Fungal Diversity
DOI 10.1007/s13225-011-0117-x
Pleosporales
Ying Zhang & Pedro W. Crous & Conrad L. Schoch &
Kevin D. Hyde
Received: 31 March 2011 / Accepted: 13 June 2011
# The Author(s) 2011. This article is published with open access at Springerlink.com
Abstract One hundred and five generic types of Pleosporales
are described and illustrated. A brief introduction and detailed
history with short notes on morphology, molecular phylogeny
as well as a general conclusion of each genus are provided.
For those genera where the type or a representative specimen
is unavailable, a brief note is given. Altogether 174 genera of
Pleosporales are treated. Phaeotrichaceae as well as Kriegeriella, Zeuctomorpha and Muroia are excluded from
Pleosporales. Based on the multigene phylogenetic analysis,
the suborder Massarineae is emended to accommodate five
families, viz. Lentitheciaceae, Massarinaceae, Montagnulaceae, Morosphaeriaceae and Trematosphaeriaceae.
Y. Zhang
Division of Microbiology, School of Biological Sciences,
The University of Hong Kong,
Pokfulam Road,
Hong Kong, SAR, People’s Republic of China
P. W. Crous
CBS-KNAW Fungal Biodiversity Centre,
P.O. Box 85167, 3508 AD Utrecht, The Netherlands
C. L. Schoch
National Center for Biotechnology Information,
National Library of Medicine, National Institutes of Health,
45 Center Drive, MSC 6510,
Bethesda, MD 20892-6510, USA
K. D. Hyde
School of Science, Mae Fah Luang University,
Tasud, Muang,
Chiang Rai 57100, Thailand
K. D. Hyde (*)
Botany and Microbiology Department, College of Science,
King Saud University,
Riyadh 11442, Saudi Arabia
e-mail: kdhyde3@gmail.com
Keywords Generic type . Massarineae . Molecular
phylogeny . Morphology . Pleosporales . Taxonomy
Introduction
Historic overview of Pleosporales
Pleosporales is the largest order in the Dothideomycetes,
comprising a quarter of all dothideomycetous species (Kirk
et al. 2008). Species in this order occur in various habitats,
and can be epiphytes, endophytes or parasites of living
leaves or stems, hyperparasites on fungi or insects, lichenized, or are saprobes of dead plant stems, leaves or bark
(Kruys et al. 2006; Ramesh 2003).
The Pleosporaceae was introduced by Nitschke (1869),
and was assigned to Sphaeriales based on immersed
ascomata and presence of pseudoparaphyses (Ellis and
Everhart 1892; Lindau 1897; Wehmeyer 1975; Winter
1887). Taxa in this family were then assigned to Pseudosphaeriaceae (Theissen and Sydow 1918; Wehmeyer 1975).
Pseudosphaeriales, represented by Pseudosphaeriaceae,
was introduced by Theissen and Sydow (1918), and was
distinguished from Dothideales by its uniloculate, perithecioid ascostromata. Subsequently, the uni- or pluri-loculate
ascostromata was reported to be an invalid character to
separate members of Dothideomycetes into different orders
(Luttrell 1955). In addition, the familial type of Pseudosphaeriales together with its type genus, Pseudosphaeria, was
transferred to Dothideales, thus Pseudosphaeriales became a
synonym of Dothideales. The name “Pseudosphaeriales”
has been applied in different senses, thus Pleosporales (as an
invalid name due to the absence of a Latin diagnosis) was
proposed by Luttrell (1955) to replace the confusing name,
Pseudosphaeriales, which included seven families, i.e.
Fungal Diversity
Botryosphaeriaceae, Didymosphaeriaceae, Herpotrichiellaceae, Lophiostomataceae, Mesnieraceae, Pleosporaceae
and Venturiaceae. Müller and von Arx (1962) however,
reused Pseudosphaeriales with 12 families included, viz.
Capnodiaceae, Chaetothyriaceae, Dimeriaceae, Lophiostomataceae, Mesnieraceae, Micropeltaceae, Microthyriaceae,
Mycosphaerellaceae, Pleosporaceae, Sporormiaceae, Trichothyriaceae and Venturiaceae.
Familial circumscriptions of the Pleosporales were based
on characters of ascomata, morphology of asci and their
arrangement in locules, presence and type of hamathecium,
shape of papilla or ostioles, morphology of ascospores and
type of habitats (Luttrell 1973) (Table 1). Based on these
characters, Luttrell (1973) included eight families, i.e.
Botryosphaeriaceae, Dimeriaceae, Lophiostomataceae, Mesnieraceae, Mycoporaceae, Pleosporaceae, Sporormiaceae
and Venturiaceae in Pleosporales. In their review of bitunicate
ascomycetes, von Arx and Müller (1975) accepted only a
single order, Dothideales, with two suborders, i.e. Dothideineae (including Atichiales, Dothiorales, Hysteriales and
Myriangiales) and Pseudosphaeriineae (including Capnodiales, Chaetothyriales, Hemisphaeriales, Lophiostomatales,
Microthyriales, Perisporiales, Pleosporales, Pseudosphaeriales and Trichothyriales). This proposal has however, rarely
been followed. Three existing families, i.e. Lophiostomataceae, Pleosporaceae and Venturiaceae plus 11 other families
were accepted in Pleosporales as arranged by Barr (1979a)
(largely using Luttrell’s concepts, Table 1), and she assigned
these families to six suborders. The morphology of pseudoparaphyses was given much prominence at the ordinal level in
this classification (Barr 1983). In particular the Melanommatales was introduced to accommodate taxa with trabeculate
pseudoparaphyses (Sporormia-type centrum development)
(Barr 1983), distinguished from cellular pseudoparaphyses
(Pleospora-type centrum development) possessed by members of Pleosporales sensu Barr. The order Melanommatales
included Didymosphaeriaceae, Fenestellaceae, Massariaceae, Melanommataceae, Microthyriaceae, Mytilinidiaceae,
Platystomaceae and Requienellaceae (Barr 1990a).
Pleosporales was formally established by Luttrell and Barr
(in Barr 1987b), characterised by perithecioid ascomata,
usually with a papillate apex, ostioles with or without
periphyses, presence of cellular pseudoparaphyses, bitunicate
asci, and ascospores of various shapes, pigmentation and
septation (Table 1). Eighteen families were included, i.e.
Arthopyreniaceae, Botryosphaeriaceae, Cucurbitariaceae,
Dacampiaceae, Dimeriaceae, Hysteriaceae, Leptosphaeriaceae, Lophiostomataceae, Parodiellaceae, Phaeosphaeriaceae, Phaeotrichaceae, Pleomassariaceae, Pleosporaceae,
Polystomellaceae, Pyrenophoraceae, Micropeltidaceae,
Tubeufiaceae and Venturiaceae. Recent phylogenetic analysis
based on DNA sequence comparisons, however, indicated
that separation of the orders (Pleosporales and Melanomma-
tales) based on the Pleospora or Sporormia centrum type, is
not a natural grouping, and Melanommatales has therefore
been combined under Pleosporales (Liew et al. 2000;
Lumbsch and Lindemuth 2001; Reynolds 1991). Six more
families, i.e. Cucurbitariaceae, Diademaceae, Didymosphaeriaceae, Mytilinidiaceae, Testudinaceae and Zopfiaceae,
were subsequently added to Pleosporales (Lumbsch and
Huhndorf 2007). After intensive sampling and multigene
phylogenetic studies, 20 families were accepted in Pleosporales, namely Aigialaceae, Amniculicolaceae, Delitschiaceae,
Didymellaceae, Didymosphaeriaceae, Hypsostromataceae,
Lentitheciaceae, Leptosphaeriaceae, Lindgomycetaceae,
Lophiostomataceae, Massarinaceae, Melanommataceae,
Montagnulaceae, Morosphaeriaceae, Phaeosphaeriaceae,
Pleosporaceae, Pleomassariaceae, Sporormiaceae, Tetraplosphaeriaceae and Trematosphaeriaceae (Boehm et al.
2009a, b; Mugambi and Huhndorf 2009b; Schoch et al. 2009;
Shearer et al. 2009; Suetrong et al. 2009; Tanaka et al. 2009;
Zhang et al. 2009a) (Table 1). In addition, another five
families, i.e. Arthopyreniaceae, Cucurbitariaceae, Diademaceae, Teichosporaceae and Zopfiaceae are tentatively included (Kruys et al. 2006; Plate 1). In the most recent issue of
Myconet, 28 families were included in Pleosporales
(Lumbsch and Huhndorf 2010).
Species included in Pleosporales have different ecological or morphological characters. For instance, members of
the Leptosphaeriaceae have saprobic or parasitic lifestyles
and lightly pigmented, multi-septate ascospores. Members
of the Lophiostomataceae are mostly saprobic with ascomata that usually possess a compressed apex. Members of
Sporormiaceae are coprophilous, and are characterized by
heavily pigmented, multi-septate ascospores with germ
slits, and with or without non-periphysate ostioles. The
lack of DNA sequence data for representatives of numerous
families means that their inter-relationships are unclear and
many genera or species are artificially placed based on
morphological classification. The most recent study on
Venturiaceae indicated that this group had a set of unique
morphological and ecological characters, which is distinct
and distantly related to other members of Pleosporales
(Kruys et al. 2006; Zhang et al. unpublished). Molecular
phylogenetic results indicated that members of Venturiaceae form a robust clade separate from the core members of
Pleosporales, and the clade of Venturiaceae was uncertainly placed but outside of the two currently designated
dothideomycetous subclasses, i.e. Pleosporomycetidae and
Dothideomycetidae (Schoch et al. 2009). In addition,
phylogenetic analysis of rDNA sequence data indicates
that members of Zopfiaceae (as Testudinaceae) seem to
lack affinity with Pleosporales (Kodsueb et al. 2006b).
Thus, 26 families are temporarily accepted in Pleosporales
in this study, although some such as Zopfiaceae, still
require extensive DNA sequence sampling (Table 4).
Fungal Diversity
Table 1 Major circumscription changes of Pleosporales from 1955 to 2011
References
Circumscriptions distinguishing Pleosporales from other orders of Dothideomycetes
Luttrell 1955
Müller and von Arx 1962
Pleospora-type centrum development.
Ascomata perithecoid, with rounded or slit-like ostiole; asci produced within a locule, arranged regularly
in a single layer or irregularly scattered, surrounded with filiform pseudoparaphyses, cylindrical,
ellipsoidal or sac-like.
Ascocarps perithecioid, immersed, erumpent to superficial on various substrates, asci ovoid to mostly
clavate or cylindrical, interspersed with pseudoparaphyses (sometimes form an epithecium) in mostly
medium- to large-sized locules.
Saprobic, parasitic, lichenized or hypersaprobic. Ascomata perithecioid, rarely cleistothecioid or
hysterothecioid, peridium pseudoparenchymatous, pseudoparaphyses cellular, narrow or broad,
deliquescing early at times, not forming an epithecium, asci oblong, clavate or cylindrical, interspersed
with pseudoparaphyses, ascospores mostly asymmetric.
Saprobic, biotrophic or hemibiotrophic. Ascomata globose, subglobose or conical, asci bitunicate, oblong,
clavate or cylindrical, cellular pseudoparaphyses, ascospores hyaline or pigmented, asymmetric or
symmetric, with or without septa.
Ascomata perithecioid or rarely cleistothecioid, sometimes clypeate, mostly globose, thick-walled,
immersed or erumpent, black, sometimes setose, peridium composed of pseudoparenchymatous cells,
pseudoparaphyses trabeculate or cellular, asci cylindrical, fissitunicate, with a well-developed ocular
chamber, rarely with a poorly defined ring (J-), ascospores hyaline to brown, septate, thin or thickwalled, sometimes muriform, usually with sheath, anamorphs hyphomycetous or coelomycetous.
Hemibiotrophic, saprobic, hypersaprobic, or lichenized. Habitats in freshwater, marine or terrestrial
environment. Ascomata perithecioid, rarely cleistothecioid, immersed, erumpent to superficial, globose
to subglobose, or lenticular to irregular, with or without conspicuous papilla or ostioles. Ostioles with or
without periphyses. Peridium usually composed of a few layers of cells with various shapes and
structures. Hamathecium persistent, filamentous, very rarely decomposing. Asci bitunicate, fissitunicate,
cylindrical, clavate to obclavate, with or without pedicel. Ascospores hyaline or pigmented, ellipsoidal,
broadly to narrowly fusoid or filiform, mostly septate.
Luttrell 1973
Barr 1979a
Barr 1987b
Kirk et al. 2001, 2008
Boehm et al. 2009a, b; Mugambi
and Huhndorf 2009b; Schoch et
al. 2009; Shearer et al. 2009;
Suetrong et al. 2009; Tanaka et
al. 2009; Zhang et al. 2009a
Morpho-characters used in taxonomy of Pleosporales
Sexual characters
According to the Linnean classification system, reproductive structures are the most important criteria in plant
taxonomy, and this proposal is widely applied in fungal
taxonomy (Gäumann 1952). In the classification of Dothideomycetes, reproductive characters such as the uni- or
multilocular nature and shape of ascomata, presence and
shape of ostioles/papillae, shape and apical structures of
asci and shape, pigmentation and septation of ascospores
play important roles at different ranks (Clements and Shear
1931; Luttrell 1951, 1955, 1973). Besides the common
morphological characters possessed by Dothideomycetes
(bitunicate and fissitunicate asci as well as the perithecioidlike ascostromata), most pleosporalean fungi also have
pseudoparaphyses among their well-arranged asci (Zhang et
al. 2009a). Currently, classification of Pleosporales at the
family level focuses mostly on morphological characters of
ascomata (such as size, shape of ostiole or papilla), presence
or absence of periphyses, characters of centrum (such as asci,
pseudoparaphyses and ascospores) as well as on lifestyle or
habitat (Barr 1990a; Shearer et al. 2009; Suetrong et al.
2009; Tanaka et al. 2009; Zhang et al. 2009a), whilst relying
extensively on DNA sequence comparisons.
Ascomata
Most species of Pleosporales have uniloculate ascomata.
The presence (or absence) and forms of papilla and ostiole
are the pitoval character of ascomata, which serve as
important characteristics in generic or higher rank classification
(Clements and Shear 1931). The vertically flattened papilla has
recently been shown as an effective criterion for familial level
classification, e.g. in the Amniculicolaceae and the Lophiostomataceae (Zhang et al. 2009a). Papillae and ostioles are
present in most species of Pleosporales, except in the
Diademaceae and Sporormiaceae. Members of Diademaceae
have apothecial ascomata, and some genera of Sporormiaceae
have cleistothecioid ascomata. Another coprophilous pleosporalean family, Delitschiaceae, can be distinguished from
Sporormiaceae by the presence of periphysate ostioles.
Pseudoparaphyses
Presence of pseudoparaphyses is a characteristic of
Pleosporales (Kirk et al. 2008; Liew et al. 2000). Although
pseudoparaphyses may be deliquescing in some families
when the ascomata mature (e.g. in Didymellaceae), they are
persistent in most of other pleosporalean members. According to the thickness, with or without branching and density
of septa, pseudoparaphyses were roughly divided into two
types: trabeculate and cellular, and their taxonomic significance need to be re-evaluated (Liew et al. 2000).
Fungal Diversity
Asci
The asci of Pleosporales are bitunicate, usually fissitunicate, mostly cylindrical, clavate or cylindro-clavate, and
rarely somewhat obclavate or sphaerical (e.g. Macroventuria anomochaeta Aa and Westerdykella dispersa). There are
ocular chambers in some genera (e.g. Amniculicola and
Asteromassaria), or sometimes with a large apical ring (J-)
(e.g. Massaria).
Ascospores
Ascospores of Pleosporales can be hyaline or colored
to varying degrees. They may be amerosporous (e.g.
species of Semidelitschia), phragmosporous (e.g. Phaeosphaeria and Massariosphaeria), dictyosporous (e.g. most
species of Pleospora and Bimuria), or scolecosporous
(e.g. type species of Cochliobolus, Entodesmium or
Lophionema). Although ascospore morphology had been
regarded as a key factor in differentiating genera under
some families, e.g. Arthopyreniaceae (Watson 1929) and
Testudinaceae (Hawksworth 1979), it has been proven
variable even within a single species. For instance, two
types of ascospores are produced by Mamillisphaeria
dimorphospora, i.e. one type is large and hyaline, and the
other is comparatively smaller and brown. Numerous
studies have shown the unreliability of ascospore characters above genus level classification (e.g. Phillips et al.
2008; Zhang et al. 2009a).
Plate 1 The best scoring likelihood tree of representative Pleospor-
ales obtained with RAxML v. 7.2.7 for a concatenated set of
nucleotides from LSU, SSU, RPB2 and TEF1. Family and suborder
names are indicated where possible. The percentages of nodes present
in 250 bootstrap pseudo replicates are shown above branches. Culture
and voucher numbers are indicated after species names and the
presence of the genes used in the analysis are indicated by pluses in
this order: LSU, SSU, RPB2, TEF1
Cucurbitariaceae
Based on the molecular phylogenetic analysis, some
species of Coniothyrium, Pyrenochaeta, Phoma, Phialophorophoma and Pleurophoma belong to Cucurbitariaceae (de
Gruyter et al. 2010; Hyde et al. 2011). Other reported
anamorphs of Cucurbitaria are Camarosporium, Diplodialike and Pleurostromella (Hyde et al. 2011; Sivanesan 1984).
The generic type of Cucurbitaria (C. berberidis Fuckel) is
linked to Pyrenochaeta berberidis (Farr et al. 1989). Curreya
has a Coniothyrium-like anamorphic stage (von Arx and van
der Aa 1983; Marincowitz et al. 2008). The generic type of
Curreya is C. conorum (Fuckel) Sacc., which is reported to
be linked with Coniothyrium glomerulatum Sacc. (von Arx
and van der Aa 1983). The generic type of Rhytidiella (R.
moriformis, Cucurbitariaceae) can cause rough-bark of
Populus balsamifera, and has a Phaeoseptoria anamorphic
stage (Zalasky 1968). Rhytidiella baranyayi Funk & Zalasky,
another species of Rhytidiella associated with the cork-bark
disease of aspen is linked with Pseudosporella-like anamorphs (Funk and Zalasky 1975; Sivanesan 1984).
Asexual states of Pleosporales
Anamorphs of pleosporalean families
Anamorphs of Pleosporales are mostly coelomycetous,
but may also be hyphomycetous. Phoma or Phoma-like
anamorphic stages and its relatives are most common
anamorphs of Pleosporales (Aveskamp et al. 2010; de
Gruyter et al. 2009, 2010; Hyde et al. 2011). Some of the
reported teleomorph and anamorph connections (including
some listed below) are, however, based on the association
rather than single ascospore isolation followed by induction
of the other stage in culture (Hyde et al. 2011).
Pleosporales suborder Pleosporineae
Pleosporineae is a phylogenetically well supported
suborder of Pleosporales, which temporarily includes
seven families, namely Cucurbitariaceae, Didymellaceae,
Didymosphaeriaceae, Dothidotthiaceae, Leptosphaeriaceae, Phaeosphaeriaceae and Pleosporaceae, and
contains many important plant pathogens (de Gruyter et
al. 2010; Zhang et al. 2009a). De Gruyter et al. (2009,
2010) systematically analyzed the phylogeny of Phoma
and its closely related genera, and indicated that their
representative species cluster in different subclades of
Pleosporineae.
Didymellaceae, Didymosphaeriaceae and Dothidotthiaceae
As has been mentioned before, Phoma sensu lato species
have been proved to be highly polyphyletic, and they cluster
in six distinct familial clades within the Pleosporales
(Aveskamp et al. 2010). Most Phoma species, including the
generic type (P. herbarum), clustered in Didymellaceae
(Aveskamp et al. 2010). The clade of Didymellaceae also
comprises other sections, such as Ampelomyces, Boeremia,
Chaetasbolisia, Dactuliochaeta, Epicoccum, Peyronellaea,
Phoma-like, Piggotia, Pithoascus, as well as the type species
of Ascochyta and Microsphaeropsis (Aveskamp et al. 2010;
de Gruyter et al. 2009; Kirk et al. 2008; Sivanesan 1984).
Leptosphaerulina is another genus of Didymellaceae, which
has hyphomycetous anamorphs with pigmented and
muriform conidia, such as Pithomyces (Roux 1986).
The other reported anamorphs of Didymosphaeria are
Fusicladiella-like, Dendrophoma, Phoma-like (Hyde et al.
2011). Hyphomycetous Thyrostroma links to Dothidotthiaceae (Phillips et al. 2008).
Some important plant pathogens are included within
Didymellaceae, such as Phoma medicaginis Malbr. &
Roum., which is a necrotrophic pathogen on Medicago
truncatula (Ellwood et al. 2006). Phoma herbarum is
another plant pathogen, which has potential as a
Fungal Diversity
Fungal Diversity
Plate 1 (continued)
Fungal Diversity
biocontrol agent of weeds (Neumann and Boland 2002).
Ascochyta rabiei is a devastating disease of chickpea in
most of the chickpea producing countries (Saxena and
Singh 1987).
Leptosphaeriaceae
The anamorphic stages of Leptosphaeriaceae can be
Coniothyrium, Phoma, Plenodomus and Pyrenochaeta. All
are coelomycetous anamorphs, and they may have phialidic
or annellidic conidiogenous cells. Phoma heteromorphospora Aa & Kesteren, the type species of Phoma sect.
Heterospora and Coniothyrium palmarum, the generic type
of Coniothyrium, reside in Leptosphaeriaceae (de Gruyter
et al. 2009).
Pleosporaceae
Various anamorphic types can occur in Pleosporaceae,
which can be coelomycetous or hyphomycetous, and the
ontogeny of conidiogenous cells can be phialidic, annellidic
or sympodial blastic. Both Ascochyta caulina and Phoma
betae belong to Pleosporaceae (de Gruyter et al. 2009).
Some species of Bipolaris and Curvularia are anamorphs of Cochliobolus. Many species of these two
genera cause plant disease or even infect human beings
(Khan et al. 2000). They are hyphomycetous anamorphs
with sympodial proliferating conidiogenous cells, and
pigmented phragmosporous poroconidia. The generic type
of Lewia (L. scrophulariae) is linked with Alternaria
conjuncta E.G. Simmons (Simmons 1986), and the
generic type of Pleospora (P. herbarum) is linked with
Stemphylium botryosum Sacc. (Sivanesan 1984). Both
Alternaria and Stemphylium are hyphomycetous anamorphs
characterized by pigmented, muriform conidia that develop
at a very restricted site in the apex of distinctive conidiophores (Simmons 2007).
The generic type of Pleoseptum (P. yuccaesedum) is
linked with Camarosporium yuccaesedum (Ramaley and
Barr 1995), the generic type of Macrospora (M. scirpicola)
with Nimbya scirpicola (Fuckel) E.G. Simmons (Simmons
1989), and the generic type of Setosphaeria (S. turcica)
with Drechslera turcica (Pass.) Subram. & B.L. Jain
(Sivanesan 1984). Pyrenophora has the anamorphic stages
of Drechslera, and the anamorphic stage of Wettsteinina
can be species of Stagonospora (Farr et al. 1989).
Most common anamorphs in Pleosporaceae are Alternaria, Bipolaris, Phoma-like and Stemphylium, and they
can be saprobic or parasitic on various hosts. Phoma betae
A.B. Frank is a notorious pathogen on sugar beet, which
causes zonate leaf spot or Phomopsis of sugar beet.
Alternaria porri (Ellis) Cif., Stemphylium solani G.F.
Weber, S. botryosum and S. vesicarium (Wallr.) E.G.
Simmons can cause leaf blight of garlic (Zheng et al.
2009). Phoma incompta Sacc. & Martelli is a pathogen on
olive, and Stemphylium botryosum, the anamorph of
Pleospora herbarum, causes leaf disease of olive trees
(Malathrakis 1979).
Phaeosphaeriaceae
The type species of Phoma sect. Paraphoma (Phoma
radicina (McAlpine) Boerema) as well as several pathogens
on Gramineae, i.e. Stagonospora foliicola (Bres.) Bubák, S.
neglecta var. colorata and Wojnowicia hirta Sacc. belong to
Phaeosphaeriaceae (de Gruyter et al. 2009). Other anamorphs reported for Phaeosphaeriaceae are Amarenographium, Ampelomyces, Chaetosphaeronema, Coniothyrium,
Hendersonia, Neosetophoma, ?Parahendersonia, Paraphoma, Phaeoseptoria, Rhabdospora, Scolecosporiella, Setophoma, Sphaerellopsis and Tiarospora.
These anamorphic fungi can be saprobic, but mostly
pathogenic on herbaceous plants. For instance, Stagonospora foliicola and Coniothyrium concentricum (Desm.)
Sacc. can cause leaf spots on herbaceous plants (Zeiders
1975), and Ampelomyces quisqualis Ces. is a hyperparasite
of powdery mildews.
Pleosporales suborder Massarineae
Massarineae species are mostly saprobic in terrestrial or
aquatic environments. Five families are currently included
within Massarineae, viz. Lentitheciaceae, Massarinaceae,
Montagnulaceae, Morosphaeriaceae and Trematosphaeriaceae. Anamorphs of the five families are summarized as
follows.
Lentitheciaceae
Stagonospora macropycnidia Cunnell nests within the
clade of Lentitheciaceae (Plate 1). A relatively broad genus
concept of Stagonospora is currently accepted, which
comprises parasitic or saprobic taxa. Keissleriella cladophila (Niessl) Corbaz is another species nesting within
Lentitheciaceae (Zhang et al. 2009a), and is linked with
Dendrophoma sp., which has branching conidiogenous
cells, and 1-celled, hyaline conidia (Bose 1961; Sivanesan
1984).
Massarinaceae
A relatively narrow concept tends to be accepted for
Massarinaceae, which seems only to comprise limited
species such as Byssothecium circinans, Massarina eburnea, M. cisti S.K. Bose, M. igniaria (C. Booth) Aptroot
(anamorph: Periconia igniaria E.W. Mason & M.B. Ellis)
and Neottiosporina paspali (G.F. Atk.) B. Sutton & Alcorn
(Zhang et al. 2009a; Plate 1). Similarly, a relatively narrow
generic concept of Massarina was accepted, containing
only M. eburnea and M. cisti (Zhang et al. 2009b), and both
species have been linked with species of Ceratophoma
(Sivanesan 1984).
Fungal Diversity
Montagnulaceae
Montagnula has an Aschersonia anamorph, and Kalmusia and Paraphaeosphaeria have Coniothyrium-like, Cytoplea, Microsphaeropsis and Paraconiothyrium anamorphs.
The generic type of Paraphaeosphaeria (P. michotii) is
linked with Coniothyrium scirpi Trail (Webster 1955). The
Coniothyrium complex is highly polyphyletic, and was
subdivided into four groups by Sutton (1980), viz.
Coniothyrium, Microsphaeropsis, Cyclothyrium and Cytoplea. Paraconiothyrium was introduced to accommodate
Coniothyrium minitans W.A. Campb. and C. sporulosum
(W. Gams & Domsch) Aa, which are closely related to
Paraphaeosphaeria based on 18S rDNA sequences phylogeny (Verkley et al. 2004).
Morosphaeriaceae
Based on the multigene phylogenetic analysis in this
study, Asteromassaria is tentatively included in Morosphaeriaceae. Asteromassaria macrospora is linked with
Scolicosporium macrosporium (Berk.) B. Sutton, which is
hyphomycetous. No anamorphic stages have been reported
for other species of Morosphaeriaceae.
Trematosphaeriaceae
Three species from three different genera were included
in Trematosphaeriaceae, i.e. Falciformispora lignatilis,
Halomassarina thalassiae and Trematosphaeria pertusa
(Suetrong et al. data unpublished; Plate 1). Of these, only
Trematosphaeria pertusa, the generic type of Trematosphaeria, produces hyphopodia-like structures on agar
(Zhang et al. 2008a).
Other families of Pleosporales
2009a). Leuchtmann (1985) studied cultures of some
Lophiostoma species, and noticed that L. caulium (Fr.)
Ces. & De Not., L. macrostomum, L. semiliberum (Desm.)
Ces. & De Not., Lophiostoma sp. and Lophiotrema nucula
produced Pleurophomopsis anamorphic stages, which are
similar to those now in Melanomma (Chesters 1938), but
Lophiostoma and Melanomma has no proven phylogenetic
relationship (Zhang et al. 2009a, b; Plate 1). Species of
Aposphaeria have also been reported in Massariosphaeria
(Farr et al. 1989; Leuchtmann 1984), but the polyphyletic
nature of Massariosphaeria is well documented (Wang et
al. 2007).
Melanommataceae
The anamorphs of the Melanommataceae are mostly
coelomycetous and rarely hyphomycetous with various
ontogenic structures, such as annellidic or sympodial for
hyphomycetes (Exosporiella and Pseudospiropes) and
coelomycetes (Aposphaeria-like and Pyrenochaeta).
Herpotrichia is reported as having a Pyrenochaeta
anamorphic stage with or without seta on the surface of
pycnidia (Sivanesan 1984). Aposphaeria and Phoma-like
have been reported in Melanomma species (Chesters
1938; Sivanesan 1984). Similarly, the anamorphs of
Karstenula are reported as coelomycetous, i.e. Microdiplodia (Constantinescu 1993). The anamorphic stage of
Anomalemma is Exosporiella (Sivanesan 1983), and that
of Byssosphaeria is Pyrenochaeta (Barr 1984). Ohleria
brasiliensis Starbäck has been linked with Monodictys
putredinis (Wallr.) S. Hughes (Samuels 1980). Astrosphaeriella is a contentious genus as its familial status is
not determined yet. Here we temporarily assigned it under
Melanommataceae, which is linked with the anamorph
genus Pleurophomopsis.
Amniculicolaceae
Three anamorphic species nested within the clade of
Amniculicolaceae, i.e. Anguillospora longissima (Sacc. &
P. Syd.) Ingold, Repetophragma ontariense (Matsush.) W.
P. Wu and Spirosphaera cupreorufescens Voglmayr
(Zhang et al. 2009a). Sivanesan (1984, p. 500) described
the teleomorphic stage of Anguillospora longissima as
Massarina sp. II, which fits the diagnostic characters of
Amniculicola well. Thus this taxon may be another species of
Amniculicola.
Hypsostromataceae
A Pleurophomopsis-like anamorph is reported in the
subiculum of the generic type of Hypsostroma (H. saxicola
Huhndorf) (Huhndorf 1992).
Lophiostomataceae
The concept of Lophiostomataceae was also narrowed,
and presently contains only Lophiostoma (Zhang et al.
Pleomassariaceae
Shearia and Prosthemium are all anamorphs of Pleomassaria, and Prosthemium betulinum is linked with the generic
type of Pleomassaria (P. siparia) (Barr 1982b; Sivanesan
1984; Sutton 1980; Tanaka et al. 2010). Splanchnonema is a
genus of Pleomassariaceae, the teleomorphic morphology of
which is difficult to distinguish from two other genera, i.e.
Asteromassaria and Pleomassaria, and the reported anamorphs of Splanchnonema are Ceuthodiplospora, Myxocyclus and Stegonsporium, which are comparable with those of
Asteromassaria and Pleomassaria.
Tetraplosphaeriaceae
Tetraplosphaeriaceae was introduced to accommodate
the Massarina-like bambusicolous fungi that produce
Tetraploa sensu stricto anamorphs (Tanaka et al. 2009).
Tetraploa aristata Berk. & Broome, the generic type of
Tetraploa is widely distributed, associated with various
Fungal Diversity
substrates and many occur in freshwater or has been
isolated from air. The polyphyletic nature of T. aristata
has been well documented (Tanaka et al. 2009). Anamorphic stages can serve as a diagnostic character for this
family.
Diademaceae, Massariaceae, Sporormiaceae
and Teichosporaceae
The Sporormiaceae is coprophilous having Phoma or
Phoma-related anamorphic states (Cannon and Kirk 2007).
Comoclathris (Diademaceae) is linked with Alternaria-like
anamorphs (Simmons 1952). Myxocyclus links to Massaria
(Massariaceae) (Hyde et al. 2011). The anamorphic stage of
Chaetomastia (Teichosporaceae) is Aposphaeria- or Coniothyrium-like (Barr 1989c).
Generally speaking, the morphologically simple conidiophores are usually considered phylogenetically uninformative
(Seifert and Samuels 2000). Phoma-like anamorphs commonly occur in Pleosporales, while their colorless and unicellular
conidia are also not phylogenetically informative (Seifert and
Samuels 2000).
All of the above mentioned anamorphic taxa of
Pleosporales have phialidic, annellidic or sympodial conidiogenous cells, representing apical wall-building type
(compared to ring wall-building and diffused wallbuilding) (Nag Raj 1993), which may indicate that the
wall-building type probably has phylogenetic significance.
Molecular phylogeny of Pleosporales
Numerous genes have been applied in phylogenetic
studies of Pleosporales, mostly including LSU, SSU,
mtSSU and ITS as well as the protein genes, such as
RPB1, RPB2, TEF1, β-tubulin (TUB1) and actin (ACT1).
A single gene such as ITS or LSU, has been used to study
phylogenetic relationships between Leptosphaeria and
Phaeosphaeria (Câmara et al. 2002) or Pleosporaceae
and Tubeufiaceae (Kodsueb et al. 2006a, b) (Table 2). The
use of these phylogenetic markers, although making
important contributions, has not been successful in
resolving numerous relationships in single gene dendrograms. One exception is the use of SSU sequences to
demonstrate the phylogenetic significance of pseudoparaphyses (Liew et al. 2000) whilst rejecting the phylogenetic utility of pseudoparaphyses morphology (cellular or
trabeculate). Analyses with combined genes have had
more success. For instance combined analyses with LSU
and SSU sequence data could be used to define family
level classification in a few cases (Dong et al. 1998; de
Gruyter et al. 2009; Lumbsch and Lindemuth 2001; Pinnoi
et al. 2007; Zhang et al. 2009b) (Table 2). The addition of
more than two genes has been used to determine relationships between orders. For instance, genes such as LSU,
SSU and mtSSU have been used to analyze ordinal
relationships in Loculoascomycetes (Lindemuth et al.
2001), and to analyze phylogenetic relationships of
coprophilous families in Pleosporales (Kruys et al.
2006). Phaeocryptopus gaeumannii (T. Rohde) Petr. was
shown to belong in Dothideales based on LSU, SSU and
ITS sequence analysis (Winton et al. 2007), while Schoch
et al. (2006) used four genes, i.e. LSU, SSU, RPB2 and
TEF1 to evaluate the phylogenetic relationships among
different orders of the Dothideomycetes. Five genes, viz.
LSU, SSU, TEF1, RPB1 and RPB2, were used to study the
phylogenetic relationships of different orders within
Dothideomycetes (Schoch et al. 2009) and of different
families within Pleosporales (Zhang et al. 2009a)
(Table 2). It is clear that even more genes will be required
to address the remaining issues and the promise of genome
analyses is within reach (www.jgi.doe.gov/sequencing/
why/dothideomycetes.html) for Dothideomycetes.
The importance of generic type specimens
The type specimen (collection type) is a fundamental
element in the current Code of Botanical Nomenclature at
familial or lower ranks (Moore 1998). A type specimen
fixes the name to an exact specimen at family, genera,
species and variety/subspecies rank and is ultimately based
on this single specimen, i.e. a family name is based on a
genus, the genus name is based on a species, and the
species name is based on a specimen (Kirk et al. 2008).
The generic type is of great importance in defining
generic circumscriptions in fungal taxonomy. The generic
types of Pleosporales have been studied previously by
many mycologists. For instance, Müller and von Arx
(1962) studied the generic types of “Pyrenomycetes”, and
described and illustrated them in detail. Sivanesan (1984)
described and illustrated the generic representatives of
Loculoascomycetes for both their teleomorphs and anamorphs, and their links were emphasized. A large number
of pleosporalean genera have been studied by Barr (1990a,
b). Almost all of the previous work was conducted more
than 20 years ago, when no molecular phylogenetic studies
could be carried out and thus had been carried out in a
systematic fashion.
Aim and outline of present study
The present study had two principal objectives:
1. To explore genera under Pleosporales based on the
generic types and provide a detailed description and
illustration for the type species of selected genera,
discuss the study history of those genera, and explore
their ordinal, familial, and generic relationships;
Fungal Diversity
Table 2 List of phylogenetic studies on Pleosporales
Year
Author(s)
Loci used
Target fungi
General conclusion
1998
Dong et al.
LSU, SSU
2000
Liew et al.
SSU
Leptosphaeriaceae, Pleosporaceae
and three other families
Pleosporales and Melanommatales
2001
2001
LSU, SSU, mtSSU
LSU, SSU
loculoascomycetes
Dothideomycetes
2002
2006
Lindemuth et al.
Lumbsch and
Lindemuth
Câmara et al.
Kodsueb et al.
ITS
LSU
Leptosphaeria and Phaeosphaeria
Pleosporaceae
2006
Kodsueb et al.
LSU
Tubeufiaceae
2006
Kruys et al.
LSU, SSU, mtSSU
coprophilous familes of
Pleosporales
2006
Schoch et al.
Dothideomycetes
2007
Pinnoi et al.
LSU, SSU, TEF1,
RPB2
LSU, SSU
Pleosporales
2007
2007
Wang et al.
Winton et al.
LSU, SSU, RPB2
LSU, SSU, ITS
Massariosphaeria
Phaeocryptopus gaeumannii
2008a
Zhang et al.
LSU, SSU
Melanomma and Trematosphaeria
2009
de Gruyter et al.
LSU, SSU;
Phoma and related genera
2009a
Zhang et al.
LSU, SSU, TEF1,
RPB1, RPB2
Pleosporales
2009
Mugambi and
Huhndorf
LSU, TEF1
Melanommataceae,
Lophiostomataceae
2009
Nelsen et al.
LSU and mtSSU
lichenized Dothideomycetes
2009
Suetrong et al.
LSU, SSU, TEF1, RPB1
marine Dothideomycetes
2009
Shearer et al.
LSU, SSU
freshwater Dothideomycetes
2009
Tanaka et al.
LSU, SSU, TEF1, ITS, BT
bambusicolous Pleosporales
2009
Kruys and Wedin
ITS-nLSU, mtSSU rDNA
and β-tubulin
Sporormiaceae
2010
Hirayama et al.
LSU, SSU
Massarina ingoldiana sensu lato
2010
Aveskamp et al.
LSU, SSU, ITS and β-tubulin
2010
de Gruyter et al.
LSU, SSU
Phoma and related genera within
Didymellaceae
Phoma and related genera within
Pleosporineae
Leptosphaeriaceae is paraphyletic and
Pleosporaceae is monophyletic.
Pleosporales and Melanommatales are not
naturial groups.
Loculoascomycetes are not monophyletic.
Presence of pseudoparaphyses is a major
character at order level classification
Accepted Leptosphaeria sensu stricto.
Wettsteinina should be excluded from the
Pleosporaceae.
Tubeufiaceae is more closely related to the
Venturiaceae.
coprophilous familes of Pleosporales form
phylogenetic monophyletic groups
respectively
Proposed the subclasses
Pleosporomycetidae
phylogenetic relationships of different
families of Pleosporales, introduced a
new fungus–– Berkleasmium crunisia
Massariosphaeria is not monophyletic
Phaeocryptopus gaeumannii nested in
Dothideales.
Melanomma and Trematosphaeria belong
to different families
They are closely related with
Didymellaceae, Leptosphaeriaceae,
Phaeosphaeriaceae and Pleosporaceae
Amniculicolaceae and Lentitheciaceae were
introduced, and Pleosporineae
recircumscribed.
Recircumscribed Melanommataceae and
Lophiostomataceae, and reinstated
Hypsostromataceae.
Pyrenocarpous lichens with bitunicate asci
are not monophyletic, but belong to at
least two classes (Dothideomycetes and
Erotiomycetes).
Two new families are introduced
Aigialaceae and Morosphaeriaceae.
Freshwater Dothideomycetes are related to
terrestrial taxa and have adapted to
freshwater habitats numerous times.
Introduced Tetraplosphaeriaceae with
Tetraploa-like anamorphs.
