
Pleospora-like species are now transferred to other genera
(e.g. Curreya).
Based on our phylogenetic assessment we proposed to
confine Pleospora to one well-supported clade based on the
type strains of P. herbarum (CBS 191.86) along with the
other alternative strains of P. herbarum used in this study
(P. herbarum MFLUCC 13–0344, P. herbarum MFLUCC
13–0266 and P. herbarum (MFLUCC 14–0261), sister to
Paradendryphiella clade applying GCPSR (Taylor et al.
2000; Dettman et al. 2003). By implementing this, we treat
some putative strains, such as P. tomatonis (CBS109844),
and Pleospora sedicola (CBS109843) as alternative strains
of P. herbarum. Furthermore we treated the putative strains
of Pleospora papaveracea (CBS432.50), Pleospora
herbarum (DAOM195299), Pleospora tarda (CBS
714.68) and Pleospora halophila (CBS410.73) as separate
species which clustered outside the P. herbarum sensu
stricto.
Wijayawardene et al. (2014) proposed to conserve
Stemphyllium over Pleospora because Stemphyllium has priority as it is an older name, is better established in the literature
than Pleospora (more than double the number of hits on Google Scholar), and is a well-known genus to the plant pathology
community, and is well circumscribed on a molecular basis
(Lawrence et al. 2012). In this study the first author prefers to
keep Pleospora because the entire family Pleosporaceae, as
well as the order Pleosporales, is based on the Pleospora-type
of centrum development.
Porocercospora Amaradasa et al., Mycologia 106(1): 81
(2014)
Facesoffungi number: FoF 00536
Type species: Porocercospora seminalis (Ellis & Everh.)
Amaradasa et al., Mycologia 106(1): 81 (2013), Fig. 23
≡ Cercospora seminalis Ellis & Everh., J. Mycol. 4(1): 4
(1888)
≡ Sporidesmium seminale (Ellis & Everh.) U. Braun,
Cryptogamie Mycologie 20 (3): 175 (1999).
Facesoffungi number: FoF 588.
Parasitic on grasses. Sexual morph: undetermined. Asexual
morph: Conidiophores 450–500 × 5–8 μm, intermingled
among hyphae, subcylindrical, medium brown, smooth to
finely verruculose, branched above, thin-walled, septate.
Conidiogenous cells 20–30×5–7 μm, subcylindrical, medium
brown, smooth to finely verruculose, apex rounded,
monotretic, with a central pore, indistinct, not darkened or
thickened. Conidia 60–90×7–8 μm solitary, medium brown,
thick-walled, finely verruculose, obclavate to cylindroobclavate, with short conidia obovoid to subcylindrical, transversely multi-distoseptate; apex subobtuse, base obconically
truncate, with hila having a distinct central brown pore, thickened and darkened (Description obtained from Amaradasa
et al. 2014).
Notes: Porocercospora was introduced by Amaradasa et al.
(2014) to accommodate Cercospora seminalis which causes
false smut in buffalograss (Buchloe¨ dactyloides). The genus
is characterised by densely aggregated and repeatedly
branched conidiophores arising from a brown stroma,
monotretic conidiogenous cells with inconspicuous loci, and
scolecosporous conidia with distosepta, and thickened, darkened hila. DNA sequence data in the present study as well as
in the previous studies (Amaradasa et al. 2014) indicated that
Porocercospora seminalis is phylogenetically close to but distinct from the genera Bipolaris and Curvularia. In our phylogeny the type strain of Porocercospora seminalis (CPC
21332) forms a distinct clade sister to Johnalcornia. Therefore
we accept the placement of Porocercospora as separate genus
in Pleosporaceae.
Pseudoyuconia Lar.N. Vassiljeva, Nov. sist. Niz. Rast. 20: 71
(1983).
= Barrella Ahn & Shearer, Canadian Journal of Botany 73
(4): 578 (1995).
Facesoffungi number: FoF 00537
Saprobic on dead wood. Sexual morph: Ascomata semiimmersed to superficial, globose to subglobose. Ostiole central when mature, ostiolar canal filled with periphyses.
Peridium comprising of thick-walled cell, forming a textura
angularis. Hamathecium comprising hyaline, cellular, septate
Fungal Diversity
Fig. 23 Porocercospora
seminalis (Redrawn from
Amaradasa et al. 2014, Fig. 2.).
Scale bars: 5 μm
pseudoparaphyses. Asci 8-spored, bitunicate, broadly cylindrical or clavate to obovoid, with a minute pedicel, apically
rounded end, with an ocular chamber. Ascospores broadly
ellipsoidal, pale brown to light brown, 2-septate. Asexual
morph: undetermined
Type species: Pseudoyuconia thalictri (G. Winter) Lar.N.
Vassiljeva [as ‘thalicti’], Nov. sist. Niz. Rast. 20: 71 (1983), Fig. 24
≡ Leptosphaeria thalictri G. Winter, Hedwigia 10: 40
(1872)
≡ Metasphaeria thalictri (G. Winter) Sacc., Sylloge
Fungorum 2: 156 (1883)
≡ Scleropleella thalictri (G. Winter) Höhn., Annales
Mycologici 18 (1–3): 76 (1920)
≡ Buergenerula thalictri (G. Winter) E. Müll., Sydowia 4
(1–6): 307 (1950)
Fungal Diversity
Fig. 24 Pseudoyuconia thalictri (VLA, P/187) a. Fungus on host substrate. b. Section through ascoma. c. Close up of the peridium. d.
Pseudoparaphyses. e, f. Asci with eight ascospores. g, j. Ascospores with two septa. Scale bars: b=100 μm, c, g–j=10 μm, d=5 μm, e–f=50 μm
Fungal Diversity
≡ Pseudoyuconia thalicti (G. Winter) Lar.N. Vassiljeva
(1983)
≡ Barrella thalictri (G. Winter) Ahn & Shearer, Can. J. Bot.
73(4): 578 (1995)
Facesoffungi number: FoF 00538
Saprobic on dead wood. Sexual morph: Ascomata 137–
186×221–137 μm semi-immersed to superficial, globose to
subglobose, black. Ostiole central present when mature,
ostiolar canal filled with periphyses. Peridium 19–25 μm
wide, 3–4 layered, comprising of thick-walled cell, forming
a textura angularis. Hamathecium 2–3 μm wide, comprising
hyaline, cellular, septate pseudoparaphyses anastomosing
above the asci. Asci 80–119×17–23 μm (x=97×20 μm, n=
20), 8-spored, bitunicate, broadly cylindrical or clavate to obovoid, with a minute pedicel, apically rounded end, with an
ocular chamber. Ascospores 21–25×9–12 μm (x=23×11 μm,
n=20), broadly ellipsoidal, pale brown to light brown, 2-septate, slightly constricted at each septum. Asexual morph: undetermined.
Material examined: RUSSIA, Magadan region, Magadan
vicinity, Snow Valley, 8 July 1974, on Thalictrum contortum
L. (Ranunculaceae), L. Vasilyeva (VLA P/187).
Notes: Pseudoyuconia, a monotypic genus, was introduced
by Vassiljeva (1983) to accommodate Leptosphaeria thalictri
G. Winter, and was typified by P. thalictri. Currently,
Pseudoyuconia is included in Pleosporaceae (Lumbsch and
Huhndorf 2010). No molecular data is available for this genus.
Therefore, fresh collections of the type species of the genus
are needed so that molecular data can be obtained to verify the
natural taxonomic affinities of this genus. Based on the similarities with the other genera of the family Pleosporaceae,
such as, in having globose to subglobose ascomata with central ostiole filled with periphyses, peridium composed cells of
textura angularis we tentatively place Pseudoyuconia in
Pleosporaceae.
In this study we could not locate the type material therefore
our understanding is based on the collection of Vasilyeva
(1983), which was observed by Vasilyeva (1983) when she
was introduced this genus.
Pyrenophora Fr., Summa veg. Scand., Section Post. (Stockholm): 397 (1849).
= Drechslera S. Ito, Proc. Imp. Acad. Japan 6: 355 (1930).
Facesoffungi number: FoF 00009
Saprobic or pathogenic on leaves and stems. Sexual morph:
Ascomata immersed, becoming erumpent to near superficial,
solitary or scattered, globose to subglobose, broadly or narrowly conical, smooth-walled, ostiolate. Ostiole papillate,
covered with brown to reddish-brown setae, which are darkened at the base. Peridium comprising 2–4 layers of brown,
thick-walled cells of textura angularis. Pseudoparaphyses not
observed. Asci 8-spored, bitunicate, fissitunicate, clavate to
sub-cylindrical, with a short, broad pedicel, with a distinct
ocular chamber surrounded by a large apical ring. Ascospores
2–3-seriate, muriform, constricted at the septum, smoothwalled, surrounded by a mucilaginous sheath. Asexual
morph: hyphomycetous Conidiophores macronematous,
mononematous, sometimes caespitose, straight or flexuous,
often geniculate, unbranched or in a few species loosely
branched, brown, smooth in most species. Conidiogenous
cells polytretic, integrated, terminal, frequently becoming intercalary, sympodial, cylindrical, cicatrized. Conidia solitary,
in certain species also sometimes catenate or forming secondary conidiophores which bear conidia, acropleurogenous, simple, straight or curved, clavate, cylindrical, rounded at the
ends, ellipsoidal, fusiform or obclavate, straw-coloured or pale
to dark brown or olivaceous brown, sometimes with cells
unequally coloured, the end cells then being paler than intermediate ones, mostly smooth, rarely verruculose,
pseudoseptate (description of asexual morph obtained from
Ellis 1971).
Type species: Pyrenophora phaeocomes (Rebent.) Fr., Summa veg. Scand., Section Post. (Stockholm): 397 (1849),
Fig. 25
Basionym: Sphaeria phaeocomes Rebent., Prodr. fl.
neomarch. (Berolini): 338 (1804)
≡ Ceuthospora phaeocomes (Rebent.) Fr., J. Taihoku Soc.