Analyzed the inter-generic relationships as
well as evaluated the morphological significance used in this family.
Massarina ingoldiana sensu lato is
polyphyletic, and separated into two
clades within Pleosporales.
Rejected current Boeremaean subdivision.
Introduced Pyrenochaetopsis, Setophoma
and Neosetophoma and reinstated
Cucurbitariaceae within Pleosporineae
Fungal Diversity
2. To investigate the phylogeny of Pleosporales, its interfamilial relationships, and the morphological circumscription of each family;
In order to clarify morphological characters, the generic
types of the majority of teleomorphic pleosporalean genera
(> 60%) were studied. Most of them are from the “core
families” of Pleosporales, i.e. Delitschiaceae, Lophiostomataceae, Massariaceae, Massarinaceae, Melanommataceae,
Montagnulaceae, Phaeosphaeriaceae, Phaeotrichaceae, Pleomassariaceae, Pleosporaceae, Sporormiaceae and Teichosporaceae. Notes are given for those where type
specimens could not be obtained during the timeframe
of this study. A detailed description and illustration of
each generic type is provided. Comments, notes and
problems that need to be addressed are provided for each
genus. Phylogenetic investigation based on five nuclear loci,
viz. LSU, SSU, RPB1, RPB2 and TEF1 was carried out using
available strains from numerous genera in Pleosporales. In
total, 278 pleosporalean taxa are included in the phylogenetic
analysis, which form 25 familial clades on the dendrogram
(Plate 1). The suborder, Massarineae, is emended to
accommodate Lentitheciaceae, Massarinaceae, Montagnulaceae, Morosphaeriaceae and Trematosphaeriaceae.
Materials and methods
Molecular phylogeny
Four genes were used in this analysis, the large and small
subunits of the nuclear ribosomal RNA genes (LSU, SSU)
and two protein coding genes, namely the second largest
subunit of RNA polymerase II (RPB2) and translation
elongation factor-1 alpha (TEF1). All sequences were
downloaded from GenBank as listed in Table 3. Each of
the individual ribosomal genes was aligned in SATé under
default settings with at least 20 iterations. The protein coding
genes were aligned in BioEdit (Hall 2004) and completed by
manual adjustment. Introns were removed and all genes were
concatenated in a single nucleotide alignment with 43%
missing and gap characters out of a total set of 5081. The
alignment had 100% representation for LSU, 75% for SSU,
48% for RPB2 and 65% for TEF1. The final data matrix had
280 taxa including outgroups (Table 3).
Previous results indicated no clear conflict amongst the
majority of the data used (Schoch et al. 2009). A phylogenetic
analysis of the concatenated alignment was performed on
CIPRES webportal (Miller et al. 2009) using RAxML v. 7.2.7
(Stamatakis 2006; Stamatakis et al. 2008) applying unique
model parameters for each gene and codon (8 partitions). A
general time reversible model (GTR) was applied with a
discrete gamma distribution and four rate classes. Fifty
thorough maximum likelihood (ML) tree searches were done
in RAxML v. 7.2.7 under the same model, each one starting
from a separate randomized tree and the best scoring tree
selected with a final likelihood value of −95238.628839. Two
isolates of Hysterium angustatum (Hysteriales, Pleosporomycetidae) were used as outgroups based on earlier work
(Boehm et al. 2009a). Bootstrap pseudo-replicates were run
with the GTRCAT model approximation, allowing the
program to halt bootstraps automatically under the majority
rule criterion (Pattengale et al. 2010). The resulting 250
replicates were plotted on to the best scoring tree obtained
previously. The phylogram with bootstrap values on the
branches is presented in Plate 1 by using graphical options
available in TreeDyn v. 198.3 (Chevenet et al. 2006).
Morphology
Type specimens as well as some other specimens were loaned
from the following herbaria: BAFC, BISH, BPI, BR, BRIP,
CBS, E, ETH, FFE, FH, G, H, Herb. J. Kohlmeyer, HHUF,
IFRD, ILLS, IMI, K(M), L, LPS, M, MA, NY, PAD, PC, PH,
RO, S, TNS, TRTC, UB, UBC, UPS and ZT. Attempts were
made to trace and borrow all the type specimens from herbaria
worldwide, but only some of them could be obtained. Some of
the type specimens are in such bad condition that little
information could be obtained. In order to obtain the location
of specimens, original publications were searched.
Ascostroma and ascomata were examined under an
Olympus SZ H10 dissecting microscope. Section of the
fruiting structures was carried out by cryotome or by handcutting. Measurements and descriptions of sections of the
ascomata, hamathecium, asci and ascospores were carried out
by immersing ascomata in water or in 10% lactic acid.
Microphotography was taken with material mounted in water,
cotton blue, Melzer’s reagent or 10–100% lactic acid.
Terminologies are as in Ulloa and Hanlin (2000). In
addition, ascomata size is defined as: small-sized: < 300 μm
diam., medium-sized: from 300 μm to 600 μm diam., largesized: > 600 μm diam.
Question mark (“?”) before family (or genus) name
means its familial (or generic) status within Pleosporales
(or some particular family) is uncertain. Other question
marks after habitats, latin names or other substantives mean
the correctness of their usages need verification.
Results
Molecular phylogeny
In total, 278 pleosporalean taxa are included in the phylogenetic analysis. These form 25 familial clades in the dendrogram, i.e. Aigialaceae, Amniculicolaceae, Arthopyreniaceae,
Fungal Diversity
Table 3 Taxa used in the phylogenetic analysis and their corresponding GenBank numbers. Culture and voucher abbreviations are indicated were
available
Species
Culture/voucher1
LSU
SSU
RPB2
Acrocordiopsis patilii
Acrocordiopsis patilii
Aigialus grandis
Aigialus grandis
Aigialus mangrovis
Aigialus mangrovis
Aigialus parvus
Aigialus parvus
Aigialus rhizophorae
Aigialus rhizophorae
Alternaria alternata
Amniculicola immersa
Amniculicola parva
BCC 28166
BCC 28167
BCC 18419
JK 5244A
BCC 33563
BCC 33564
A6
BCC 32558
BCC 33572
BCC 33573
CBS 916.96
CBS 123083
CBS 123092
GU479772
GU479773
GU479774
GU301793
GU479776
GU479777
GU301795
GU479779
GU479780
GU479781
DQ678082
FJ795498
FJ795497
GU479736
GU479737
GU479738
GU296131
GU479741
GU479742
GU296133
GU479743
GU479745
GU479746
DQ678031
GU456295
GU296134
GU479811
GU479812
GU479813
GU371762
GU479815
GU479816
GU371771
GU479818
GU479819
GU479820
DQ677980
GU456358
Anteaglonium abbreviatum
Anteaglonium abbreviatum
Anteaglonium globosum
Anteaglonium latirostrum
Arthopyrenia salicis
Arthopyrenia salicis
Ascochyta pisi
Ascocratera manglicola
Ascocratera manglicola
Asteromassaria pulchra
Astrosphaeriella aggregata
Astrosphaeriella aggregata
Astrosphaeriella bakeriana
Astrosphaeriella stellata
Beverwykella pulmonaria
Biatriospora marina
Bimuria novae-zelandiae
Byssolophis sphaerioides
ANM 925.1
GKM 1029
ANM 925.2
L100N 2
1994 Coppins
CBS 368.94
CBS 126.54
BCC 09270
JK 5262 C
CBS 124082
MAFF 239485
MAFF 239486
CBS 115556
MAFF 239487
CBS 283.53
CY 1228
CBS 107.79
IFRDCC2053
GQ221877
GQ221878
GQ221879
GQ221876
AY607730
AY538339
DQ678070
GU479782
GU301799
GU301800
AB524590
AB524591
GU301801
AB524592
GU301804
GQ925848
AY016356
GU301805
Byssosphaeria jamaicana
Byssosphaeria rhodomphala
Byssosphaeria salebrosa
Byssosphaeria schiedermayeriana
Byssosphaeria schiedermayeriana
Byssosphaeria villosa
Byssothecium circinans
Chaetosphaeronema hispidulum
Cochliobolus heterostrophus
Cochliobolus sativus
Corynespora cassiicola
Corynespora olivacea
Corynespora smithii
Cucurbitaria berberidis
Decaisnella formosa
Decaisnella formosa
Decorospora gaudefroyi
SMH1403
GKM L153N
SMH2387
GKM1197
GKM152N
GKM204N
CBS 675.92
CBS 216.75
CBS 134.39
DAOM 226212
CBS 100822
CBS 114450
CABI 5649b
CBS 394.84
BCC 25616
BCC 25617
CBS 332.63
GU385152
GU385157
GU385162
GU385161
GU385168
GU385151
AY016357
EU754144
AY544645
DQ678045
GU301808
GU301809
GU323201
GQ387605
GQ925846
GQ925847
EF177849
TEF1
GU479838
GU479840
GU479841
GU349064
GU479843
GU479844
GU479845
DQ677927
GU456273
GU349065
GQ221924
GQ221915
GQ221925
GQ221938
AY538333
DQ678018
GU479747
GU296136
GU296137
AB524449
AB524450
DQ677967
GU479821
GU371763
GU371772
DQ677913
GU479846
AB539105
AB539092
GU349015
GU349066
AB524451
GQ925835
AY016338
GU296140
AY016339
EU754045
AY544727
DQ677995
GU296144
GQ387544
GQ925833
GQ925834
AF394542
GU371768
GU479823
DQ470917
GU456348
DQ767646
GU371777
DQ247790
DQ677939
GU371742
GU479848
DQ471087
GU456263
GU327746
GU327747
GU327748
GU327750
GU327749
GU327751
GU349061
DQ497603
GU371783
GU349052
GU349014
GU349018
GU479825
GU479824
GU479851
GU479850
Fungal Diversity
Table 3 (continued)
Species
Culture/voucher1
LSU
Delitschia cf. chaetomioides
Delitschia cf. chaetomioides
Delitschia chaetomioides
Delitschia chaetomioides
Delitschia winteri
Didymella exigua
Didymocrea sadasivanii
Didymosphaeria futilis
Didymosphaeria futilis
Dothidotthia aspera
Dothidotthia symphoricarpi
GKM 1283
GKM 3253.2
GKM1283
SMH3253.2
CBS 225.62
CBS 183.55
CBS 438 65
CMW 22186
HKUCC 5834
CPC 12933
CBS119687
GU385172
GU390656
GU385172
GU390656
DQ678077
EU754155
DQ384103
EU552123
GU205219
EU673276
EU673273
Entodesmium rude
Falciformispora lignatilis
Falciformispora lignatilis
Floricola striata
Floricola striata
Halomassarina thalassiae
Halomassarina thalassiae
Halotthia posidoniae
Helicascus nypae
Helicascus nypae
Herpotrichia diffusa
Herpotrichia juniperi
Herpotrichia macrotricha
Herpotrichia macrotricha
Hypsostroma caimitalense
Hypsostroma saxicola
Hysterium angustatum
Hysterium angustatum
CBS 650.86
BCC 21117
BCC 21118
JK 5603 K
JK 5678I
BCC 17055
JK 5262D
BBH 22481
BCC 36751
BCC 36752
CBS 250.62
CBS 200.31
GKM196N
SMH269
GKM 1165
SMH 5005
CBS 123334
CBS 236.34
GU301812
GU371826
GU371827
GU479785
GU301813
GQ925850
GU301816
GU479786
GU479788
GU479789
DQ678071
DQ678080
GU385176
GU385177
GU385180
GU385181
FJ161207
FJ161180
Julella avicenniae
Julella avicenniae
Julella avicenniae
Kalmusia scabrispora
Kalmusia scabrispora
Karstenula rhodostoma
Katumotoa bambusicola
Keissleriella cladophila
Keissleriella rara
Kirschsteiniothelia elaterascus
Lentithecium aquaticum
Lentithecium arundinaceum
Lentithecium arundinaceum
Lentithecium fluviatile
Lepidosphaeria nicotiae
Leptosphaeria biglobosa
Leptosphaeria biglobosa
Leptosphaeria doliolum
BCC 18422
BCC 20173
JK 5326A
MAFF 239517
NBRC 106237
CBS 690.94
MAFF 239641
CBS 104.55
CBS 118429
A22-5A/HKUCC7769
CBS 123099
CBS 123131
CBS 619.86
CBS 122367
CBS 101341
CBS 298.36
CBS 303.51
CBS 505.75
GU371823
GU371822
GU479790
AB524593
AB524594
GU301821
AB524595
GU301822
GU479791
AY787934
GU301823
GU456320
GU301824
GU301825
DQ678067
GU237980
GU301826
GU301827
Leptosphaeria dryadis
CBS 643.86
GU301828
Leptosphaerulina argentinensis
CBS 569.94
GU301829
SSU
DQ678026
EU754056
DQ384066
RPB2
TEF1
DQ677975
GU327752
GU327753
DQ677922
GU205236
EU673228
EU673224
GU371834
GU371835
GU479751
GU296149
GQ925843
GU349012
GU371819
GU371820
GU371758
GU349011
GU479752
GU479754
GU479755
DQ678019
DQ678029
GU479826
GU479827
DQ677968
DQ677978
GU479854
GU479855
DQ677915
DQ677925
GU327755
GU327756
FJ161167
GU397359
FJ161129
FJ161117
FJ161111
FJ161096
GU371831
GU371830
GU479756
AB524452
AB524453
GU296154
AB524454
GU296155
GU479757
AF053727
GU296156
GU456298
GU296157
GU296158
GU371787
GU371786
GU371816
GU371815
AB539093
AB539094
GU371788
AB539095
GU371735
AB539106
AB539107
GU349067
AB539108
GU349043
GU371789
GU349068
GU456281
FJ795473
DQ677963
GU349074
DQ677910
GU238207
GU349010
GU349069
GU296159
GU371733
GU349009
GU349008
Fungal Diversity
Table 3 (continued)
Species
Culture/voucher1
LSU
SSU
Leptosphaerulina australis
Leptosphaerulina australis
Leptosphearia maculans
Letendraea helminthicola
Letendraea padouk
Lindgomyces breviappendiculatus
Lindgomyces cinctosporae
Lindgomyces cinctosporae
Lindgomyces ingoldianus
Lindgomyces rotundatus
Lophiostoma alpigenum
CBS 311.51-T
CBS 317.83
DAOM 229267
CBS 884.85
CBS 485.70
KT 1399
R56-1
R56-3
KH 100 JCM 16479
KH 114 JCM 16484
GKM 1091b
FJ795500
GU301830
DQ470946
AY016362
AY849951
AB521749
AB522431
GU266245
AB521737
AB521742
GU385193
GU296160
DQ470993
AY016345
GU296162
AB521734
AB522430
GU266238
AB521720
AB521725
Lophiostoma arundinis
Lophiostoma caulium
Lophiostoma compressum
Lophiostoma crenatum
Lophiostoma fuckelii
Lophiostoma fuckelii
Lophiostoma fuckelii
Lophiostoma macrostomum
Lophiostoma macrostomum
Lophiostoma macrostomum
Lophiostoma quadrinucleatum
Lophiostoma sagittiforme
Lophiotrema brunneosporum
Lophiotrema lignicola
Massarina arundinariae
Massarina arundinariae
Lophiotrema nucula
Loratospora aestuarii
CBS 621.86
CBS 623.86
IFRD 2014
CBS 629.86
CBS 101952
CBS 113432
GKM 1063
CBS 122681
HHUF 27293
KT 635
GKM1233
HHUF 29754
CBS 123095
CBS 122364
MAFF 239461
NBRC 106238
CBS 627.86
JK 5535B
DQ782384
GU301833
GU301834
DQ678069
DQ399531
EU552139
GU385192
EU552141
AB433274
AB433273
GU385184
AB369267
GU301835
GU301836
AB524596
AB524597
GU301837
GU301838
GU296165
GU296166
AB524455
AB524456
GU296167
GU296168
Macroventuria anomochaeta
Massaria anomia
Massaria anomia
Massaria ariae
Massaria aucupariae
Massaria campestris
Massaria conspurcata
Massaria gigantispora
Massaria inquinans
Massaria lantanae
Massaria macra
Massaria mediterranea
Massaria platanoidea
Massaria pyri
Massaria vindobonensis
Massaria vomitoria
Massarina cisti
Massarina eburnea
CBS
CBS
CBS
M52
M49
M28
M14
M26
M19
M18
M3
M45
M7
M21
M27
M13
CBS
CBS
266.62
473.64
GU456315
GU301792
GU301839
HQ599382
HQ599384
HQ599385
HQ599393
HQ599397
HQ599402
HQ599406
HQ599408
HQ599417
HQ599420
HQ599424
HQ599429
HQ599437
FJ795447
GU301840
GU296130
GU296169
HQ599456
HQ599455
HQ599449
HQ599441
HQ599447
HQ599444
HQ599443
HQ599450
HQ599452
HQ599457
HQ599445
HQ599448
HQ599440
FJ795490
GU296170
Massarina igniaria
Massarina ricifera
CBS 845.96
JK 5535 F
GU301841
GU479793
GU296171
GU479759
525.71
123109
591.78
DQ782383
GU296163
GU296164
DQ678017
RPB2
TEF1
GU456357
GU371790
DQ470894
GU456272
GU349070
DQ471062
DQ782386
GU371791
FJ795457
DQ677965
DQ782387
DQ677912
AB521731
GU327760
AB539096
AB539097
GU371792
GU371760
GU456346
GU349071
GU349072
AB524817
AB524818
GU349073
GU456262
GU349062
GU371769
HQ599459
HQ599460
HQ599462
HQ599464
HQ599466
FJ795464
GU371732
GU371793
HQ599322
HQ599324
HQ599325
HQ599333
HQ599337
HQ599342
HQ599346
HQ599348
HQ599357
HQ599359
HQ599363
HQ599368
HQ599375
GU349040
Fungal Diversity
Table 3 (continued)
Species
Culture/voucher1
LSU
SSU
RPB2
TEF1
Massariosphaeria phaeospora
Mauritiana rhizophorae
Mauritiana rhizophorae
Melanomma pulvis-pyrius
Melanomma pulvis-pyrius
Melanomma pulvis-pyrius
Melanomma rhododendri
Misturatosphaeria aurantonotata
Misturatosphaeria aurantonotata
Misturatosphaeria claviformis
Misturatosphaeria kenyensis
CBS 611.86
BCC 28866
BCC 28867
CBS 124080
CBS 371.75
SMH 3291
ANM 73
GKM1238
GKM1280
GKM1210
GKM1195
GU301843
GU371824
GU371825
GU456323
GU301845
GU385197
GU385198
GU385173
GU385174
GU385212
GU385194
GU296173
GU371832
GU371833
GU456302
GU371794
GU371796
GU371797
GU456350
GU371798
GU371817
GU371818
GU456265
GU349019
Misturatosphaeria kenyensis
Misturatosphaeria minima
Misturatosphaeria tennesseensis
Misturatosphaeria uniseptata
Monascostroma innumerosum
Monotosporella tuberculata
Montagnula anthostomoides
Montagnula opulenta
Morosphaeria ramunculicola
Morosphaeria ramunculicola
Morosphaeria velataspora
Morosphaeria velataspora
Massariosphaeria grandispora
Massariosphaeria typhicola
Neophaeosphaeria filamentosa
Neotestudina rosatii
Neottiosporina paspali
Ophiosphaerella herpotricha
GKM L100Na
GKM169N
ANM911
SMH4330
CBS 345.50
CBS 256.84
CBS 615.86
CBS 168.34
BCC 18405
JK 5304B
BCC 17059
BCC 17058
CBS 613 86
CBS 123126
CBS 102202
CBS 690.82
CBS 331.37
CBS 240.31
GU385189
GU385165
GU385207
GU385167
GU301850
GU301851
GU205223
DQ678086
GQ925854
GU479794
GQ925852
GQ925851
GU301842
GU301844
GQ387577
Ophiosphaerella herpotricha
Ophiosphaerella sasicola
Paraconiothyrium minitans
Paraphaeosphaeria michotii
Paraphaeosphaeria michotii
Phaeosphaeria ammophilae
Phaeosphaeria avenaria
Phaeosphaeria avenaria
Phaeosphaeria brevispora
Phaeosphaeria brevispora
Phaeosphaeria caricis
Phaeosphaeria elongata
Phaeosphaeria eustoma
Phaeosphaeria luctuosa
Phaeosphaeria nigrans
Phaeosphaeria nodorum
Phaeosphaeria oryzae
Phaeosphaeriopsis musae
CBS 620.86
MAFF 239644
CBS 122788
CBS 591.73
CBS 652.86
CBS 114595
CBS 602.86
DAOM 226215
MAFF 239276
NBRC 106240
CBS 120249
CBS 120250
CBS 573.86
CBS 308.79
CBS 576.86
CBS 259.49
CBS 110110
CBS 120026
DQ678062
AB524599
EU754173
GU456326
GU456325
GU301859
AY544684
AY544684
AB524600
AB524601
GU301860
GU456327
DQ678063
GU301861
GU456331
GU456332
GQ387591
GU301862
Phoma apiicola
Phoma betae
CBS 285.72
CBS 109410
GU238040
EU754178
GU238211
EU754079
EU754172
DQ767656
GU327761
GU327762
GU327763
GU327767
GU327766
GU327768
GU327769
GU327770
GU349033
GU349006
GU296179
GU205246
AF164370
GQ925839
GU479760
GQ925841
GQ925840
GU296172
GU296174
GQ387516
DQ384069
EU754073
DQ767650
DQ677984
GU479831
GU371725
GU371795
GU371773
GU349084
GU371779
DQ767645
GU349079
DQ767639
DQ678010
AB524458
EU754074
GU456305
GU456304
GU296185
AY544725
AY544725
AB524459
AB524460
DQ677958
AB539098
GU371776
GU456352
GU456351
GU371724
DQ677941
DQ677941
AB539099
AB539100
GU456306
DQ678011
GU456345
DQ677959
DQ677905
AB539111
GU349083
GU456267
GU456266
GU349035
DQ677885
DQ677885
AB539112
AB539113
GU349005
GU456261
DQ677906
GU349004
GU456271
GU456285
GU456356
GQ387530
GU296186
GU349036
GU349037
GU371774
GU349075
Fungal Diversity
Table 3 (continued)
Species
Culture/voucher1
LSU
SSU
RPB2
TEF1
Phoma complanata
Phoma cucurbitacearum
Phoma exigua
Phoma glomerata
Phoma herbarum
Phoma radicina
Phoma valerianae
Phoma vasinfecta
Phoma violicola
Phoma zeae-maydis
Platychora ulmi
CBS
CBS
CBS
CBS
CBS
CBS
CBS
CBS
CBS
CBS
CBS
EU754180
GU301863
EU754183
EU754184
DQ678066
EU754191
GU238150
GU238151
GU238156
EU754192
EF114702
EU754081
GU371778
GU371767
GU371780
GU371781
DQ677962
GU349078
Lophiostoma compressum
Lophiostoma scabridisporum
Lophiostoma scabridisporum
Pleomassaria siparia
Pleospora ambigua
Pleospora herbarum
Polyplosphaeria fusca
Polyplosphaeria fusca
Preussia funiculata
Preussia lignicola
Preussia terricola
Prosthemium betulinum
Prosthemium canba
Prosthemium orientale
Prosthemium stellare
Pseudotetraploa curviappendiculata
Pseudotetraploa curviappendiculata
Pseudotetraploa javanica
GKM1048
BCC 22836
BCC 22835
CBS 279.74
CBS 113979
CBS 191.86
CBS 125425
MAFF 239687
CBS 659.74
CBS 264.69
DAOM 230091
CBS 127468
JCM 16966
JCM 12841
CBS 126964
CBS 125426
MAFF 239495
MAFF 239498
GU385204
GQ925845
GQ925844
DQ678078
AY787937
DQ247804
AB524607
AB524606
GU301864
GU301872
AY544686
AB553754
AB553760
AB553748
AB553781
AB524610
AB524608
AB524611
Pseudotetraploa longissima
Pseudotrichia guatopoensis
Pyrenochaeta acicola
Pleurophoma cava
Pyrenochaeta corn
Pyrenochaeta nobilis
Pyrenochaeta nobilis
Pyrenochaeta quercina
Pyrenochaeta unguis-hominis
Pyrenochaetopsis decipiens
Pyrenophora phaeocomes
Pyrenophora tritici-repentis
Quadricrura bicornis
Quadricrura meridionalis
Quadricrura septentrionalis
Quintaria lignatilis
Quintaria lignatilis
Quintaria submersa
MAFF 239497
SMH4535
CBS 812.95
CBS 257.68
CBS 248.79
CBS 292.74
CBS 407.76
CBS 115095
CBS 378.92
CBS 343.85
DAOM 222769
OSC 100066
CBS 125427
CBS 125684
CBS 125428
BCC 17444
CBS 117700
CBS 115553
AB524612
GU385202
GQ387602
EU754199
GQ387608
GQ387615
DQ678096
GQ387619
GQ387621
GQ387624
DQ499596
AY544672
AB524613
AB524614
AB524617
GU479797
GU301865
GU301866
Repetophragma ontariense
Rimora mangrovei
HKUCC 10830
JK 5246A
DQ408575
GU301868
268.92
133.96
431.74
528.66
276.37
111.79
630.68
539.63
306.68
588.69
361.52
EU754084
EU754085
DQ678014
EU754092
GU238229
GU238230
GU238231
EU754093
EF114726
GU349080
GU349081
DQ677909
GU349076
GU371782
GU349082
GQ925832
GQ925831
DQ678027
GU479829
GU479830
DQ677976
GU327772
GU479856
GU479857
DQ677923
DQ247812
AB524466
AB524465
GU296187
GU296197
AY544726
AB553644
AB553646
AB553641
AB553650
AB524469
AB524467
AB524470
DQ247794
DQ471090
AB524822
GU371799
GU371765
DQ470895
GU349032
GU349027
DQ471063
AB524825
AB524826
AB524471
AB524827
GU327774
GQ387541
EU754100
GQ387547
GQ387554
DQ677991
DQ677936
GQ387558
GQ387560
GQ387563
DQ499595
DQ497614
AB524472
AB524473
AB524476
GU479764
GU296188
GU479832
GU371761
DQ497607
DQ677882
AB524828
AB524829
AB524832
GU479859
GU296193
DQ435077
GU371759
GU349003
Fungal Diversity
Table 3 (continued)
Species
Culture/voucher1
LSU
SSU
Rimora mangrovei
Roussoella hysterioides
Roussoella hysterioides
Roussoella pustulans
Roussoellopsis tosaensis
Saccothecium sepincola
Salsuginea ramicola
Salsuginea ramicola
Setomelanomma holmii
Setosphaeria monoceras
Massaria platani
JK 5437B
CBS 125434
MAFF 239636
MAFF 239637
MAFF 239638
CBS 278.32
KT 2597.1
KT 2597.2
CBS 110217
AY016368
CBS 221.37
GU479798
AB524622
AB524621
AB524623
AB524625
GU301870
GU479800
GU479801
GU301871
AY016368
DQ678065
GU479765
AB524481
AB524480
AB524482
Sporormiella minima
Stagonospora macropycnidia
Tetraploa aristata
Tetraplosphaeria nagasakiensis
Lophiostoma macrostomoides
Lophiostoma macrostomoides
Thyridaria rubronotata
Tingoldiago graminicola
Trematosphaeria pertusa
Trematosphaeria pertusa
Trematosphaeria pertusa
Triplosphaeria cylindrica
Triplosphaeria maxima
Triplosphaeria yezoensis
Ulospora bilgramii
Verruculina enalia
Verruculina enalia
Westerdykella cylindrica
CBS 524.50
CBS 114202
CBS 996.70
MAFF 239678
GKM1033
GKM1159
CBS 419.85
KH 68
CBS 122368
CBS 122371
SMH 1448
MAFF 239679
MAFF 239682
CBS 125436
CBS 110020
BCC 18401
BCC 18402
CBS 454.72
DQ678056
GU301873
AB524627
AB524630
GU385190
GU385185
GU301875
AB521743
FJ201990
GU301876
GU385213
AB524634
AB524637
AB524638
DQ678076
GU479802
GU479803
AY004343
Westerdykella dispersa
Westerdykella ornata
Wicklowia aquatica
Wicklowia aquatica
Xenolophium applanatum
Xenolophium applanatum
Zopfia rhizophila
CBS 508.75
CBS 379.55
AF289-1
CBS 125634
CBS 123123
CBS 123127
CBS 207.26
DQ468050
GU301880
GU045446
GU045445
GU456329
GU456330
DQ384104
RPB2
TEF1
GU296195
GU479767
GU479768
GU296196
AB539102
AB539101
AB539103
AB539104
GU371745
GU479833
GU479834
GU371800
AB539115
AB539114
AB539116
AB539117
GU349029
GU479861
GU479862
GU349028
DQ678013
DQ677961
DQ677908
DQ678003
GU296198
AB524486
AB524489
DQ677950
GU371728
DQ677897
GU349026
AB524836
AB524837
GU327776
GU327778
GU349002
AB521726
FJ201991
GU348999
FJ795476
GU371801
GU456276
GU349085
AB524493
AB524496
AB524497
DQ678025
GU479770
GU479771
AY016355
DQ677974
GU479835
GU479836
DQ470925
AB524844
DQ677921
GU479863
GU479864
DQ497610
U42488
GU296208
GU371803
GU349021
GU456354
GU456355
GU456269
GU456270
GU266232
GU456312
GU456313
L76622
1
BCC Belgian Coordinated Collections of Microorganisms; CABI International Mycological Institute, CABI-Bioscience, Egham, Bakeham Lane,
U.K.; CBS Centraalbureau voor Schimmelcultures, Utrecht, The Netherlands; DAOM Plant Research Institute, Department of Agriculture
(Mycology), Ottawa, Canada; DUKE Duke University Herbarium, Durham, North Carolina, U.S.A.; HHUF Herbarium of Hirosaki University,
Japan; IFRDCC Culture Collection, International Fungal Research & Development Centre, Chinese Academy of Forestry, Kunming, China;
MAFF Ministry of Agriculture, Forestry and Fisheries, Japan; NBRC NITE Biological Resource Centre, Japan; OSC Oregon State University
Herbarium, U.S.A.; UAMH University of Alberta Microfungus Collection and Herbarium, Edmonton, Alberta, Canada; UME Herbarium of the
University of Umeå, Umeå, Sweden; Culture and specimen abbreviations: ANM A.N. Miller; CPC; P.W. Crous; EB E.W.A. Boehm; EG E.B.G.
Jones; GKM G.K. Mugambi; JK J. Kohlmeyer; KT K. Tanaka; SMH S.M. Huhndorf
Cucurbitariaceae/Didymosphaeriaceae, Delitschiaceae,
Didymellaceae, Dothidotthiaceae, Hypsostromataceae, Lentitheciaceae, Leptosphaeriaceae, Lindgomycetaceae,
Lophiostomataceae, Massariaceae, Massarinaceae, Melanommataceae, Montagnulaceae, Morosphaeriaceae,
Phaeosphaeriaceae, Pleomassariaceae, Pleosporaceae,
Sporormiaceae, Testudinaceae/Platystomaceae, Tetraplosphaeriaceae, Trematosphaeriaceae and Zopfiaceae (Plate 1).
Of these, Lentitheciaceae, Massarinaceae, Montagnulaceae,
Morosphaeriaceae and Trematosphaeriaceae form a robust
Fungal Diversity
clade in the present study and in previous studies (Schoch et
al. 2009; Zhang et al. 2009a, b). We thus emended the
suborder, Massarineae, to accommodate them.
Pleosporales suborder Massarineae Barr, Mycologia 71:
948. (1979a). emend.
Habitat freshwater, marine or terrestrial environment,
saprobic. Ascomata solitary, scattered or gregarious, globose, subglobose, conical to lenticular, immersed, erumpent
to superficial, papillate, ostiolate. Hamathecium of dense or
rarely few, filliform pseudoparaphyses. Asci bitunicate,
fissitunicate, cylindrical, clavate or broadly clavate, pedicellate. Ascospores hyaline, pale brown or brown, 1 to 3 or
more transverse septa, rarely muriform, narrowly fusoid,
fusoid, broadly fusoid, symmetrical or asymmetrical, with
or without sheath.
Accepted genera of Pleosporales
Acrocordiopsis Borse & K.D. Hyde, Mycotaxon 34: 535
(1989). (Pleosporales, genera incertae sedis)
Generic description
Habitat marine, saprobic. Ascomata seated in blackish
stroma, scattered or gregarious, superficial, conical to
semiglobose, ostiolate, carbonaceous. Hamathecium of
dense, long trabeculate pseudoparaphyses. Asci 8-spored,
cylindrical with pedicels and conspicuous ocular chambers.
Ascospores hyaline, 1-septate, obovoid to broadly fusoid.
Anamorphs reported for genus: none.
Literature: Alias et al. 1999; Barr 1987a; Borse and Hyde
1989.
Type species
Acrocordiopsis patilii Borse & K.D. Hyde, Mycotaxon 34:
536 (1989). (Fig. 1)
Ascomata 1–2 mm high×1.8–3 mm diam., scattered or
gregarious, superficial, conical or semiglobose, with a
flattened base not easily removed from the substrate, ostiolate,
black, very brittle and carbonaceous and extremely difficult to
cut (Fig. 1a and b). Peridium 250–310 μm thick, to 600 μm
thick near the apex, thinner at the base, comprising three
types of cells; outer cells pseudoparenchymatous, small
heavily pigmented thick-walled cells of textura epidermoidea, cells 0.6–1×6–10 μm diam., cell wall 5–9 μm thick;
cells near the substrate less pigmented, composed of cells of
textura prismatica, cell walls 1–3(−5) μm thick; inner cells
less pigmented, comprised of hyaline to pale brown thinwalled cells, merging with pseudoparaphyses (Fig. 1c, d and
e). Hamathecium of dense, long trabeculate pseudoparaphyses, ca. 1 μm broad, embedded in mucilage, hyaline,
Fig. 1 Acrocordiopsis patilii (from IMI 297769, holotype). a
Ascomata on the host surface. b Section of an ascoma. c Section of
lateral peridium. d Section of the apical peridium. e Section of the
basal peridium. Note the paler cells of textura prismatica. f
Cylindrical ascus. g Cylindrical ascus in pseudoparaphyses. h, i
One-septate ascospores. Scale bars: a=3 mm, b=0.5 mm, c=200 μm,
d, e =50 μm, f–i=20 μm
anastomosing and sparsely septate. Asci 140–220×13–
17 μm (x ¼ 165:3 15:6mm, n=10), 8-spored, bitunicate,
fissitunicate, cylindrical, with short pedicels, 15–25(−40) μm
long, with a large and conspicuous ocular chamber (Fig. 1f
and g). Ascospores 17.5–25 × 12.5–15(−20) μm
(x ¼ 21:5 13:6mm, n=10), uniseriate to partially overlapping, ovoid or ellipsoidal, hyaline, 1-septate, not constricted at the septum, smooth-walled (Fig. 1h and i).
Anamorph: none reported.
Material examined: INDIA, Indian Ocean, Malvan
(Maharashtra), on intertidal wood of Avicennia alba Bl.,
30 Oct. 1981 (IMI 297769, holotype).
Notes
Morphology
Acrocordiopsis was formally established by Borse and
Hyde (1989) as a monotypic genus represented by A. patilii
based on its “conical or semiglobose superficial carbonaceous ascomata, trabeculate pseudoparaphyses, cylindrical,
bitunicate, 8-spored asci, and hyaline, 1-septate, obovoid or
ellipsoid ascospores”. Acrocordiopsis patilii was first
collected from mangrove wood (Indian Ocean) as a marine
fungus, and a second marine Acrocordiopsis species
was reported subsequently from Philippines (Alias et al.
1999). Acrocordiopsis is assigned to Melanommataceae
(Melanommatales sensu Barr 1983) based on its ostiolate
ascomata and trabeculate pseudoparaphyses (Borse and
Hyde 1989). Morphologically, Acrocordiopsis is similar to
Astrosphaeriella sensu stricto based on the conical ascomata
and the brittle, carbonaceous peridium composed of thickwalled black cells with rows of palisade-like parallel cells at
the rim area. Ascospores of Astrosphaeriella are, however,
elongate-fusoid, usually brown or reddish brown and
surrounded by a gelatinous sheath when young; as such
they are readily distinguishable from those of Acrocordiopsis.
A new family (Acrocordiaceae) was introduced by Barr
(1987a) to accommodate Acrocordiopsis. This proposal,
however, has been rarely followed and Jones et al. (2009)
assigned Acrocordiopsis to Melanommataceae.
Phylogenetic study
Acrocordiopsis patilii nested within an unresolved clade
within Pleosporales (Suetrong et al. 2009). Thus its familial
placement is unresolved, but use of the Acrocordiaceae
could be reconsidered with more data.
b
Fungal Diversity
Fungal Diversity
Concluding remarks
Acrocordiopsis, Astrosphaeriella sensu stricto, Mamillisphaeria, Caryospora and Caryosporella are morphologically
similar as all have very thick-walled carbonaceous ascomata,
narrow pseudoparaphyses in a gelatinous matrix (trabeculae)
and bitunicate, fissitunicate asci. Despite their similarities, the
shape of asci and ascospores differs (e.g. Mamillisphaeria has
sac-like asci and two types of ascospores, brown or hyaline,
Astrosphaeriella has cylindro-clavate asci and narrowly fusoid
ascospores, both Acrocordiopsis and Caryosporella has
cylindrical asci, but ascospores of Caryosporella are reddish
brown). Therefore, the current familial placement of Acrocordiopsis cannot be determined. All generic types of Astrosphaeriella sensu stricto, Mamillisphaeria and Caryospora
should be recollected and isolated for phylogenetic study.
Aigialus Kohlm. & S. Schatz, Trans. Br. Mycol. Soc. 85:
699 (1985). (Aigialaceae)
Generic description
Habitat marine, saprobic. Ascomata mostly subglobose in
front view, fusoid in sagittal section, rarely subglobose,
scattered, immersed to erumpent, papillate, ostiolate, ostiole
rounded or slit-like, periphysate. Peridium 2-layered. Hamathecium of trabeculate pseudoparaphyses. Asci 8-spored,
cylindrical, pedicellate, with an ocular chamber and conspicuous apical ring. Ascospores ellipsoidal to fusoid, muriform, yellow brown to brown, with terminal appendages.
Anamorphs reported for genus: none.
Literature: Eriksson 2006; Jones et al. 2009; Kohlmeyer
and Schatz 1985; Lumbsch and Huhndorf 2007.
Type species
Aigialus grandis Kohlm. & S. Schatz, Trans. Br. Mycol.
Soc. 85: 699 (1985). (Fig. 2)
Ascomata 1–1.25 mm high×1–1.3 mm diam. in front view,
250–400 μm broad in sagittal section, vertically flattened
subglobose, laterally compressed, scattered, immersed to semiimmersed, papillate, with an elongated furrow at the top of the
papilla, wall black, carbonaceous, ostiolate, ostiole filled with
branched or forked septate periphyses (Fig. 2a). Peridium 70–
100 μm thick laterally, up to 150 μm thick at the apex, thinner
at the base, comprising two cell types, outer layer composed of
small heavily pigmented thick-walled pseudoparenchymatous
cells, cells 1–2 μm diam., cell wall 2–5 μm thick, inner layer
thin, composed of small hyaline cells (Fig. 2b). Hamathecium
of dense, very long trabeculate pseudoparaphyses, 0.8–1.2 μm
broad, embedded in mucilage, anastomosing and branching
above the asci. Asci 450–640×22–35 μm (x ¼ 505 30mm,
n=10), 8-spored, bitunicate, fissitunicate, cylindrical to
cylindro-clavate, with a long furcate pedicel, 90–180 μm long,
with a low truncate ocular chamber and a refractive apical
apparatus (to 12 μm wide×4 μm high) (Fig. 2e and f).