Agric.: 119 (1825)
≡ Pleospora phaeocomes (Rebent.) Fr., Summa veg.
Scand., Section Post. (Stockholm): 398 (1849).
Facesoffungi number: FoF 00010
Saprobic on dead leaves. Sexual morph: Ascomata 380–
450 × 370–430 μm (x = 395 × 380 μm, n = 10), solitary or
scattered, initially immersed, becoming erumpent to near superficial, globose to subglobose, broadly or narrowly conical,
coriaceous, smooth-walled, ostiolate. Ostiole usually broadly
papillate, central ostiolar canal filled with periphyses and covered with setae. Setae brown to reddish brown, darkened at the
base, septate and tapered towards the apex. Peridium 40–
70 μm (x=45 μm, n=20) wide, comprising two types of cells,
outer cells of 1–2 layers of heavily pigmented cells of textura
angularis, inner layer composed of small, light brown to hyaline cells of textura angularis. Pseudoparaphyses not observed. Asci 300–400×130–160 μm (x=345×140 μm, n=
20), 8-spored, bitunicate, fissitunicate, clavate to
subcylindrical, with a short, broad pedicel, thickened and
rounded at apex with a distinct ocular chamber surrounded
by a large, distinct, apical ring. Ascospores 78–96× 27–
34 μm (x = 88 × 30 μm, n = 40), biseriate to overlapping
triseriate, ellipsoidal with broadly rounded ends, hyaline to
light brown when immature, becoming brown to chestnut
Fungal Diversity
Fig. 25 Pyrenophora phaeocomes (neotype) a. Fungus on host
substrate. b. Ascomata on substrate. c. Section through ascoma. d.
Brown setae. e. Peridium. f, g. Asci with a distinct ocular chamber
surrounded by a large, distinct, apical ring. h–j. Immature and mature
ascospores. Scale bars: c=100 μm, d–e=50 μm, f–g=40 μm, h–j=
10 μm
brown when mature, muriform with 5–6 transverse septa, and
single longitudinal septa in one or all cells, constricted at the
septa, smooth-walled, relatively thick-walled, with a 5–9 μm
thick mucilaginous sheath. Asexual morph of Drechslera
teres (Fig. 26); Conidiophores 150–200×7–11 μm, solitary
or in groups of 2–3, straight or flexuous, sometimes geniculate, often swollen at the base, pale to mid brown or olivaceous
brown. Conidia 70–160×16–23 μm, straight, cylindrical,
rounded at the ends, subhyaline to straw-coloured, smooth,
1–10 (4–6), pseudoseptate, frequently with constriction.
Notes: The asexual morph of Pyrenophora teres,
Drechslera teres is described and illustrated here in Fig. 25
because in original description of P. phaeocomes only
mentored that it has Drechslera- like asexual morph without
giving a species name (Sivanasen 1984).
Material examined: SWEDEN, on leaves of Anthoxanthum
(Poaceae), 7 August 1951, J. Ax. Nannfeldt (UPS 170980,
neotype).
Notes: Based on both morphology and molecular data,
Pyrenophora has been assigned to Pleosporaceae (Zhang
et al. 2012). The genus is characterized by immersed, erumpent
to nearly superficial ascomata, asci usually with a large apical
ring with a clear ocular chamber, lack of pseudoparaphyses and
muriform, terete ascospores (Sivanesan 1984). Currently 199
species of Pyrenophora are listed in Index Fungorum (2015).
Over 3000 hits are found in GenBank including the putative
strain of Pyrenophora phaeocomes (DAOM 222769), which is
the type species for this genus. In the present study the genus
Pyrenophora clusters in the suborder Pleosporineae of the family Pleosporaceae with a relatively high bootstrap support
Fungal Diversity
Fig. 26 Morphology of
Drechslera teres (Redrawn from
Ellis 1971, Fig 299.). Scale bars:
50 μm
(Fig. 1, 60 %). Further phylogenetic analysis shows that sexual
Pyrenophora morphs cluster with asexual Drechslera morphs,
i.e. Pyrenophora dictyoides (DAOM 75616) clusters with
Drechslera dictyoides (DAOM 63666). The putative strain of
Pyrenophora phaeocomes (DAOM 222769), which is the type
species of the genus, clusters with other Pyrenophora species.
As a genus can now only have one name, Ariyawansa et al.
(2014c) proposed to synonymise Drechslera under
Pyrenophora by giving the priority for the oldest name. This
was followed by Wijayawardene et al. (2014).
Excluded genera
Austropleospora R.G. Shivas & L. Morin, in Morin et al.,
Fungal Diversity 40(1): 70 (2010).
Facesoffungi number: FoF 00539
Type species: Austropleospora osteospermi R.G. Shivas & L.
Morin, Morin et al., Fungal Diversity 40(1): 70 (2010), Fig. 27
Facesoffungi number: FoF 00540
Parasitic on stem and leaves of Chrysanthemoides monilifera
(Asteraceae). Sexual morph: Ascomata 75–110 × 130–
200 μm (x=100×146 μm, n=10), subglobose, sometimes
slightly flattened, solitary or in groups, scattered, immersed
immediately below the stem epidermis, ostiole 60–90 μm
long, with a protruding neck. Peridium 8–18 μm (x=12 μm,
n=10) wide, composed of dark brown to black cells of textura
angularis. Hamathecium of 2–3 μm wide, dense, filamentous,
anastomosing, aseptate, hyaline pseudoparaphyses. Asci 75–
120 × 13–18 μm (x = 92 × 16 μm, n = 20), bitunicate,
fissitunicate, 6–8-spored, cylindrical to clavate, rounded at
the apex and minute ocular chamber with a short, broad, pedicel. Ascospores 16.5−21× 6–8.4 μm (x=18×7.7 μm, n=25),
biseriate to overlapping uniseriate, ellipsoidal, yellowish
brown, muriform, mostly with 3-transverse septa, 0–2 longitudinal septa, slightly constricted at median septum, not or
very slightly constricted at other septa, apex rounded to slightly tapered, base tapered to rounded, smooth. Asexual morph:
Conidiomata 75–110×100–130 μm (x=88×115 μm, n=10),
pycnidial, globose, superficial on stem, immersed in the host
tissue and becoming erumpent at maturity, globose, dark
brown in the erumpent part, with a single ostiole.
Conidiomata wall 9–16 μm wide, brown to reddish-brown,
thin-walled, comprising several layers with cells of textura
angularis. Conidiophores reduced to conidiogenous cells.
Conidiogenous cells 10–12×2.5−3.5 μm (x=11×2 μm, n=
15), inconspicuously annellidic, discrete, cylindrical. Conidia
14–18×4.8−6.5 μm (x=15.8×5.7 μm, n=25), cylindrical to
narrowly ellipsoidal, initially hyaline and aseptate, becoming
yellowish-brown at maturity, mostly transversely 1–3-septate,
ends rounded.
Material examined: AUSTRALIA, NSW Bellinger Head,
Kalang river mouth, on stem and leaves of Chrysanthemoides
monilifera ssp. rotundata (L.) Norlindh (Asteraceae). 5 March
2008, L. Morin (BRIP 52234, holotype).
Notes: Austropleospora was introduced by Morin et al.
2010 to accommodate A. osteospermi on Chrysanthemoides
monilifera ssp. rotundata (Asteraceae) and it is monotypic. In
the same study Shivas and Morin (in Morin et al. 2010) observed Hendersonia osteospermi Wakef. on the same host and
identified it as the asexual morph of A. osteospermi both in
culture and by DNA sequence analysis. Based on ITS
sequence analysis, Morin et al. (2010) placed Austropleospora
Fungal Diversity
Fig. 27 Austropleospora
osteospermi (holotype) a.
Herbarium material. b, c.
Ascomata on host surface. d.
Section through ascoma. e.
Section through ostiole. f.
Peridium. g. Pseudoparaphyses h.
Immature ascus. i. Mature
bitunicate asci. j. Apex of ascus.
k. Ascospores. k. Pycnidia on
host. l. Section of pycnidium. m.
Pycnidial wall. o. Conidiogenous
cells and developing conidia. n.
Brown 3-septate conidia. Scale
bars: d=50 μm, e–f=25 μm, g=
10 μm, h–i=25 μm, k=15 μm, l,
m=20 μm, n=5 μm
Fungal Diversity
under Pleosporales without assigning it to any family.
Austropleospora is classified in Dothideomycetes genera
incertae sedis in Index Fungorum (2015). After reexamineing the holotype of A. osteospermi, Thambugala
et al. (2014b) tentatively referred Austropleospora in
Pleosporaceae based on morphological similarities. During
our preliminary phylogenetic analysis (data not shown), we
observed that the type strain of A. osteospermi (BRIP 51628)
forms a distinct clade in family Didymosphaeriaceae. Therefore we exclude Austropleospora from Pleosporaceae and
refer it to Didymosphaeriaceae.
Dendryphion Wallr., Fl. crypt. Germ. (Norimbergae) 2: 300
(1833)
Dendryphiella Bubák & Ranoj., in Ranojevic, Annls
mycol. 12(4): 417 (1914)
Dwayamala Subram., J. Indian bot. Soc. 35: 475 (1956)
Entomyclium Wallr., Fl. crypt. Germ. (Norimbergae) 2: 189
(1833)
Facesoffungi number: FoF 00541
Type species: Dendryphion comosum Wallr., Fl. crypt.
Germ. (Norimbergae) 2: 300 (1833)
= Dendryphion curtum Berk. & Broome, Ann. Mag. nat.
Hist., Ser. 2 7: 176 (1851)
= Dendryphion curtum Berk. & Broome, Ann. Mag. nat.
Hist., Ser. 2 7: 176 (1851) var. curtum
= Dendryphion curtum var. ramosius Sacc., Syll. fung.