Ascospores 75–95×15–26 μm (x ¼ 84:3 17:5mm, n=10),
obliquely uniseriate and partially overlapping, broadly fusoid
to fusoid with narrowly rounded ends in front view, flat on one
side from side view (14–20 μm thick), yellowish brown, apical
cells usually hyaline, muriform, with 14–17(−18) transversal
septa, 1–3 longitudinal septa in most cells, slightly constricted
at the septa, with a gelatinous cap at each end (Fig. 2c and d).
Anamorph: none reported.
Material examined: BELIZE, Wee-Wee Cay, on submerged wood of roots and branches of Rhizophora mangle
L., Mar. 1983, leg. J. Kohlmeyer (NY, J.K. 4332b, isotype).
Notes
Morphology
Aigialus was formally established by Kohlmeyer and Schatz
(1985) based on its immersed or semi-immersed ascomata
with periphysate ostiole, trabeculate pseudoparaphyses, cylindrical and fissitunicate asci, and distinctive muriform ascospores with gelatinous sheath or caps. There are five accepted
species in the genus, namely A. grandis, A. mangrovei Borse,
A. parvus S. Schatz & Kohlm., A. rhizophorae Borse and A.
striatispora K.D. Hyde (Jones et al. 2009). Aigialus was first
assigned to the Melanommatales, but its familial status was
uncertain (Kohlmeyer and Schatz 1985). Barr (1990b)
included Aigialus in Massariaceae based on its conspicuous
apical ring in the asci and ascospore characters, and this has
subsequently been widely followed (Eriksson 2006; Hawksworth
et al. 1995; Kirk et al. 2001; Lumbsch and Huhndorf 2007).
Phylogenetic study
The generic type of Aigialus (A. grandis) together with
other three marine species, i.e. A. mangrovei, A. parvus as well
as A. rhizophorae form a robust clade on the phylogenetic
tree. Thus a new family, Aigialaceae, was introduced to
accommodate Aigialus together with Ascocratera and Rimora
(Suetrong et al. 2009).
Concluding remarks
The pleosporalean status of Aigialus has been phylogenetically verified, and the single branch containing Aigialus,
Ascocratera and Rimora represents a familial rank of
Aigialaceae (Suetrong et al. 2009).
Amniculicola Yin. Zhang & K.D. Hyde, Mycol. Res. 112:
1189 (2008). (Amniculicolaceae)
Generic description
Habitat freshwater, saprobic. Ascomata solitary, scattered,
or in small groups, initially immersed, becoming erumpent,
Fungal Diversity
Fig. 2 Aigialus grandis (from NY, J.K. 4332b, isotype). a Ascomata
on the host surface. Note the longitudinal slit-like furrow which is the
ostiole. b Section of the peridium. c, d Released ascospores. e
Ascospores in ascus. Note the conspicuous apical ring. f Cylindrical
ascus with a long pedicel. Scale bars: a=1 mm, b=200 μm, c–f=
20 μm
Fungal Diversity
to nearly superficial, globose, subglobose to conical, wall
black, roughened; apex well differentiated into two tuberculate
flared lips surrounding a slit-like ostiole. Peridium thin, 2layered, outer layer composed of small heavily pigmented
thick-walled cells of textura angularis, inner layer composed
of hyaline thin-walled cells of textura angularis. Hamathecium of dense, long trabeculate pseudoparaphyses, embedded
in mucilage, anastomosing between and above the asci. Asci
8-spored, bitunicate, fissitunicate, cylindrical to narrowly
fusoid, short pedicellate, with an ocular chamber and a small
apical apparatus. Ascospores fusoid, hyaline, 1-septate,
constricted at the septum, surrounded by an irregular hyaline
gelatinous sheath.
Anamorphs reported for genus: Anguillospora longissima, Spirosphaera cupreorufescens and Repetophragma
ontariense (Zhang et al. 2008c, 2009c).
Literature: Zhang et al. 2008c, 2009a, c.
Type species
shorter and broader than the lower one, smooth, containing
four refractive globules, surrounded by an irregular hyaline
gelatinous sheath 4–8.5 μm thick, best seen in India ink,
released senescent ascospores are greyish and 3-septate,
strongly constricted at all septa (Fig. 3g).
Anamorph: none reported.
Colonies slow growing, reaching 4 cm diam. after 70 d
growth on Malt Extract Agar (MEA) at 25°C, flat, with
irregular to rhizoidal margin, off-white to grey, reverse
reddish purple to deep reddish purple, the medium is
stained pale yellow.
Material examined: FRANCE, Ariège, Prat Communal,
Ruisseau de Loumet, 1000 m, on partly submerged wood of
Fraxinus excelsior, 8 Aug. 2006, leg. Jacques Fournier (PC
0092661, holotype); 3 Sept. 2004 (BPI 877774; CBS: H17932); Rimont, Ruisseau de Peyrau, 400 m, on driftwood
of Alnus glutinosa (L.) Gaertn., 23 Jul. 2006 (HKU(M)
17515, isotype).
Amniculicola lignicola Ying Zhang & K.D. Hyde, Mycol.
Res. 112: 1189 (2008). (Fig. 3)
Ascomata 350–450 μm high × 300–500 μm diam.,
solitary, scattered, or in small groups of 2–3, initially
immersed, becoming erumpent, to nearly superficial, with
basal wall remaining immersed in host tissue, globose,
subglobose, broadly or narrowly conical, often laterally
flattened, with a flattened base not easily removed from the
substrate, wall black, roughened, often bearing remnants of
wood fibers; apex well differentiated into two tuberculate
flared lips surrounding a slit-like ostiole, 150–250 μm long,
filled with a purplish amorphous matter, oriented in the axis
of the wood fibers; underlying wood stained pale purple
(Fig. 3a and b). Peridium 40–55 μm thick laterally, up to
120 μm thick at the apex, thinner at the base, coriaceous,
2-layered, outer layer composed of small heavily pigmented
thick-walled cells of textura angularis, cells 4–9 μm diam.,
cell wall 2–3 μm thick, apex cells smaller and walls thicker,
inner layer composed of hyaline thin-walled cells of textura
angularis, 8–16 μm diam., in places with columns of textura
prismatica, oriented perpendicular to the ascomatal surface,
and larger, paler cells of textura prismatica towards the
interior and at the base, 10–25 μm (Fig. 3c, d and e).
Hamathecium of dense, long trabeculate pseudoparaphyses
<1 μm broad, embedded in mucilage (Indian ink), anastomosing between and above the asci. Asci 140–184×9–10 μm,
8-spored, bitunicate, fissitunicate, cylindrical to narrowly
fusoid, with a short, narrowed, twisted, furcate pedicel which
is 15–25 μm long, with a low truncate ocular chamber and a
small inconspicuous apical apparatus barely seen in water
(Fig. 3f). Ascospores (20.5-)28–32×(6-)8(−9) μm, obliquely
uniseriate and partially overlapping, broadly fusoid to fusoid
with broadly to narrowly rounded ends, hyaline, 1-septate,
deeply constricted at the median septum, the upper cell often
Notes
Morphology
Amniculicola is a freshwater genus which stains the woody
substrate purple (Zhang et al. 2008c, 2009a, c). This genus
appears only to be reported from Europe. A detailed description
of the generic type was provided by Zhang et al. (2008c).
Phylogenetic study
Three species of Amniculicola cluster together with
Anguillospora longissima, Spirosphaera cupreorufescens and
Repetophragma ontariense as well as Pleospora rubicunda
Niessl (current name Murispora rubicunda (Niessl) Y. Zhang
ter, J. Fourn. & K.D. Hyde) and Massariosphaeria typhicola
(P. Karst.) Leuchtm. (current name Neomassariosphaeria
typhicola (P. Karst.) Yin. Zhang, J. Fourn. & K.D. Hyde). A
new family, i.e. Amniculicolaceae, was introduced to accommodate these taxa (Zhang et al. 2008c, 2009a, c).
Concluding remarks
All of the five teleomorphic taxa within Amniculicolaceae are from freshwater in Europe and their ascomata stain
the woody substrate purple. Purple staining makes taxa of
this family easily recognized in the field.
Anomalemma Sivan., Trans. Br. Mycol. Soc. 81: 328
(1983). (?Melanommataceae)
Generic description
Habitat terrestrial, fungicolous. Ascomata gregarious, superficial, papillate, ostiolate. Peridium composed cells of
pseudoparenchymatous. Asci clavate, 8-spored. Hamathecium of dense, filliform pseudoparaphyses. Ascospores
Fungal Diversity
Fig. 3 Amniculicola lignicola (from PC 0092661, holotype). a
Superficial ascomata gregarious on the host surface. b An erumpent
ascoma with elongated papilla and slit-like ostiole. c Habitat section of
a superficial ascoma. d, e Section of an ascoma and the partial
peridium. f Cylindrical 8-spored ascus with a short pedicel. g Hyaline,
1-septate broadly fusoid ascospores. Scale bars: a=1 mm, b–d=
100 μm, e=50 μm, f, g=20 μm
1- (rarely 2- to 3-) septate, fusoid, reddish brown, constricted
at the main septum.
Anamorphs reported for genus: Exosporiella (= Phanerocorynella) (Sivanesan 1983).
Fungal Diversity
Fig. 4 Anomalemma epochnii (from K(M):143936, syntype). a
Gregarious ascomata on the host surface. b, c Bitunicate asci. Note the
wide pseudoparaphyses. d Section of the apical peridium comprising
thick-walled cells of textura angularis. e–h Fusoid to broadly fusoid
ascospores. Scale bars: a=0.5 mm, b–h=20 μm
Fungal Diversity
Literature: Berkeley and Broome 1866; Keissler 1922; Massee
1887; Saccardo 1878a; Sivanesan 1983.
Type species
Anomalemma epochnii (Berk. & Broome) Sivan., Trans.
Br. Mycol. Soc. 81: 328 (1983). (Fig. 4)
≡ Sphaeria epochnii Berk. & Broome, Ann. Mag. nat.
Hist., Ser. 3 18: 128 (1866).
Ascomata 340–500 μm high×170–286 μm diam., gregarious on the intertwined hyphae, superficial, papillate, wall
black, coriaceous, roughened (Fig. 4a). Peridium composed of
two types of cells, outer layer 17–22 μm wide, composed of
heavily pigmented thick-walled cells of textura angularis,
cells up to 8×13 μm diam., cell wall 1–1.5 μm thick, inner
layer 30–34 μm thick, composed of hyaline thin-walled cells
(Fig. 4d). Hamathecium of dense, long cellular pseudoparaphyses, 2–4 μm broad, septate. Asci 75–108×9.5–12.5 μm
(x ¼ 92:8 11:1mm, n=10), 8-spored, bitunicate, fissitunicate, dehiscence not observed, cylindro-clavate to clavate, with
a furcate pedicel up to 6–25 μm long, with a small ocular
chamber best seen in immature asci (ca. 2 μm wide×1 μm
high) (Fig. 4b and c). Ascospores 20–25(−30)×5–7.5 μm
(x ¼ 23:1 6:3mm, n=10), obliquely uniseriate and partially
overlapping to biseriate, fusoid to narrowly fusoid with narrowly
rounded ends, brown, 1-septate, rarely 2- to 3-septate, deeply
constricted at the median septum, smooth (Fig. 4e, f, g and h).
Anamorph: Exosporiella fungorum (Fr.) P. Karst.
(Sivanesan 1983).
= Epochnium fungorum Fr., Syst. mycol. 3: 449 (1832).
Mycelium composed of branched, septate, pale brown
hyphae. Stroma none. Conidiophores macronematous or
semi-macronematous, mononematous, hyaline, smooth,
branched towards the apex. Conidiogenous cells monoblastic, cylindrical or doliform. Conidia cylindrical or
ellipsoidal, dry, 3-4-septate, smooth, hyaline or pale brown.
Material examined: UK, England, Warleigh near Bath,
on fungus on bark (Epochnium sp.), Mar. 1866, leg.
Warbright? (K(M):143936, syntype, ex herb. C.E.
Broome).
reported having Aposphaeria or Pseudospiropes anamorphs,
which differs from Exosporiella (Sivanesan 1983). In addition, the presence of well developed prosenchymatous stroma
in Sphaeria epochnii can also readily distinguish it from
Melanomma (Sivanesan 1983).
The gregarious ascomata and formation of prosenchymatous stroma of Anomalemma resembles those of Cucurbitaria,
but the pleosporaceous dictyosporous ascospores of Cucurbitaria readily distinguish it from Anomalemma epochnii. In
addition, the pseudoparenchymatous peridium, fungicolous
habitat and brown 1-septate ascospores, which later becoming
3-septate differ from any other pleosporalean genus. Thus a
new genus, Anomalemma, was introduced to accommodate it
(Sivanesan 1983). Anomalemma is presently monotypic.
Phylogenetic study
None.
Concluding remarks
Anomalemma epochnii certainly resembles Byssosphaeria in its ascomata clustering together in groups on closely
intertwined hyphae and brown ascospores, and may well be
included in this genus. Its fungicolous habitat, however,
distinguishes it from Byssosphaeria.
Appendispora K.D. Hyde, Sydowia 46: 29 (1994a).
(?Didymellaceae)
Generic description
Habitat terrestrial, saprobic. Ascomata small, clustered,
immersed, subglobose or irregularly pyriform. Peridium
thin. Hamathecium of dense, long trabeculate pseudoparaphyses. Asci 8-spored, bitunicate, fissitunicate, cylindrical,
apical rounded with ocular chamber and faint ring, with
short pedicels. Ascospores uniseriate to partially overlapping,
fusoid, brown, 1-septate, slightly constricted at the septum.
Anamorphs reported for genus: none.
Literature: Hyde 1994a.
Notes
Type species
Morphology
Sphaeria epochnii was first described and illustrated by
Berkeley and Broome (1866) from Britain and the anamorphic
stage is the hyphomycetous Epochniella fungorum. Sphaeria
epochnii has subsequently been transferred to Melanomma (as
M. epochnii (Berk. & Broome) Sacc.; Saccardo 1878a),
Byssosphaeria (as B. epochnii (Berk. & Broome) Cooke;
Massee 1887) and Chaetosphaeria (as C. epochnii (Berk. &
Broome) Keissl.; Keissler 1922). The deposition of Sphaeria
epochnii in Chaetosphaeria is obviously unacceptable, as
Chaetosphaeria has unitunicate asci. Melanomma has been
Appendispora frondicola K.D. Hyde, Sydowia 46: 30
(1994a). (Fig. 5)
Ascomata 120–280 μm high×180–280 μm diam., clustered, immersed with minute ostioles visible through cracks
or blackened dots on the host surface, subglobose or
irregularly pyriform (Fig. 5a and b). Peridium 40 μm thick,
comprising two types of cells; outer cells, small heavily
pigmented thick-walled cells of textura angularis, inner cells
compressed, hyaline. Hamathecium of dense, very long
trabeculate pseudoparaphyses, ca. 1 μm broad, embedded in
mucilage, hyaline, anastomosing (Fig. 5e). Asci 130–144×11–
Fungal Diversity
Fig. 5 Appendispora frondicola (from BRIP 21354, holotype). a Immersed ascomata on
host surface. b Valsoid configuration of the ascomata. c Cylindrical ascus. d Squash showing
asci and numerous pseudoparaphyses. e Thin strands of
anastomosing pseudoparaphyses.
f, g Ascospores with one or two
appendages. Scale bars:
a, b=0.5 mm, c–g=30 μm
13 μm, 8-spored, bitunicate, fissitunicate, cylindrical, with an
ocular chamber and faint ring, with short pedicels (Fig. 5c and
d). Ascospores 21–30×7–9 μm, uniseriate to partially overlapping, fusoid, brown, 1-septate, slightly constricted at the
septum, with an irregular ridged ornamentation and 3–5
narrow appendages at each end (Fig. 5f and g).
Anamorph: none reported.
Material examined: BRUNEL, Jalan, Muara, Simpang
835, on dead rachis of Oncosperma horridum on forest floor,
Nov. 1992, K.D. Hyde 1652 (BRIP 21354, holotype).
Notes
Morphology
Appendispora was described as a saprobe of palm, and is
characterized by small, immersed ascomata, bitunicate,
fissitunicate asci, trabeculate pseudoparaphyses, brown, 1septate, appendaged ascospores with irregular wall striations (Hyde 1994a). Based on its trabeculate pseudoparaphyses embedded within gel matrix and its brown
ascospores, Appendispora was assigned to Didymosphaeriaceae (Barr 1987b; Hyde 1994a).
Phylogenetic study
None.
Concluding remarks
The saprobic habitat and association with monocots, cylindrical asci, trabeculate pseudoparaphyses as
well as its brown, 1-septate ascospores make it
difficult to determine a better phylogenetic position than
Didymellaceae.
Fungal Diversity
Ascorhombispora L. Cai & K.D. Hyde, Cryptog. Mycol.
28: 294 (2007). (Pleosporales, genera incertae sedis)
Generic description
Habitat freshwater, saprobic. Ascomata solitary or gregarious,
superficial, globose to subglobose, dark brown to black, short
papillate, ostiolate, coriaceous. Peridium relatively thin,
textura angularis in longitudinal section, 2-layered. Hamathecium not observed. Asci 8-spored, obpyriform, broadly clavate
to saccate, pedicellate, bitunicate, apex rounded, persistent.
Ascospores overlapping 2-3-seriate, broadly fusoid to rhomboid, thick-walled, surrounded by mucilaginous sheath, 3euseptate, not constricted at septa, median septum wide,
forming a darker band, central cells large, trapezoid, dark
brown to black, verruculose, polar end cells small and paler.
Anamorphs reported for genus: none.
Literature: Cai and Hyde 2007.
and trapezoid (rhombic), 3-septate, dark brown to black
ascospores with smaller end cells which are subhyaline to
pale brown. Ascorhombispora is most comparable with
Caryospora and Zopfia. But the globose to subglobose
ascomata and thin peridium, saccate asci lacking interascal
pseudoparaphyses, and the 3-septate, rhomboid ascospores
with the paler end cells of Ascorhombispora differs from
those of Caryospora (Cai and Hyde 2007).
Phylogenetic study
Phylogenetic analysis based on either SSU or LSU
rDNA sequences indicated that Ascorhombispora aquatica
belongs to Pleosporales, but its familial placement was left
undetermined (Cai and Hyde 2007).
Concluding remarks
The sac-shaped asci and absence of pseudoparaphyses
are uncommon in Pleosporales, especially among those
from freshwater.
Type species
Ascorhombispora aquatica L. Cai & K.D. Hyde, Cryptog.
Mycol. 28: 295 (2007). (Fig. 6)
Ascomata 140–170 μm high×150–185 μm diam., solitary
or gregarious, superficial, globose to subglobose, dark brown
to black, short papillate, ostiolate, ostioles rounded, small,
coriaceous. Peridium relatively thin, 10–18 μm wide, textura
angularis in longitudinal section, composed of two layers of
angular cells, outer later dark brown to black, relatively thickwalled, inner layer hyaline, relatively thin-walled (Fig. 6a and
b). Hamathecium not observed. Asci 100–198×72–102 μm
(x ¼ 186 88mm, n=15), 8-spored, obpyriform, broadly
clavate to saccate, pedicellate, bitunicate, apex rounded,
deliquescent (Fig. 6c, d and e). Ascospores 30.5–45×16–
26.5 μm (x ¼ 38:5 21mm, n=25), overlapping 2-3-seriate,
broadly fusoid to rhomboid, thick-walled, surrounded by
mucilaginous sheath, 3-euseptate, not constricted at septa,
median septum wide, forming a darker band, central cells
large, trapezoid, 11–18 μm long, dark brown to black,
verruculose, polar end cells small, hemispherical, 3.5–4 μm
long, subhyaline to pale brown, smooth (Fig. 6f).
Anamorph: none reported.
Material examined: CHINA, Yunnan, Jinghong, on
submerged bamboo in a small forest stream, 26 Jan. 2003,
leg. det. L. Cai, CAI-1H31 (HKU(M) 10859, holotype).
Notes
Morphology
Ascorhombispora was introduced as a monotypic
genus from freshwater by Cai and Hyde (2007), and is
characterized by superficial, coriaceous, non-stromatic
ascomata, large, saccate asci; lack of interascal filaments
Asteromassaria Höhn., Sber. Akad. Wiss. Wien, Math.naturw. Kl., Abt. I 126: 368 (1917). (?Morosphaeriaceae)
Generic description
Habitat terrestrial, saprobic. Ascomata medium-sized,
clustered, at first immersed and then breaking through
the host surface and becoming superficial, globose,
subglobose, coriaceous. Peridium 2-layered, thicker near
the base. Hamathecium of dense, septate, cellular pseudoparaphyses which branch and anastomosing frequently
between and above asci. Asci (4-)8-spored, bitunicate,
cylindro-clavate to clavate, with a short truncated pedicel
and a small ocular chamber. Ascospores obliquely uniseriate and partially overlapping to biseriate, fusoid to
fusoid-ellipsoidal, pale brown when mature, 1-septate,
some becoming 3-septate when old, constricted at the
median septum.
Anamorphs reported for genus: Scolicosporium (Sivanesan
1984).
Literature: Barr 1982a; b; 1993a; Boise 1985; Shoemaker
and LeClair 1975; Sivanesan 1987; Tanaka et al. 2005.
Type species
Asteromassaria macrospora (Desm.) Höhn., F. von, Sber.
Akad. Wiss. Wien, Math.-naturw. Kl., Abt. I 126: 368
(1917). (Fig. 7)
≡ Sphaeria macrospora Desm., Ann. Sci. Nat. Bot. 10:
351 (1849).
Ascomata 400–600 μm high×450–650 μm diam., 4–20
clustered together, at first immersed and then breaking
through the host surface and becoming superficial, globose,
Fungal Diversity
Fig. 6 Ascorhombispora aquatica (from HKU(M) 10859, holotype).
a Section of an ascoma. b Section of a partial peridium. c Immature
ascus. d–f Mature asci with ascospores. Note the deliquescent ascal
wall in f. Note the wide, dark band in the medium septum of
ascospores in d and e and the mucilaginous sheath and paler end cells
in e and f. Scale bars: a=20 μm, b–f=15 μm (figures referred to Cai
and Hyde 2007)
subglobose, not easily removed from the substrate, wall black,
coriaceous, roughened, apex usually widely porate, with or
without papilla (Fig. 7a). Peridium 70–90 μm wide, thicker
near the base where it is up to 180 μm wide, comprising two
cell types, outer cells composed of heavily pigmented
small cells, cells 3–5 μm diam., inner layer composed of
less pigmented cells of textura angularis, 10–20 μm
diam. (Fig. 7b and c). Hamathecium of dense, septate, 2–
3 μm broad, pseudoparaphyses which branch and anasto-
mosing frequently between and above asci (Fig. 7d). Asci
(180-)200–280×28–43 μm (x ¼ 230 35mm, n=10), 8Fig. 7 Asteromassaria macrospora (from L, 1004). a Ascomata
clustered in a group breaking through the host surface. b Section of an
ascoma. c Section of a partial peridium. Note the cells of textura
angularis. d Pseudoparaphyses. Note the branches. e Upper part of the
ascus illustrating the ocular chamber. f Ascus with a short pedicel. g–j
Ascospores. Note the mucilaginous sheath in G and minutely verruculose
ornamentation in J. Scale bars: a=0.5 mm, b, c=100 μm, d–j=10 μm
b
Fungal Diversity
Fungal Diversity
spored (sometimes 4-spored), bitunicate, fissitunicate dehiscence not observed, cylindro-clavate to clavate, with a short
truncated pedicel up to 30 μm, with a small ocular chamber
(ca. 3 μm wide×3 μm high) (Fig. 7e and f). Ascospores 50–
58×(14-)18–21 μm (x ¼ 55:3 18:2mm, n=10), obliquely
uniseriate and partially overlapping to biseriate, fusoid to
fusoid-ellipsoidal, with narrowly rounded ends, lightly
brown when mature, 1-septate, some becoming 3-septate
when old, constricted at the median septum, the upper
cell often broader and longer than the lower one,
minutely verrucose (Fig. 7g, h, i and j).
Anamorph: Scolicosporium macrosporium (Berk.) B.
Sutton.
Acervuli immersed in bark, brown, discrete, up to
250 μm diam., opening by irregular rupture of the
overlaying tissues. Peridium of thin-walled angular cells.
Conidiophores cylindrical, 1-2-septate, up to 30 μm long
and 3–5 μm wide. Conidiogenous cells holoblastic, 1-2annellate, cylindrical, hyaline. Conidia 100–190 × 12–
15 μm, fusoid, pale brown with paler or hyaline ends, 7–
17 transverse septate, smooth-walled, with a tapered apex
and truncate base (adapted from Sivanesan 1984).
Material examined: CZECH REPUBLIC, MährischWelвkirchen (Hranice), Wsetin (Vsetin), Berg Čap., on
Fagus sylvatica L., Aug. 1938, F. Petrak (L, 1004).
angularis peridium and Scolicosporium anamorphic stage
(see under Pleomassaria).
Astrosphaeriella Syd. & P. Syd., Annls mycol. 11: 260
(1913). (?Melanommataceae)
Generic description
Habitat terrestrial, saprobic. Ascomata densely scattered or
in small groups, erumpent through the outer layers of the
host tissues to nearly superficial, reflexed pieces of the
ruptured host tissue usually persisting around the base of
the ascomata, often star-like, conical to semiglobose, with a
central papilla. Peridium upper wall usually comprising a
thick dark brittle pseudoparenchymatous layer, base usually
flattened and thin-walled. Hamathecium of dense, filliform,
trabeculate pseudoparaphyses, embedded in mucilage. Asci
8-spored, bitunicate, fissitunicate, cylindro-clavate to narrowly fusoid. Ascospores narrowly fusoid with acute ends,
hyaline, pale brown or brown, 1-3-septate.
Anamorphs reported for genus: Pleurophomopsis (Hyde
et al. 2011).
Literature: von Arx and Müller 1975; Barr 1990a; Chen
and Hsieh 2004; Hawksworth 1981; Hawksworth and
Boise 1985; Hyde and Fröhlich 1998; Hyde et al. 2000;
Kirk et al. 2001; Sydow and Sydow 1913; Tanaka and
Harada 2005a; b; Tanaka et al. 2009.
Notes
Type species
Morphology
In this study we were unable to obtain the holotype,
so we used a collection of Petrak’s. The main morphological characters of Asteromassaria are the medium- to
large-sized, globose to depressed ascomata opening with a
pore, clavate to oblong asci, narrowly cellular pseudoparaphyses, pale to dark brown, bipolar symmetric, mostly
fusoid, distoseptate or euseptate ascospores (Barr 1993a).
The bipolar symmetric ascospores of Asteromassaria can
readily be distinguished from other genera of this family
(Barr 1993a; Tanaka et al. 2005). Currently, it comprises
12 species (Tanaka et al. 2005; http://www.mycobank.org,
28-02-2009).
Phylogenetic study
Asteromassaria pulchra (Harkn.) Shoemaker & P.M.
LeClair is basal to Morosphaeriaceae in the phylogenetic tree based on four genes, but its placement is
influenced by taxon sampling that was different in
several analyses.
Concluding remarks
Asteromassaria can be distinguished from other comparable genera, i.e. Pleomassaria and Splanchnonema by 1septate and pale brown ascospores, thick-walled textura
Astrosphaeriella stellata Syd. & P. Syd., Annls mycol. 11:
260 (1913). (Fig. 8)
Ascomata 360–570 μm high×860–1150 μm diam.,
densely scattered or in small groups, erumpent through the
outer layers of the host tissues to nearly superficial, reflexed
pieces of the ruptured host tissue usually persisting around
the base of the ascomata, forming star-like flanges around the
ascomata from the surface view; ascomata broadly conical,
with a flattened base not easily removed from the substrate,
wall black; apex with a central papilla which is black and
shiny at maturity, scarcely projecting (Fig. 8a). Peridium 40–
70 μm thick, carbonaceous and crisp, 1-layered, composed of
very small dark brown thick-walled pseudoparenchymatous
cells, cells 2–5 μm diam., cell wall 2–6 μm thick, in places at
the base composed of hyaline cells of textura prismatica,
cells 5×8 μm diam. (Fig. 8b). Hamathecium of dense, very
long trabeculate pseudoparaphyses, <1 μm broad, embedded
in mucilage (Indian ink), anastomosing between and above
the asci. Asci 130–190×11.5–15 μm (x ¼ 161:5 12:8mm,
n=10), 8-spored, bitunicate, fissitunicate, cylindro-clavate to
narrowly fusoid, with a short, narrowed pedicel which is 10–
35 μm long, with a large ocular chamber (Fig. 8c).
Ascospores 35–50×5–7.5 μm (x ¼ 43:4 6mm, n=10),
biseriate, elongate- fusoid, gradually tapering towards the
Fungal Diversity
Fig. 8 Astrosphaeriella fusispora (BISH 145726). a Ascomata forming a small group on
host surface. Note the remains
of the host forming flanges
around the ascomata. b Section
of the partial peridium. Note the
black peridium and wedge of
palisade cells between the lateral
and basal walls. c Asci in trabeculate pseudoparaphyses. d–f
Narrowly fusoid ascospores.
Scale bars: a=1 mm, b=
100 μm, c=50 μm, d=15 μm,
f=10 μm
ends, hyaline, turning pale brown when mature, 1(−3)septate, constricted at the median septum (Fig. 8d,e and f).
Anamorph: none reported.
Material examined: USA, Hawaii, Kapano Gulch, in
bamboo culms, 5 Jun. 1947, leg. Kopf & Rogers, det. Miller
(BISH 145726, as Astrosphaeriella fusispora Syd. & P. Syd.).
Notes
Morphology
Astrosphaeriella has been treated as a synonym of Microthelia (von Arx and Müller 1975), but the large conical
ascomata, numerous trabeculate pseudoparaphyses and 1septate and elongated ascospores of Astrosphaeriella all
disagree with those of Microthelia (Hawksworth 1981). It
was assigned to Platystomaceae by Barr (1990a) in Pleosporales or Melanommataceae by Kirk et al. (2001).
Following a systematic study of Astrosphaeriella, only four
species were accepted, i.e. A. aosimensis I. Hino & Katum.,
A. stellata, A. trochus (Penz. & Sacc.) D. Hawksw. and A.
venezuelensis M.E. Barr & D. Hawksw. (Hawksworth 1981),
and it was defined as a tropical genus, occurring exclusively
on palms or bamboo. Astrosphaeriella stellata was selected
as the type of Astrosphaeriella, and A. fusispora was regarded
Fungal Diversity
as a synonym of A. stellata (Hawksworth 1981). More taxa
were subsequently added (Barr 1990a; Hawksworth and
Boise 1985; Hyde and Fröhlich 1998), and the generic
concept extended to include three elements: 1. typical semiimmersed to superficial ascomata with flattened base,
cylindro-clavate asci with fusoid ascospores and trabeculate
pseudoparaphyses, i.e. Astrosphaeriella sensu stricto (e.g.
A. fusispora and A. vesuvius (Berk. & Broome) D. Hawksw.
& Boise); 2. Trematosphaeria-like with rounded ascomata (e.
g. A. africana D. Hawksw.); and 3. Massarina-like species
with immersed ascomata (e.g. A. bakeriana (Sacc.) K.D.
Hyde & J. Fröhl.) (Chen and Hsieh 2004; Tanaka and Harada
2005a; b). Currently, a broad generic concept of Astrosphaeriella is accepted, and 47 taxa are included in Astrosphaeriella.
Phylogenetic study
Phylogenetic analysis based on LSU and SSU nurDNA
sequence data indicates that Astrosphaeriella is polyphyletic,
and located in the basal region of the Pleosporales between
Testudinaceae and Zopfiaceae/Delitschiaceae (Tanaka et al.
2009), or basal to Aigialaceae (Schoch et al. 2009). The
genus is, however, clearly not related to Trematosphaeria as
previously understood (Boise 1985).
Concluding remarks
Astrosphaeriella is currently polyphyletic and new
collections of the different elements listed above are needed
in order to understand the placement of various species. We
suggest that some immersed bambusicolous species may
belong in Tetraplospheariaceae.
Ascomata 675–950 μm high×875–1500 μm diam.,
solitary or in small groups of 2–10, immersed and forming
slightly protruding domes on the substrate surface, with nearwhite rim around the central ostiole; in vertical view lenticular,
multi- or rarely unilocular, individual locules 175–270 μm
high×320–400 μm diam., with a flattened base, ostiole a
central opening without tissue differentiation (Fig. 9a). Upper
peridium 32–70 μm wide, carbonaceous, composed of a few
layers of black walled cells of textura angularis. Lower
peridium thinner, composed of hyaline cells of textura
globulosa or textura prismatica (Fig. 9b). Hamathecium of
long trabeculate pseudoparaphyses, 1.2–1.6(−2) μm wide,
branching and anastomosing between and above asci,
embedded in mucilage. Asci 137.5–207.5×26–35 μm
(x ¼ 172:8 31:5mm, n=20), 8-spored, bitunicate, fissitunicate dehiscence not observed, clavate, with short pedicel (to
25 μm), with ocular chambers (ca. 3 μm wide×4 μm high)
(Fig. 9c and d). Ascospores 35–55 × 10.5–15 μm
(x ¼ 44:7 12:4mm, n=50), biseriate, navicular to obovoid,
hyaline, becoming pale brown when senescent, straight or
usually curved, smooth, asymmetric, 1-septate, the upper cell
larger with a rounded end, basal cell with a tapering end,
constricted at the septum, with spreading mucilaginous sheath
(Fig. 9e, f and g) (data from Fröhlich and Hyde 1998).
Anamorph: none reported.
Material examined: AUSTRALIA, North Queensland,
Palmerston, Palmerston National Park, on dead rattan of
Calamus caryotoides A.Cunn. ex Mart., Mar. 1994, J.
Fröhlich (HKU(M) 7794, holotype).
Notes
Asymmetricospora J. Fröhl. & K.D. Hyde, Sydowia 50:
183 (1998). (?Melanommataceae)
Generic description
Habitat terrestrial, saprobic. Ascomata solitary or in small
groups, immersed, black, lenticular in section, uni- or often
multi-locular, with a central ostiole without tissue differentiation. Upper peridium carbonaceous, thicker at sides and apex.
Lower peridium composed of irregular-shaped, hyaline cells.
Hamathecium of trabeculate pseudoparaphyses, branching and
anastomosing between and above asci, embedded in mucilage.
Asci 8-spored, bitunicate, fissitunicate unknown, clavate, short
pedicellate. Ascospores 1-septate, hyaline, constricted at the
septum, with a broad, spreading mucilaginous sheath.
Anamorphs reported for genus: none.
Literature: Fröhlich and Hyde 1998.
Morphology
Asymmetricospora was introduced as a monotypic genus
represented by A. calamicola based on its “absence of a
subiculum, the absence of short dark setae around the papilla
and its asymmetric ascospores” (Fröhlich and Hyde 1998).
Because of the immersed ascomata, ostiole and peridium
morphology, fissitunicate asci and trabeculate pseudoparaphyses, Asymmetricospora was assigned to Melanommataceae
(sensu Barr 1990a; Fröhlich and Hyde 1998).
Morphologically Asymmetricospora can be distinguished
from its most comparable genus, Astrosphaeriella, by its
ostiole, which is a simple opening without tissue differentiation, asymmetric ascospores, and the usually multiloculate fruiting body (Fröhlich and Hyde 1998).
Phylogenetic study
None.
Type species
Asymmetricospora calamicola J. Fröhl. & K.D. Hyde,
Sydowia 50: 184 (1998). (Fig. 9)
Concluding remarks
The placement of Asymmetricospora under Melanommataceae remains to be confirmed.
Fungal Diversity
Fig. 9 Asymmetricospora calamicola (from HKU(M) 7794, holotype). a Ascomata immersed in the substrate. b Section of the
peridium. c Mature and immature asci in pseudoparaphyses (in cotton
blue). d Clavate ascus with a small ocular chamber. e–g Ascospores
with sheath. Scale bars: a, b=0.5 mm, c=50 μm, d–g=20 μm
Barria Z.Q. Yuan, Mycotaxon 51: 313 (1994).
(Phaeosphaeriaceae)
Asci bitunicate, fissitunicate, cylindrical to clavate, with a short,
furcate pedicel. Ascospores ellipsoid, hyaline at first, turning
brown at maturity, 1-septate, strongly constricted at the septum.
Anamorphs reported for genus: none.
Literature: Yuan 1994.
Generic description
Habitat terrestrial, parasitic. Ascomata small- to medium-sized,
solitary or scattered, immersed, globose, subglobose, ostiolate,
coriaceous. Apex with or without papilla and with a pore-like
ostiole. Peridium 2-layered. Hamathecium of dense, long
cellular pseudoparaphyses, septate, embedded in mucilage.
Type species
Barria piceae Z.Q. Yuan, Mycotaxon 51: 314 (1994).
(Fig. 10)
Fungal Diversity
Fig. 10 Barria piceae (from
NY 92003, isotype). a Ascoma
on the host surface. Note the
wide opening ostiole. b Section
of the partial peridium with two
types of cells. c, d Asci with
ocular chambers and short pedicels. e, f Ellipsoid ascospores
which are turning brown with
thin sheath around them. Scale
bars: a=0.5 mm, b=50 μm, c,
d=20 μm, e, f=10 μm
Ascomata 240–370 μm high × 200–320 μm diam.,
solitary, scattered, immersed, globose, subglobose, coriaceous, apex with or without papilla and with a pore-like
ostiole (Fig. 10a). Peridium 20–35 μm thick, comprising
two cell types, the outer cells comprising 3–4 layers of
brown pseudoparenchymatous cells, cells 4–5 μm diam.,
cell wall 2–3 μm thick, inner cells comprising 3–4 layers of
pale brown compressed cells, cells 2×16 μm diam., cell
wall 0.5–1.5 μm thick (Fig. 10b). Hamathecium of dense,
long cellular pseudoparaphyses, 2–3 μm broad, septate.
Asci 135–200(−220)×14–20 μm (x ¼ 156 16:6mm, n=
10), 8-spored, bitunicate, fissitunicate, cylindrical to cla-
vate, with a short, furcate pedicel, up to 22 μm long, with a
large ocular chamber (ca. 4 μm wide×3 μm high) (Fig. 10c
and d). Ascospores 19–21.5×10–12 μm (x ¼ 20:4 11mm,
n=10), uniseriate to partially overlapping, ellipsoid, hyaline
or greenish with numerous small guttules at first and olive
green to smoky brown at maturity, 1-septate, strongly
constricted at the septum, foveolate, surrounded with sheath
(Fig. 10e and f).
Anamorph: none reported.
Material examined: CHINA, Xinjiang Province, Uygur,
Urumqi, Tianshan Mountain, on needles of Picea schrenkiana, 1 Jul. 1992, Z.Q. Yuan (NY 92003, isotype).
Fungal Diversity
Notes
Type species
Morphology
Barria was established by Yuan (1994) as a monotypic
genus represented by B. piceae according to its “two-celled,
pigmented ascospores, pseudoparenchymatous peridium and
narrowly cellular pseudoparaphyses” thus differing in its
combination of characters from all of the morphologically
related dothideomycetous genera, such as Didymosphaeria,
Didymopleella or Stegasphaeria. The taxon was considered to belong in Phaeosphaeriaceae. Ascomata and
colour or shape of ascospores, however, readily distinguish it from other 1-septate Phaeosphaeriaceae genera,
i.e. Didymella, Lautitia and Metameris (Yuan 1994).
Barria piceae causes blight of spruce needles.
Belizeana tuberculata Kohlm. & Volkm.-Kohlm., Bot.
Mar. 30: 196 (1987). (Fig. 11)
Ascomata 170–300 μm high×160–290 μm diam., solitary, scattered, or in small groups of 2–3, immersed to semiimmersed, subglobose to broadly ampulliform, carbonaceous,
black, pale brown on the sides, ostiolate, epapillate or shortly
papillate, ostiolar canal filled with a tissue of hyaline cells
(Fig. 11a). Peridium 25–35 μm wide, comprising several
layers thin-walled cells of textura angularis, which are
hyaline inwardly, near the base composed of a hyaline
hyphal mass producing asci, up to 20 μm thick (Fig. 11b, c
and e). Hamathecium of dense, ca. 2 μm broad, filliform
pseudoparaphyses, rarely branched, embedded in mucilage
(Fig. 11g). Asci 145–170×20–30 μm (x ¼ 163 25mm, n=
10), 8-spored, bitunicate, fissitunicate, broadly cylindrical to
clavate with a short pedicel, thick-walled, with a small ocular
chamber (Fig. 11d, f and h). Ascospores 21–26 × 13–18 μm
(x ¼ 22 15mm, n = 10), uniseriate, broadly ellipsoidal,
hyaline, turn pale brown when senescent, 1-septate, constricted at the septum, thick-walled, 2-layered, mature spores
with tuberculate ornamentation between the two layers
(Fig. 11i and j).
Anamorph: Phoma-like (Kohlmeyer and VolkmannKohlmeyer 1987).
Material examined: BELIZE, Twin Cays, on Laguncularia sp., 7 Apr. 1983, leg. & det. J. Kohlmeyer (Herb. J.
Kohlmeyer No. 4398, holotype); AUSTRALIA, Towra
Point, New South Wales, trunk of eroded tree with oysters
and shipworms, intertidal zone, Botany Bay, 23 Aug. 1981
(Herb. J. Kohlmeyer No. 4209, paratype).
Phylogenetic study
None.
Concluding remarks
The status of Barria with its unusual verrucose ascospores and thick gel coating is uncertain. In many ways it
resembles Belizeana, with its cylindrical asci, 1-septate,
ellipsoid ascospores with sheath and verruculose surface
(Kohlmeyer and Volkmann-Kohlmeyer 1987). However,
the latter is a marine genus while Barria causes leaf blight
of terrestrial Picea (Yuan 1994). The placement in
Phaeosphaeriaceae seems logical based on the parasitic
life style, thin and simple peridium, wide cellular pseudoparaphyses and brown ascospores. However, molecular
data are needed to confirm this.
Belizeana Kohlm. & Volkm.-Kohlm., Bot. Mar. 30: 195
(1987). (Pleosporales, genera incertae sedis)
Generic description
Habitat marine, saprobic. Ascomata solitary, scattered, or
in small groups, medium-sized, immersed to semiimmersed, subglobose to broadly ampulliform, black,
ostiolate, carbonaceous. Peridium thin, comprising several
layers of brown thin-walled cells of textura angularis.
Hamathecium of dense, filliform pseudoparaphyses, rarely
branched. Asci 8-spored, bitunicate, fissitunicate, broadly
cylindrical to clavate, with a short pedicel and an ocular
chamber. Ascospores uniseriate, broadly ellipsoidal, hyaline, turn pale brown when senescent, 1-septate, constricted at the septum, thick-walled, 2-layered, mature
spores with tuberculate ornamentation between the two
layers.
Anamorphs reported for genus: Phoma-like (Kohlmeyer
and Volkmann-Kohlmeyer 1987).
Literature: Kohlmeyer and Volkmann-Kohlmeyer 1987.
Notes
Morphology
Belizeana was formally established to accommodate B.
tuberculata, an obligate marine fungus, which is characterized by verrucose ascospores (Kohlmeyer and VolkmannKohlmeyer 1987). Belizeana tuberculata can be assigned to
Pleosporaceae (Pleosporales) according to Luttrell’s (1973)
treatment and keys of von Arx and Müller (1975), but cannot
resolve a proper family based on Barr (1979a, 1983). The
unique morphology together with obligate marine habitat
makes B. tuberculata readily distinguishable from all other
taxa of Pleosporaceae.
Phylogenetic study
None.
Concluding remarks
The ascospores of Belizeana tuberculata are most comparable with those of Acrocordiopsis patilii, but the superficial
Fungal Diversity
Fig. 11 Belizeana tuberculata
(from Herb. J. Kohlmeyer No.
4398, holotype). a Immersed to
semi-immersed ascomata. b, e
Vertical section of an ascoma. c
Section of a partial peridium. d
Squash mounts with a large
number of asci. f Broadly
cylindrical ascus with a large
ocular chamber. g Filliform
pseudoparaphyses. h Apical part
of an ascus. Note the large
ocular chamber. i, j One-septate
ascospores. Scale bars:
a=0.3 mm, b=100 μm,
c=20 μm, d, e=50 μm,
f–j=10 μm
conical ascomata of A. patilii are distinct from B. tuberculata.
Thus, the familial placement of Belizeana is still undetermined.
Biatriospora K.D. Hyde & Borse, Mycotaxon 26: 263
(1986). (Pleosporales, genera incertae sedis)
Fungal Diversity
Generic description
Habitat marine, saprobic. Ascomata large, solitary or gregarious, immersed, subglobose to pyriform, ostiolate, papillate,
periphysate, black, branching, carbonaceous. Hamathecium
of dense, long trabeculate pseudoparaphyses, embedded in
mucilage. Asci 8-spored, bitunicate, fissitunicate, cylindrical,
with apical apparatus. Ascospores uniseriate to partially
overlapping, fusoid, hyaline when young, becoming brown
to dark brown at maturity, multi-septate towards each end,
with a hyaline, globose refractive chamber or appendage at
each end, not constricted at the septum.
Fig. 12 1 Biatriospora marina
(from IMI 297768, holotype). a,
b Cylindrical asci. Note the
mucilage pseudoparaphyses in
(a) and the conspicuous ocular
chamber in (b). c, d Ascospores
with hyaline end chambers
(arrowed). Scale bars: a, b=
50 μm, c, d=20 μm. 2 Line
drawings of Biatriospora marina (based on holotype). a Section through ascocarp showing
asci and pseudoparaphyses. b
Asci and pseudoparaphyses. c
Ascospores. Scale bars: a=
200 μm, b=40 μm, c=30 μm
(figure with permission from
Hyde and Borse 1986)
Anamorphs reported for genus: none.
Literature: Hyde and Borse 1986; Suetrong et al. 2009.
Type species
Biatriospora marina K.D. Hyde & Borse, Mycotaxon 26:
264 (1986). (Fig. 12)
Ascomata 650–860 μm high×350–510 μm diam., solitary
or gregarious, immersed, subglobose to pyriform, ostiolate,
papillate, periphysate, black, carbonaceous (Fig. 12.2a). Hamathecium of dense, long trabeculate pseudoparaphyses, 1–
1.5 μm broad, branching, embedded in mucilage. Asci 175–
Fungal Diversity
Fig. 12 2 (continued)
400×22–40 μm, 8-spored, bitunicate, fissitunicate, cylindrical,
with long pedicels and apical apparatus (Fig. 12.1a, b, 2b).
Ascospores 55–82×16–25 μm, uniseriate to partially overlapping, fusoid, hyaline when young, becoming brown to
dark brown at maturity, 2-4-septate towards each end, and
with a hyaline, globose refractive chamber or appendage
at each end, 6–8×4–6 μm diam., not constricted at the
septum (Fig. 12.1c, d, 2c).
Fungal Diversity
Anamorph: none reported.
Material examined: SEYCHELLES, 2 Jan. 1984 (Herb.
IMI 297768 holotype).
Notes
Morphology
Biatriospora was introduced to accommodate a
marine fungus B. marina, which is characterized by
horizontal ascomata and ascospores with polar, globose
refractive chambers and polar septa (Hyde and Borse
1986). Polar refractive chambers can also occur in other
marine fungi, such as Lulworthia and Aigialus. The
chambers have been proposed as important for spore
attachment to substrates in a liquid environment (Hyde
and Borse 1986).
Phylogenetic study
Multigene phylogenetic analysis indicated that Biatriospora
marina forms a separate branch, sister to other families of
Pleosporales (Suetrong et al. 2009), and maybe related to
species in Roussoella (Plate 1).
Concluding remarks
The familial status of Biatriospora can not be determined.
Bicrouania Kohlm. & Volkm.-Kohlm., Mycol. Res. 94:
685 (1990). (?Melanommataceae)
Generic description
Habitat marine, saprobic. Ascomata immersed gregarious,
erumpent to superficial, globose to subglobose, black,
periphysate, coriaceous, epapillate or papillate, ostiolate.
Peridium thin, 2-layered. Hamathecium of dense, long
trabeculate pseudoparaphyses, branching and anastomosing between and above the asci. Asci 8-spored, bitunicate, fissitunicate, cylindrical, with a thick, furcate pedicel
lacking ocular chamber. Ascospores obliquely uniseriate
and partially overlapping, ellipsoidal with broadly rounded
ends, reddish brown, 1-septate, thick-walled, constricted at the
septum.
Anamorphs reported for genus: none.
Literature: Jones et al. 2009; Kohlmeyer and VolkmannKohlmeyer 1990.
Ascomata 320–440 μm high × 370–460 μm diam.,
gregarious, immersed, mostly erumpent to superficial,
globose to subglobose, black, coriaceous, with a rough
surface, papillate or epapillate, ostiolate, periphysate
(Fig. 13a). Peridium 40–50 μm thick laterally, up to
75 μm thick at the apex, thinner at the base, 2-layered,
outer layer composed of small heavily pigmented
pseudoparenchymatous cells, inner layer very thin,
composed of hyaline thin-walled small cells, merging into
pseudoparaphyses (Fig. 13a and b). Hamathecium of
dense, very long trabeculate pseudoparaphyses, 0.8–
1.2 μm broad, branching and anastomosing between and
above the asci. Asci 170–225 × 17.5–22.5 μm
(x ¼ 199:6 20mm, n=10), 8-spored, bitunicate, fissitunicate, cylindrical, with a thick, furcate pedicel which is up
to 70 μm long, lacking ocular chamber (Fig. 13c, d and e).
Ascospores 22–26× 12–15 μm (x ¼ 24:5 13:3mm, n=
10), obliquely uniseriate and partially overlapping, ellipsoidal with broadly rounded ends, reddish brown, 1septate, slightly constricted at the septum, thick-walled,
with a thick darkened band around the septum, smooth
(Fig. 13c, d and e).
Anamorph: none reported.
Material examined: FRANCE, Finistère, on Halimone portulacoides (IMI 330806, isotype, as Sphaeria
maritima).
Notes
Morphology
When Kohlmeyer and Volkmann-Kohlmeyer (1990) studied the four marine Didymosphaeria species, the monotypic
Bicrouania was established to accommodate B. maritima (as
Didymosphaeria maritima (P. Crouan & H. Crouan) Sacc.),
which could be distinguished from Didymosphaeria by its
superficial ascomata lacking a clypeus, thick-walled asci and
its association with algae (Kohlmeyer and VolkmannKohlmeyer 1990). Jones et al. (2009) agreed that it cannot
be placed in Didymosphaeria based on its superficial
ascomata, but that it does have many similarities with
Didymosphaeria. Molecular data are required to determine
its relationship with Didymosphaeria and to resolve its
higher level placement.
Phylogenetic study
None.
Type species
Bicrouania maritima (P. Crouan & H. Crouan) Kohlm. &
Volkm.-Kohlm., Mycol. Res. 94: 685 (1990). (Fig. 13)
≡ Sphaeria maritima P. Crouan & H. Crouan, Florule du
Finistére, Paris: 27 (1867) non Sphaeria maritima Cooke &
Plowright, Grevillia 5: 120 (1877).
Concluding remarks
Besides the morphological differences, its marine and
substrate habitats also differ from Didymosphaeria.
Bimuria D. Hawksw., Chea & Sheridan, N. Z. J. Bot. 17:
268 (1979). (Montagnulaceae)
Fungal Diversity
Fig. 13 Bicrouania maritima
(from IMI 330806, isotype). a
Section of an ascoma. b Section
of papilla. Note the periphyses.
c–e Eight-spored asci. Note the
furcated pedicel. Scale bars: a,
b=100 μm, c–e=20 μm
Generic description
Habitat terrestrial, saprobic. Ascomata solitary, superficial,
globose, dark brown, epapillate, ostiolate. Peridium thin,
pseudoparenchymatous. Hamathecium of few, cellular
pseudoparaphyses, embedded in mucilage, rarely anastomosing and branching. Asci bitunicate, fissitunicate, broadly
clavate with short pedicels, 2-3-spored. Ascospores muriform, broadly ellipsoid, dark brown with subhyaline end
cells, verrucose.
Anamorphs reported for genus: none.
Literature: Barr 1987b; Hawksworth et al. 1979;
Lumbsch and Huhndorf 2007.
Type species
Bimuria novae-zelandiae Hawksworth, Chea & Sheridan,
N. Z. J. Bot. 17: 268 (1979). (Fig. 14)
Ascomata (185-)200×310(-330) μm diam., solitary, scattered, semi-immersed or superficial, globose, hyaline when
Fungal Diversity
Fig. 14 Bimuria novae-zelandiae (from CBS 107.79, isotype). a–c
Asci with a short pedicel and small ocular chamber. d Immature ascus
(in cotton blue). e Partial ascospore. Note the convex verrucae on the
ascospore surface. f Released ascospores. Note the lighter end cells,
germ pore and the longiseptum (arrowed). g Fissitunicate ascus
dehiscent. Scale bars: a–g=20 μm
young, turning dark brown to black when mature, ostiolate,
the ostiole more or less sessile or raised into a very short neck.
Peridium 5–8(-12) μm thick, comprising 2–3 layers of
radically compressed pseudoparenchymatous cells, cells 10–
15 μm diam. in surface view, cell wall 2–3 μm thick.
Hamathecium consisting of few, 2.5–4 μm broad cellular
pseudoparaphyses, embedded in mucilage, rarely anastomosing and branching, septate, 7–13 μm long between
two septa. Asci (65-)80–95 × 20–32.5 μm (x ¼ 75:6
29:4mm, n=10), (1-)2(-3)-spored, bitunicate, fissitunicate,
broadly clavate, with a short and small knob-like pedicel
which is up to 13 μm long, ocular chamber best seen in
immature asci (Fig. 14a, b, c, d and g). Ascospores
accumulating in a subglobose black shiny mass adhering
together outside the ostiole, 55–68×25–28 μm (x ¼ 59
26mm, n=10), broadly ellipsoid but becoming narrowed
towards the poles, muriform with (5-)7 transverse septa, cells
with (0-)l(-2) longitudinal septa in each cell, no constriction
Fungal Diversity
at the septa, dark brown, the apical cells paler with no
longitudinal septa, verruculose (Fig. 14e and f).
Anamorph: none reported.
Material examined: NEW ZEALAND, North Island,
Wairarapa District, Nutty Farm, isolated from soil, 3 Mar.
1978, Chea Chark Yen & J.E. Sheridan (CBS 107.79,
isotype).
Notes
Morphology
Bimuria novae-zelandiae was first isolated from soil of a
barley field in New Zealand (Hawksworth et al. 1979). Based
on B. novae-zelandiae, the genus is characterized by a very
thin peridium, mostly 2-spored and fissitunicate asci as well as
the muriform, dark brown, verrucose ascospores (Hawksworth
et al. 1979). Because of its unique morphological characters,
the familial placement of this genus has been debatable and it
has been placed in Pleosporaceae (Hawksworth et al. 1979),
in Phaeosphaeriaceae (Barr 1987b) and in Melanommataceae
(Lumbsch and Huhndorf 2007).
Morphologically, Bimuria is most comparable with
some superficially similar or allied genera, in particular
Montagnula (Hawksworth et al. 1979). However, the thick
carbonaceous peridium distinguishes Montagnula from
that of Bimuria (Hawksworth et al. 1979). In addition, the
ascospores of Montagnula are discharged forcibly through
the ostiole instead of forming a mass outside of the ostiole
as in Bimuria (Hawksworth et al. 1979). Ascomauritiana
lignicola V.M. Ranghoo & K.D. Hyde has somewhat
similar ascospores in 4-spored asci, but this taxon has
unitunicate asci (Ranghoo and Hyde 1999). The morphological characters of Bimuria, such as ascospore release
and large, thick-walled ascospores may be an adaptation to
its soil-borne habitat (Hawksworth et al. 1979).
Phylogenetic study
Bimuria novae-zelandiae was found to be closely related
to Phaeodothis winteri (Niessl) Aptroot (syn. Didymosphaerella opulenta (De Not.) Checa & M.E. Barr) and
Montagnula opulenta (De Not.) Aptroot in analysis of
combined sequences, i.e. SSU rDNA, LSU rDNA, RPB2
and TEF1 sequences (Schoch et al. 2006, 2009). These two
species had been included by Barr (2001) in her new family
Montagnulaceae.
Concluding remarks
We agree with Barr (2001) and include the genus in
Montagnulaceae based on both morphological and phylogenetic characters.
Bricookea M.E. Barr, Mycotaxon 15: 346 (1982).
(?Phaeosphaeriaceae)
Generic description
Habitat terrestrial, saprobic (or parasitic?). Ascomata small- to
medium-sized, solitary, scattered, or in small groups, immersed, erumpent to superficial, depressed globose, papillate,
ostiolate. Peridium thin. Hamathecium filliform, cellular
pseudoparaphyses, embedded in mucilage, anastomosing,
septate. Asci bitunicate, fissitunicate, cylindrical, cylindroclavate or slightly obclavate, with a short knob-like pedicel,
with an ocular chamber. Ascospores hyaline, ellipsoid to
narrowly obovoid, 3-septate, constricted at each septum.
Anamorphs reported for genus: none.
Literature: Barr 1982a; Berlese 1896; Holm 1957; Shoemaker
and Babcock 1989a.
Type species
Bricookea sepalorum (Vleugel) M.E. Barr, Mycotaxon 15:
346 (1982). (Fig. 15).
≡ Metasphaeria sepalorum Vleugel, Svensk bot. Tidskr.
2: 369 (1908).
Ascomata 120–250 μm high×170–440 μm diam., solitary,
scattered, or in small groups, or forming locules in massive
stromatic tissues, initially immersed, becoming erumpent, to
nearly superficial, depressed globose, black, membraneous,
roughened; apex rounded, sometimes very short and almost
inconspicuous, with a somewhat slit-like or Y-shaped ostiole
(Fig. 15a). Peridium 16–30 μm wide, comprising two types of
cells, outer cells heavily pigmented thick-walled textura
angularis, cells 4.5–8 μm diam., cell wall 1–1.5 μm thick,
inner cells of subhyaline thin-walled textura angularis, cells
larger than outer cells (Fig. 15b). Hamathecium of long
cellular pseudoparaphyses, 1.5–2 μm broad, embedded in
mucilage, anastomosing, septate. Asci 63–83×9.5–11 μm
(x ¼ 73:8 10:8mm, n=10), 8-spored, bitunicate, fissitunicate,
oblong, cylindro-clavate or slightly obclavate, with a short
knob-like pedicel which is 5–13 μm long, with an ocular
chamber (Fig. 15c, d and e). Ascospores (14-)15.5–19×5–
7 μm (x ¼ 16:9 5:9mm, n=10), obliquely uniseriate and
partially overlapping to biseriate, ellipsoid to narrowly
obovoid, hyaline, 3-septate, constricted at each septum, the
cells above central septum often broader than the lower ones,
smooth (Fig. 15f, g, h, i and j).
Anamorph: none reported.
Material examined: SWEDEN, on Juncus filliformis,
Stockholm, J. Vleugel. Jul. 1907 (S, type as Metasphaeria
sepalorum Vleugel).
Notes
Morphology
Bricookea was formally established by Barr (1982a) as a
monotypic genus represented by B. sepalorum based on its
Fungal Diversity
Fig. 15 Bricookea sepalorum (from S, type). a Ascomata on host
surface (arrowed). b Section of partial peridium. Note thick-walled
out layer and thin-walled inner layer. c–e Cylindrical to slightly
obclavate asci with short knob-like pedicels. f–j Hyaline, 3-septate
smooth-walled ascospores. Scale bars: a=0.5 mm, b=50 μm, c–j=
10 μm
Fungal Diversity
“globose to depressed ascomata, slit-like ostiole with labial
cells, bitunicate asci, cellular pseudoparaphyses, and hyaline
septate ascospores”. Bricookea was morphologically assigned
to Phaeosphaeriaceae. Holm (1957) checked the authentic
collections from North America and type material from
Europe, and observed that the ascospores of collections from
North America were significantly larger than those from the
type material from Sweden. Thus, Shoemaker and Babcock
(1989a) considered that the collections from North America
represented a new species, which they introduced as B. barrae
Shoemaker & C.E. Babc. Although the short slit-like
ostiole has previously been reported (Shoemaker and
Babcock 1989a), it is inconspicuous in the type specimen
from Sweden. Currently, only two species are accommodated in this genus.
Phylogenetic study
None.
erumpent, coriaceous, ovoid with a flattened base and
apex with a elongate slit-like ostiole, up to 700 μm long and
200 μm wide (Fig. 16a). Peridium not observed. Hamathecium of dense, long pseudoparaphyses, up to 1.5–2.5 μm
broad, anastomosing and branching between and above the
asci (Fig. 16b). Asci 80–105 × (5-)7.5–10 μm
(x ¼ 91 8mm; n=10), 8-spored, bitunicate, fissitunicate,
cylindrical or cylindro-clavate, with a furcate pedicel and a
small ocular chamber (J-) (Fig. 16d). Ascospores 18–20
(−28)×4.5–6(−7.5) μm (x ¼ 20:8 5:7mm, n=10), uniseriate to biseriate, fusoid, hyaline, turning faintly brown when
old, 1-septate, with 1–2 distinct oil drops in each cell and
usually with a short terminal appendage at each end
(Fig. 16c).
Anamorph: none reported.
Material examined: on decaying wood (K(M):164030,
isotype).
Notes
Concluding remarks
The knob-shaped pedicel, slit-like ostiole, hyaline
ascospores as well as the herbaceous substrate all disagree
with any current pleosporalean family. Thus, we temporarily retain this genus under Phaeosphaeriaceae until DNA
sequence comparisons can be carried out.
Byssolophis Clem., in Clements & Shear, Gen. fung., Edn 2
(Minneapolis): 286 (1931). (Pleosporales, genera incertae
sedis)
Generic description
Habitat terrestrial, saprobic. Ascomata medium-sized, gregarious, semi-immersed to erumpent, coriaceous, ovoid, with a
conspicuous elongate slit-like ostiole on the top. Peridium not
observed. Hamathecium of dense, long pseudoparaphyses,
anastomosing and branching between and above the asci. Asci
8-spored, bitunicate, fissitunicate, cylindrical or cylindroclavate, with a furcate pedicel. Ascospores fusoid, hyaline,
turning faintly brown when old, 1-septate, with a short
terminal appendage at each end.
Anamorphs reported for genus: none.
Literature: Clements and Shear 1931; Holm 1986; Müller
and von Arx 1962.
Type species
Byssolophis byssiseda (Flageolet & Chenant.) Clem., Gen.
Fung. (Minneapolis): 286 (1931). (Fig. 16)
≡ Schizostoma byssisedum Flageolet & Chenant., in
Chenantaise, Bull. Soc. mycol. Fr. 35: 125 (1919).
Ascomata 300–450 μm high × 600–750 μm long ×
350–420 μm broad, gregarious, semi-immersed to
Morphology
Byssolophis was introduced as a monotypic genus based
on B. byssiseda, which is characterized by its semiimmersed, gregarious, ovoid ascomata, with a conspicuous
central apical ostiolar slit (Holm 1986). Subsequently, two
more species were introduced, viz. B. ampla (Berk. &
Broome) L. Holm and B. sphaerioides (P. Karst.) E. Müll.
(Holm 1986; Müller and von Arx 1962).
Phylogenetic study
The current phylogeny places Byssolophis sphaerioides
in proximity of Hypsostromataceae without resolving any
sister taxa (Plate 1).
Concluding remarks
The slit-like ostiole, cylindrical asci, hyaline and 1septate ascospores as well as the form of pseudoparaphyses
are similar to species in Lophiostoma. Thus, Byssolophis
may be a synonym of Lophiostoma.
Byssosphaeria Cooke, Grevillea 7: 84 (1879).
(Melanommataceae)
Generic description
Habitat terrestrial, saprobic. Ascomata medium-sized,
scattered to gregarious, superficial, globose, subglobose
to turbinate, non papillate with white, orange, red or green
ostiolar region, wall black. Hamathecium of dense,
long trabeculate pseudoparaphyses, embedded in mucilage, anastomosing between and above the asci. Asci
bitunicate, fissitunicate, clavate to nearly cylindrical,
with a furcate pedicel. Ascospores fusoid with narrow
Fungal Diversity
Fig. 16 Byssolophis byssiseda
(from K(M):164030, isotype). a
Ascomata gregarious on the host
surface. b Numerous pseudoparaphyses. c Fusoid ascospores
with or without terminal appendages. d Clavate ascus with a
short furcate pedicel. Scale bars:
a=1 mm, b–d=10 μm
ends, straight or slightly curved, brown, 1-septate when
young.
Anamorphs reported for genus: Pyrenochaeta or Chaetophoma-like (Barr 1984; Hawksworth et al. 1995; Samuels
and Müller 1978).
Literature: von Arx and Müller 1975; Barr 1984; Boise
1984; Bose 1961; Chen and Hsieh 2004; Cooke and
Plowright 1879; Hyde et al. 2000; Luttrell 1973; Mugambi
and Huhndorf 2009b; Müller and von Arx 1962; Samuels
and Müller 1978.
Type species
Byssosphaeria keitii (Berk. & Broome) Cooke [as ‘Byssosphaeria keithii’], (1879). (Fig. 17)
≡ Sphaeria keitii Berk. & Broome [as ‘Sphaeria keithii’],
Ann. Mag. Nat. Hist., IV 17: 144 (1876).
Ascomata 360–500(−600) μm high × 420–640 μm
diam., scattered or in small groups, superficial with
basal subiculum anchoring on the substrate, globose,
subglobose to turbinate, non-papillate with pore-like
Fungal Diversity
Fig. 17 Byssosphaeria schiedermayriana (from K(M):108784,
holotype). a Superficial ascomata on the host surface. b Brown, 1septate ascospores. c Section of the lateral peridium. Note the outer
textura angularis and inner textura epidermoidea cells. d, e Furcate
asci with a long pedicel. f Dehiscent ascus. Scale bars: a=0.5 mm, c=
50 μm, b, d–f=15 μm
ostiole, ostiolar region sometimes with orange and
greenish tint, wall black, roughened, coriaceous
(Fig. 17a). Peridium 55–85 μm thick, peridium outside
of the substrate comprising two cell types, outer layer
composed of brown thick-walled cells of textura epidermoidea, cells 1–3 μm diam., inner layer composed of
small hyaline cells, cells 3–5 μm diam., merging into
pseudoparaphyses; peridium inside the substrate one
layer, composed of large pale brown cells of textura
angularis, cells 6–13 μm diam. (Fig. 17c). Hamathecium
of dense, long trabeculate pseudoparaphyses, 1–2 μm
broad, embedded in mucilage, anastomosing between and
above the asci. Asci 90–120(−148)×10–14 μm, 8-spored,
bitunicate, fissitunicate, cylindro-clavate to clavate,
biseriate above and uniseriate below, pedicel 15–20
(−53) μm long, the immature asci usually with longer
and furcate pedicel (−68 μm) (Fig. 17d,e and f).
Ascospores 29–34(−38) × 5.5–8(−10) μm, fusoid with
narrow ends, mostly straight, sometimes slightly curved,
smooth, pale brown, 1-septate, becoming 3-septate after
discharge, with hyaline appendages at each acute to
subacute end; in some mature spores the appendage may
be absent (Fig. 17b).
Anamorph: Pyrenochaeta sp. (Barr 1984; Samuels and
Müller 1978).
Pycnidia 70–500 μm diam. Conidiogenous cells phialidic, lining cavity, 5–8×4–6 μm to 5–10×3–6 μm. Conidia
2.5–3.5(−4)×1.5–2(−3) μm, hyaline, ellipsoid or subglobose (Barr 1984).
Material examined: ERIE, Dublin, Glasnevin Botanic
Garden, on old rope, Jun. 1872, W. Keit (K(M):108784,
holotype, as Sphaeria keitii Berk. & Broome).
Fungal Diversity
Notes
Morphology
Byssosphaeria was introduced by Cooke and Plowright
(1879) based on its superficial ascomata seated on a “tomentose
subiculum of interwoven threads”, which includes various
species in Sphaeria and Byssisedae, and was validly typified by
B. keitii (Cooke 1878). Byssosphaeria keitii was treated as a
synonym of B. schiedermayeriana (Fuckel) M.E. Barr by
Sivanesan (1971), and B. schiedermayeriana exclusively occurs
in tropical regions or greenhouse environments in temperate
regions (Barr 1984). Morphologically, B. keitii is characterized
by its large ascomata with orange to reddish plain apices, and is
closely related to B. rhodomphala (Berk.) Cooke (Barr 1984).
For a long time, Byssosphaeria was assigned to
Herpotrichia sensu lato, and Byssosphaeria schiedermayeriana was renamed as H. schiedermayeriana Fuckel
(von Arx and Müller 1975; Bose 1961; Luttrell 1973;
Müller and von Arx 1962; Sivanesan 1971). After studying
Herpotrichia in North America, Barr (1984) accepted a
relatively narrow generic concept, Herpotrichia sensu
stricto, and revived Byssosphaeria; this proposal is
supported by phylogenetic study (Mugambi and Huhndorf
2009b). Currently Byssosphaeria comprises 32 species
(http://www.mycobank.org, 08-01-2009).
Phylogenetic study
The monophyletic nature of Byssosphaeria is well
demonstrated, as well as its familial status in Melanommataceae (Mugambi and Huhndorf 2009b).
Concluding remarks
Orange and greenish plain apices exist in the specimen
we examined, which is different from records as “orange,
bright or dull reddish plain apices” by Barr (1984). This
might be because different specimens have different
colours, or there may be a variation of apical colour within
a single species, as both orange and green can coexist on
the same ascoma (see Fig. 17a). The coloured apical rim,
together with the trabeculate pseudoparaphyses as well as
the presence of subiculum make Byssosphaeria readily
distinguishable from other morphologically comparable
genera, e.g. Herpotrichia and Keissleriella (Hyde et al. 2000).
lenticular to subglobose, papillate, ostiolate. Hamathecium
of long, filliform pseudoparaphyses, branching and anastomosing, embedded in mucilage. Asci 4- to 8-spored,
bitunicate, fissitunicate, cylindrical to cylindro-clavate, with
a short, furcate pedicel. Ascospores muriform, broadly
fusoid to fusoid with broadly to narrowly rounded ends,
hyaline.
Anamorphs reported for genus: none.
Literature: Barr 1983; Rossman et al. 1999; Spegazzini
1909.
Type species
Calyptronectria platensis Speg., Anal. Mus. nac. Hist. nat.
B. Aires 19: 412 (1909). (Fig. 18)
Ascomata 120–270 μm high×170–400 μm diam., solitary,
scattered, immersed, lenticular to subglobose, papillate,
ostiolate (Fig. 18a and b). Apex with a small and slightly
protruding papilla. Peridium 18–30 μm wide, comprising two
types of cells, outer layer composed of pseudoparenchymatous cells, cells 3–6 μm diam., cell wall 1–2 μm thick, inner
layer comprising less pigmented cells, merging with pseudoparaphyses (Fig. 18b and c). Hamathecium of long, filliform
pseudoparaphyses, 1–2 μm broad, branching and anastomosing, embedded in mucilage. Asci 98–140×12.5–20 μm
(x ¼ 107 15:4mm, n=10), 8-spored, sometimes 4-spored,
bitunicate, fissitunicate, cylindrical to cylindro-clavate, with
a short, furcate pedicel, 12–20 μm long, with an ocular
chamber (to 4 μm wide×3 μm high) (Fig. 18e and f).
Ascospores 17–22.5 μm × (6.3-)7.5–10 μm
(x ¼ 19:8 7:6mm, n=10), biseriate above and uniseriate
below, ellipsoid to broadly fusoid with broadly to narrowly
rounded ends, hyaline, usually with (3-)5 transverse septa,
with or without 1–3 longitudinal septa in the central cells,
constricted at the median septum, the upper cell often broader
than the lower one, smooth, surrounded by an irregular
hyaline gelatinous sheath up to 3 μm thick (in dry specimen)
(Fig. 18d).
Anamorph: none reported.
Material examined: ARGENTINA, La Plata, on decaying branches of Manihot carthaginensis (Jacq.) Müll., Sept.
1906, Spegazzini (LPS 1209, holotype).
Notes
Calyptronectria Speg., Anal. Mus. nac. Hist. nat. B. Aires
19: 412 (1909). (Melanommataceae)
Generic description
Habitat terrestrial, saprobic. Ascomata small- to mediumsized, solitary, scattered, or in small groups, immersed,
Morphology
Calyptronectria is a relatively poorly studied genus,
which was formally established based on C. argentinensis
Speg. and C. platensis, with C. platensis being chosen as
the generic type (Spegazzini 1909). Morphologically,
Calyptronectria is characterized by its immersed ascomata,
Fungal Diversity
Fig. 18 Calyptronectria platensis (from LPS 1209, holotype). a
Appearance of ascomata scattered in the substrate (after removing the
out layer of the substrate). Note the protruding papilla. b Section of an
ascoma. c Section of the partial peridium. Note the lightly pigmented
pseudoparenchymatous cells. d Released ascospores with mucilaginous sheath. e Eight-spored asci in hamathecium and embedded in gel
matrix. f Ascus with a short pedicel. Scale bars: a=0.5 mm, b=
100 μm, c=50 μm, d–f=10 μm
trabeculate pseudoparaphyses and hyaline, muriform ascospores as well as its peridium that turns reddish brown in KOH
(Rossman et al. 1999) (not shown here). Subsequently, C.
indica Dhaware was introduced from India, and Barr (1983)
transferred Teichospora ohiensis Ellis & Everh. to Calyptronectria as C. ohiensis (Ellis & Everh.) M.E. Barr.
However, this proposal is inappropriate as the type specimen
of T. ohiensis is “unitunicate” (Barr 1983; Rossman et al.
1999). Subsequently, Rossman et al. (1999) transferred
Calyptronectria ohiensis to Thyridium (as T. ohiense (Ellis
& Everh.) Rossman & Samuels).
Phylogenetic study
None.
Concluding remarks
The immersed ascomata, trabeculate pseudoparaphyses,
bitunicate asci, hyaline and muriform ascospores as well as
Fungal Diversity
the reaction of peridium to KOH (turns reddish brown)
make it distinguishable from all other reported genera
(Rossman et al. 1999). Thus Calyptronectria is a morphologically well defined genus.
Carinispora K.D. Hyde, J. Linn. Soc., Bot. 110: 97 (1992).
(Pleosporales, genera incertae sedis)
Generic description
Habitat marine, saprobic. One or two ascomata per
stroma. Ascomata scattered or in small groups, developing beneath the host epidermis, erumpent, lenticular,
ostiolate, lacking periphyses. Peridium pale brown,
composed of thin-walled elongated cells at the sides
and thick-walled cells of textura epidermoidea at the
base. Hamathecium of dense, long filliform pseudoparaphyses, embedded in mucilage, anastomosing between
and above the asci, rarely septate. Asci 8-spored,
bitunicate, fissitunicate, clavate to cylindrical, with a
short furcate pedicel, apex with an ocular chamber and
apical ring. Ascospores biseriate, narrowly fusoid, yellow
to pale brown, multi-septate, constricted at the septa, the
two central cells being the largest, surrounded by a
gelatinous sheath.
Anamorphs reported for genus: none.
Literature: Hyde 1992a, 1994b.
Type species
Carinispora nypae K.D. Hyde, J. Linn. Soc., Bot. 110: 99
(1992). (Fig. 19)
One or two ascomata per stroma. Ascomata up to 0.8 mm
diam., scattered or in small groups, developing beneath the
host epidermis, crust-like, as circular spots, wall brown, with
a small central ostiole, in section 225–285 μm high×510–
750 μm diam., lenticular, ostiolar canal lacking periphyses
(Fig. 19a and b). Peridium 35–45 μm wide at sides, pale
brown, at sides composed of a thin layer of thin-walled
elongate cells, fusing with the stromatic tissue and host cells,
at the base composed of thick-walled cells, forming a textura
epidermoidea and fusing with host cells. A wedge of pale
brown hyphae forming a textura porrecta is present at the
rim (Fig. 19c). Hamathecium of dense, long filliform
pseudoparaphyses 1–3 μm broad, embedded in mucilage,
anastomosing between and above the asci, rarely septate.
Asci 142–207×14.2–19.8 μm, 8-spored, bitunicate, fissitunicate, clavate to cylindrical, with a furcate pedicel, up to
40 μm long, apex with an ocular chamber and apical ring (to
2 μm wide×3 μm high, J-), developing from ascogenous
tissue at the base of the ascocarp (Fig. 19d, e, f, g and h).
Ascospores 42–66×7–10.6 μm, biseriate, narrowly fusoid
with broadly to narrowly rounded ends, somewhat curved,
yellow to pale brown, yellow in mass, 7-8-septate, constricted at the septa, the two central cells being the largest,
surrounded by a gelatinous sheath; the sheath has a central
“spine” and curved polar extrusions (Fig. 19i and j).
Anamorph: none reported.
Material examined: BRUNEI DARUSSALAM, Tungit
Api Api mangrove, from decaying intertidal fronds of Nypa
fruticans Wurmb., 14 Apr. 1987, K.D. Hyde (BRIP 17106,
holotype).
Notes
Morphology
Carinispora is distinguished from Phaeosphaeria by its
saprobic life style and lenticular ascomata formed under the
host epidermis, peridium structure and sheath surrounding the
ascospores (Hyde 1992a, 1994b). Two species were reported,
i.e. C. nypae and C. velatispora K.D. Hyde.
Phylogenetic study
Suetrong et al. (2009) could not resolve Carinispora
nypae in a phylogeny based on four genes.
Concluding remarks
Both Carinispora nypae and C. velatispora are reported
as marine fungi, which should be taken into consideration
for their familial placement.
Caryosporella Kohlm., Proc. Indian Acad. Sci., Pl. Sci. 94:
355 (1985). (?Melanommataceae)
Generic description
Habitat marine, saprobic. Ascomata densely scattered or
gregarious, superficial, subglobose, black, papillate, ostiolate,
periphysate, carbonaceous. Peridium carbonaceous. Hamathecium of dense, long trabeculate pseudoparaphyses, anastomosing
and branching above the asci. Asci 8-spored, bitunicate,
fissitunicate, cylindrical. Ascospores ellipsoidal to broadly fusoid
with narrowly hyaline rounded ends, deep reddish brown, thickwalled, 1-septate with hyaline germ pore at each end.
Anamorphs reported for genus: suspected spermatia
(Kohlmeyer 1985).
Literature: Eriksson 2006; Kohlmeyer 1985; Lumbsch and
Huhndorf 2007.
Type species
Caryosporella rhizophorae Kohlm., Proc. Indian Acad.
Sci., Pl. Sci. 94: 356 (1985). (Fig. 20)
Ascomata 0.8–1.1 mm high×0.9–1.2 mm diam., densely
scattered or gregarious, superficial with a flattened base, not
easily removed from the host surface, subglobose, black, short
Fungal Diversity
papillate, ostiolate, periphysate, carbonaceous (Fig. 20a
and b). Peridium 120–150 μm thick at sides, up to
200 μm thick at the apex, thinner at the base, 3-layered,
outer layer composed of golden-yellow, very thickwalled cells of textura epidermoidea, mixed with subglobose, large cells near the surface, cells 7–15 μm
diam., middle layer composed of deep brown, very thickwalled cells of textura epidermoidea, inner layer composed of hyaline, thin-walled cells of textura prismatica,
up to 50×5 μm diam., merging with pseudoparaphyses
(Fig. 20b, c and d). Hamathecium of dense, long
trabeculate pseudoparaphyses, 1.5-2 μm wide, anastomosing and branching above the asci. Asci 225–250
(−275)× 14–17 μm (x ¼ 137 16:3mm, n= 10), 8-spored,
bitunicate, fissitunicate, cylindrical, with a long, narrowed, pedicel which is up to 75 μm long, apical
characters not observed (Fig. 20e). Ascospores 25–28
(−30)×9–13 μm (x ¼ 26:8 11mm, n= 10), uniseriate to
partially overlapping, ellipsoidal to broadly fusoid with
narrow hyaline rounded ends, deep reddish brown, thickwalled, 1-septate with hyaline germ pore at each end, slightly
constricted at the septum, verruculose, sometimes with “net”like ridged ornamentations (Fig. 20f and g).