(Abellini) 4: 490 (1886)
Facesoffungi number: FoF 00542
Notes: Dendryphion was introduced by Wallroth (1833)
and is typified by Dendryphion comosum. The genus
Dendryphion is characterised by having apically branched
conidiophores with polytretic dark scars, and chains of
brown, septate (didymo or cheiro) conidia (Crous et al.
2014). The genus Brachycladium had been synonymised
with Dendryphion by Ellis (1971), was resurrected for the
asexual morph based on polyphyly within Dendryphion and
morphological distinction from type species, D. comosum
(Crous et al. 2014). The type species of Brachycladium,
B. penicillatum, resides in Alternaria sect. Crivellia in this
s t u d y a s w e l l a s i n Wo u d e n b e r g e t a l . ( 2 0 1 3 ) .
Brachycladium is therefore a synonymy of Alternaria instead of Dendryphion. Although there are more than 80
names recorded in Index Fungorum (2015), Seifert et al.
(2011) confined them to six species. Inderbitzin et al.
(2006) showed that Dendryphion was polyphyletic thus
the natural classification of the genus has become confused.
Currently, no sequence data is available in GenBank for the
generic type of Dendryphion. Recently, Crous et al. (2014)
introduced a new species of Dendryphion, D. europaeum
with molecular and morphological data, and placed it species in the suborder Pleosporineae, family Pleosporaceae.
Considering the morphological key provided by Siboe et al.
(1999), Crous et al. (2015) confirmed that D. europaeum
appears to be distinct from the currently recognised
Dendryphion species based on its conidial morphology.
During our preliminary analysis we observed that
D. europaeum forms a robust clade outside the suborder
Pleosporineae. Therefore we tentatively exclude
Dendryphion from Pleosporaceae and place it in
Pleosporales, genera incertae sedis. However, the placement of Dendryphion in Pleosporales can only be confirmed by phylogenetic analysis, including sequencing the
generic type of Dendryphion (D. europaeum).
Edenia M.C. González et al., in González et al., Mycotaxon
101: 254 (2007)
Type species: Edenia gomezpompae M.C. González et al., in
González et al., Mycotaxon 101: 254 (2007)
Facesoffungi number: FoF 00543
Notes: Edenia was introduced by González et al. (2007)
and is typified by Edenia gomezpompae. This endophytic
fungus was isolated from living leaves of Callicarpa acuminta
Kunth (González et al. 2007). González et al. (2007) placed
Edenia in Pleosporaceae based on the ITS sequence data. In
our phylogeny we observed that the type strain of Edenia
gomezpompae (C1c) along with putative strains JLCC
34533, CBS 124106 and 11G048 form a robust clade in family Phaeosphaeriaceae outside Pleosporaceae. Therefore we
exclude Edenia from Pleosporaceae and placed it in the
Phaeosphaeriaceae.
Kriegeriella Höhn., Annls mycol. 16(1/2): 39 (1918).
Type species: Kriegeriella mirabilis Höhn., Annls mycol.
16(1/2): 39 (1918).
Facesoffungi number: FoF 00544
Notes: Kriegeriella was introduced by von Höhnel (1918b)
in order to accommodate K. mirabilis and K. transiens Höhn.
The genus Kriegeriella was typified with K. mirabilis and
referred to Microthyriaceae by Höhnel 1918b. It is
characterised by superficial, subglobose, black thyrothecialike ascomata, cellular pseudoparaphyses, bitunicate,
obpyriform asci bearing hyaline, multi-septate ascospores
which turn brown when mature. Subsequently, Kriegeriella
was assigned to the subfamily of Aulographiodeae
(Microthyriaceae) (Batista et al. 1959), Asterinaceae
(Hemisphaeriales) (Luttrell 1973) and Pseudosphaeriaceae
(Dothideales) (Barr 1975). Barr (1987b) introduced a family,
i.e. Kriegeriellaceae (Dothideales) to accommodate
Kriegeriella and Extrawettsteinina. This scheme is rarely
followed, and Kriegeriella is generally classified under
Pleosporaceae (Pleosporales) (Eriksson 2006; Lumbsch and
Huhndorf 2007, 2010).
Fungal Diversity
Zhang et al. (2012) excluded Kriegeriella from the family
Pleosporaceae and concluded that it is not a typical genus of
the family. Furthermore, Zhang et al. (2012) suggested that
Kriegeriella might belong to Microthyriaceae, as it would be
unusual in Pleosporaceae in having thyrothecium-like
ascomata and 5–6-septate ascospores. Therefore we exclude
Kriegeriella from Pleosporaceae, following the opinion of
Zhang et al. (2012).
Macrospora Fuckel, Jb. nassau. Ver. Naturk. 23–24: 139
(1870) [1869–70].
= Nimbya E.G. Simmons, Sydowia 41: 316 (1989)
Facesoffungi number: FoF 00545
Saprobic on stems or wood. Sexual morph: Ascomata immersed to nearly superficial, subglobose. Ostiole central, present when mature, ostiolar canal filled with periphyses.
Peridium comprising thick-walled cells, forming a textura
angularis. Hamathecium comprising hyaline, unbranched,
septate, pseudoparaphyses. Asci 8-spored, bitunicate, broadly
cylindrical or clavate to obovoid, with a knob-like pedicel at
the base, apically rounded, with an ocular chamber.