Anamorph: suspected spermatia (Kohlmeyer 1985).
Material examined: BELIZE, Twin Cays, tip of prop
root of Rhizophora mangle, 18 Mar. 1984, J. Kohlmeyer
(NY. Herb. J. Kohlmeyer No. 4532a, holotype).
Notes
Morphology
Caryosporella was formally established by Kohlmeyer
(1985) based on the obligate marine fungus, C. rhizophorae, which is characterized by its superficial ascomata,
3-layered peridium, filliform trabeculate pseudoparaphyses, and brown, 1-septate ascospores. Caryosporella was
originally assigned to Massariaceae despite several major
differences, such as the superficial ascomata, reddish
brown ascospores (Kohlmeyer 1985). Subsequently,
Caryosporella was assigned to Melanommataceae
(Eriksson 2006; Lumbsch and Huhndorf 2007).
Phylogenetic study
Suetrong et al. (2009) showed that a single isolate of
Caryosporella rhizophorae does not reside in Pleosporales,
but is related to Lineolata rhizophorae (Kohlm. & E.
Kohlm.) Kohlm. & Volkm.-Kohlm. and placed in Dothideomycetidae incertae sedis.
Concluding remarks
As an obligate marine fungus, the familial placement of
Caryosporella rhizophorae is uncertain but it may not
belong to Pleosporales.
Fig. 19 Carinispora nypae (from BRIP 17106, holotype). a
Ascomata on the host surface. b Section of an ascoma. c Section of
a partial peridium. d, e, g, h Asci with ocular chambers and short
pedicels. f The ocular chamber and apical ring of ascus. i–j Narrowly
fusoid ascospores. Scale bars: a=1 mm, b, c=100 μm, d, g, h=50 μm,
e, f, i, j =10 μm
Chaetomastia (Sacc.) Berl., Icon. fung. (Abellini) 1: 38
(1890). (Teichosporaceae)
≡ Melanomma subgen. Chaetomastia Sacc., Syll. fung.
(Abellini) 2: 113 (1883).
Generic description
Habitat terrestrial, saprobic. Ascomata relatively small,
scattered, or in small groups, superficial, globose or
subglobose, black, papillate, ostiolate, coriaceous. Peridium relatively thin, 1-layered, composed of heavily
pigmented cells of textura angularis. Hamathecium of
dense, long cellular pseudoparaphyses, embedded in
mucilage. Asci mostly 4-spored, bitunicate, fissitunicate,
broadly cylindrical with a furcate pedicel, with a large
ocular chamber, especially apparent in immature asci.
Ascospores ellipsoid to broadly fusoid with broadly to
narrowly rounded ends, brown, 3-septate, constricted at all
septa.
Anamorphs reported for genus: coelomycetous where
known: conidia hyaline or brown, aseptate or 1-septate
(Aposphaeria- or Coniothyrium-like) (Barr 1989c).
Literature: Barr 1987b, 1989c; 1993a; b; 2002; Berlese
1890; Clements and Shear 1931; Eriksson 1999; Eriksson
and Hawksworth 1987, 1998; Holm 1957; Leuchtmann
1985; Saccardo 1883.
Type species
Chaetomastia hirtula (P. Karst.) Berl., Icon. fung. (Abellini)
1: 38 (1890). (Fig. 21)
≡ Sphaeria hirtula P. Karst., Fungi Fenn. Exs. N. 825 (1869).
Ascomata 214–286 μm high × 210–258 μm diam.,
scattered or in groups, superficial, globose, wall black;
apex often opening with a broad pore within slightly
raised papilla, up to 30 μm diam., coriaceous (Fig. 21a).
Peridium 20–26 μm thick, 1-layered, composed of
heavily pigmented cells of textura angularis, cells up to
5× 15 μm diam., cell wall up to 3.5 μm thick (Fig. 21b).
Hamathecium of dense, long cellular pseudoparaphyses,
embedded in mucilage. Asci 90–130×12.5–17.5(−22.5)
μm (x ¼ 111 16:3mm, n =10), mostly 4-spored, bitunicate, fissitunicate, broadly cylindrical, with a furcate
pedicel, 18–48 μm long, with a large ocular chamber
best seen in immature asci (to 3 μm wide× 3 μm high)
(Fig. 21c and d). Ascospores 20.5–27 × 7–10 μm
b
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Fungal Diversity
Fig. 20 Caryosporella rhizophoriae (from NY. Herb. J. Kohlmeyer
No. 4532a, holotype). a Gregarious ascomata on host surface. b
Section of an ascoma. c, d Section of partial peridium at sides (c) and
base (d). Note the three layers. e Asci with long peduncles in
pseudoparaphyses. f, g Ascospores. Note the “net”-like ridged
ornamentation of spore surface and hyaline germ pores. Scale bars:
a=1 mm, b=200 μm, c–e=100 μm, f, g=10 μm
(x ¼ 23:5 8:2mm, n =10), uniseriate to partially overlapping, ellipsoid to broadly fusoid with broadly to
narrowly rounded ends, brown, 3-septate, verruculose,
constricted at all septa, constricted at the median septum,
the cell above the central septum often broader than the
others (Fig. 21e and f).
Fungal Diversity
Fig. 21 Chaetomastia hirtula (from H, FFE 825, kleptotype). a
Superficial ascomata gregarious on the host surface. b Section of a
partial peridium. Note the cells of textura angularis with relatively
thick wall. c, d Cylindrical asci with long and furcate pedicels. e, f
Brown, 3-septate ascospores. Scale bars: a=0.5 mm, b=50 μm, c–f=
15 μm
Anamorph: none reported.
Material examined: FINLAND, ETELÄ-HÄME (EH/
Ta), Tammela, Mustiala, På Rub. id., 8 May 1866. P.A.
Karsten (H, FFE 825, kleptotype).
ed, i.e. M. canescens Speg., M. cucurbitarioides Speg.,
M. hirtulum (P. Karst.) Sacc., M. hispidulum Sacc. and
M. pilosellum P. Karst. Berlese (1890) promoted it to
genus rank. Subsequently, Chaetomastia hirtula (P.
Karst.) Berl. was selected as the lectotype species of
the genus (Clements and Shear 1931). Chaetomastia has
been regarded as having unitunicate asci (Eriksson and
Hawksworth 1986, 1998; Eriksson 1999). However its
bitunicate status was confirmed by Holm (1957). Holm
(1957) treated C. hirtula as Melanomma hirtulum (P.
Notes
Morphology
Chaetomastia was introduced by Saccardo (1883) as a
subgenus of Melanomma, and five species were includ-
Fungal Diversity
Karst.) Sacc., and Leuchtmann (1985) transferred this
species to Montagnula sensu lato based on the ascospore
morphology and the hyphae surrounding the ascomata.
Barr (1987b) suggested that ascoma, peridium structure
and ascospore characters pointed Montagnula sensu
stricto to Phaeosphaeriaceae, while the characters of
ascomata and peridium structure of Chaetomastia were
thought to fit the definition of Dacampiaceae (Barr
1987b). In particular, the peridium and ascospore characters of C. hirtula are comparable with those of the generic
type of Massariosphaeria (M. phaeospora). Thus, Barr
(1989c) accepted Massariosphaeria sensu stricto and
assigned the phragmosporous species of Massariosphaeria sensu lato to Chaetomastia.
Barr (2002) later assigned Chaetomastia to Teichosporaceae based on its saprobic or hypersaprobic lifestyle,
occurring on woody stems and peridium structure, and this
is widely followed (Eriksson 2006; Lumbsch and Huhndorf
2007). Currently, 11 species are accepted in this genus
(http://www.indexfungorum.org/).
Phylogenetic study
None.
Concluding remarks
Familial placement of Chaetomastia is undetermined
currently but has been included in the Teichosporaceae
by authoritative sources (Eriksson 2006; Lumbsch and
Huhndorf 2007) or the Dacampiaceae (http://www.
indexfungorum.org/).
Fig. 22 Chaetoplea calvescens (from FH-81113, isotype). a, b Fourspored and 8-spored asci. c Released ascospores. Scale bars: a–c=
10 μm
Type species
Chaetoplea calvescens (Fr.) Clem., Gen. Fung. (Minneapolis):
275 (1931). (Fig. 22)
≡ Sphaeria calvescens Fr. Scleromyc. Sueciae 401.
Ascomata not examined. Peridium not examined.
Hamathecium of dense, long, narrow cellular pseudoparaphyses, 2–3 μm broad, septate, branching and anastomosing. Asci 90–110×10–12 μm, 8-spored, rarely 4-spored,
bitunicate, fissitunicate, cylindro-clavate, with a thick,
furcate pedicel which is up to 30 μm long (Fig. 22a
and b). Ascospores 13–18×5.5–7 μm, obliquely uniseriate and partially overlapping, broadly fusoid to oblong
with broadly rounded ends, pale brown, 2-3-septate,
constricted at the septa, containing four refractive globules
(Fig. 22c).
Note: The specimen is only a slide, and no peridium or
ascomata information could be obtained.
Anamorph: coelomycetous, conidia yellowish, 1septate, 9–13×4–5(−8) μm (Webster and Lucas 1959);
Microdiplodia henningsii Staritz=Chaetodiplodia caudina
Karst. (Sutton 1980) (referred to Barr 1990b (p50)).
Material examined: SWEDEN, sub-collection: Curtis
Herbarium, verified by R.A. Shoemaker, leg. E.M. Fries
401 (FH-81113, isotype, microscope slide).
Notes
Chaetoplea (Sacc.) Clem., Gen. Fung. (Minneapolis): 275
(1931). (?Phaeosphaeriaceae)
≡ Pyrenophora subgen. Chaetoplea Sacc., Syll. fung.
(Abellini) 2: 279 (1883).
Generic description
Habitat terrestrial, saprobic. Ascomata small to medium,
immersed, erumpent to superficial, globose to subglobose,
papillate, ostiolate. Peridium not examined. Hamathecium
of dense, long, narrowly cellular pseudoparaphyses. Asci 8spored or 4-spored, bitunicate, fissitunicate, cylindroclavate, with a thick, furcate pedicel. Ascospores ellipsoid
or fusoid, pale brown to brown, phragmosporous or
muriform.
Anamorphs reported for genus: Microdiplodia-like (Barr
1990b).
Literature: Barr 1981; 1987a; b; 1990b; Clements and
Shear 1931; Ramaley and Barr 1995; Yuan and Barr
1994.
Morphology
Chaetoplea was introduced based on C. calvescens,
which has been regarded as similar to Pleospora or
Leptosphaeria (Eriksson and Hawksworth 1987;
Wehmeyer 1961; von Arx and Müller 1975). Based on
the differences in ascomata, peridium structure, pseudoparaphyses as well as its anamorphic stage, Chaetoplea was maintained as a separate genus (Barr 1990b;
Yuan and Barr 1994). Chaetoplea sensu lato was
accepted by Barr (1990b), which included some species
of Teichospora as well as the subgenus Pleospora subg.
Cylindrosporeae.
The following is from the label of specimen.
“Sphaeria calvescens, Scler. Suecicae (Ed. 2) 401. No
specimen of Scler. Suecicae 401 is now at Uppsala
according to R. Santesson 1966. This Curtis Herbarium specimen in the Farlow Herbarium is isotype.
Wehmeyer (1961) in his Pleospora monograph did
not study any portion of the Scler. Suecicae exsiccatus
b
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Fungal Diversity
401, nor did Webster & Lucas in the taxonomic and
life-history study (Trans. Brit. Myc. Soc. 42, 332–
342. 1959) of this species.
The specimen has most of the features described by
Webster & Lucas including the presence of the
conidial state Microdiplodia henningsii Staritz. I did
not see vertical septa in the ascospores. Webster &
Lucas note that vertical septa may be occasionally be
lacking. The fungus is otherwise as they describe it
although some perithecia collapse and appear cupulate.”—by R.A. Shoemaker.
Phylogenetic study
None.
Concluding remarks
The substrate of Chaetoplea sensu Barr (1990b) can
be herbaceous stalks, decorticated wood or periderm, or
old cotton cloth and string, which may indicate its
heterogeneous nature. The ascospores seem very much
like Phaeosphaeria which may be an earlier name; more
details concerning the ascomatal, peridial and hamathecial structures are needed to make any conclusion.
Cilioplea Munk, Dansk botanisk Arkiv 15: 113 (1953).
(Pleosporales, genera incertae sedis)
Generic description
Habitat terrestrial, saprobic. Ascomata small- to medium-sized,
solitary, scattered or in small groups, immersed, globose or
subglobose, papilla covered with short and blackish setae,
coriaceous. Peridium thin, comprising small heavily pigmented thick-walled cells of textura angularis. Hamathecium
of cellular pseudoparaphyses. Asci 8-spored, bitunicate,
fissitunicate, broadly clavate, with a short, furcate pedicel, and
small ocular chamber. Ascospores fusoid to narrowly fusoid
with narrowly rounded ends, pale brown to reddish brown,
multi-transverse septa, usually with one longitudinal septum in
some central cells, constricted at the primary septum.
Anamorphs reported for genus: none.
Literature: Barr 1990b, 1992b; Crivelli 1983; Lumbsch
and Huhndorf 2007; Müller 1951; Munk 1953, 1957.
Type species
Cilioplea coronata (Niessl) Munk, Dansk botanisk Arkiv
15: 113 (1953). (Fig. 23)
≡ Pleospora coronata Niessl, Notiz. Pyr.: 16 (1876).
Ascomata 170–290 μm high×200–410 μm diam., solitary,
scattered, or in small groups, immersed, globose or subglobose, wall black, papilla raised, 50–80 μm high, with short
and blackish setae, coriaceous (Fig. 23a). Peridium 9–15 μm
thick laterally, up to 28 μm thick at the apex, thinner at the
base, 1-layered, composed of small heavily pigmented thickwalled cells of textura angularis, cells up to 4×2.5 μm diam.,
cell wall 2–3 μm thick, apex cells smaller and walls thicker
(Fig. 23b). Hamathecium of long cellular pseudoparaphyses,
2–3 μm broad. Asci (60-)80–108 × 10–15 μm
(x ¼ 85:3 12:1mm, n=10), 8-spored, bitunicate, fissitunicate, broadly clavate, with a short, thick, furcate pedicel, 5–
15 μm long, and a small ocular chamber (to 3 μm wide×
2 μm high) (Fig. 23c and d). Ascospores 21–27.5×5.5–
7.5 μm (x ¼ 24 6:7mm, n=10), biseriate to uniseriate at
base, fusoid to narrowly fusoid with narrowly rounded ends,
pale reddish brown, 5–7 transverse septa (mostly 5), usually
with one longitudinal septum in some central cells, deeply
constricted at the median septum, the part above the primary
septum shorter and broader, smooth-walled.
Anamorph: none reported.
Material examined: GERMANY, Hadiberg. on Reseda
lutea Hadiberg, 20 Sept. 1875, Niessl (M 175-89-290,
lectotype; M 175-89-291, type).
Notes
Morphology
Cilioplea was introduced by Müller (1951) as a
subgenus of Pleospora, and this was followed by Munk
(1957), who had earlier proposed it as a separate genus
typified by C. coronata based on its hairy papilla, clavate
asci as well as its “perfectly paraphysoid” (see Munk
1953). A relatively narrow concept of Pleospora was
accepted by Crivelli (1983), and four species was
assigned under the separate genus Cilioplea, viz. C.
coronata, C. genisticola (Fautrey & Lambotte) Crivelli,
C. kansensis (Ellis & Everh.) Crivelli and C. nivalis
(Niessl) Crivelli. Subsequently, another six species were
added (Barr 1990b, 1992b). Currently, ten species are
included under Cilioplea.
Phylogenetic study
None.
Concluding remarks
The most striking character of Cilioplea is its setose
papilla, which has been shown to have no phylogenetic
significance in Lentitheciaceae (Zhang et al. 2009a).
Cilioplea was assigned under Lophiostomataceae (Lumbsch
and Huhndorf 2007), but there is little morphological
similarity with the Lophiostomataceae sensu stricto (Zhang
et al. 2009a). Thus its familial placement needs further study.
Crivellia Shoemaker & Inderb., in Inderbitzin, Shoemaker,
O’Neill, Turgeon & Berbee, Can. J. Bot. 84: 1308 (2006).
(Pleosporaceae)
Fungal Diversity
Fig. 23 Cilioplea coronata (M 175-89-290, lectotype). a Immersed
ascomata in small groups on the host surface (the covering host tissue
was removed). b Section of a partial ascoma. Note the thin peridium. c
Clavate asci within pseudoparaphyses. d Ascus with a small ocular
chamber. Scale bars: a=0.5 mm, b=100 μm, c=50 μm, d=10 μm
Generic description
Literature: Inderbitzin et al. 2006.
Habitat terrestrial, hemibiotrophic or parasitic. Ascomata
small- to medium-sized, scattered, immersed, erumpent to
nearly superficial, papillate, ostiolate. Peridium thin, composed of two cells types, outer cells of thick walled and
textura angularis, inner cells thin-walled, yellow. Hamathecium of dense, long and thin pseudoparaphyses. Asci (4-)
8-spored, bitunicate, fissitunicate dehiscence not observed, broadly cylindrical to cylindrical, with a short,
furcate pedicel and an ocular chamber. Ascospores
fusoid to broadly fusoid, pale brown, septate, sometimes
with one or two vertical septa in the middle cells, constricted
at the septa.
Anamorphs reported for genus: Brachycladium (Inderbitzin
et al. 2006).
Type species
Crivellia papaveracea (De Not.) Shoemaker & Inderb.,
Can. J. Bot. 84: 1308 (2006). (Fig. 24)
≡ Cucurbitaria papaveracea De Not., Sfer. Ital.: 62
(1863).
Ascomata 210–260 μm high × 300–380 μm diam.,
densely scattered, immersed, erumpent to nearly superficial,
flattened globose, dark brown, papillate, ostiolate
(Fig. 24a). Peridium 25–30 μm thick, thicker near the apex
and thinner at the base, composed of two cell types, outer
cells of thick-walled and textura angularis, cells up to 10×
5 μm diam., cell wall 2–4 μm thick, inner cells thin-walled,
yellow (Fig. 24b). Hamathecium of dense, long, 1–2 μm
Fungal Diversity
broad, rarely septate pseudoparaphyses. Asci 85–125×10–
13 μm (x ¼ 106 11mm, n=10), (4-)8-spored, bitunicate,
fissitunicate dehiscence not observed, broadly cylindrical to
cylindrical, with a short, furcate pedicel, with a relatively large
ocular chamber (Fig. 24c and d). Ascospores (16-)19–24×5–
7.5 μm (x ¼ 20:4 6:3mm, n=10), overlapping uniseriate to
rarely biseriate, fusoid to broadly fusoid, pale brown, 3septate, sometimes with one or two vertical septa in the
middle cells, constricted at the septa, the upper cell often
broader than the lower one, smooth-walled.
Anamorph: Brachycladium penicillatum (Corda) Fr.
(Inderbitzin et al. 2006).
Fig. 24 Crivellia papareracea
(from UBC F14995, epitype).
a Gregarious ascomata immersed within the host surface.
b Section of an ascoma. c Asci
within pseudoparaphyses. d
Cylindrical ascus with a
short pedicel. Scale bars:
a=1 mm, b=100 μm, c,
d=30 μm
Material examined: AUSTRIA, Vienna, on decaying
stems of Papaver rhoeas L., 28 Oct. 2001, W. Jaklitsch
(UBC F14995, epitype).
Notes
Morphology
Crivellia was separated from Pleospora and introduced
as a new genus by Inderbitzin et al. (2006) based on their
differences in ascospore morphology and anamorphic
stages. Crivellia is characterized by having small- to
medium-sized ascomata, and yellow, 3-septate ascospores
Fungal Diversity
with one or two vertical septa in central cells. Its
Brachycladium anamorphic stage with phragmosporous
conidia also differs from that of Stemphylium, which is
the anamorphic stage of Pleospora (Inderbitzin et al. 2006).
Currently, two species are included within Crivellia, i.e. C.
homothallica Inderb. & Shoemaker and C. papaveracea.
Phylogenetic study
Crivellia papaveracea was shown to be closely related to
some species of Alternaria, and its pleosporaceous status was
confirmed following molecular studies (Inderbitzin et al. 2006).
Concluding remarks
Crivellia seems to belong to Pleosporaceae, and may be
closely related to Pleospora.
Decaisnella Fabre, Annls Sci. Nat., Bot., sér. 6 9:112
(1878). (Pleosporales, genera incertae sedis)
Generic description
Habitat terrestrial, saprobic. Ascomata medium to large,
immersed to erumpent, clypeate, papillate, ostiolate. Hamathecium of dense, long, cellular pseudoparaphyses, rarely
septate, embedded in mucilage. Asci mostly 4- or 8-spored,
rarely 2-spored, cylindrical to cylindro-clavate, with a
furcate pedicel. Ascospores muriform, dark brown, oblong
with broadly rounded ends.
Anamorphs reported for genus: none.
Literature: Barr 1986; 1990a; b; Fabre 1878; Saccardo
1883.
Type species
Decaisnella spectabilis Fabre, Annls Sci. Nat., Bot., sér. 6
9: 112 (1879). (Fig. 25)
Ascomata 520–680 μm high × 430–600 μm diam.,
solitary, scattered, or in small groups of 2–3, immersed
to erumpent, clypeate, globose or subglobose, black,
roughened, with a blunt papilla up to 170 μm high,
apex with a round ostiole, coriaceous (Fig. 25a).
Peridium 70–90 μm thick at sides, thicker near the apex,
comprising two types of cells; part immersed in host
tissue, outer layer pseudoparenchymatous, 55–65 μm
thick, pigmented, inner layer composed of lightly pigmented to hyaline thin-walled compressed cells, 15–
23 μm thick, cells 3.5–7 μm diam., part above host
tissue heavily pigmented covered by clypeus tissues
(Fig. 25b). Hamathecium of dense, long, cellular pseudoparaphyses, 1.5–3 μm broad, rarely septate, embedded in
mucilage. Asci 150–200 × 15–25(−33) μm
(x ¼ 181 20:6mm, n= 10), (2-)4-spored, bitunicate, fissitunicate, broadly cylindrical, with a short, thick, furcate
pedicel which is 20–40 μm long, no apical apparatus
observed (Fig. 25e). Ascospores 37–45 × 12–17 μm
(x ¼ 43 15mm, n=10), uniseriate and sometimes slightly
overlapping, oblong with broadly rounded ends, dark
brown, verrucose or smooth, 7–9 transverse septa and 1–
3 longitudinal septa in some of the cells, no constriction at
the septa (Fig. 25c and d).
Anamorph: none reported.
Material examined: GERMANY, Valsalpe in der Ramsau,
Bayer, Alpen, on Rhamnus pumila Turra., Jul. 1913, Karl Arnold
(NY2082, syntype as Teichospora megalocarpa Rehm).
Notes
Morphology
Decaisnella was formally established by Fabre (1879),
but was treated as a synonym of Teichospora by
Saccardo (1883). This was followed by several mycologists over a long time. The main morphological differences between Decaisnella and Teichospora include the
size and septation of ascospores, shape of ascomata,
structure of peridium and type of pseudoparaphyses (Barr
1986). Thus Barr (1986) revived Decaisnella and assigned it
to Massariaceae based on the shape of ascomata and large,
distoseptate ascospores. Currently, 15 species are accepted
under Decaisnella (http://www.mycobank.org/MycoTaxo.
aspx). Neither the size of ascomata nor the ascospore
characters have proven sufficient to place taxa at the family
level in Pleosporales (Zhang et al. 2009a), and therefore
familial placement of Decaisnella remains uncertain.
Phylogenetic study
Decaisnella formosa resided in the clade of Lophiostomataceae and in proximity to Lophiostoma macrostomoides De Not. (Plate 1).
Concluding remarks
The muriform ascospores, saprobic life style and 4spored asci point Decaisnella spectabilis to Montagnulaceae, but this can only be confirmed following a molecular
phylogenetic study.
Delitschia Auersw., Hedwigia 5: 49 (1866). (Delitschiaceae)
Generic description
Habitat terrestrial, saprobic (coprophilous). Ascomata
medium- to large-sized, solitary or scattered, immersed to
erumpent, globose or subglobose, apex with or without
papilla, ostiolate. Peridium thin, composed of compressed
cells. Hamathecium of dense, long pseudoparaphyses,
anastomosing and branching. Asci 8-spored, cylindrical to
cylindro-clavate, with short pedicel. Ascospores uni- to
Fungal Diversity
Fig. 25 Decaisnella spectabilis (NY2082, syntype). a Appearance of
ascomata on the host surface. b Section of a partial peridium
(immersed in the substrate). Note the pseudoparenchymatous out
layer. c, d Muriform ascospores. Note the minuitely verrucose
ornamentation. e Ascus with a short pedicel. Scale bars: a=0.5 mm,
b=100 μm, c–e=20 μm
triseriate, pale to dark brown, ellipsoid, 1-septate, usually
constricted at the septum, smooth, with a full length germ
slit in each cell.
Anamorphs reported for genus: none.
Literature: Auerswald 1866; Barr 2000; Cain 1934;
Dennis 1968; Eriksson 2006; Griffiths 1901; Hyde and
Steinke 1996; Kirschstein 1911; Kruys et al. 2006; LuckAllen and Cain 1975; Lumbsch and Huhndorf 2007;
Fungal Diversity
Moreau 1953; Munk 1957; Romero and Samuels 1991;
Schoch et al. 2006; Winter 1887.
Type species
Delitschia didyma Auersw., Hedwigia 5: 49 (1866).
(Fig. 26)
Ascomata 400–800 μm diam., solitary or scattered,
immersed, globose or subglobose, black, papilla short, 70–
130 μm broad, central, with a wide opening, coriaceous
(Fig. 26a). Peridium ca. 15 μm thick laterally, up to 35 μm
thick at the apex, up to 30 μm at the base, comprising a
single layer of small lightly pigmented thin-walled cells of
textura angularis, cells 4–10 μm diam., cell wall <1 μm
thick, apex cells smaller and wall thicker (Fig. 26b).
Hamathecium of dense, very long pseudoparaphyses, 1.5–
2 μm broad, anastomosing and branching. Asci 290–380×
35–45 μm (x ¼ 357:5 40:6mm, n=10), 8-spored, bitunicate, fissitunicate, cylindrical to cylindro-clavate, with
short, narrowed pedicels which are rounded at the base,
25–60 μm long, apex with a wide ocular chamber
(Fig. 26d and e). Ascospores 50–58 × 20–22.5 μm
(x ¼ 54 21:3mm, n=10), obliquely uniseriate and partially overlapping, ellipsoid with narrowly rounded ends,
reddish brown, 1-septate, slightly constricted at the
septum, smooth-walled, each cell with a full length germ
slit (Fig. 26c).
Anamorph: none reported.
Material examined: GERMANY, Near Königstein, in
forest, rare, Oct. 1904, W. Krieger (L, 1950).
Notes
Morphology
Delitschia was established by Auerswald (1866), and
assigned to Sphaeriaceae. It was considered to be closely
related to Sordariaceae and Amphisphaeriaceae. Winter
(1887) assigned Delitschia under Sordariaceae, and this
placement is followed in several subsequent studies
(Griffiths 1901; Kirschstein 1911). Cain (1934) suggested
that Delitschia might belong in Pleosporaceae, and this
proposal was supported by Moreau (1953) and Dennis
(1968). Finally, Munk (1957) established Sporormiaceae
(Pseudosphaeriales), and Delitschia was assigned therein.
Luck-Allen and Cain (1975) reviewed and redefined the
genus as having bitunicate asci, pigmented and 1-septate
ascospores with an elongated germ slit in each cell and
surrounded by a gelatinous sheath, and in particular, the
coprophilous habitat. Luck-Allen and Cain (1975) accepted 46 species. Subsequently, some wood-inhabiting
species were also described (Hyde and Steinke 1996;
Romero and Samuels 1991). Three genera, i.e. Delitschia,
Ohleriella and Semidelitschia were separated from Spor-
ormiaceae, and a new family, Delitschiaceae, was
introduced by Barr (2000) to accommodate them. Delitschiaceae is characterized by a periphysate ostiole, wide
endotunicate asci with a wide ocular chamber and
ascospores having cells with germ slits. Delitschiaceae
has been subsequently accepted (Eriksson 2006; Lumbsch
and Huhndorf 2007).
The genus comprises 83 names (Index Fungorum) and is
estimated to comprise 51 species (Kirk et al. 2008). Keys to
Delitschia can be found in Luck-Allen and Cain (1975) and
Hyde and Steinke (1996).
Phylogenetic study
Delitschia didyma and D. winteri (W. Phillips & Plowr.)
Sacc. form a robust phylogenetic clade within Delitschiaceae, which is basal to other members of Pleosporales
(Kruys et al. 2006; Schoch et al. 2006) except for
Massariaceae (Voglmayr and Jaklitsch 2011). This might
indicate its early derivation (Zhang et al. 2009a).
Concluding remarks
Morphologically, Delitschia is a well defined genus, and
each cell of the ascospore has a full length germ slit.
Currently, most species of this genus are coprophilous,
although a few species are reported from wood (Hyde and
Steinke 1996; Luck-Allen and Cain 1975). Whether the
lignicolous habitat is an important character that might
separate these taxa from the main coprophilous group,
needs to be addressed, however, the morphological characters are similar.
Didymosphaeria Fuckel, Jb. nassau. Ver. Naturk. 22–23:
140 (1870). (Didymosphaeriaceae)
Generic description
Habitat terrestrial, saprobic or parasitic. Ascomata
solitary, scattered, or in small groups, immersed to
erumpent, globose to ovoid, papillate, ostiolate, periphysate. Ostiole with a pore-like opening. Peridium 1layered, thin, composed of brown pseudoparenchymatous
cells of textura angularis. Hamathecium of dense, trabeculate, anastomosing mostly above the asci. Asci (2-)4spored or 8-spored, bitunicate, cylindrical, with a furcate
pedicel. Ascospores uniseriate, ellipsoid, brown, 1distoseptate.
Anamorphs reported for genus: Dendrophoma, Fusicladiella and Phoma (Aptroot 1995).
Literature: Aptroot 1995; Barr 1989a, b, 1990a, 1992a, b;
1993a; b; Fuckel 1870; Hawksworth 1985a, b; Hawksworth
and Boise 1985; Hawksworth and Diederich 1988; Hyde
et al. 2000; Lumbsch and Huhndorf 2007; Saccardo 1882;
Scheinpflug 1958; Sivanesan 1984.
Fungal Diversity
Fig. 26 Delitschia didyma
(from L, 1950). a Ascomata on
the substrate surface. Note the
ostiolar opening. b Section of
peridium. Note the small cells of
textura angularis. c Released
and unreleased ascospores. Note
the germ slit in each cell. d, e
Asci with ascospores and short
pedicels with rounded ends.
Scale bars: a=0.5 mm,
b =30 μm, c–e=70 μm
Type species
Didymosphaeria futilis (Berk. & Broome) Rehm, Hedwigia
18: 167 (1879). (Fig. 27)
≡ Sphaeria futilis Berk. & Broome, Ann. Mag. nat.
Hist., Ser. 2 9: 326 (1852).
Ascomata 190–230 μm high×240–340 μm diam., scattered, or in small groups, immersed to slightly erumpent,
subglobose to ovoid, membraneous, near-hyaline, under
clypeus, papillate, periphysate (Fig. 27a and c). Papilla
central, up to 100 μm high, black, with a pore-like ostiole
(Fig. 27a and c). Peridium 30–40 μm wide upper part, 6–
23 μm wide near the base, 1-layered, composed of brown
pseudoparenchymatous cells of textura angularis, cell wall 2–
3 μm thick (Fig. 27b). Hamathecium of dense, long
trabeculate pseudoparaphyses, 0.8–1.5 μm broad, anastomosing mostly above the asci, embedded in mucilage (Fig. 27d).
Asci 90–110×7.5–10 μm (x ¼ 97 9mm, n=10), 2–4-spored,
rarely 8-spored, bitunicate, fissitunicate, cylindrical, with a
furcate pedicel, 17.5–27.5 μm long, with a large ocular (to
2.5 μm wide×4 μm high) (Fig. 27d, e and f). Ascospores 14–
15.5×(5.5-) 6–7.5 μm (x ¼ 14:8 6:9mm, n=10), uniseriate,
ellipsoid with obtuse ends, brown, 1-septate, distoseptate,
slightly to not constricted, capitate (Fig. 27g).
Anamorph: Dendrophoma sp., Fusicladiella sp. vel aff.
(Sivanesan 1984).
Fungal Diversity
Material examined: UK, England, Norfolk, King’s
Cliffe; on dead stem (in ramis emortuis) Rosa sp., Mar.
1850, M.J. Berkeley (K(M): 147683, holotype).
Notes
Morphology
Didymosphaeria is a widely distributed genus with wide
host range (Aptroot 1995). Didymosphaeria was formally
established by Fuckel (1870) based on six ascomycetous
species, and D. epidermidis (Fries) Fuckel (or D. peltigerae
Fig. 27 Didymosphaeria futilis
(from K(M): 147683, holotype).
a Two immersed ascomata on
the host surface (one of them is
cut horizontally). b Section of
an ascoma. Note the thin peridium. c Hand cut portion of
ascoma showing habitat in
wood. d Asci in pseudoparaphyses. Note the trabeculate pseudonparaphyses anastomosing
above the asci. e, f Four-spored
asci with long pedicels which
are rounded at their bases. g
Brown, 1-septate ascospores
with spinulose ornamentation.
Scale bars: a=0.3 mm, b, c=
100 μm, d–g=10 μm
Fuckel) has been chosen as the lectotype species (see
comments by Aptroot 1995). Hawksworth and David
(1989: 494) proposed to conserve the genus with a lectotype
specimen, Fungi Rhenani 1770. The genus had been
considered as a depository to accommodate all types of
didymosporous pyrenocarpous ascomycetes. Many workers
have tried to redefine the genus and excluded some species.
Saccardo (1882) restricted the genus to brown-spored species,
and about 100 species have been excluded subsequently (Barr
1989a, b, 1990a, 1992a, b, 1993b; Hawksworth 1985a, b;
Hawksworth and Boise 1985; Hawksworth and Diederich
Fungal Diversity
1988; Scheinpflug 1958). Over 400 epithets of Didymosphaeria were included until the monograph of Aptroot (1995).
Aptroot (1995) examined more than 3000 specimens under
the name Didymosphaeria. The type specimen of Didymosphaeria (Fungi Rhenani 1770) represents the widespread and
common D. futilis (Aptroot 1995). In this study, we did not
get the lectotype specimen, but described the type of D. futilis
(Sphaeria futilis). Using a narrow concept (ignoring differences of host or country of origin), Aptroot (1995) accepted
only seven species, which were closely related with the
generic type of Didymosphaeria with over 100 synonyms
distributed among them. Many taxa were found to belong to
other groups, i.e. Aaosphaeria, Amphisphaeria, Astrosphaeriella, Dothidotthia, Flagellosphaeria, Kirschsteiniothelia, Megalotremis, Montagnula, Munkovalsaria, Mycomicrothelia,
Parapyrenis or Phaeodothis. Didymosphaeria is mainly characterized by a peridium consisting of flattened or irregular cells
or completely hyphae; a hamathecium consisting of narrow,
trabeculate paraphysoids or paraphyses, richly anastomosing
above the asci; and brown thinly distoseptate ascospores.
Didymosphaeriaceae was maintained as a separated family
within Pleosporales by Aptroot (1995) because of the
distoseptate ascospores and trabeculate pseudoparaphyses
mainly anastomosing above the asci. This proposal, however,
has not received much support (Lumbsch and Huhndorf 2007).
Phylogenetic study
There have been few molecular investigations of
Didymosphaeria when compared to the morphological
studies. Didymosphaeria futilis resided in the clade of
Cucurbitariaceae (or Didymosphaeriaceae) (Plate 1). The
correct identification of the Didymosphaeria strain used for
sequencing, however, has not been verified.
Concluding remarks
Didymosphaeria is a well established genus represented
by D. futilis. Of particular significance are the narrow
pseudoparaphyses which anastomose above the asci and
brown 1-septate ascospores with indistinct distosepta. Familial placement of Didymosphaeria is unclear yet because
of insufficient molecular data.
Dothidotthia Höhn., Ber. Deutsch. Bot. Ges. 36: 312
(1918). (Didymellaceae)
Generic description
Habitat terrestrial, saprobic. Ascomata medium-sized, solitary,
clustered or somewhat gregarious, erumpent, subglobose,
apex somewhat papillate to depressed, coriaceous. Peridium
composed of a few layers of dark brown cells of textura
angularis, and giving rise dark brown, thick-walled hyphae in
the basal region, 2-layered. Hamathecium septate pseudopar-
aphyses branched in upper part above asci. Asci 8-spored,
bitunicate, clavate, straight to curved. Ascospores biseriate to
obliquely uniseriate, ellipsoid, pale brown, 1-septate.
Anamorphs reported for genus: Dothiorella and Thyrostroma (Hyde et al. 2011; Phillips et al. 2008).
Literature: Barr 1989b; Phillips et al. 2008.
Type species
Dothidotthia symphoricarpi (Rehm) Höhn., Ber. Deutsch.
Bot. Ges. 36: 312 (1918). (Fig. 28)
≡ Pseudotthia symphoricarpi Rehm, Ann. Mycol. 11:
169 (1913).
Ascomata up to 500 μm high×550 μm diam., gregarious
clustered, rarely solitary, erumpent, subglobose, apex somewhat
papillate to depressed, coriaceous (Fig. 28a). Peridium 20–
80 μm thick, composed of 3–6 layers of dark brown cells of
textura angularis, giving rise dark brown, thick-walled hyphae
in the basal region, 2-layered, outer layer wall thicker and inner
layer wall thinner (Fig. 28b). Hamathecium hyaline, septate
pseudoparaphyses, 2–3 μm wide, branched in upper part above
asci. Asci 70–120×15–22 μm, 8-spored, bitunicate, clavate,
straight to curved (Fig. 28c, d and e). Ascospores (20-)22–23
(−26)×(8-)9–10(−11) μm, biseriate to obliquely uniseriate and
partially overlapping, ellipsoid tapering towards subacutely
rounded ends, pale brown, 1-septate, constricted at the septum,
smooth (Fig. 28f) (description referred to Phillips et al. 2008).
Anamorph: Thyrostroma negundinis (Phillips et al. 2008).
Material examined: USA, North Dakota, on branches
of Symphoricarpos occidentalis Hook. (NY, holotype);
Colorado, San Juan Co, c. 0.5 mile up Engineer Mountain
Trail from turnoff at mile 52.5, Hwy 550, dead twigs of
Symphoricarpos rotundifolius A. Gray, 24 Jun. 2004, A.W.
Ramaley 0410 (BPI 871823, epitype).
Notes
Morphology
Dothidotthia was formally established to accommodate
Pseudotthia symphoricarpi (Montagnellaceae, Dothideales)
(von Höhnel 1918a). Many mycologists considered Dothidotthia closely related to a genus of Venturiaceae such as
Dibotryon by Petrak (1927), or Gibbera by von Arx and
Müller (1954) and Müller and von Arx (1962). Dothidotthia
had been treated as a synonym of Gibbera (von Arx 1954;
Müller and von Arx 1962), which was followed by
Shoemaker (1963) and Eriksson and Hawksworth (1987).
Based on the coelomycetous anamorphic stage and peridium structure, shape of asci, as well as morphology of
pseudoparaphyses, Barr (1987b, 1989b) retrieved Dothidotthia, and considered it closely related to Botryosphaeria
(Botryosphaeriaceae). Currently, 11 species are included
within Dothidotthia (http://www.mycobank.org, 01–2011).
Fungal Diversity
Fig. 28 Dothidotthia symphoricarpi (from NY, holotype). a
Clustered ascomata on the host
stubstrate. b Longitudinal section through an ascoma. c, d
Asci with pale brown, 1-septate
ascospores. e Immature asci. f
Pale brown, 1-septate ascospores within asci. g Conidia of
Thyrostroma anamorph in association with ascomata. Scale
bars: a=0.5 mm, b=100 μm,
c–g=10 μm. (figure with permission from Phillips et al.
2008)
Phylogenetic study
Based on a multi-gene phylogenetic analysis, Dothidotthia formed a separate familial clade (Phillips et al. 2008).
Thus Dothidotthiaceae was introduced to accommodate it
(Phillips et al. 2008).
Concluding remarks
By comparing the morphological characters and phylogenetic
dendrograms by Phillips et al. (2008) and de Gruyter et al.
(2009), Dothidotthia seems closely related to Didymellaceae,
but Dothidotthiaceae should still be treated as a separate family.
Fungal Diversity
Dubitatio Speg., Anal. Soc. cient. argent. 12: 212
(1881). (Arthopyreniaceae (or Massariaceae))
Generic description
Habitat terrestrial, saprobic. Ascomata medium-sized, solitary, densely scattered, or in small groups of 2–4,
immersed, covered with white crystaline rim, papillate,
ostiolate. Hamathecium of dense pseudoparaphyses, long,
2–3 μm broad, branching and anastomosing. Asci cylindrical, pedicellate, with furcate pedicel. Ascospores 1-septate,
asymmetrical, reddish to dark brown.
Anamorphs reported for genus: Aplosporella-like (Rossman
et al. 1999).
Literature: Barr 1979b, 1987b; Müller and von Arx 1962;
Rossman et al. 1999; Spegazzini 1881.
Type species
Dubitatio dubitationum Speg., Anal. Soc. cient. argent. 12:
212 (1881). (Fig. 29)
Ascomata 350–530 μm high × 550–700 μm diam.,
solitary, densely scattered, or in small groups of 2–4,
immersed, with a protruding papilla, 110–160 μm high,
160–250 μm diam., globose or subglobose, black,
covered with white crystalline material which becomes
hyaline and gel-like in water, ostiolate (Fig. 29a and b).
Peridium 18–25 μm thick laterally (excluding the rim), up
to 35 μm thick at the apex, thinner at the base, 1-layered,
composed of small pale brown thin-walled cells of textura
prismatica, cells 5–12×3–5 μm diam., cell wall up to
1 μm thick, apex cells smaller and walls thicker
(Fig. 29b). Hamathecium of dense, long pseudoparaphyses, 2–3 μm broad, branching and anastomosing between
and above the asci. Asci 150–190(−230)×12.5–15 μm
(x ¼ 172:5 13:4mm, n = 10), (6-)8-spored, rarely 4spored, bitunicate, fissitunicate, cylindrical, with a furcate
pedicel which is up to 40 μm long, ocular chamber not
observed (Fig. 29c, d and e). Ascospores 19–22.5×10–
12 μm (x ¼ 20:2 11:4mm, n=10), uniseriate to obliquely uniseriate and partially overlapping, broadly ellipsoid
with broadly to narrowly rounded ends, reddish brown,
1-septate, constricted at septum, asymmetric with a
larger upper cell, thick-walled, possibly distoseptate
(Fig. 29f, g and h).
Anamorph: Aplosporella-like (for detailed description
see Rossman et al. 1999).
Conidiomata globose, ca. 300 μm diam. Conidia
holoblastic, broadly fusoid, 13–15×7–10 μm, dark brown,
finely spinulose (Rossman et al. 1999).
Material examined: ARGENTINA, Buenos Aires,
Tuyu, on Celtis tala Gill., Jan. 1881, leg. det. C. Spegazzini
(NY, isotype; LPS, holotype).
Notes
Morphology
When established Dubitatio, Spegazzini (1881) considered it as intermediate between Sphaeriaceae and Nectriaceae as has been mentioned by Rossman et al. (1999). Müller
and von Arx (1962) treated Dubitatio as a synonym of
Passerinula, while the differences of ascomata and ascospores
could easily distinguish these two genera (Rossman et al.
1999). After checking the type specimen, Dubitatio was assigned
to Dothideomycetes, and considered closely related to Dothivalsaria in the Massariaceae (Barr 1979b, 1987b). Dubitatio
chondrospora was assigned to Pseudomassaria (as P. chondrospora (Ces.) Jacz.) (Barr 1964; Müller and von Arx 1962).
Phylogenetic study
None.
Concluding remarks
The black ascomata with white crystalline covering and
central white ostiolar region as well as the asymmetrical
reddish brown ascospores are striking characters of Dubitatio dubitationum. The genus cannot be assigned to any
family with certainty based on morphological characters
and fresh collections are needed for sequencing.
Entodesmium Reiss, Hedwigia 1: 28 (1854).
(Phaeosphaeriaceae)
Generic description
Habitat terrestrial, saprobic (or parasitic?). Ascomata
scattered or in small groups, immersed, papillate, ostiolate,
periphysate. Peridium thin, comprising one cell type of
pigmented pseudoparenchymatous cells. Hamathecium of
dense, long pseudoparaphyses, septate, embedded in mucilage. Asci 8-spored, bitunicate, fissitunicate, cylindrical, with
furcate pedicel. Ascospores ellipsoid to filliform, multiseptate, deeply constricted at the primary septum (usually
near apex), breaking into partspores.
Anamorphs reported for genus: none.
Literature: von Arx and Müller 1975; Barr 1992b; Eriksson
1967a; b; Holm 1957; Liew et al. 2000; Shoemaker 1984a, b.
Type species
Entodesmium rude Reiss, Hedwigia 1: 28 (1854). (Fig. 30)
Ascomata 160–250 μm high×150–300 μm diam., in groups,
immersed with long and protruding cylindrical papilla,
globose to subglobose, black, coriaceous (Fig. 30a). Papilla
100–220 μm long, 70–120 μm broad, cylindrical, with
periphysate ostiole. Peridium 25–33 μm wide, comprising
pseudoparenchymatous cells, cells up to 10×7.5 μm diam., cell
wall up to 2 μm thick, beak cells smaller and wall thicker
Fungal Diversity
Fig. 29 Dubitatio dubitationum (from NY, isotype; LPS, holotype).
a Appearance of ascomata scattered on the host surface. Note the
exposed white covering around the ostioles. b, c Section of an ascoma.
Note the white covering (see arrow). d–f Cylindrical asci with short
furcate pedicels. g, h Asymmetrical, 1-septate reddish-brown ascospores. Scale bars: a=1 mm, b=100 μm, c-e=50 μm, f–h=20 μm
(Fig. 30b and c). Hamathecium of dense, long pseudoparaphyses, septate, 2–3 μm wide, embedded in mucilage. Asci
100–175 × 8–13 μm (x ¼ 147:5 11:3mm, n = 10), 8spored, bitunicate, fissitunicate, cylindrical, with a furcate
pedicel which is 18–50 μm long, and with a low ocular
chamber (ca. 1 μm wide×1 μm high) (Fig. 30e,f, g and h).
Ascospores 108–138×3–3.5 μm (x ¼ 123 3:2mm, n=10),
filliform, brown, multi-septate, breaking into 22–28 partspores, 5–7×3–3.5 μm diam. (Fig. 30d).
Anamorph: none reported.
Material examined: GERMANY, Königstein, on stems of
Coronilla varia L., 20 May 1895, W. Krieger (H, Krieger 1070).
Fungal Diversity
Fig. 30 Entodesmium rude (from H, Krieger 1070). a Ascomata in
groups on the host surface. Note the erumpent papilla which is
cylindrical and has an inconspicuous ostiole. b Section of part of an
ascoma. Note the arrangement of asci and pseudoparaphyses. c Section
of the peridium comprising cells of textura angularis. d Part-spores
inside the ascus. e Relatively immature ascus with filliform ascospores
and low ocular chamber. f–h Mature and immature asci with pedicels.
Scale bars: a=0.5 mm, b, c=50 μm, d–h=10 μm
Notes
sonii L. Holm, E. lapponicum (L. Holm) L. Holm, E.
mayorii (E. Müll.) L. Holm, E. niessleanum (Rabenh. ex
Niessl) L. Holm and E. rude are accepted in this genus
(Holm 1957; Shoemaker 1984b). Von Arx and Müller
(1975) assigned Entodesmium to the Pleosporaceae sensu
lato, and Shoemaker (1976) assigned E. rude (as
Ophiobolus rudis) to Ophiobolus sensu lato based on
the fragmenting filliform ascospores. According to the
short, blackish beak and periphysate ostiole, Barr (1992b)
Morphology
Entodesmium is characterized by having immersed
ascomata dark cylindrical, periphysate papillae, numerous clavate to cylindrical asci surrounded by narrowly
cellular pseudoparaphyses, and ellipsoidal to filliform
multi-septate ascospores (Barr 1992b; Shoemaker
1984b). Currently, five species, viz. Entodesmium elias-
Fungal Diversity
assigned Entodesmium to Lophiostomataceae. The hosts
of Entodesmium are restricted to stems of legumes (Barr
1992b; Shoemaker 1984b).
Phylogenetic study
Limited phylogenetic studies indicate that Entodesmium
rude may have affinities to Phaeosphaeriaceae (Liew et al.
2000; Plate 1).
Concluding remarks
Species of Entodesmium share several morphological
characters, such as immersed, papillate ascomata, periphysate ostioles, pale yellow to light yellowish brown, multiseptate (≥ 3), narrowly fusoid to filliform ascospores, and
are specific to legumes. All of the above similarities
indicate a close relationship among members of Entodesmium. We do not agree with Barr (1992b) who assigned
Entodesmium to Lophiostomataceae because the ascomata
are immersed, the papilla are not laterally compressed
and the peridium comprises a single type of cells of
textura angularis. These characters plus multi-septate,
lightly pigmented ascospores, which break up into
partspores and host specificity to legumes support inclusion in Phaeosphaeriaceae. Entodesmium multiseptatum
(G. Winter) L. Holm and E. niessleanum were originally
described as Leptosphaeria species (Shoemaker 1984b)
indicating their similarity to Phaeosphaeria with which
Leptosphaeria is commonly confused (Shoemaker 1984a;
Shoemaker and Babcock 1989b). Phylogenetic study has
also shown that Entodesmium rude is related to members
of Phaeosphaeriaceae (Liew et al. 2000). Thus we assign
Entodesmium to Phaeosphaeriaceae as a separate genus
until further phylogenetic analysis is carried out on
verified specimens.
Eudarluca Speg., Revta Mus. La Plata 15: 22 (1908).
(?Phaeosphaeriaceae)
Generic description
Habitat terrestrial, parasitic. Ascomata small, solitary,
scattered, immersed to erumpent, subglobose, ostiolate,
papillate. Peridium thin, composed of a few layers cells of
textura prismatica. Hamathecium of dense, cellular pseudoparaphyses, septate. Asci 8-spored, bitunicate, fissitunicate, cylindrical to fusoid, with a furcate pedicel.
Ascospores broadly fusoid to fusoid, hyaline to pale yellow,
rarely 1- or 3- septate, mostly 2-septate, constricted at the
primary septum.
Anamorphs reported for genus: Sphaerellopsis (Sivanesan
1984).
Literature: Bayon et al. 2006; Eriksson 1966; Katumoto
1986; Ramakrishnan 1951; Spegazzini 1908.
Type species
Eudarluca australis Speg., Revta Mus. La Plata 15: 22
(1908). (Fig. 31)
Ascomata 160–190 μm high×180–290 μm diam., solitary,
scattered, or in small groups, semi-immersed to erumpent,
subglobose to broadly ellipsoid, wall black, ostiolate, apex
with a short papilla, 40–70 μm broad (Fig. 31a and b).
Peridium<10 μm wide laterally, up to 25 μm thick at the
apex, thinner at the base, composed of lightly pigmented thinwalled cells of textura prismatica, cells up to 12×4 μm diam.,
cell wall <1 μm thick, apex cells heavily pigmented, smaller
and walls thicker (Fig. 31b and c). Hamathecium of dense,
long cellular pseudoparaphyses, 1.5–2.5 μm broad, septate.
Asci 50–70×7.5–10 μm (x ¼ 61:4 8:4mm, n=10), 8spored, with a short, thick, furcate pedicel, up to 12.5 μm
long, bitunicate, fissitunicate, cylindrical to fusoid, no obvious
ocular chamber (Fig. 31d, e, f and g). Ascospores 16–20×4–
6 μm (x ¼ 17:3 5mm, n=10), obliquely uniseriate and
partially overlapping to biseriate, broadly fusoid to fusoid,
hyaline to pale yellow, 2-septate, sometimes 1- or 3-septate,
constricted at the two main septa, the medium cell often
broader than the others, smooth (Fig. 31h).
Anamorph: Sphaerellopsis filum (Biv.) B. Sutton
(Sivanesan 1984).
Material examined: BRAZIL, Sao Paulo, on leaves of
Canna sp., 1905, leg. Usteri, nro; det. Ove Eriksson (LPS
5.415, type).
Notes
Morphology
Eudarluca was introduced based on E. australis
(Spegazzini 1908), and E. australis was subsequently
treated as a synonym of E. caricis (Biv.) O.E. Erikss.
(Eriksson 1966). The most striking character of E.
australis is its 2-septate ascospores, which is quite rare
in Pleosporales. Sphaerellopsis filum, anamorph of E.
caricis, is a cosmopolitan hyperparasite associated with
a large number of rust species (Płachecka 2005).
Phylogenetic study
A detailed phylogenetic study was conducted on Sphaerellopsis filum, the anamorphic stage of Eudarluca australis based on
both AFLP and ITS sequences, and only limited variation
between different isolates was detected (Bayon et al. 2006).
Concluding remarks
By blasting within GenBank, ITS sequences of E. caricis
(= E. australis, strain MullMK, GB, access AY836374) are
most comparable with species in Leptosphaeria and
Phoma. Thus Eudarluca appears to be related to Leptosphaeriaceae pending further study.
Fungal Diversity
Fig. 31 Eudarluca australis (from LPS 5.415, type). a Ascomata on
the host surface. b Section of an ascoma. c Section of a partial
peridium. Note the thin peridium with cells of textura angularis. d–g
Asci with short pedicels. h Ascospores. Note the 2-septate hyaline
ascospore. Scale bars: a, b=100 μm, c=50 μm, d–h=10 μm
Falciformispora K.D. Hyde, Mycol. Res. 96: 26 (1992).
(Trematosphaeriaceae)
globose to ovoid, black, ostiolate, epapillate, coriaceous.
Peridium thin, comprising two cells types, outer layer
composed of thick-walled cells of textura angularis, inner
layer composed of hyaline compressed cells. Hamathecium
long and cellular pseudoparaphyses, septate, embedded in
mucilage. Asci 8-spored, bitunicate, fissitunicate, broadly
clavate to fusoid, with a short, thick pedicel. Ascospores
Generic description
Habitat freshwater, saprobic. Ascomata small, scattered to
gregarious, erumpent to nearly superficial, depressed
Fungal Diversity
fusoid to somewhat clavate, hyaline, usually slightly
curved, multi-septate.
Anamorphs reported for genus: none.
Literature: Hyde 1992b; Raja and Shearer 2008.
Type species
Falciformispora lignatilis K.D. Hyde, Mycol. Res. 96: 27
(1992). (Fig. 32)
Ascomata 180–270 μm high×250–340 μm diam., scattered
to gregarious, erumpent and eventually superficial, depressed
globose to ovoid, black, ostiolate, epapillate, coriaceous
(Fig. 32a). Peridium up to 35 μm wide, comprising two cell
types, outer layer composed of thick-walled cells of textura
angularis, up to 8 μm diam., cell wall up to 5 μm thick, inner
layer composed of hyaline compressed cells, cells 12×3 μm
diam., cell wall 1–1.5 μm thick (Fig. 32a). Hamathecium long
and cellular pseudoparaphyses, 2–3 μm broad, septate,
embedded in mucilage. Asci 115–130×23–31 μm, 8-spored,
bitunicate, fissitunicate, broadly clavate to fusoid, with a short,
thick pedicel, 8–15 μm long, with an ocular chamber (to 5 μm
wide×3 μm high) (Fig. 32b and c). Ascospores 42–50×8–
10 μm, 2–3 seriate, fusoid to somewhat clavate, hyaline,
usually slightly curved, 6–8-septate, mostly 7-septate, slightly
constricted at all septa, smooth-walled, surrounded by a thin
mucilaginous sheath which is longer at the base (up to 20–
30 μm) (Fig. 32d, e and f).
Anamorph: none reported.
Material examined: MEXICO, Nova Hispania, mangrove near Boca de Pascuales, saprobic on immersed
intertidal mangrove wood, Mar. 1988, K.D. Hyde (BRIP
16972, holotype).
Notes
Morphology
Falciformispora was formally established by Hyde (1992b)
as a monotypic genus and was assigned to Pleosporaceae by
comparing with Setosphaeria, but Setosphaeria has the
anamorphic stage of Exserohilum and is exclusively parasitic
on Gramineae unlike Falciformispora. The setae on the
ascomata of Setosphaeria could also serve as a distinguishing character from Falciformispora. Raja and Shearer (2008)
also collected this species from freshwater in Florida. They
considered that the species was more closely related to
Chaetomastia than Setosphaeria, but that Falciformispora
differed in having hyaline ascospores.
Phylogenetic study
Phylogenetic analyses in Schoch et al. (2009) and Suetrong et
al. (2009) placed Falciformispora lignatilis in Trematosphaeriaceae in proximity to another marine species associated with mangroves, Halomassarina thalassiae.
Concluding remarks
Phylogenetic work confirmed that the saprobic habitat of
Falciformispora is inconsistent with most other members of
Pleosporaceae. The hyaline multi-septate ascospores with a
mucilaginous sheath indicate affinities to Lophiostomataceae
but this is not supported in DNA sequence comparisons.
Carinispora is also similar and may be related.
Hadrospora Boise, Mem. N. Y. bot. Gdn 49: 310 (1989).
(?Phaeosphaeriaceae)
Generic description
Habitat terrestrial (or freshwater?), saprobic. Ascomata
small- to medium-sized, solitary, scattered, or in groups,
immersed to nearly superficial, globose to subglobose,
papillate. Peridium thin, comprising pseudoparenchymatous cells. Hamathecium dense, narrowly cellular, embedded in mucilage. Asci bitunicate, fissitunicate, oblong to
ovoid, with a short pedicel. Ascospores ellipsoid to broadly
fusoid with narrow ends, reddish brown, multi-septate,
constricted at the primary septum.
Anamorphs reported for genus: Zalerion (Tanaka and
Harada 2003a).
Literature: Boise 1984, 1989; Fisher and Webster 1992; Shearer
and Crane 1971; Tanaka and Harada 2003a; Webster 1993.
Type species
Hadrospora fallax (Mouton) Boise, Mem. N. Y. bot. Gdn
49: 310 (1989). (Fig. 33)
≡ Trematosphaeria fallax Mouton, Bull. Soc. R. Bot.
Belg. 25: 155, (1886).
Ascomata 130–240 μm high×200–330 μm diam., solitary,
scattered or in groups, initially immersed, becoming erumpent to
nearly superficial, with basal wall remaining immersed in host
tissue, not easily removed from the substrate, globose or
subglobose, roughened, papillate, coriaceous (Fig. 33a). Peridium 30–45 μm wide, comprising cells of pseudoparenchymatous, up to 12.5×9 μm diam. (Fig. 33b and c). Hamathecium of
dense, narrowly cellular pseudoparaphyses, 1–2 μm broad,
embedded in mucilage. Asci 150–200 × 40–60 μm
(x ¼ 171:5 48mm, n=10), 8-spored, bitunicate, fissitunicate, oblong to ovoid, with a short pedicel, 10–24 μm long,
with a ocular chamber (to 5 μm wide×6 μm high) (Fig. 33d).
Ascospores 55–80×16–22 μm (x ¼ 67:1 18:1mm, n=10),
biseriate to 4-seriate, ellipsoid to broadly fusoid with narrow
ends, reddish brown with paler end cells, 8-septate,
constricted at the primary septum, smooth-walled (Fig. 33e,
f, g, h and i).
Anamorph: Zalerion sp. (Tanaka and Harada 2003a).
Material examined: BELGIUM, Beaufays, on cut off,
still hard wood. Oct. 1922, V. Mouton (BR, Capsa: K 7534,
Fungal Diversity
Fig. 32 Falciformispora lignatilis (from BRIP 16972, holotype). a Section of a superficial
ascoma. The peridium comprises two layers. b, c Squash
mounts showing asci with wide
pseudoparaphyses. The asci are
cylindro-clavate with very short
pedicels. d–f Hyaline multiseptate ascospores. Note the elongated appendage at the base
(arrow head). Scale bars:
a, b=100 μm, c=50 μm,
d–f=10 μm
holotype). (Note: The specimen is not in good condition,
only a few ascomata left).
Phylogenetic study
None.
Notes
Concluding remarks
Hadrospora seems not closely related to Phaeosphaeriaceae.
Morphology
Boise (1989) formally established Hadrospora to accommodate Trematosphaeria fallax and T. clarkia (Sivan.) Boise,
and Hadrospora fallax (syn. T. fallax) was selected as the
generic type. Hadrospora is a widely distributed species that
has been reported from Belgium, China, Italy, Japan,
Switzerland and the United States (Boise 1989; Fisher and
Webster 1992; Shearer and Crane 1971; Tanaka and Harada
2003a; Webster 1993). Hadrospora was temporarily
assigned to Phaeosphaeriaceae based on its “small, thinwalled ascomata and narrow cellular pseudoparaphyses”
(Boise 1989), which is distinguished from other genera of
Phaeosphaeriaceae by its “large, stout ascospores that form
within oblong-ovoid asci” (Boise 1989). Currently, Hadrospora includes two species, i.e. H. fallax and H. clarkii
(Sivan.) Boise differentiated by ascospore size.
Halotthia Kohlm., Nova Hedwigia 6: 9 (1963). (?Zopfiaceae)
Generic description
Habitat marine, saprobic. Ascomata large, solitary, gregarious or confluent, broadly conical to subglobose,
flattened at the base, carbonaceous, immersed to erumpent, ostiolate, epapillate. Peridium plectenchymatous.
Hamathecium of dense, long, cellular pseudoparaphyses,
septate, branching. Asci 8-spored, bitunicate, cylindrical,
with a short pedicel. Ascospores uniseriate, ellipsoidal,
subcylindrical or obtuse-fusoid, dark brown, 1-septate,
constricted at the septum.
Anamorphs reported for genus: none.
Literature: Kohlmeyer 1963; Suetrong et al. 2009.
Fungal Diversity
Fig. 33 Hadrospora fallax
(from BR, Capsa: K 7534,
holotype). a Ascomata forming
a cluster on the host surface. b
Section of an ascoma. Note the
peridium structure. c Section of
a partial peridium. Note the
pseudoparenchymatous cells. d
Asci in pseudoparaphyses. e–i
Reddish brown multiseptate
ascospores. Scale bars: a=
0.5 mm, b=100 μm, c, d=
50 μm, e–i=20 μm
Type species
Halotthia posidoniae (Durieu & Mont.) Kohlm., Nova
Hedwigia 6: 9 (1963). (Fig. 34)
≡ Sphaeria posidoniae Durieu & Mont. Exploration
scientifique de l’Algérie, pp. 502–503, Taf. 25, Abb. 8a-i, 1849.
Ascomata 0.8–1.1 mm high×1.5–2.1 mm diam., solitary, gregarious or confluent, broadly conical to subglobose, flattened at the base, carbonaceous, immersed to
erumpent, ostiolate, epapillate (Fig. 34a). Peridium 165–
275 μm thick at sides, thicker near the apex, plectenchymatous. Hamathecium of dense, long cellular pseudo-
Fungal Diversity
paraphyses, 1.5–2 μm broad, septate, branching. Asci
275–290×25–35 μm, 8-spored, bitunicate, cylindrical,
with a short pedicel (Fig. 34b, c and d). Ascospores 37–
60.5×16.5–26 μm, uniseriate, ellipsoidal, subcylindrical
or obtuse-fusoid, dark brown, 1-septate, constricted at the
septum (Fig. 34e, f, g and h) (adapted from Kohlmeyer
and Kohlmeyer 1979).
Anamorph: none reported.
Material examined: ITALY, in rhizomes of Posidonia
oceanica (Posidoniaceae), 1861, Caldesi (S, isotype of
Sphaeria posidoniae)
Anamorphs reported for genus: none.
Literature: Kohlmeyer 1969; Kohlmeyer and Kohlmeyer
1979; Suetrong et al. 2009.
Type species
Helicascus Kohlm., Can. J. Bot. 47: 1471 (1969).
(Morosphaeriaceae)
Helicascus kanaloanus Kohlm., Can. J. Bot. 47: 1471
(1969). (Fig. 35)
Ascostromata 0.6–0.78 mm high×1.25–2.75 mm diam.,
lenticular, immersed, black, carbonaceous, enclosing 3–4(−5)
loculi, pseudoclypeus composed of host cells enclosed in
black stromatic fungus material (Fig. 35a). Ascomata 235–
370 μm high×440–800 μm diam., depressed ampulliform,
horizontally arranged under a black pseudoclypeus, ostioles
70–170 μm diam., torsellioid ostiole (Adams et al. 2005),
papilla slightly rising over the surface of the pseudoclypeus,
subconical,canal filled with thick, bright orange to yellowish
periphyses, 270–435 μm high, 255–300 μm diam. Peridium
absent, partitions between loculi formed of brown, isodiametric or elongated cells of the stroma. Hamathecium of
dense, very long pseudoparaphyses. Asci 250–335×25–
30 μm, 8-spored, subcylindrical, finally oblong-clavate
(400–480 μm long), with a short pedicel, bitunicate,
thick-walled, physoclastic, apically multi-layered and
annulate, ectoascus forming a third, thin permeable outer
layer around the base, endoascus swelling in water and
becoming coiled at maturity, finally stretching and
pushing the ascus into the ostiolar canal (Fig. 35b).
Ascospores 36.5–48.5 × 18–22.5 μm, uniseriate, obovoid,
brown, 1-septate, at each end with a germ pore,
surrounded with dissolving sheath, 2.7–5.4 μm thick,
with funnel-shaped, apical indentations (Fig. 35c and d)
(adapted from Kohlmeyer and Kohlmeyer 1979).
Anamorph: none reported.
Material examined: USA, Hawaii, Oahu, Kaneohe Bay,
Heeia Swamp, on Rhizophora mangle, 4 Jun. 1968 (Herb. J.
Kohlmeyer No. 2566, holotype; No. 2565, 2567, paratype).
Generic description
Notes
Habitat marine, saprobic. Ascostromata lenticular, immersed, black, carbonaceous, enclosing several loculi,
pseudoclypeus composed of host cells enclosed in black
stromatic fungus material. Ascomata depressed ampulliform, horizontally arranged under a black pseudoclypeus,
ostiolate, torsellioid ostioles, papillate. Peridium absent,
partitions between loculi formed of brown, isodiametric or
elongated cells of the stroma. Hamathecium of dense, long
pseudoparaphyses. Asci 8-spored, bitunicate, subcylindrical
to oblong clavate, with a short pedicel and conspicuous
apical ring. Ascospores uniseriate, obovoid, brown, 1septate, at each end with a germ pore, surrounded with
dissolving sheath.
Morphology
Helicascus is another marine genus, which is characterized by its thin additional sheath around the base of the asci,
the coiling and stretching mechanism of the basal part of the
endoascus and its conspicuous apical apparatus which is not
that common in bitunicate asci (Kohlmeyer 1969). The
immersed stroma comprising several loculi sharing one
common ostiole is another striking character of Helicascus.
Notes
Morphology
Halotthia was introduced to accommodate the marine
fungus, H. posidoniae (as Sphaeria posidoniae), which is
characterized by immersed to erumpent, large, carbonaceous ascomata, thick peridium, bitunicate, 8-spored,
cylindrical asci, ellipsoidal, 1-septate, and dark brown
ascospores (Kohlmeyer 1963). Morphologically, Halotthia
is most comparable with Bicrouania maritima, but the
conical ascomata with flattened base of H. posidoniae can
be readily distinguished from B. maritima.
Phylogenetic study
Phylogenetically, Halotthia posidoniae, Pontoporeia
biturbinata and Mauritiana rhizophorae form a robust
clade, which may represent a potential family (Suetrong et
al. 2009).
Concluding remarks
Currently the familial status of Halotthia is unresolved
(Suetrong et al. 2009).
Phylogenetic study
Multigene phylogenetic analysis indicated that both
Helicascus kanaloanus and H. nypae K.D. Hyde nested
within Morosphaeriaceae (Suetrong et al. 2009).
Fungal Diversity
Fig. 34 Halotthia posidoniae (from S, isotype of Sphaeria posidoniae). a Ascomata gregarious on the host surface. b–d Mature or immature
cylindrical asci. e–h Ellipsoidal, dark-brown, 1-septate ascospores. Scale bars: a=1 mm, b–d=50 μm, e–h=5 μm
Concluding remarks
Helicascus is a well defined marine genus.
Herpotrichia Fuckel, Fungi rhenani exsic.: no. 2171
(1868). (Melanommataceae)
Generic description
Habitat terrestrial, parasitic, hyperparasitic or saprobic.
Ascomata medium-sized, immersed, erumpent to nearly
superficial, scattered to gregarious, globose to subglobose
Fungal Diversity
Fig. 35 Helicascus kanaloanus
(from Herb. J. Kohlmeyer No.
2566, holotype). a Section of
ascostroma immersed in the host
tissue. Note the torsellioid ostiole. b One-septate, brown,
asymmetrical ascospores within
the asci. c, d Released thickwalled ascospores. Note the
germ pore at the lower end of
the ascospores. Scale bars: a=
0.5 mm, b–d=20 μm
with a broad pore. Peridium composed of pseudoparenchymatous cells. Hamathecium of dense, long pseudoparaphyses, embedded in mucilage, septate, branching. Asci
cylindrical to cylindro-clavate, with a furcated pedicel.
Ascospores fusoid, ellipsoid or oblong with broadly to
narrowly round ends, 1-septate, constricted at the septum,
uni- to biseriate.
Anamorphs reported for genus: Pyrenochaeta or Pyrenochaeta-like (Sivanesan 1984).
Literature: von Arx and Müller 1975; Barr 1984; Cannon
1982; Freyer and van der Aa 1975; Mugambi and
Huhndorf 2009b; Samuels 1973; Samuels and Müller
1978; Sivanesan 1971, 1984.
Type species
Herpotrichia rubi Fuckel, Fungi rhenani exsic 2171.
(1868). (Fig. 36)
Ascomata 220–430 μm high × 240–390(-530) μm
diam., scattered to gregarious, immersed to erumpent,
rarely superficial, globose to subglobose, wall black,
coriaceous, apex with a small sometimes inconspicuous
papilla, usually with a pore, lacking periphyses (Fig. 36a
and b). Peridium 32–45 μm wide at the sides, up to 60 μm
wide at the apex, basal wall thinner, all walls comprising
cells of textura angularis, cells 2.5–4 μm diam., cell wall
2–4(−7) μm thick, exterior cells more thick-walled and
pigmented, inner cells thin-walled and less pigmented,
comprising thin-walled cells up to 9 μm diam., apex cells
smaller and walls thicker (Fig. 36b and c). Hamathecium
of dense, long pseudoparaphyses, 2–3 μm broad, embedded in mucilage, septate, branching (Fig. 36e). Asci 105–
150×12.5–15 μm (x ¼ 137:5 13:8mm, n=10), 8-spored,
bitunicate, fissitunicate, cylindrical to cylindro-clavate,
with a furcate pedicel that is 20–42.5 μm long, and
ocular chamber up to 2.5 μm wide × 2.5 μm high
(Fig. 36d and f). Ascospores 17.5–25×(5.5-)6.3–9 μm
(x ¼ 20:5 7:3mm, n=10), biseriate to partially overlapping uniseriate near the base, fusoid with narrowly
rounded ends, hyaline when immature and becoming pale
brown, 1-septate, deeply constricted at the septum, the
upper cell often broader than the lower one, verruculose
(Fig. 36g and h).
Anamorph: Pyrenochaeta rhenana Sacc. (Sivanesan
1984).
Material examined: AUSTRIA, on Rubus idaeus L.,
very rarely in the spring, in the Oestreicher meadow forest
(G, F. rh. 2171, type).
Fungal Diversity
Notes
Morphology
Herpotrichia was established by Fuckel (1868) comprising two species H. rhenana Fuckel and H. rubi Fuckel, but
no generic type was assigned. Bose (1961) designated H.
rhenana as the lectotype species with H. rubi as a synonym.
This proposal was followed by Müller and von Arx (1962)
and Sivanesan (1971). Herpotrichia rubi was later assigned
as the generic type (Holm 1979) as it was found to be validly
published 2 years earlier than H. rhenana, thus having
priority (Cannon 1982). However, Cannon (1982) reported
that Sphaeria herpotrichoides Fuckel (1864, cited as a
synonym of H. rhenana) was the earliest name. Thus he
made a new combination as H. herpotrichoides (Fuckel) P.F.
Cannon and cited H. rubi as the synonym. Herpotrichia rubi
is maintained as the type of the genus (Holm 1979; Cannon
1982), but the current name is H. herpotrichoides.
Herpotrichia is a morphologically well studied genus
(Barr 1984; Bose 1961; Müller and von Arx 1962;
Pirozynski 1972; Samuels and Müller 1978; Sivanesan
1971, 1984), and Herpotrichia sensu lato is characterized
by having subglobose, pyriform to obpyriform ascomata
and a peridium of textura angularis or comprising thickwalled polygonal cells with thin-walled hyaline cells
towards the centre. Asci are clavate to cylindrical, 4–8spored and ascospores are hyaline at first, becoming pale to
dark brown, one to many septate, constricted or not at the
septa and often surrounded by a mucilaginous sheath.
Several morphologically distinct genera were synonymized
under Herpotrichia using the above broad circumscription
(Barr 1984; Müller and von Arx 1962; Sivanesan 1984). In
particular, Barr kept Lojkania as a separate genus after
studying its type material (Barr 1984, 1990a). Sivanesan
(1984) was also of the opinion that Lojkania and Neopeckia
were distinct genera as several of their characters differed.
Byssosphaeria and Pseudotrichia have subsequently been
assigned to Melanommataceae, Lojkania to Fenestellaceae
and Neopeckia to Coccoideaceae (Barr 1984). Herpotrichia
sensu stricto is represented by H. rubi and has erumpent to
superficial ascomata or immersed in a subiculum, clavate to
cylindrical, 4–8-spored, stalked asci with a conspicuous
apical “nasse”, hyaline, 1-septate ascospores, usually becoming pale brown and several septate, constricted or not
constricted at septa, usually surrounded by sheath (Sivanesan
1984). Currently, about 90 species are included in this genus
(http://www.indexfungorum.org/, 12/01/2009).
Phylogenetic study
Herpotrichia diffusa (Schwein.) Ellis & Everh., H.
juniperi (Duby) Petr., H. herpotrichoides and H. macrotricha have been shown to have phylogenetic affinity with
the generic types of Byssosphaeria schiedermayeriana,
Melanomma pulvis-pyrius and Pleomassaria siparia, which
had been assigned under Melanommataceae (Kruys et al.
2006; Mugambi and Huhndorf 2009b; Schoch et al. 2006,
2009; Zhang et al. 2009a). In this study, Pleomassaria
siparia together with its closely related species of Prosthemium is kept in a separate family, viz Pleomassariaceae.
Concluding remarks
Even species under Herpotrichia sensu stricto (according to Sivanesan 1984) have diverse hosts (such as
gymnosperms (H. coulteri (Peck) S.K. Bose and H. parasitica (R. Hartig) Rostr.) and angiosperms (H. diffusa and
H. villosa Samuels & E. Müll.)) or substrates (like dead or
living leaves, bark or decorticated wood) (Sivanesan 1984).
Species of Herpotrichia sensu stricto are also reported from
various locations such as Europe, Asia or America, and
they have various life styles, e.g. parasitic, hyperparasitic or
saprobic (Sivanesan 1984). Additional factors (like hosts or
locations) may need to be considered in order to get a
natural concept for Herpotrichia.
Immotthia M.E. Barr, Mycotaxon 29: 504 (1987).
(Teichosporaceae)
Generic description
Habitat terrestrial, hyperparasitic. Ascomata gregarious,
globose, superficial, ostiolate, periphysate. Hamathecium
of cellular pseudoparaphyses. Asci 8-spored, bitunicate,
cylindrical, with a short pedicel. Ascospores 1-seriate,
ellipsoidal, brown to reddish brown, 1-septate, constricted
at the septum, smooth.
Anamorphs reported for genus: none.
Literature: Barr 1987a, 2002; Wang et al. 2004.
Type species
Immotthia hypoxylon (Ellis & Everh.) M.E. Barr, Mycotaxon 29: 504 (1987). (Fig. 37)
≡ Amphisphaeria hypoxylon Ellis & Everh., J. Mycol. 2:
41 (1886).
Ascomata gregarious, globose, superficial, ostiolate,
periphysate, papillate (Fig. 37a). Hamathecium of cellular
pseudoparaphyses, 2–2.5 μm broad, septate. Asci 60–82×
7–9 μm, 8-spored, bitunicate, cylindrical, with a short
pedicel (Fig. 37b, c and d). Ascospores 10–13×4.4–5.4 μm,
1-seriate, ellipsoidal, brown to reddish brown, 1-septate,
constricted at the septum, smooth (Fig. 37f, g and h)
(adapted from Wang et al. 2004).
Anamorph: none reported.
Material examined: USA, Louisiana, Pointe a la
Hache, on decaying wood, a branch of Carya oliviformis
(Juglandaceae) lying on the ground in grass (parasitic on
Fungal Diversity
Fig. 36 Herpotrichia rubi (from g, f. rh. 2171, type). a Numerous
ascomata gregariously immersed in the host tissue. b Section of an
ascoma. Note the central ostiole and peridium structure and also note
the arrangement of asci and pseudoparaphyses. c Section of partial
lateral peridium which comprises cells of textura angularis. d Part of a
mature squashed ascus. e Relatively wide, septate pseudoparaphyses. f
Immature ascus. Note the furcate pedicel. g–h One-septate ascospores.
Note the verruculose ornamentation which is visible at the sides. Scale
bars: a=0.5 mm, b=100 μm, c=50 μm, d=20 μm, e–h=10 μm
Fungal Diversity
some effused Hypoxylon), 30 Dec. 1885, A.B. Langlois, No. 138
(NY, holotype of Amphisphaeria hypoxylon Ellis & Everh.).
Notes
Morphology
Immotthia was introduced to accommodate a species of
Amphisphaeria (A. hypoxylon), which has bitunicate asci,
and is characterized by superficial, ostiolate, periphysate,
papillate ascomata, cellular pseudoparaphyses, bitunicate, 8spored, cylindrical asci, ellipsoid, smooth, brown to reddish
brown, 1-septate ascospores (Barr 1987a; Wang et al. 2004).
Phylogenetic study
None.
Concluding remarks
It seems that those Amphisphaeria species with bitunicate
asci should be assigned to Pleosporales. Morphologically,
Immotthia is somewhat comparable with Herpotrichia.
Isthmosporella Shearer & J.L. Crane, Mycologia 91: 141
(1999). (Pleosporales, genera incertae sedis)
Generic description
Habitat freshwater, saprobic. Ascomata small- to mediumsized, scattered, immersed, erumpent to superficial, globose,
papillate, ostiolate, periphysate, membranous. Peridium 2layered, outer layer composed of brown, pseudoparenchymatic, fusoid-cylindric cells, inner layer composed of fusoid,
subhyaline to pale brown, compressed cells. Hamathecium of
rare, broad, septate, interascal pseudoparaphyses. Asci 8spored, bitunicate, fissitunicate, oblong to clavate, with a
short pedicel, ocular chamber not observed. Ascospores 3–4
seriate, cylindrical to fusoid, isthmoid at centre, constricted at
septa, isthmus 1-septate, surrounded by a gelatinous sheath.
Anamorphs reported for genus: none.
Literature: Shearer and Crane 1999.