Ascospores broadly ellipsoidal, brown to dark brown, multi
septate. Asexual morph: undetermined.
Type species: Macrospora scirpicola (DC.) Fuckel, Jb. nassau. Ver. Naturk. 23–24: 139 (1870) [1869–70], Fig. 28
Basiyonym: Sphaeria scirpicola DC., in Lamarck & de
Candolle, Fl. franç., Edn 3 (Paris) 2: 300(1805)
≡ Clathrospora scirpicola (DC.) Höhn., Annls mycol.
18(1/3): 77 (1920)
≡ Macrospora scirpicola (DC.) Fuckel, Jb. nassau. Ver.
Naturk. 23–24: 139 (1870)
≡ Pleospora scirpicola (DC.) P. Karst., Bidr. Känn. Finl.
Nat. Folk 23: 72 (1873)
= Pleospora scirpicola var. scirpicola (DC.) P. Karst., Bidr.
Känn. Finl. Nat. Folk 23: 72 (1873)
≡ Pyrenophora scirpicola (DC.) E. Müll., Sydowia 5(3–6):
256 (1951)
= Sphaeria culmorum Wallr., Fl. crypt. Germ.
(Norimbergae) 2: 770 (1833)
Facesoffungi number: FoF 00546
Saprobic on dead stems. Sexual morph: Ascomata 83–101×
72–79 μm, immersed in host tissues, subglobose, brown to
dark brown, ostiolate. Ostiole central when mature, ostiolar
canal 31–36 μm wide at the base, with periphyses. Peridium
9–25 μm wide, comprising 2–3 layers of thick-walled cells,
forming a textura angularis, brown to dark brown at outside
and hyaline at the inner layer. Hamathecium of 2–4 μm wide,
thick-walled, cellular, hyaline, unbranched, septate,
pseudoparaphyses. Asci (26–) 64–88×9–13 μm, 8-spored,
bitunicate, broadly cylindrical or clavate to obovoid, with a
knob-like pedicel at the base, apically rounded, with an ocular
chamber. Ascospores 16–28×8–12 μm., 2–4 overlapping seriate, broadly ellipsoidal, ends rounded, straight or curved,
brown to dark brown, 3-septate, constricted at the central cell,
dark brown at the septum, second cell from above widest.
Asexual morph: undetermined.
Material examined: FRANCE, collector unspecified,
Exsiccati title: Plantes Cryptogames De France, Exsiccati
Number: 1778 (NY 0098598, holotype).
Notes: Macrospora had been assigned to Diademaceae
based on its applanate and muriform ascospores with 1-row
of longitudinal septa, with a sheath, 2–3 μm wide and constricted at the first septum and ascospores that are paler and
larger than those of Comoclathris (Shoemaker and Babcock
1992). Macrospora was however, considered as a synonym of
Pyrenophora by Eriksson and Hawksworth (1991) which was
assigned to Pleosporaceae, and this proposal has been widely
followed (Eriksson 2006; Lumbsch and Huhndorf 2010).
Ariyawansa et al. (2014a) suggested that Macrospora
scirpicola is neither typical of Diademaceae nor
Pleosporaceae. Macrospora is compatible with Thyridaria
in having immersed, subglobose ascomata with 2–3 layers,
of thick-walled cells of textura angularis and cylindrical asci
bearing phragmospores, but differs in having cellular
pseudoparaphyses and asci with a with a knob-like or furcate
pedicel.
Currently no molecular data is available for any
Macrospora species in GenBank. Therefore based on
its similarities with Thyridaria, we tentatively place
Macrospora in Thyridariaceae, but fresh collections of
the type species of the genus are needed so that molecular data can be obtained to verify its natural taxonomic
affinities.
Monascostroma Höhn., Annls mycol. 16(1/2): 160 (1918).
Facesoffungi number: FoF 00547
Type species: Monascostroma innumerosum (Desm.) Höhn.
[as ‘innumerosa’], Annls mycol. 16(1/2): 160 (1918).
Basionym: Hendersonia innumerosa Desm., Annls Sci.
Nat., Bot., sér. 3 16: 10 [repr.] (1851)
≡ Stagonospora innumerosa (Desm.) Sacc., Sylloge
Fungorum 3: 451 (1884)
Facesoffungi number: FoF 00548
Notes: Monascostroma was introduced by Desmazières
(1851) to accommodate M. innumerosum (= Hendersonia
innumerosa). Eriksson and Hawksworth (1998) and Kodsueb
et al. (2006) included Monascostroma in Pleosporaceae,
while later researchers classified Monascostroma in
Didymellaceae (Zhang et al. 2009, 2012; Schoch et al. 2009;
Lumbsch and Huhndorf 2010).
Fungal Diversity
Fungal Diversity
The putative strain of Monascostroma innumerosum (CBS
345.50) forms a robust clade in family Didymellaceae (Zhang
et al. 2009, 2012; Schoch et al. 2009; Hyde et al. 2013). Thus
we also accept the genus Monascostroma in the
Didymellaceae following other published work (Zhang et al.