Type species
Isthmosporella pulchra Shearer & J.L. Crane, Mycologia
91: 142 (1999). (Fig. 38)
Ascomata 240–330 μm diam., scattered on decorticated
wood, immersed, erumpent to superficial, globose, black,
papillate, papilla short, cylindrical, 60 μm long×55 μm
wide, ostiolate, periphysate, membranous (Fig. 38a). Peridium 2-layered, outer 3–4 cell layers composed of brown,
pseudoparenchymatic, fusoid-cylindric cells, 2–6.5 μm
long; inner layer composed of 5–7 rows of fusoid, subhyaline to pale brown compressed cells, 11–20×2–3.5 μm
diam. (Fig. 38a and b). Hamathecium of rare, broad,
septate, interascal pseudoparaphyses (Fig. 38f). Asci (95-)
135–160(−175)×(25-)30–45(−60) μm, 8-spored, bitunicate,
fissitunicate, oblong to clavate, with a short pedicel, ocular
chamber not observed (Fig. 38c, d and e). Ascospores 80–
105(−110)×(7-)8–10 μm, 3–4-seriate, cylindrical to fusoid,
isthmoid at centre, sometimes bent at isthmus and becoming u- or v- shaped, end cells tapering, 12–17-phragmoseptate, constricted at septa, isthmus 1-septate, 2–5.5×2–
4.5 μm diam., hyaline, frequently fragmenting to form
partspores; filled with lipid droplets that merge to form
large guttules; surrounded by a gelatinous sheath with a
dense region near the isthmus, sheath greatly enlarging in
water (Fig. 38g, h, i and j).
Anamorph: none reported.
Colonies on yeast soluble starch agar containing balsa
wood sticks effuse, white. Hyphae hyaline, septate.
Material examined: USA, New York, Adirondack Park.
Piercefield. Tupper Lake at public boat launch from Rt. 30,
UTM Zone 18, 539840 mE, 4892100mN; 44°10″59″N, 80°
31′6″W, on submerged, decorticated wood, 7 Jul. 1994, J.L.
Crane & C.A. Shearer A-254-1 (ILLS 53086, holotype).
Notes
Morphology
Isthmosporella was described as a freshwater genus
typified by I. pulchra, and is characterized by globose,
pseudoparenchymatous ascomata, sparse, septate pseudoparaphyses, fissitunicate asci and hyaline, cylindrical to
fusoid, phragmoseptate, isthmoid ascospores surrounded
with a gelatinous sheath (Shearer and Crane 1999). Based
on the morphological characters, i.e. small, globose
ascomata, peridium with small pseudoparenchymatous cells
and sparse pseudoparaphyses, Isthmosporella was assigned
to the Phaeosphaeriaceae (Shearer and Crane 1999). The
aquatic habitat of Isthmosporella, however, disagree with
the Phaeosphaeriaceae. Isthmosporella seems less likely to
belong to Pleosporineae.
Phylogenetic study
None.
Concluding remarks
Molecular phylogenetic studies should be conducted to
explore its familial placement within Pleosporales.
Kalmusia Niessl, Verh. nat. Ver. Brünn 10: 204 (1872).
(Montagnulaceae)
Generic description
Habitat terrestrial, saprobic. Ascomata small- to mediumsized, solitary, scattered or in small groups, immersed to
Fungal Diversity
Fig. 37 Immotthia hypoxylon (from holotype of Amphisphaeria hypoxylon). a Ascomata gregarious on host surface. b–d Bitunicate asci. e–
h Released 1-septate ascospores. Scale bars: a=0.5 mm; b–h=10 μm
erumpent, globose or subglobose, often laterally flattened, coriaceous, wall black, with or without papilla.
Hamathecium of dense, filliform, delicate, septate pseudoparaphyses, branching and anastomosing between and
above asci, embedded in mucilage. Asci bitunicate,
fissitunicate unknown, clavate, with a long, furcate
pedicel. Ascospores narrowly ovoid to clavate, pale
brown, 3-septate, distoseptate.
Anamorphs reported for genus: Cytoplea (Petrak and
Sydow 1926).
Fungal Diversity
Fig. 38 Isthmosporella pulchra (from ILLS 53086, holotype). a Section of an ascoma. b
Section of a partial peridium. c–
e Broadly clavate asci with short
pedicels. f Pseudoparaphyses.
g–j Ascospores. Note the 2celled isthmus in J and mucilaginous sheath in G and H.
Scale bars: a=50 μm, b–j=
20 μm (figure with permission
from http://fungi.life.illinois.
edu/search/fwa_lit_refs)
Literature: Barr 1987b, 1990a, 1992a; Lindau 1897; von
Niessl 1872.
Type species
Kalmusia ebuli Niessl, Verh. nat. Ver. Brünn 10: 204
(1872). (Fig. 39)
Ascomata 290–360 μm high×300–520 μm diam., solitary, scattered, or in small groups, immersed to erumpent,
globose or subglobose, coriaceous, wall black, with or
without papilla, ostiolate (Fig. 39a). Papilla small, up to
100 μm high, with small ostioles (Fig. 39a). Peridium 15–
40 μm wide, comprising one cell type of small, pigmented,
thick-walled cells of textura prismatica to textura angularis,
cells ca. 5×3 μm diam., cell wall 2–3 μm thick (Fig. 39b
and c). Hamathecium of dense, delicate pseudoparaphyses, 1–
1.5 μm broad, septate, branching and anastomosing between
and above asci, embedded in mucilage. Asci 75–125×10–
15 μm (x ¼ 90:5 12mm, n=10), 8-spored, bitunicate,
fissitunicate unknown, clavate, with a long, narrowed, furcate
Fungal Diversity
pedicel which is up to 45 μm long, and a low ocular chamber
(ca. 2 μm wide×1 μm high) (Fig. 39d, e and f). Ascospores
15–18×5.5–6.5 μm (x ¼ 16:3 5:8mm, n=10), biseriate,
narrowly ovoid to clavate, pale brown, 3-distoseptate, without
constriction, smooth-walled (Fig. 39g, h and i).
Anamorph: none reported.
Material examined: BELGIUM, Dolembreux, on branchlets and pieces of stumps of Sarothamnus scoparius from
woodland, Oct. 1922, V. Mouton (BR 101525–63, holotype).
Notes
Morphology
Kalmusia was formally established by von Niessl (1872),
and is mainly characterized as “immersed, sphaeroid ascoma
with central, stout papilla, surrounded by hyphae in the
substrate, stipitate asci with septate pseudoparaphyses, and
brown, 3-septate, inequilateral ascospores” (Barr 1992a).
The most morphologically comparable genus to Kalmusia is Thyridaria, which had been treated as a subgenus
under Kalmusia (Lindau 1897), and was subsequently
transferred to Platystomaceae in Melanommatales (Barr
1987b, 1990a). Compared to Thyridaria, Kalmusia has
sphaeroid ascomata, a peridium of small pseudoparenchymatous cells, basal asci and very thin pseudoparaphyses,
thus it was assigned to Phaeosphaeriaceae of the Pleosporales by Barr (1990a), and the genus is utilized to
accommodate both K. ebuli and K. clivensis (Berk. &
Broome) M.E. Barr, as well as closely related species, i.e.
K. utahensis (Ellis & Everh.) Huhndorf & M.E. Barr and K.
coniothyrium (Fuckel) Huhndorf (Barr 1992a). But this
proposal is questionable, as the clavate, distoseptate ascospores, as well as the clavate asci with very long pedicels are
uncommon in Phaeosphaeriaceae, and most recent phylogenetic study indicated that some species of Kalmusia reside
outside of Phaeosphaeriaceae (Zhang et al. 2009a).
Phylogenetic study
Both Kalmusia scabrispora Teng Kaz. Tanaka, Y.
Harada & M.E. Barr and K. brevispora (Nagas. & Y.
Otani) Yin. Zhang, Kaz. Tanaka & C.L. Schoch reside in
the clade of Montagnulaceae (Zhang et al. 2009a). Familial
placement of Kalmusia can only be verified after the DNA
sequences of the generic type (K. ebuli) are obtained.
Concluding remarks
Kalmusia is distinct amongst the Pleosporales as it has
pale brown ascospores with indistinct distosepta and clavate
asci with long pedicels. Although both K. scabrispora and
K. brevispora reside in the clade of Montagnulaceae, they
both lack the distoseptate ascospores that are possessed by
the generic type (K. ebuli). Thus, the familial placement of
Kalmusia is still undetermined.
Karstenula Speg., Decades Mycologicae Italicae ad no. 94
(in sched.) (1879). (Montagnulaceae)
Generic description
Habitat terrestrial, saprobic. Ascomata rarely small-,
usually medium-sized, immersed usually under thin
clypeus, scattered to gregarious, with flattened top and
rounded pore-like ostiole, coriaceous. Peridium 2-layered,
outer layer composed of reddish brown to dark brown
small cells, inner layer of pale compressed cells. Hamathecium of dense, cellular pseudoparaphyses. Asci cylindrical to cylindro-clavate with short furcate pedicel.
Ascospores muriform, ellipsoid to fusoid, reddish brown
to dark brown.
Anamorphs reported for the genus: Microdiplodia
(Constantinescu 1993).
Literature: Barr 1990a; Eriksson and Hawksworth 1991;
Kodsueb et al. 2006a; Munk 1957; Zhang et al. 2009a.
Type species
Karstenula rhodostoma (Alb. & Schwein.) Speg., Decades
Mycologicae Italicae no. 94. (1879). (Fig. 40)
≡ Sphaeria rhodostoma Alb. & Schwein., Consp. fung.
(Leipzig): 43 (1805).
Ascomata 250–430 μm high×450–650 μm diam., scattered
or gregarious, immersed in the subiculum which sometimes
sloths off, globose or subglobose, black, flattened top often
white or reddish and sometimes slightly protruding out of the
substrate surface, usually with a wide opening ostiole after
removing the cover, coriaceous (Fig. 40a and b). Peridium
30–40 μm wide, comprising two cell types, outer region 1layered, composed of relatively small heavily pigmented
thick-walled compressed cells, cells 2–4×5–10 μm diam.,
cell wall 2–4 μm thick, inner layer cells larger and wall
thinner, comprising cells of textura angularis, merging with
pseudoparaphyses (Fig. 40c and d). Hamathecium of dense,
long cellular pseudoparaphyses 2–3.5 μm broad, septate,
branching or anastomosing not observed. Asci 150–210×
12.5–15 μm (x ¼ 182 13:1mm, n=10), 8-spored, bitunicate,
fissitunicate, cylindrical, with a broad, furcate pedicel which is
12–35 μm long, and with an ocular chamber (to 4 μm wide×
3 μm high) (Fig. 40e and f). Ascospores 20–26×7.5–10 μm
(x ¼ 22:4 8mm, n=10), obliquely uniseriate and partially
overlapping, ellipsoid, reddish brown, with 3 transverse septa
and a vertical septum in one or two central cells, constricted at
the septa, verruculose (Fig. 40g, h and i).
Anamorph: Microdiplodia frangulae Allesch. (Constantinescu
1993).
Conidiomata globose to subglobose, 330–495 μm diam.,
in subiculum. Conidia 9–13×4–5 μm, reddish brown, 1septate (information obtained from Barr 1990a).
Fungal Diversity
Fig. 39 Kalmusia ebuli (from BR 101525–63, holotype). a Immersed to erumpent ascomata scattered on the host surface. b Section
of a partial peridium. Note the compressed peridium cells. c Section of
an ascoma. d–f Eight-spored asci with long pedicels. g Partial ascus in
pseudoparaphyses. h, i Ascospores with 3 thick-walled septa. Scale
bars: a=0.5 mm, b=50 μm, c=100 μm, d–g=20 μm, h, i=10 μm
Material examined: Fries, Suecia (received by herbarium
in 1834) (PH 01048835, type, as Sphaeria rhodostoma
Alb. & Schwein.).
rable with those of Didymosphaeria, such as ascomata
seated in subiculum or beneath a clypeal thickening, the
development of apex vary in a large degree, even to the
occasional formation of a blackened internal clypeus, and
sometimes apical cells become reddish or orange-brown
(Barr 1990a). Barr (1990a) redefined the concept of
Karstenula (sensu lato), which encompasses some species
of Thyridium. In her concept, however, Barr (1990a)
treated Karstenula as having trabeculate pseudoparaphyses and this is clearly not the case. In most cases the
Notes
Morphology
Karstenula is an ambiguous genus, which has been
synonymized under Pleomassaria (Lindau 1897; Winter
1885). Some of the ascomata characters are even compa-
Fungal Diversity
Fig. 40 Karstenula rhodostoma (from PH 01048835, type). a Line
of ascomata on host surface (after remove the decaying cover). Note
the wide ostiolar opening and light colored region around the ostiole.
b Immersed ascoma under the decaying cover (see arrow). c, d
Section of the peridium. The peridium comprises small thick-walled
cells in the outer layer. The outside comprises defuse hyphae which is
probably part of the subiculum. e Ascus with a short furcate pedicel. f
Partial ascus showing arrangement of ascospores. g–i Released
ascospores. Note the transverse and rarely vertical septa. Scale bars:
a, b=0.5 mm, c=50 μm, d–f=20 μm, g–i=10 μm
Fungal Diversity
ascospores were brown with transverse septa and sparse
longitudinal septa.
The ascomata of this species are similar to those found in
Byssosphaeria and Herpotrichia, especially in the paler area
around the ostiole and even in peridial structure and
development under a subiculum. The numerous wide cellular
pseudoparaphyses and cylindrical asci (in Herpotrichia) are
also similar. The main difference of Karstenula from other
two genera are the 3-septate ascospores with rare longitudinal septa (1-septate in Byssosphaeria and Herpotrichia).
Phylogenetic study
Karstenula forms a robust phylogenetic clade with
Phaeodothis winteri (Niessl) Aptroot, Didymocrea sadasivanii, Bimuria novae-zelandiae, Montagnula opulenta, Curreya pityophila (J.C. Schmidt & Kunze) Arx & E. Müll. and
some species of Letendraea and Paraphaeosphaeria (Kodsueb et al. 2006a; Zhang et al. 2009a). Consequently,
Karstenula might be included in Montagnulaceae.
Concluding remarks
The description of the type of Karstenula here clearly
excludes it from Melanommataceae as it has wide
pseudoparaphyses. But its Montagnulaceae status can only
be confirmed by more phylogenetic work including
sequencing the generic type of Karstenula (K. rhodostoma).
Katumotoa Kaz. Tanaka & Y. Harada, Mycoscience 46:
313 (2005). (Lentitheciaceae)
Generic description
Habitat terrestrial or freshwater, saprobic. Ascomata smallto medium-sized, scattered or in small groups, immersed to
erumpent, with a central protruding hairy papilla, subglobose. Peridium thin, comprising several layers of thinwalled compressed cells. Hamathecium of dense, cellular,
filliform, embedded in mucilage, branching and anastomosing. Asci 8-spored, bitunicate, fissitunicate, clavate with
short furcate pedicels. Ascospores apiosporous and hyaline
when young, becoming 2-septate with reddish brown
echinate central cell at maturity, with long gelatinous
terminal appendages.
Anamorphs reported for genus: none.
Literature: Tanaka and Harada 2005b; Tanaka et al. 2009;
Zhang et al. 2009a.
Type species
Katumotoa bambusicola Kaz. Tanaka & Y. Harada,
Mycoscience 46: 313 (2005). (Fig. 41)
Some information for the following description is from
Tanaka and Harada (2005).
Ascomata 240–330 μm high × 260–420 μm diam.,
scattered or in small groups, immersed, becoming erumpent, with a slightly protruding papilla covered with brown
hyphae, subglobose (Fig. 41a). Peridium 13–30 μm thick,
composed of a few layers of lightly pigmented, depressed
cells. Hamathecium of dense, long cellular pseudoparaphyses,
1.5–3 μm broad, embedded in mucilage, branching and
anastomosing. Asci 110–160×17.5–24 μm (x ¼ 139 21mm,
n=10), 8-spored, bitunicate, fissitunicate, cylindro-clavate
with a short furcate pedicel which is up to 25 μm long
(Fig. 41b and d). Ascospores 39–50(−57)×7–10 μm
(x ¼ 45:8 8:2mm, n=10), biseriate, fusoid to narrowly
fusoid with acute ends, usually curved, apiosporus and
hyaline when young, constricted at the primary septum, the
upper cell longer and broader than the lower one, smooth,
surrounded by a bipolar sheath which is up to 15 μm long,
best seen in India ink, senescent ascospores yellowish
brown, 2–4-septate (Fig. 41c).
Anamorph: none reported.
Material examined: JAPAN, Mt. Iwate, near Yakebashiri,
Hirakasa, Nishine, Iwate, on culms of Oryza sativa L., 19
Oct. 2003, K. Tanaka (HHUF 28663, holotype).
Notes
Morphology
Katumotoa was formally established by Tanaka and
Harada (2005b) to accommodate the monotypic species,
K. bambusicola, which is characterized by immersed
ascomata with a thin peridium comprising thin-walled
compressed cells, cellular pseudoparaphyses, cylindroclavate and fissitunicate asci and fusoid ascospores with
an elongated bipolar mucilaginous sheath. Based on its
immersed ascomata, psuedoparenchymatous peridium
cells and cellular pseudoparaphyses, Katumotoa was
assigned to Phaeosphaeriaceae (Tanaka and Harada
2005b; Tanaka et al. 2009), but this classification has
been shown to be incorrect in subsequent phylogenetic
studies (Tanaka et al. 2009; Zhang et al. 2009a).
Phylogenetic study
Phylogenetic analysis based on five genes (LSU, SSU,
RPB1, RPB2 and EF1) indicates that Katumotoa bambusicola resides in Lentitheciaceae, and this receives high
bootstrap support (Zhang et al. 2009a). In particular, K.
bambusicola forms a robust clade with Ophiosphaerella
sasicola (Nagas. & Y. Otani) Shoemaker & C.E. Babc.,
which has filliform ascospores (Shoemaker and Babcock
1989b).
Concluding remarks
The hyaline, apiosporous ascospores which become
2–4-celled with central reddish brown cells and large
Fungal Diversity
Fig. 41 Katumotoa bambusicola (from HHUF 28663, holotype). a Ascomata scattered on the host surface. b Asci in pseudoparaphyses. c
Hyaline ascospore with long terminal appendages. d Clavate ascus with a short pedicel. Scale bars: a=0.5 mm. b–d=20 μm
unraveling appendages are the most striking features of
this species and readily distinguish it from other
pleosporalean taxa. Both Katumotoa bambusicola and
Ophiosphaerella sasicola are associated with bambusicolous hosts, which might indicate that host spectrum in
this case, has greater phylogenetic significance than some
morphological characters (Zhang et al. 2009a).
Keissleriella Höhn., Sber. Akad. Wiss. Wien, Math.-naturw.
Kl., Abt. 1 128: 582 (1919). (Lentitheciaceae)
Generic description
Habitat terrestrial or freshwater, saprobic. Ascomata smallto medium-sized, immersed, erumpent to nearly superficial,
globose, papillate, ostiolate. Papilla covered by dark setae
or small blackened cells. Peridium thick, composed of cells
of pseudoparenchymatous and inner layer composed of pale
cells. Hamathecium of dense, long pseudoparaphyses,
rarely septate, anastomosing and branching. Asci 4- or 8spored, bitunicate, fissitunicate, cylindro-clavate, with a
Fungal Diversity
furcate pedicel and a small ocular chamber. Ascospores
hyaline to pale brown, ellipsoid to fusoid, 1-septate,
constricted at the septum (Barr 1990a).
Anamorphs reported for genus: Dendrophoma (Bose 1961).
Literature: von Arx and Müller 1975; Bose 1961; Barr
1990a; Dennis 1978; Eriksson 1967a; von Höhnel 1919;
Luttrell 1973; Munk 1957; Zhang et al. 2009a.
Type species
Keissleriella aesculi (Höhn.) Höhn., Sber. Akad. Wiss.
Wien, Math.-naturw. Kl., Abt. 1 128: 582 (1919). (Fig. 42)
≡ Pyrenochaeta aesculi Höhn., Ber. dt. bot. Ges. 35: 249
(1917).
Ascomata ca. 250 μm high × 450 μm diam., gregarious,
immersed to erumpent, globose or subglobose, with a
small black papilla, ca. 75 μm high and 110 μm broad,
with short black external setae (Fig. 42a). Peridium ca.
25–40 μm wide laterally, up to 70 μm near the apex,
thinner at the base, comprising two types of cells which
merge in the middle; outer cells composed of small
heavily pigmented thick-walled cells, cells ca. 4 μm
diam., cell wall up to 4 μm thick, and thick near the apex
and thinner laterally and absent in the immersed part of
the ascoma, inner cells less pigmented, comprising
lightly pigmented to hyaline cells, 5–7 μm thick
(Fig. 42a). Hamathecium of dense, long pseudoparaphyses, 0.8–1.2 μm broad, rarely septate, anastomosing and
branching, thicker near the base, ca. 2 μm, constricted
near the septum (Fig. 42b). Asci 80–120 × 6–11 μm (x ¼
101 8:5mm, n = 10), 4- or 8-spored, bitunicate, fissitunicate, cylindro-clavate, with a furcate pedicel which is
up to 20–40 μm long, with a small ocular chamber
(Fig. 42e and f). Ascospores 13–18 × 4–5.5 μm
(x ¼ 14:5 4:8mm, n = 10), obliquely uniseriate and partially overlapping, fusoid with narrowly rounded ends,
hyaline, 1-septate, constricted at the septum, smooth
(Fig. 42c and d).
Anamorph: none reported.
Material examined: AUSTRIA, Brentenmaistal in the
Viennese forest, Aesculus hippocastanum L., 1916, Höhnel
(FH, holotype of Otthiella aesculi). (Note: only two slides;
setae cannot be seen from the slides but could be seen from
the drawings on the cover).
Notes
Morphology
Keissleriella is characterized by ascomata with setae in
and over the papilla, asci are cylindrical and ascospores are
hyaline, 1-septate. Based on the morphological characters,
K. aesculi was regarded as conspecific with K. sambucina;
as an earlier epithet, K. sambusina typifies the genus (see
comments by Barr 1990a). Munk (1957) placed Trichometasphaeria and Keissleriella in Massarinaceae, and distinguished them by their substrates (Trichometasphaeria
occurs on herbaceous plants and Keissleriella on woody
substrates). Bose (1961) combined Trichometasphaeria
under Keissleriella, which was followed by some workers
(von Arx and Müller 1975; Dennis 1978; Eriksson 1967a;
Luttrell 1973). Barr (1990a), however, maintained these as
distinct genera based on the differences of peridium
structure and pseudoparaphyses.
Phylogenetic study
The phylogeny of Keissleriella is poorly studied.
Limited phylogenetic information indicates that K. cladophila forms a robust clade with other species of Lentitheciaceae (Zhang et al. 2009a).
Concluding remarks
The presence of black setae on the surface of papilla is a
striking character of Keissleriella, but phylogenetic significance of setae is undetermined yet.
Lentithecium K.D. Hyde, J. Fourn. & Yin. Zhang, Fungal
Divers. 38: 234 (2009). (Lentitheciaceae)
= Tingoldiago K. Hirayama & Kaz. Tanaka, Mycologia
102: 740 (2010) syn. nov.
Generic description
Habitat freshwater, saprobic. Ascomata small, scattered
or gregarious, immersed, slightly erumpent, depressed
spherical to lenticular, ostiolate, papillate or epapillate.
Peridium thin. Hamathecium of cellular pseudoparaphyses. Asci 8-ascospored, bitunicate, fissitunicate, clavate,
short-stipitate. Ascospores broadly fusoid with broadly
rounded ends, 1-septate, constricted, hyaline, usually
with sheath.
Anamorphs reported for genus: none.
Literature: Shearer et al. 2009; Zhang et al. 2009a, b.
Type species
Lentithecium fluviatile (Aptroot & Van Ryck.) K.D. Hyde, J.
Fourn. & Yin. Zhang, Fungal Divers. 38: 234 (2009). (Fig. 43)
≡ Massarina fluviatilis Aptroot & Van Ryck., Nova
Hedwigia 73: 162 (2001).
Ascomata 230–260 μm high×280–325 μm diam., scattered
or gregarious, immersed, slightly erumpent, subglobose to
depressed spherical, under a small black pseudostroma
originating from the apical part of the peridium, apex slightly
papillate, ostiole rounded, 60–70 μm diam. (Fig. 43a and b).
Peridium 15–20 μm thick at sides and at base, comprising 4–5
layers of angular cells more thick-walled outwards, 50–55 μm
Fungal Diversity
Fig. 42 Keissleriella sambucina (from FH, holotype of Otthiella
aesculi). a Section of an ascoma. b Pseudoparaphyses which are
narrow (less than 1.5 μm) and branch and anastomosing as
trabeculate. c, d Hyaline ascospores with distinct constrictions at the
septa. e Asci amongst narrow pseudoparaphyses. F. Ascus with a
pedicel and ocular chamber. Scale bars: a=100 μm, b–f=10 μm
thick at apex, of small very thick-walled cells. Hamathecium
of cellular pseudoparaphyses, 2–2.5 μm broad (Fig. 43c and
d). Asci 89–100×19–21 μm, 8-spored, bitunicate, fissitunicate, clavate, bumpy, short-stipitate, apex without obvious
apical chamber (Fig. 43e). Ascospores 27–35×8.5–9.4 μm,, 2-
3-seriate, broadly fusoid with broadly rounded ends, straight to
slightly curved, 1-septate, slightly constricted, with four large
guttules, hyaline, smooth-walled, a very thin mucilaginous
sheath can be occasionally observed in India ink but in most
cases no sheath can be observed (Fig. 43f and g).
Fungal Diversity
Anamorph: none reported.
Material examined: FRANCE, Haute Garonne: Avignonet, Lac de Rosel, artificial lake, on bark and wood of a
submerged branch Populus sp., 23 Nov. 2006, leg. Michel
Delpont, det. Jacques Fournier (IFRD 2039, holotype).
Notes
Morphology
Lentithecium was introduced to accommodate some freshwater fungi previous assigned under Massarina, such as M.
arundinacea (Sowerby) Leuchtm. and M. fluviatilis (Zhang et
al. 2009a). It is characterized by its immersed and
lenticular ascomata, thin peridium which is almost equal
in thickness, short pedicellate asci and fusoid or filliform,
hyaline or rarely lightly pigmented, 1- to multi-septate
ascospores (Zhang et al. 2009b). Lentitheciaceae was
introduced to accommodate Lentithecium and some other
related taxa (Zhang et al. 2009a).
Phylogenetic study
The clade of Lentitheciaceae comprises the generic type
Lentithecium fluviatile, as well as L. arundinaceum
(Sowerby) K.D. Hyde, J. Fourn. & Yin. Zhang, Stagonospora macropycnidia, Wettsteinina lacustris (Fuckel) Shoemaker & C.E. Babc., Keissleriella cladophila, and the
bambusicolous species Katumotoa bambusicola and
Ophiosphaerella sasicola, which receive high bootstrap
support (Zhang et al. 2009a).
Concluding remarks
Tingoldiago graminicola K. Hirayama & Kaz. Tanaka
form a robust clade with species of Lentithecium (Shearer et
al. 2009). Tingoldiago has lenticular immersed to erumpent
ascomata, numerous and septate pseudoparaphyses,
cylindro-clavate asci and hyaline, 1-septate ascospores with
sheath. All of these characters fit Lentithecium well. We
treat Tingoldiago as a synonym of Lentithecium.
Leptosphaeria Ces. & De Not., Comm. Soc. crittog. Ital. 1:
234 (1863). (Leptosphaeriaceae)
Generic description
Habitat terrestrial, saprobic or parasitic. Ascomata small- to
medium-sized, solitary, scattered or in small groups,
erumpent to superficial, subglobose, broadly or narrowly
conical, papillate, ostiolate. Peridium thick, comprising
layers of cells of textura angularis. Hamathecium of dense
cellular pseudoparaphyses, embedded in mucilage, anastomosing and branching. Asci 8-spored, bitunicate, fissitunicate unknown, cylindrical with a furcate pedicel and a large
ocular chamber. Ascospores fusoid or narrowly fusoid,
brown or reddish brown, 3-septate, constricted at each
septum.
Anamorphs reported for genus: Coniothyrium and
Phoma (Hyde et al. 2011; Sivanesan 1984).
Literature: von Arx and Müller 1975; Barr 1987a, b; Cesati
and de Notaris 1863; Crane and Shearer 1991; Dong et al.
1998; Eriksson 1967a; Eriksson and Hawksworth 1986,
1991; de Greuter et al. 1988; Hedjaroude 1969; von Höhnel
1907; Holm 1957, 1975; Huhndorf et al. 1990; Luttrell 1973;
Müller 1950; Munk 1957; Saccardo 1878b, 1883, 1891,
1895; Schoch et al. 2009; Shearer 1993; Shearer et al. 1990;
Shoemaker 1984a; Sivanesan 1984; Zhang et al. 2009a.
Type species
Leptosphaeria doliolum Ces. & De Not., Comm. Soc.
crittog. Ital. 1: 234 (1863). (Fig. 44)
≡ Sphaeria doliolum Pers., Icon. Desc. Fung. Min.
Cognit. (Leipzig) 2: 39 (1800).
Ascomata 340–450 μm high×380–500 μm diam., solitary,
scattered or in small groups, superficial, subglobose, broadly
or narrowly conical, with a flattened base on the host surface,
black, usually with 2–4 ring-like ridges surrounding the
ascomata surface, apex with a conical, usually shiny papilla
(Fig. 44a). Peridium 85–110 μm wide at sides, thinner at the
apex, comprising two types of cells, outer layer composed of
small thick-walled cells of textura angularis, cells <2 μm
diam., cell wall up to 8 μm thick, surface heavily pigmented
and inner lightly pigmented, apex cells smaller, walls thicker,
and cells more heavily pigmented, inner layer composed of
subhyaline relatively thin-walled cells of textura angularis,
3–6 μm diam., wall up to 5 μm, cells near the base larger and
wall thinner and paler (Fig. 44b). Hamathecium of dense,
long cellular pseudoparaphyses, 1.5–3 μm broad, embedded
in mucilage, anastomosing and branching. Asci 110–150×7–
9(−10) μm (x ¼ 130:6 8:5mm, n=10), 8-spored, bitunicate,
fissitunicate unknown, cylindrical, furcate pedicel which is
usually less than 25 μm long, with a large ocular chamber
(Fig. 44c, d and e). Ascospores 25–31 × 4.5–6 μm
(x ¼ 27:7 5:3mm, n=10), uniseriate and somewhat partially overlapping, narrowly fusoid with sharp to narrowly
rounded ends, reddish brown, 3-septate, constricted at each
septum, smooth (Fig. 44f).
Anamorph: Phoma hoehnelii (Sivanesan 1984).
Material examined: Herb., Persoon 910270–650 (L,
lectotype).
Notes
Morphology
Leptosphaeria was first established by Cesati and de
Notaris (1863) with 26 species included; L. doliolum (Pers.:
Fr.) Ces. & De Not. was subsequently selected as the
Fungal Diversity
lectotype species (de Greuter et al. 1988; Holm 1975;
Shearer et al. 1990). Leptosphaeria was originally
defined based mainly on the characters of ascospores
being ellipsoid or fusoid, one to many septa, hyaline to
dark brown. These few common characters meant that
Leptosphaeria comprised many species, and some of
them should be assigned to either Euascomycetes or
Loculoascomycetes (Crane and Shearer 1991). Leptosphaeria had been divided based on host and habitat
(Saccardo 1878b, 1891, 1895) as well as the pseudothecium (glabrous, hairy, setose) and ascospore septation
(see comments by Crane and Shearer 1991). von Höhnel
(1907) used centrum structure in the classification of
Leptosphaeria, and divided Leptosphaeria into three
genera, viz. Leptosphaeria, Scleropleella and Nodulosphaeria. Müller (1950) subdivided Leptosphaeria into
four sections based on pseudothecial and centrum
structure as well as ascospore characters. This classification was modified by Munk (1957), who named these
four sections as section I (Eu-Leptosphaeria), section II
(Para-Leptosphaeria), section III (Scleropleella) and
section IV (Nodulosphaeria). Holm (1957) used a
relatively narrow concept for Leptosphaeria, which
included species closely related to the generic type, L.
doliolum. This viewpoint was accepted by some workers
(Eriksson 1967a; Hedjaroude 1969; Shoemaker 1984a).
Nevertheless, it still seems a heterogeneous group of
fungi (see comments by Crane and Shearer 1991). Its position
among the Loculoascomycetes is also debated. It has been
placed in the Pleosporaceae (von Arx and Müller 1975;
Luttrell 1973; Sivanesan 1984) or Leptosphaeriaceae (Barr
1987a, b; Eriksson and Hawksworth 1991) or Phaeosphaeriaceae (Eriksson and Hawksworth 1986).
Phylogenetic study
Molecular phylogenetic analysis based on multigenes
indicated that species of Leptosphaeria (including the
generic type L. doliolum) and Neophaeosphaeria form a
paraphyletic clade with moderate bootstrap support (Dong
et al. 1998; Schoch et al. 2009; Zhang et al. 2009a), which
is sister to other families of Pleosporales (Zhang et al.
2009a). Thus the familial rank of the Leptosphaeriaceae
could be temporarily verified, but further molecular
phylogenetic study is needed in which more related taxa
should be included.
Concluding remarks
Morphologically, Leptosphaeria is mostly comparable
with Amarenomyces, Bricookea, Diapleella, Entodesmium,
Melanomma, Nodulosphaeria, Paraphaeosphaeria, Passeriniella, Phaeosphaeria and Trematosphaeria. While it
prefers non-woody parts of dicotyledonous hosts, its
cylindrical ascus with short pedicel and smooth, fusoid
Fig. 43 Lentithecium fluviatile (from IFRD 2039). a Erumpent
ascomata scattering on the host surface. b Habitat section of the
immersed ascomata. c, d Section of an ascoma and a partical peridium.
Note the peridium cells of textura angularis. e Clavate 8-spored ascus
with a short pedicel. f, g Hyaline, 1-septate broadly fusoid ascospores.
Scale bars: a, b=0.5 mm, c=100 μm, d=50 μm, e–g=20 μm
and multi-septate ascospores make it readily distinguishable
from all other genera (Shoemaker 1984a).
Leptosphaerulina McAlpine, Fungus diseases of stonefruit trees in Australia and their treatment: 103 (1902).
(Didymellaceae)
Generic description
Habitat terrestrial, parasitic or saprobic. Ascomata small,
scattered, immersed, globose to subglobose, with a small,
slightly protruding papilla, ostiolate. Peridium thin. Hamathecium of rare or decomposing cellular pseudoparaphyses.
Asci bitunicate, obpyriform. Ascospores broadly clavate or
cylindrical, hyaline, turning pale brown when old, asymmetrical, multi-septate, smooth-walled.
Anamorphs reported for genus: Pithoascus and Pithomyces (Hyde et al. 2011).
Literature: Barr 1972; Chlebicki 2002; Crivelli 1983;
Kodsueb et al. 2006a; Zhang et al. 2009a.
Type species
Leptosphaerulina australis McAlpine, Fungus diseases of
stone-fruit trees in Australia and their treatment: 103 (1902).
(Fig. 45)
Ascomata 140–170 μm diam., scattered, immersed,
globose to subglobose, with a small slightly protruding
papilla, ostiolate (Fig. 45a). Peridium thin, composed of
one or two layers of large cells of textura angularis, pale
brown (Fig. 45a). Hamathecium of rare or decomposing
cellular pseudoparaphyses, up to 5 μm broad, filling
the gaps between the asci. Asci 38–53 × 55–75 μm
(x ¼ 67:5 43:3mm, n=10), 8-spored, without pedicel, bitunicate, fissitunicate dehiscence not observed, obpyriform, with a
large ocular chamber and apical ring (Fig. 45b and c).
Ascospores 30–40(-47)×11–14 μm (x ¼ 36:5 13mm, n=
10), broadly clavate, hyaline, turning pale brown when old,
asymmetrical, upper hemisphere usually with one transverse septum and with a somewhat narrowly rounded
end, lower hemisphere usually with two transverse septa
and with broadly rounded ends, slighted constricted at
the primary septum, mostly with one vertical septum in
each central cell, smooth, with thin gelatinous sheath
when young, 2–3 μm thick (Fig. 45d and e).
Anamorph: none reported.
b
Fungal Diversity
Fungal Diversity
Fig. 44 Leptosphaeria doliolum (from L, lectotype). a Ascomata on the host surface. Note the shiny black surface. b Section of the partial peridium.
Note the uneven thickness. c–e Asci with a short pedicel. f Three ascospores in ascus. Scale bars: a=0.5 mm, b=100 μm, c–f=20 μm
Fungal Diversity
Material examined: USA, Kansas, Kansas State College,
on Poa pratensis L. Grass plots, 2 Jul. 1953, leg. T.
Rogerson, det. L.E. Wehmeyer (NY, C.T. Rogerson 3836).
Notes
Morphology
Leptosphaerulina, introduced by McAlpine (1902), is
characterized by small immersed ascomata, obpyriform asci
with a large ocular chamber and apical ring as well as
muriformly septate ascospores which may be hyaline or
pigmented. Species of Leptosphaerulina may occur on
monocotyledons or dicotyledons. Leptosphaerulina is most
comparable with Pleospora, and the only difference between
them is that Leptosphaerulina has smaller ascomata and
hyaline ascospores that only become pigmented after
discharge, whereas the ascospores of Pleospora become
brown within the asci. Currently, about 60 names are
accepted in this genus, and some even reported from marine
environments, e.g. L. mangrovei (Inderbitzin et al. 2000).
Phylogenetic study
Based on multigene phylogenetic analysis, two putative
strains of Leptosphaerulina australis, the generic type of
Leptosphaerulina, from Switzerland (CBS 311.51) and
Indonesia (CBS 317.83) resided within Didymellaceae (de
Gruyter et al. 2009; Zhang et al. 2009a).
Concluding remarks
Because of its morphological confusion with Pleospora
and the diversity of habitats within the genus, Leptosphaerulina sensu lato is likely to be polyphyletic. Fresh
collections of this species are needed from Australia to
epitypify this taxon and define the genus in a strict sense.
The specimen described here is a collection from USA and
therefore may not represent the type.
Lewia M.E. Barr & E.G. Simmons, Mycotaxon 25: 289
(1986). (Pleosporaceae)
Generic description
Habitat terrestrial, parasitic or saprobic? Ascomata
small, scattered, erumpent to nearly superficial at maturity, subglobose to globose, black, smooth, papillate,
ostiolate. Papilla short, blunt. Peridium thin. Hamathecium of pseudoparaphyses. Asci (4–6-)8-spored, bitunicate, fissitunicate, cylindrical to cylindro-clavate, with a
short, furcate pedicel. Ascospores muriform, ellipsoid to
fusoid.
Anamorphs reported for genus: Alternaria (Simmons 1986).
Literature: Kwasna and Kosiak 2003; Kwasna et al. 2006;
Simmons 1986, 2007; Vieira and Barreto 2006.
Type species
Lewia scrophulariae (Desm.) M.E. Barr & E.G. Simmons,
Mycotaxon 25: 294 (1986). (Fig. 46)
≡ Sphaeria scrophulariae Desm., Plantes cryptogames du Nord de la France, ed. 1 fasc. 15:no. 718
(1834).
Ascomata ca. 150–200 μm diam., scattered, erumpent to
nearly superficial at maturity, subglobose to globose, black,
smooth, papillate. Papilla short, blunt. Peridium thin.
Hamathecium of septate pseudoparaphyses, ca. 2–2.5 μm
broad, anastomosing or branching not observed. Asci 100–
140×13–17 μm, (4–6-)8-spored, bitunicate, fissitunicate,
cylindrical to cylindro-clavate, with a short, furcate pedicel,
ocular chamber unknown (Fig. 46a). Ascospores ellipsoid,
5 (rarely 6 or 7) transversal septa and one longitudinal
septum mostly through the central cells, yellowish brown to
gold-brown, 20–24×8–10 μm (x ¼ 21:5 9:1mm, n=10),
constricted at median septum, smooth or verruculose
(Fig. 46b, e and f).