2009, 2012; Schoch et al. 2009; Hyde et al. 2013).
Zeuctomorpha Sivan et al., in Sivanesan, Bitunicate Ascomycetes and their Anamorphs (Vaduz): 572 (1984).
Facesoffungi number: FoF 00549
Type species: Zeuctomorpha arecae Sivan et al., in
Sivanesan, Bitunicate Ascomycetes and their Anamorphs (Vaduz): 572 (1984).
Facesoffungi number: FoF 00550
Notes: Zeuctomorpha, a monotypic genus typified by
Z. arecae, was formally treated as a member of the
Pleosporaceae by Sivanesan (1984), based on its superficial
setose ascomata, clavate, asci, ellipsoid and 1-septate
ascospores, and presence of pseudoparaphyses. Eriksson and
H a w k s w o r t h (1 9 9 8) c l as si fi e d Z e u c t o m o r p h a i n
Pleosporaceae and this was followed by Kirk et al. (2001),
Kodsueb et al. (2006) and Lumbsch and Huhndorf (2010).
Zhang et al. (2012) suggested that Zeuctomorpha was unusual amongst the Pleosporaceae with its hairy superficial
ascomata, few pseudoparaphyses, broadly clavate to
obclavate asci and 1-septate pigmented ascospores. Furthermore, Zhang et al. (2012) concluded that all of these morphological characters are most comparable with species of
Acantharia, which might be closely related to Venturiaceae,
but this needs to be confirmed by molecular data. Therefore,
based on the morphology we also recommend to exclude
Zeuctomorpha from Pleosporaceae without assigning it to
any family.
Concluding remarks
Molecular data coupled with morphology has provided the
basis for the modern classification of the kingdom fungi, but
have some restrictions in application (Hibbett et al. 2007;
Ariyawansa et al. 2014a; Boonmee et al. 2014; Hyde et al.
2014; Nilsson et al. 2014; Schoch et al. 2014; Thambugala
et al. 2014a). The most important and disconcerted problem is
that the phylogeny inferred from any gene may not divulge the
evolution history of the organism (Uilenberg et al. 2004;
Macrospora scirpicola (holotype) a. Herbarium material. b.
Appearance of ascomata on host. c. Neck with periphyses. d. Vertical
section through ascoma. e. Section of peridium. f. Pseudoparaphyses. g,
h. Immature asci. i–k. Mature asci. l. Immature ascospore. m–p. Mature
ascospores. Scale bars: b=500 μm, g–k=50 μm, d–e=30 μm, c=20 μm,
f=20 μm, l–p=20 μm
Fig. 28
Ariyawansa et al. 2014a). Therefore polyphasic taxonomy,
including genotypical and phenotypical characteristics are recommended in all future studies (Uilenberg et al. 2004;
Ariyawansa et al. 2014a, Udayanga et al. 2014a). The genome
also needs to be evaluated (Uilenberg et al. 2004; Ariyawansa
et al. 2014a). Another main problem is most early studies is
that they focused on techniques and used strains of fungi that
in most cases were not carefully referenced thus creating confusion as explained in Ariyawansa et al. (2014a). Most strains
in GenBank are named without attached voucher material and
it impractical to confirm their characters to ensure correct
identification.
There have only been a few molecular analysis of
Pleosporaceae as compared to morphological studies
(Kodsueb et al. 2006). Genera with bitunicate asci and brown
muriform ascospores viz Murispora, Pleospora, Pyrenophora
and Tremateia were generally classified under Pleosporaceae.
Molecular studies have shown that these particular morphological characters have evolved in different families such as
Aigialaceae, Amniculicolaceae, Didymellaceae,
Lophiostomataceae, Montagnulaceae and Sporormiaceae
(Zhang et al. 2012a, Hyde et al. 2013). Recently published
papers such as Zhang et al. (2012a), Woudenberg et al.
(2013) and Hyde et al. (2013), did not discuss details of the
family Pleosporaceae and related genera. Woudenberg et al.
(2013) was mainly focused on the genus Alternaria, while
Zhang et al., (2012a) gave a detail account of sexual morphs
only. Hyde et al. (2013) provided a key to the genera in
Pleosporaceae in work on Dothideomycetes.
Fourteen clades of Pleosporaceae have been identified by
our analysis based on ITS, LSU, SSU and RPB2 gene data,
which are treated as suitable for the delimitation of genera
(Figure 5.1). Although relationships among these clades may
be weakly supported and some may vary in detail, some tentative conclusions can be drawn. Several current taxonomic
hypotheses are supported by our molecular data, and this
makes it possible to propose some taxonomic hypotheses
about the relationships among the genera in Pleosporaceae.
By combining multi-locus DNA sequencing with detailed
morphological analyses, we were able to define and formally
propose two new species, four combinations and several sexual morphs among the taxa in the family Pleosporaceae. In
our study we accept 16 sexual or asexual genera and exclude
three genera from the family Pleosporaceae based on morphology coupled with molecular data.
In previous treatments Pleospora species were considered
as paraphyletic within the family Pleosporaceae (Kodsueb
et al. 2006, Inderbitzin et al. 2009). Several studies have confirmed this opinion and have led to the removal of Pleospora
species to other genera (Crivelli 1983, Inderbitzin et al. 2002,
2006; Leuchtmann 1984, Zhang et al. 2009b). In the present
study we were able to resolve the paraphyletic nature of the
genus by following Woudenberg et al. (2013) and
Fungal Diversity
Wijayawardene et al. (2014), where they have treated the genus based on the generic type Pleospora herbarum (CBS 191.
86T). Therefore in order to create a stable phylogenetic taxonomy, several putative strains of Pleospora which clustered
with the type strain of Pleospora herbarum (CBS 191.86)
were classified under P. herbarum sensu stricto (P. tomatonis
CBS109844 and P. sedicola CBS109843). The putative
strains of P. papaveracea CBS432.50, P. tarda CBS 714.68
and P. halophila CBS410.73 were treated under the genus
Pleospora as separate species but this needs to be clarified
with morphology because currently these strains are not attached to any herbarium material.
Some putative strains of Pleospora species other than species discussed in the above paragraph were assigned to different genera in Pleosporaceae based on their molecular data.
Pleospora fallens (Sacc.) Gruyter & Verkley was introduced
by de Gruyter et al. (2013) in order to accommodate Phoma
fallens Sacc., because during their study the strain of Phoma
fallens (CBS 284.70) clustered within the family
Pleosporaceae. Woudenberg et al. (2013) and Ariyawansa
et al. (2014a) showed that this strain forms a separate clade
in the family Pleosporaceae basel to the other genera of the
family. Importantly Pleospora fallens clustered outside the
Pleospora sensu stricto. Therefore here we propose to keep
the name Phoma fallens instead of Pleospora fallens. By taking this approach we were able to resolve the paraphyletic
nature of Pleospora within the family Pleosporaceae
In recent years, DNA based studies revealed multiple nonmonophyletic genera within the Alternaria complex, and
Alternaria species clades that do not always correlate to
species-groups based on morphological characteristics (Pryor
and Bigelow 2003; Inderbitzin et al. 2006; Pryor et al. 2009;
Runa et al. 2009; Wang et al. 2011; Lawrence et al. 2012;
Woudenberg et al. 2013). A separate phylogeny was prepared
to show the placement of the recently revised sections in
Alternaria (Figure 5.2). The well-supported node for the
Alternaria clade obtained in the present study, and the low
bootstrap support at the deeper nodes within the Alternaria
complex is similar to previous phylogenetic studies. (Pryor
and Bigelow 2003; Inderbitzin et al. 2006; Pryor et al. 2009;
Runa et al. 2009; Wang et al. 2011; Lawrence et al. 2012;
Woudenberg et al. 2013). We also observed the same result
as Woudenberg et al. (2013) where the Alternaria clade contains 24 internal clades and six monotypic lineages, the assemblage of which is recognised as Alternaria. Furthermore we
introduced one new species (A. sedi) and several collections of
the sexual morph of A. alternata from Italy are also introduced
in the study.
The explanation of Pleosporaceae in the current work is
sustained by a well-supported phylogenetic nodes in multiple
analyses coupled with morphology. We pursue the precedence
introduced by Lawrence et al. (2013) and Woudenberg et al.
(2013) to assign the taxonomic status of sections of Alternaria
for the different clades found. In here also we show the significance of having accurate DNA sequence data attached
with both correct taxonomic names and clearly annotated
specimen data. For end-users, this also results in a more stable
and understandable taxonomy and nomenclature. Furthermore we gave a well resolved back-bone tree for the family
Pleosporaceae, representing all the sexual and asexual genera,
which currently have DNA sequences.
Acknowledgments MFLU grant number 56101020032 is thanked for
supporting studies on Dothideomycetes. We are grateful to the Mushroom
Research Foundation, Chiang Rai, Thailand for supporting studies on
Dothideomycetes. Kevin D. Hyde thanks the Chinese Academy of Sciences, project number 2013T2S0030, for the award of Visiting Professorship for Senior International Scientists at Kunming Institute of Botany.
Jian-Chu Xu and Peter E Mortimer would like to thank Humidtropics, a
CGIAR Research Program that aims to develop new opportunities for
improved livelihoods in a sustainable environment, for partially funding
this work. H.A Ariyawansa and J.C. Kang are grateful to the agricultural
science and technology foundation of Guizhou province (Nos.
NY[2013]3042 ), the international collaboration plan of Guizhou province (No. G [2012]7006) and the innovation team construction for science
and technology of Guizhou province (No. [2012]4007) from the Science
and Technology Department of Guizhou province, China. Hiran
Ariyawansa is grateful to A.D Ariyawansa, D.M.K Ariyawansa and
Dhanuska Udayanga for their valuable suggestions. E.B. Gareth Jones
is supported by the Distinguished Scientist Fellowship Program (DSFP),
King Saud University, Saudi Arabia.
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