Anamorph: Alternaria conjuncta (Simmons 1986).
Primary conidiophore simple with a single conidiogenous locus; conidia produced in chains, the first conidia
in chain is larger, 30–45×10–12 μm, 7 transverse septa,
1–2 longitudinal or oblique septa in lower cells.
Secondary conidiophore with 5–7 conidiogenous loci,
sometimes branched; sporulation in chains, rarely
branched.
Material examined: (FH, slide from lectotype).
Note: The specimen contains only a slide, so limited
structures could be observed e.g. ascospores. The information about ascomata, peridium and whole asci is referred to
Simmons (1986).
Notes
Morphology
Lewia has “Pleospora-like” teleomorphs, while it has
Alternaria anamorphs, which are characterized by the
beakless conidia connected together with secondary
conidiophore (Simmons 1986). Based on these characters,
more species under this genus were subsequently reported,
i.e. Lewia avenicola Kosiak & Kwaśna (Kwasna and
Kosiak 2003); L. chlamidosporiformans B.S. Vieira & R.
W. Barreto (Vieira and Barreto 2006); L. alternarina (M.
D. Whitehead & J.G. Dicks.) E.G. Simmons and L.
daucicaulis E.G. Simmons (Simmons 2007). Currently
Lewia comprises 15 species (http://www.mycobank.org,
24-02-2009).
Phylogenetic study
Phylogenetic analysis based either on SSU rDNA
sequences or on multigenes indicated that Lewia species
Fungal Diversity
Fig. 45 Leptosphaerulina australis (from NY, C.T. Rogerson 3836).
a. Compressed ascoma. Note the obpyriform asci within the ascoma
and the thin peridium. b, c. Eight-spored asci released from the
ascomata. Note the apical apparatus (arrowed). d. Ascospores with
thin sheath. e. An old pale brown ascospore. Scale bars: a-c=50 μm,
d, e=10 μm
(Allewia eureka (E.G. Simmons) E.G. Simmons = L.
eureka) form a robust clade with other members of
Pleosporaceae (Schoch et al. 2006; Schoch et al. 2009;
Zhang et al. 2009a).
Generic description
Concluding remarks
Its position in Pleosporaceae is confirmed.
Lichenopyrenis Calat., Sanz & Aptroot, Mycol. Res. 105:
634 (2001). (?Pleomassariaceae)
Habitat terrestrial, parasitic on lichens. Ascomata mediumsized, globose or subglobose. Hamathecium of dense,
filliform, branching, septate pseudoparaphyses. Asci bitunicate, fissitunicate, clavate, with a short sometimes furcate
pedicel. Ascospores ellipsoidal with broadly rounded ends,
pale orange-brown, 1-distoseptate.
Anamorphs reported for genus: see below.
Literature: Calatayud et al. 2001.
Fungal Diversity
Fig. 46 Lewia scrophulariae
(from FH, slide from lectotype). a Cylindrical ascus with a
short pedicel. b Ascospores in
asci. c–f Released muriform
brown ascospores. Scale bars:
a=20 μm, b–f=10 μm
Type species
Lichenopyrenis galligena Calat., Sanz & Aptroot, Mycol.
Res. 105: 636 (2001). (Fig. 47)
Ascomata 140–260 μm high×140–250 μm diam., gregarious, initially immersed in galls, later becoming erumpent,
globose or subglobose, black, roughened (Fig. 47a and b).
Peridium 18–25 μm wide, composed of 2–5 layers of heavily
pigmented cells of textura angularis to compressed, cells
6–11 μm diam., cell wall 1–3 μm thick (Fig. 47c, d and e).
Hamathecium of dense, long filamentous pseudoparaphyses, 2.5–4 μm broad, branching, septate. Asci 65–85×
15–20 μm (x ¼ 74 18mm, n=10), 8-spored, bitunicate,
fissitunicate, broadly clavate, with a short, thick, some-
Fungal Diversity
times furcate pedicel, up to 13 μm long, ocular chamber
not observed (Fig. 47f, g, h and k). Ascospores 16–20×9–
11 μm (x ¼ 18 10mm, n=10), biseriate, ellipsoidal, pale
orange-brown, 1-distoseptate, with prominent swelling at
the septum, containing refractive globules, smooth (Fig. 47i,
j and l).
Anamorph: The following description is from Calatayud et
al. (2001).
Conidiomata pycnidial, arising in galls together with the
ascomata, immersed, ca. 100–200 μm diam.; wall dark
brown throughout, composed of 2–5 layers of angular to
laterally compressed cells; cells relatively large, ca. 8–
16 μm diam. in superficial view. Conidiophores formed by
1–3 cells, frequently branched and with the uppermost cells
bearing 1–4 conidiogenous cells; cells±cylindrical, hyaline
except at the base, which are sometimes pale brown, 7–15×
3–4 μm. Conidiogenous cells tapered towards the apex, 14–
18×3–4 μm. Conidia 5–7×1.5–2 μm. Vegetative hyphae
hyaline.
Material examined: SPAIN, Andalucía, Province, Jaén,
Andújar, lichenicolous on Leptochidium albociliatum
(Desm.) M. Choisy on acid volcanic rock, 19 Apr. 2000,
V. Calatayud (MA-Lichen 12715, holotype).
Notes
Morphology
Lichenopyrenis was formally established by Calatayud et
al. (2001) based on its “perithecioid ascomata with peridium
comprising compressed cells, fissitunicate and J- asci, wide
hamathecium filaments, and 1-septate pale orange-brown
ascospores with distoseptate thickenings at maturity”, and is
monotypic with L. galligena. The genus was temporarily
assigned to Pleomassariaceae. Lichenopyrenis galligena is a
parasite of lichens, occurring in galls in the thallus of the host
(Calatayud et al. 2001).
Phylogenetic study
None.
Concluding remarks
This is one of the few species that are parasitic on
lichens. The most comparable species are Parapyrenis
lichenicola Aptroot & Diederich and Lacrymospora parasitica Aptroot (both in Requienellaceae, Pyrenulales) as
well as some species from Dacampiaceae. The peridium
structure, cellular pseudoparaphyses, distoseptate and
smooth, orange-brown ascospores as well as the anamorphic stage of Lichenopyrenis can easily distinguish from all
of them (Calatayud et al. 2001).
Lineolata Kohlm. & Volkm.-Kohlm., Mycol. Res. 94: 687
(1990). (Pleosporales, genera incertae sedis)
Generic description
Habitat marine, saprobic (or perthophytic?). Ascomata
medium-sized, gregarious, immersed to erumpent, obpyriform, ostiolate, papillate. Peridium thin, comprising two
types of cells; outer cells thick stratum pseudostromatic,
inner stratum thin, composed of a few layers of hyaline
cells of textura angularis. Hamathecium of dense, long
trabeculate pseudoparaphyses, embedded in mucilage,
anastomosing and septate. Asci 8-spored, bitunicate, cylindrical, with short pedicels, with an ocular chamber.
Ascospores uniseriate to partially overlapping, ellipsoidal,
dark brown, 1-septate.
Anamorphs reported for genus: none.
Literature: Kohlmeyer and Kohlmeyer 1966; Kohlmeyer
and Volkmann-Kohlmeyer 1990.
Type species
Lineolata rhizophorae (Kohlm. & E. Kohlm.) Kohlm. &
Volkm.-Kohlm., Mycol. Res. 94: 688 (1990). (Fig. 48)
≡ Didymosphaeria rhizophorae Kohlm. & E. Kohlm.
Icones Fungorum Maris (Lehre) 4 & 5: tab. 62a (1967).
Ascomata 300–490 μm high × 200–360 μm diam.,
gregarious, immersed to erumpent, obpyriform, ostiolate,
papillate, subcarbonaceous to subcoriaceous, blackish
brown (Fig. 48a). Peridium 37–45 μm thick, comprising
two types of cells; outer cells thick stratum pseudostromatic, composed of irregular or roundish, dark brown cells,
on the outside with a more or less recognizable hyphal
structure, enclosing some decaying cells of the host, inner
stratum thin, composed of four or five layers of hyaline,
polygonal, elongate, thin-walled cells with large lumina,
merging into the pseudoparaphyses. Hamathecium of
dense, long trabeculate pseudoparaphyses, 1–1.5 μm broad,
embedded in mucilage, anastomosing and septate. Asci
150–175×14–17.5 μm, 8-spored, bitunicate, cylindrical,
with short pedicels, with an ocular chamber (Fig. 48b).
Ascospores 23–32(−33)×9–12 μm, uniseriate to partially
overlapping, ellipsoid, dark brown, 1-septate, not or slightly
constricted at the septum, striate by delicate costae that run
parallel or in a slight angle to the longitudinal axis of the
ascospore (Fig. 48c, d, e and f) (adapted from Kohlmeyer
and Kohlmeyer 1979).
Anamorph: none reported.
Material examined: US, Florida, Middle Torch Key, on
Rhizophora mangle, 21 Nov. 1965, J. Kohlmeyer (Herb. J.
Kohlmeyer No. 2390b, isotype); Pirate Grove Key, on R.
mangle, 5 Jan. 1964 (Herb. J. Kohlmeyer No. 1721 paratype); Florida, Virginia Key, on R. mangle, 1 Jan. 1964, leg.
E. Kohlmeyer (Herb. J. Kohlmeyer No. 1751 paratype);
Florida, Torch Key, on R. mangle, 20 Nov. 1965, leg. J.
Kohlmeyer (Herb. J. Kohlmeyer No. 2423 paratype).
Fungal Diversity
Fig. 47 Lichenopyrenis galligena (from MA-Lichen 12715, holotype). a, b Ascomata forming in the host tissues. c, d Sections of
ascomata. e Section of a partial peridium. f–h, k Broadly clavate asci.
Note the short rounded pedicel. i, j, l Ascospores. Note the small
swellings at the septa. Scale bars: a, b=0.5 mm, c, d=100 μm, e=
50 μm, f–h, k=20 μm, i, j, l=10 μm
Fungal Diversity
Notes
Morphology
Lineolata was monotypified by L. rhizophorae, which was
originally introduced by Kohlmeyer and Kohlmeyer (1966) as a
species of Didymosphaeria (as D. rhizophorae). Based on the
morphology of ascomata and asci, Barr (1990a) assigned it
under Lojkania (as L. rhizophorae). Kohlmeyer and VolkmannKohlmeyer (1990) restudied this species and noticed that the
absence of clypeus, almost superficial ascomata, coloured
peridium, a hamathecium with gelatinous matrix, asci with
apical ring-like structure and the ornamented ascospores are
quite different from the modified concept of Didymosphaeria.
Fig. 48 Lineolata rhizophorae
(from Herb. J. Kolmeyer No.
2390b, isotype of Didymosphaeria rhizophorae). a Ascomata immersed in the host
substrate with protruding papilla. b Ascospores within an ascus. Note the ascospore
arrangement. c–f One-septate
ascospores. Note the striate ornamentation in (c). Scale bars:
a=100 μm, a, b=20 μm, c–f=
10 μm
Thus they introduced Lineolata to accommodate D. rhizophorae (Kohlmeyer and Volkmann-Kohlmeyer 1990).
Phylogenetic study
Three isolates of Lineolata rhizophorae from varied geographic
localities were analyzed by Suetrong et al. (2009) and shown to be
related to Caryosporella rhizophorae in Dothideomycetidae and
excluded from Pleosporomycetidae and Pleosporales.
Concluding remarks
Based on initial molecular work it is likely that this species
does not belong to Pleosporales in spite of its dense
pseudoparaphyses and other characters shared with the order.
Fungal Diversity
Loculohypoxylon M.E. Barr, Mycotaxon 3: 326 (1976).
(Teichosporaceae)
Generic description
Habitat terrestrial, saprobic. Ascomata relatively small,
gregarious, immersed to erumpent, globose or subglobose, forming under a clypeus, papillate, ostiolate.
Peridium thin, a single layer comprising hyaline thinwalled cells of textura angularis or textura prismatica.
Hamathecium of septate pseudoparaphyses. Asci (2–4-)8spored, bitunicate, cylindrical to cylindro-clavate, with a
short, furcate pedicel, and wide ocular chamber. Ascospores broadly elliptic to subglobose, often apiculate at
both ends, pale to dark brown, aseptate, with a germ
slit.
Anamorphs reported for genus: none.
Literature: von Arx and Müller 1975; Barr 1976.
Type species
Loculohypoxylon grandineum (Berk. & Rav.) Barr, Mycotaxon 3: 326 (1976). (Fig. 49)
≡ Diatrype grandinea Berk. & Rav., in Berkeley,
Grevillea 4: 95 (1876).
Ascomata 85–130 μm high×75–145 μm diam., gregarious, immersed to widely erumpent, globose or subglobose, under a reddish brown to black clypeus, papillate,
ostiolate (Fig. 49a and b). Peridium 18–30 μm thick
laterally, 1-layered, composed of hyaline thin-walled cells
of textura angularis to prismatica, cells up to 5×9 μm
diam., cell wall 0.5–1 μm thick, apex cells smaller and
walls thicker (Fig. 49c). Hamathecium comprising 2–3 μm
broad, septate pseudoparaphyses. Asci 70–90 × 10–
12.5 μm (x ¼ 76:5 10:9mm, n = 10), (2–4-)8-spored,
bitunicate, cylindrical to cylindro-clavate, with a short,
furcate pedicel, up to 25 μm long, with a wide ocular chamber
(Fig. 49f, g, and h). Ascospores 7.5–10 × 5–7 μm
(x ¼ 8:3 5:9mm, n=10), uniseriate to partially overlapping
at the upper part, broadly elliptic to subglobose, often
apiculate at both ends, pale to dark brown, aseptate, with a
germ slit (Fig. 49d and e).
Anamorph: none reported.
Material examined: USA, New Jersey, Newfield, on
bark of Quercus coccinea, Sept. 1878, as Diatrype
grandinea, Ellis N.A.F. 494 (NY, MASS); on Quercus
sp. wood, Nov. 1893, as Anthostoma grandinea B. & Rav.,
Ellis & Everhart, N.A.F. 494 (NY); Newfield, Oct. 1881,
as Diatrype grandinea (NY); Newfield, Jan. 1882, on
Quercus coccinea, as Diatrype grandinea B. & Rav, Ex
Herb Ellis (NY); Newfield, Nov. 1893, as Anthostoma
grandinea, on bark of fallen trunks of Quercus coccinea
(NY).
Notes
Morphology
Loculohypoxylon grandineum is one of the rare pleosporalean species having aseptate ascospores. When emphasis is given to ascospore morphology, Semidelitschia
(monotypified by S. agasmatica Cain & Luck-Allen) is the
most comparable genus. The large ascomata and ascospores, the mucilaginous sheath surrounding the ascospores
as well as the coprophilous habitat of S. agasmatica differ
from L. grandineum greatly. Thus Loculohypoxylon was
introduced as a new genus.
Phylogenetic study
None.
Concluding remarks
Aseptate ascospores are rare in Pleosporales, and the
position of this fungus needs further verification. The familial
status of Loculohypoxylon in Teichosporaceae is questionable,
as it is simply based on the similarity of living habitat, ascomata
and asci with Immotthia and Teichospora (Barr 2002).
Lophionema Sacc., Syll. fung. (Abellini) 2: 717 (1883).
(Pleosporales, genera incertae sedis)
Generic description
Habitat terrestrial, saprobic? Ascomata solitary, scattered or
in small groups, immersed to erumpent, globose to subglobose, with a flattened base, wall black, papillate, ostiolate.
Peridium comprising two types of cells which merge in the
middle. Hamathecium of trabeculate pseudoparaphyses,
septate, rarely anastomosing and branching. Asci 8-spored,
bitunicate, fissitunicate unknown, clavate to cylindro-clavate,
with a short and furcate pedicel and a small inconspicuous
ocular chamber. Ascospores filliform, hyaline to pale yellow,
multi-septate, slightly constricted at each septum.
Anamorphs reported for genus: none.
Literature: Barr 1992b; Chesters and Bell 1970; Ellis and
Everhart 1892; Höhnel 1909; Solheim 1949.
Type species
Lophionema vermisporum (Ellis) Sacc., Syll. fung. (Abellini)
2: 717 (1883). (Fig. 50)
≡ Lophiostoma vermispora Ellis, Bull. Torrey bot. Club
9: 19 (1882).
Ascomata 320–430 μm high × 280–350 μm diam.,
solitary, scattered or in small groups of 2–3, immersed to
erumpent, globose to subglobose, black, papillate, ostiolate.
Papilla 80–120 μm high, up to 150 μm broad, cylindrical to
somewhat vertically flattened neck; mostly with a short
Fungal Diversity
Fig. 49 Loculohypoxylon grandineum (from N.A.F. 494). a Appearance of ascomata on the host surface. b Habitat section of ascomata. c
Section of an ascoma. Note the pale brown thin-walled peridium cells.
d, e Uniseriate ascospores in asci. f–f Cylindro-clavate asci with
ascospores. Note the ocular chamber in (g). Scale bars: a=100 μm,
b=200 μm, c=50 μm, d–h=10 μm
slot-like ostiole, periphysate (Fig. 50a). Peridium 30–45 μm
wide at the sides and slightly thicker at the apex,
2-layered, lateral walls and wall adjacent to neck comprising two types of cells which merge in the middle; outer
Fungal Diversity
cells small heavily pigmented thick-walled cells of textura
angularis, cells 4–7 μm diam., cell wall 3.5–5 μm thick,
inner cells less pigmented, comprising thin-walled compressed cells; apical wall cells smaller and walls thicker,
basal wall thinner (ca. 15 μm wide), composed of lightly
pigmented thin-walled compressed cells (Fig. 50b and c).
Hamathecium of trabeculate pseudoparaphyses, 1–2 μm
broad, septate, anastomosing and branching rarely between
and mostly above the asci. Asci 105–130(−150)×10–15 μm
(x ¼ 123 12mm, n=10), 8-spored, bitunicate, fissitunicate
dehiscence not observed, clavate to cylindro-clavate, with a
short, narrow, furcate pedicel which is 10–25 μm long, and
a small inconspicuous ocular chamber (to 1.5 μm wide×
1 μm high) (Fig. 50d and e). Ascospores (80-)90–115×3–
5 μm (x ¼ 95 3:5mm, n=10), filliform, gradually tapering
towards the base, hyaline to light yellow, (6-)7(−8)-septate,
slightly constricted at each septum, smooth (Fig. 50f).
Anamorph: none reported.
Material examined: USA, New Jersey, Newfield, on
dead stems of Oenothera biennis, Aug. 1881, Ellis (NY
643, holotype, NY 885, isotype).
Notes
Morphology
Lophionema is a relatively poorly studied genus, which
was formally established by Saccardo (1883) as a monotypic
genus represented by L. vermisporum based on its “globose
ascomata, compressed ostiole, cylindrical to clavate ascus,
and filamentous, septate, subhyaline to lightly pigmented
ascospores”. Lophionema vermisporum was consequently
listed as the generic type (Clements and Shear 1931). Berlese
(1890) placed the genus in Lophiostomataceae but mentioned that the genus was similar to Ophiobolus according to
the variable apex, and Shoemaker (1976) transferred Lophionema vermisporum to Ophiobolus sensu lato. Chesters and
Bell (1970) however, had regarded Lophionema as related to
Lophiostoma despite the distinct ascospore morphology. Barr
(1992b) assigned Lophionema to Entodesmium based on the
morphology of ascomata, papilla, peridium structure, pseudoparaphyses as well as the hyaline or slightly yellowish
ascospores with a terminal appendage (not observed here).
Species of Entodesmium, however, exclusively occur on
legumes, but Lophionema vermisporum does not. We also
note that the filliform ascospores, bitunicate asci, pseudoparaphyses and nature of the peridium may also be considered
as typical of genera in the Tubeufiaceae (Barr 1980;
Kodsueb et al. 2006b).
Phylogenetic study
None.
Concluding remarks
The immersed to erumpent ascomata, trabeculate pseudoparaphyses and laterally flattened papilla and periphysate
ostioles indicate that this genus should be included in
Lophiostomataceae. We do not accept the above proposals
and, consider that Lophionema should be maintained as a
separate genus with filliform ascospores in Lophiostomataceae until representative taxa can be sequenced and
analyzed. Currently Lophionema comprises 10 species
(http://www.mycobank.org, 08-01-2009). However, many
of these are poorly studied and obscure.
Lophiostoma Ces. & De Not., Comm. Soc. crittog. Ital. 1:
219 (1863). (Lophiostomataceae)
Generic description
Habitat terrestrial, saprobic. Ascomata immersed to erumpent, usually with a distinct depressed papilla and a slotlike ostiole. Hamathecium of dense, long, septate pseudoparaphyses, embedded in mucilage, anastomosing and
branching between and above the asci. Peridium unequal
in thickness, thicker near the apex and thinner at base. Asci
usually clavate. Ascospores 1-septate, multi-septate or even
muriform, hyaline to deep brown, usually with terminal
appendages.
Anamorphs reported for genus: Pleuorphomopsis-like
(Hyde et al. 2011).
Literature: Barr 1990a; Chesters and Bell 1970; Holm and
Holm 1988; Hyde and Aptroot 1998; Hyde et al. 2002;
Tanaka and Harada 2003b; Yuan and Zhao 1994.
Type species
Lophiostoma macrostomum (Tode) Ces. & De Not.,
Comm. Soc. crittog. Ital. 1: 219 (1863). (Fig. 51)
≡ Sphaeria macrostoma Tode, Fung. mecklenb. sel.
(Lüneburg) 2: 12 (1791).
Ascomata 400–600 μm high × 420–560 μm diam.,
densely scattered to gregarious, semi-immersed to erumpent, globose or subglobose, with a small to large
flattened crest-like raised area above the ascomata which
is variable in shape, up to 300 μm high and 480 μm
wide, with a slit-like ostiole along the full length of the
crest (Fig. 51a and b). Peridium 30–45 μm thick at the
sides, thicker at the apex and thinner at the base,
composed of one cell type of small lightly pigmented
thin-walled cells of textura prismatica, cells ca. 6–9×3–
4 μm diam., apex composed of pseudoparenchymatous
cells (Fig. 51b). Hamathecium of dense, filliform, up to
3 μm near the base and less than 1.5 μm broad in the
Fungal Diversity
Fig. 50 Lophionema vermisporum (from NY-643, holotype). a
Appearance of ascomata on the host surface. Note the form of the
neck. b Section of the peridium. c Peridium comprising two types of
cells which merge in the middle; outer cells small heavily pigmented
thick-walled cells of textura angularis, inner cells less pigmented, and
comprising thin-walled compressed cells. d, e Cylindro-clavate, 8spored asci. f A 7-septate filliform ascospore. Scale bars: a=0.5 mm,
b=100 μm, c=50 μm, d–f=10 μm
Fungal Diversity
upper place, septate pseudoparaphyses, embedded in
mucilage, anastomosing and branching between and
above the asci (Fig. 51f). Asci 110–145 × 10–15 μm
(x ¼ 127:5 13mm, n=10), 8-spored, bitunicate, fissitunicate (ectotunica no constriction), cylindro-clavate, with a
furcate pedicel and a small ocular chamber (to 1.5 μm wide×
2 μm high) (J-) (Fig. 51c, d and e). Ascospores 27–38(−43)×
5–7.5 μm (x ¼ 31:2 6:4mm, n=10), biseriate, fusoid,
curved, hyaline, usually 1-septate, with 3–5 septa and faintly
brown when old, with (2-)3(−4) distinct oil drops in each cell
and short terminal appendage at ends (Fig. 51h, i and j), and
ornamented with warts when spores are senescent (Fig. 51g).
Anamorph: none reported.
Material examined: SWEDEN, Smaland, Femsjö par.,
Femsjö, on Prunus, 2006, Elias Fries, det. Geir Mathiassen
(UPS, lectotype, as Sphaeria macrostoma Fr.). FRANCE,
Ariège, Rimont, Las Muros, on dead stems of Vitis vinifera,
2 Sept. 1996 (IFRD2005).
Notes
Morphology
Lophiostoma is morphologically a well studied genus
(Barr 1990a; Chesters and Bell 1970; Holm and Holm
1988; Mugambi and Huhndorf 2009b; Yuan and Zhao
1994), and currently it comprises about 30 species (Tanaka
and Harada 2003b). The genus was characterized as
having immersed to erumpent ascomata with a cylindrical
or crest-like papilla and full length, slit-like ostiole; a
peridium of unequal thickness, which was broader near the
base (Lophiostoma-type); mostly clavate, bitunicate asci
and 1- to several septate, hyaline to pigmented ascospores
with terminal appendages or surrounded by a mucilaginous sheath (Holm and Holm 1988). This definition was
followed by Barr (1990a), Yuan and Zhao (1994) and
Hyde et al. (2002).
The crest-like papilla has been regarded as a prominent
morphological character of Lophiostoma macrostomum
(Chesters and Bell 1970; Holm and Holm 1988). In the
lectotype specimen, the raised area above the ascomata is
up to 300 μm high and 480 μm long, and seen as a
flattened or even Y-shaped crest (Fig. 51a). In Lophiostoma
curtum (Fr.) De Not. and Lophiotrema boreale Math. the
raised area above the ascomata varies considerably in
height or is even lacking (Holm and Holm 1988). Thus
the variable “crest-like raised area in Lophiostomataceae”
was explained as an evolutionarily adaptation to the hard
substrate within which the ascomata develop (Holm and
Holm 1988). The ascospores of L. macrostomum usually
turn reddish brown when mature, and minutely verrucose
ornamentation was also found on the surface of the
pigmented ascospores. Hyaline ascospores that became
pigmented with age are common in Lophiostoma, such as
in L. appendiculatum Fuckel, L. massarioides (Sacc.) L.
Holm & K. Holm, L. semiliberum, L. subcorticale Fuckel
and L. winteri (Holm and Holm 1988; Tanaka and Harada
2003b). The phylogenetic significance of this character
should be observed carefully in the future but at present its
phylogenetic significance is unclear as this also occurs in
some Lophiotrema species.
Phylogenetic study
Phylogenetic affinity with some Massarina species has
been reported by Liew et al. (2002), and several Massarina
species were transferred into Lophiostoma. In a systematic
study of Lophiostoma- and Massarina-related fungi conducted by Zhang et al. (2009b), Lophiostoma taxa clustered
into two groups; one includes the type species L. macrostomum with crest-like ostioles, L. rugulosum Yin. Zhang, J.
Fourn. & K.D. Hyde with a wide, umbilicate pore surrounded
by 4–6 radial ridges, and L. glabro-tunicatus with small
ostiolar pores; the other cluster comprises Lophiostoma-like
taxa with slot-like ostioles lacking raised crests, which
includes L. arundinis (Pers.) Ces. & De Not., L. caulium, L.
compressum (Pers.) Ces. & De Not., L. crenatum (Pers.)
Fuckel, L. fuckelii (Sacc.) Sacc., L. macrostomoides, L.
semiliberum and L. viridarium Cooke, which seems to
represent a natural group at the family level. This conclusion
is tentative until verified sequences of L. macrostomum are
included in analyses (see comments of Zhang et al. 2009a).
Concluding remarks
We tend to accept a narrow concept of Lophiostomataceae, which only comprises species of Lophiostoma sensu
stricto (Zhang et al. 2009a).
Lophiotrema Sacc., Michelia 1: 338 (1878). (Pleosporales,
genera incertae sedis)
Generic description
Habitat terrestrial, saprobic. Ascomata small- to mediumsized, with or without short papilla. Hamathecium of dense,
long, septate pseudoparaphyses, anastomosing and branching between and above asci. Asci cylindrical to cylindroclavate. Ascospores hyaline, 1–3-septate, usually with
mucilaginous sheath.
Anamorphs reported for genus: none.
Literature: Barr 1990a; Chesters and Bell 1970; Holm and
Holm 1988; Saccardo 1878a; Tanaka and Harada 2003c;
Tang et al. 2003; Yuan and Zhao 1994.
Type species
Lophiotrema nucula (Fr.) Sacc., Michelia 1: 338 (1878).
(Fig. 52)
Fungal Diversity
≡ Sphaeria nucula Fr., Syst. mycol. (Lundae) 2: 466 (1823).
Ascomata 200–240 μm high×200–280 μm diam., scattered, erumpent to nearly superficial, with basal wall remaining immersed in host tissue, globose to subglobose, often
laterally flattened, with a flattened base not easily removed
from the substrate, black, roughened; with a cylindrical or
slightly compressed papilla. Papilla to 120 μm long and
150 μm high, protruding, with a pore-like ostiole (Fig. 52a).
Peridium 25–30 μm wide, very thin at the base, composed of
heavily pigmented pseudoparenchymatous cells near the apex,
cells 2–2×6 μm diam., wall 1–3(−4) μm thick, lower sides
composed of pigmented cells of textura angularis, 3–5 μm
diam., wall 0.8–1.5 μm thick, ostiole wall composed of
heavily pigmented and thick-walled small cells (Fig. 52b and
c). Hamathecium of dense, long, septate pseudoparaphyses,
1–2 μm broad, anastomosing and branching between and
above asci, embedded in mucilage (Fig. 52i). Asci 90–115×
9–11.5 μm (x ¼ 99:5 11:5mm; n=10), 8-spored, bitunicate, fissitunicate, cylindrical, with a short, narrowed, furcate
pedicel which is up to 10 μm long, with a small ocular
chamber (ca. 1.5 μm wide×1 μm high) (J-) (Fig. 52d, e, f
and h). Ascospores 17–21(−25) × (4-)5–6.5 μm
(x ¼ 19:5 5:5mm, n=10), obliquely uniseriate and partially overlapping to biseriate, broadly fusoid to fusoid, with
narrowly rounded ends, hyaline and lightly pigmented on
very rare occasions when senescent, 1-septate, 3-septate
when old, constricted at the median septum, the upper cell
often broader than the lower one (Fig. 52g).
Anamorph: none reported.
Material examined: on decaying wood (UPS, lectotype
as Sphaeria nucula Fr.).
Notes
Morphology
Holm and Holm (1988) provided a relatively strict
definition for Lophiotrema after they examined several specimens including the type material which they lectotypified.
Lophiotrema was mainly defined on the unique characters of
small to medium ascomata, a “Lophiotrema-type” peridium
and 1-septate ascospores. In Lophiotrema, Holm and Holm
(1988) considered the ascomata to be small- to medium-sized,
ca. pyriform but neck often reduced, even lacking and
sometimes cylindrical. The peridium was of approximately
equal thickness, 20–30 μm, composed of an outer textura
angularis of uniformly pigmented cells, up to 12 μm, and an
inner layer of very small hyaline cells, with somewhat
thickened walls. Asci are cylindrical, spores hyaline, at first
1-septate, becoming 3-septate, with distinct guttules, often
with a mucilaginous sheath. Much emphasis was given to the
1-septate ascospores by Holm and Holm (1988) when they
described and distinguished the three Lophiotrema species: L.
boreale, L. nucula, L. vagabundum (Sacc.) Sacc. and two
Fig. 51 Lophiostoma macrostomum (a–h, j from UPS, leptotype; i
from IFRD 2005). a Appearance of ascomata on the host surface.
Note the raised crest-like areas and full length germ slits. b Section of
the peridium. c–e Cylindro-clavate asci with ascospores arranged in a
2-3-seriate manner. f Hamathecium comprising branching and septate
pseudoparaphyses. g–j Released or unreleased ascospores. Note the
smooth young ascospores with terminal sheath, and the verrucose
senescent ascospores. Scale bars: a=0.5 mm, b=200 μm, c–j=10 μm
other unnamed species. This concept was widely accepted by
later workers (Kirk et al. 2001; Yuan and Zhao 1994). Tanaka
and Harada (2003c) considered the peridium and asci to
distinguish Lophiotrema from Lophiostoma, while Tang et al.
(2003) introduced a new Lophiotrema species with elongated
slit-like ostiole stating that the main difference between
Lophiotrema and Lophiostoma were size of ascomata,
structure of peridium, shape of asci and sheath of ascospores.
This peridium concept however, is not supported by the
lectotype specimen we examined here, which has a flattened
thin-walled base. Thus the “Lophiotrema-like peridium” sensu
Holm and Holm (1988) should not serve as a diagnostic
character of Lophiotrema, while the ostiole, asci and
ascospores might have some phylogenetic significance (Zhang
et al. 2009b). No anamorph is yet known for Lophiotrema.
Although the ascospores was reported by Holm and Holm
(1988) to be verruculose this could not be observed in the
lectotype examined under light microscope (1000 ×) in the
present study.
Phylogenetic study
In the phylogenetic study of Lophiostoma, Massarina and
related genera (Zhang et al. 2009b), Lophiotrema nucula
formed a consistent and robust clade with three other
Lophiotrema species: L. lignicola Yin. Zhang, J. Fourn. &
K.D. Hyde, L. brunneosporum Yin. Zhang, J. Fourn. & K.D.
Hyde and L. vagabundum, separate from other members of
Lophiostoma and Massarina sensu stricto. This clade might
represent Lophiotrema sensu stricto, however, the correctness of strains of L. vagabundum (CBS 628.86) and L.
nucula (CBS 627.86) used in the phylogenetic study are not
verified and warrant further study.
Concluding remarks
Holm and Holm (1988) distinguished Lophiostoma from
Lophiotrema based on the smaller ascomata, 1-septate
versus multi-septate ascospores, and peridial wall structure.
However, we doubt that these distinguishing characters
(size of ascomata, number of septa of ascospores) can be
confidently used to separate these genera and we could not
establish any characters that could reliably distinguish
between these two genera. The molecular data, however,
does separate Lophiostoma macrostomum and Lophiotrema
nucula into separate clades and provides some support that
these are separate genera. Although the strain of L. nucula
b
Fungal Diversity
Fungal Diversity
Fig. 52 Lophiotrema nucula (from UPS, lectotype). a Ascomata on
the host surface. b Section of a partial ascoma. c Peridium structure
near the apex. d, h Cylindrical asci in the pseudoparaphyses. e, f
Upper part of the asci, showing the small ocular chamber near the
apex. h Mature ascospores. i Pseudoparaphyses. Scale bars: a=
0.5 mm, b=100 μm, c, d=30 μm, e–i=10 μm
Fungal Diversity
(CBS 627.86) was isolated by K. & L. Holm, who had
examined the type specimen of L. nucula (Holm and Holm
1988), the culture of Lophiostoma macrostomum used in
the analysis are unverified (see comment by Zhang et al.
2009b). For the purpose of this monograph we tentatively
maintain Lophiotrema as distinct from Lophiostoma.
Macroventuria Aa, Persoonia 6: 359 (1971).
(Didymellaceae)
Generic description
Habitat terrestrial, saprobic. Ascomata small, solitary, scattered,
or in groups, initially immersed, becoming erumpent, to nearly
superficial, globose to subglobose, roughened with cylindrical
setae erect from apex. Peridium thin, membranous. Hamathecium of cellular pseudoparaphyses, seems to easily disappear
when mature. Asci bitunicate, somewhat obclavate to fusoid.
Ascospores fusoid with broadly to narrowly rounded ends,
hyaline, 1-septate, constricted at the septum.
Anamorphs reported for genus: none.
Literature: van der Aa 1971; von Arx and Müller 1975;
Barr 1987a.
Type species
Macroventuria wentii Aa, Persoonia 6: 361 (1971). (Fig. 53)
Ascomata 135–180 μm diam., rarely more than 200 μm
diam., solitary, scattered or in groups, initially immersed,
becoming erumpent, to nearly superficial, with basal wall
remaining immersed in host tissue, globose to subglobose,
broadly or narrowly conical, setae erect from the apical region of
the ascomata, cylindrical or tapering to the rounded or pointed
tip, brown, up to 90 μm long, 5–7.5 μm thick (Fig. 53a).
Peridium, 25–35 μm thick, 2-layered, out layer composed of
relatively thick-walled cells of textura angularis, cell wall up to
3 μm thick; inner layer cells with a thinner wall and subhyaline;
near apex cells smaller (Fig. 53a). Hamathecium of cellular
pseudoparaphyses, 1–2 μm thick, evanescing not sure. Asci 75–
93×24–30 μm, 8-spored, without pedicel, bitunicate, somewhat
obclavate to fusoid (Fig. 53b). Ascospores 22–32×8–14 μm, 1–
3 seriate, fusoid with broadly to narrowly rounded ends, hyaline,
1-septate, constricted at the septum, smooth (Fig. 53b) (description adapted from van der Aa 1971).
Anamorph: none reported.
Material referred: USA, Nevada; Death Valley, plant
litter, F.W. Went, 229, 1970 (CBS 526.71, holotype).
on its “near-hyaline, 1-septate ascospores, setose ascomata,
and saprobic life style”. Almost all of the above characters
(except the saprobic life style) point this group of fungi to
Venturiaceae. Thus Macroventuria was assigned to this
family as a relatively primitive genus (van der Aa 1971).
Subsequently, von Arx and Müller (1975) assigned Macroventuria to Pseudosphaeriaceae (Dothideales), and this
proposal was followed by Barr (1987a).
Phylogenetic study
Phylogenetic analysis based on combined SSU rDNA
and LSU rDNA sequences indicated that both of Macroventuria anomochaeta and M. wentii form a robust clade
with Leptosphaerulina argentinensis (Speg.) J.H. Graham
& Luttr., L. australis, L. trifolii (Rostr.) Petr. and
Platychora ulmi, which appear to share phylogenetic
affinities with the Leptosphaeriaceae and Phaeosphaeriaceae, but detached from other members of Venturiaceae
and Pleosporaceae (Kodsueb et al. 2006a). In addition,
culture characters also support the close relationship
between Macroventuria and Leptosphaerulina (Barr
1987a). Analysis based on five genes, i.e. SSU, LSU,
RPB1, RPB2 and TEF1, indicated Macroventuria anomochaeta resides in the well supported clade of Didymellaceae (Zhang et al. 2009a).
Concluding remarks
The morphological characters, such as small ascomata
and hyaline, 1-septate ascospores all point at Didymellaceae, thus the familial status of Macroventuria is verified.
Mamillisphaeria K.D. Hyde, S.W. Wong & E.B.G. Jones,
Nova Hedwigia 62: 514 (1996b). (?Melanommataceae)
Generic description
Habitat freshwater, saprobic. Ascomata superficial, scattered or gregarious, conical, carbonaceous, papillate.
Hamathecium of dense, filliform, trabeculate pseudoparaphyses. Asci broadly clavate to clavate, with small ocular
chambers and short pedicels. Ascospores of two types, (1):
2-4-seriate, ellipsoid, hyaline, slightly constricted at the
main septum; with apical appendages at each end and
around the ascospore; (2) 1-2-seriate, ellipsoid to fusoid,
brown, with mucilaginous sheath around the ascospore
(Hyde et al. 1996b).
Anamorphs reported for genus: none.
Literature: Hyde et al. 1996a, b.
Notes
Type species
Morphology
Macroventuria was formally established by van der Aa
(1971) represented by M. anomochaeta and M. wentii based
Mamillisphaeria dimorphospora K.D. Hyde, S.W. Wong
& E.B.G. Jones, Nova Hedwigia 62: 515 (1996b). (Fig. 54)
Fungal Diversity
Fig. 53 Macroventuria wenti.
a Ascomata. Note the setae. b
Ascus and ascospores. Scale
bars: a=50 μm, b=10 μm
(figures referred to van der Aa
1971)
Following description is adapted from Hyde et al.
1996a, b).
Ascomata 455–650 μm high×980–1430 μm diam.,
scattered or in small groups, superficial, conical, carbonaceous, papillate, under pseudostroma which forms a thin
layer on the host surface, up to 50 μm thick between the
ascomata and 125–250 μm thick on the ascomata surface
(Fig. 54a and b). Peridium 10–25 μm thick, comprising
several layers of compressed, densely packed, thin-walled,
hyaline cells. A wedge-shaped area of vertically orientated
hyaline palisade-like cells occurs at the periphery
(Fig. 54b). Hamathecium of dense, trabeculate pseudoparaphyses, ca. 1 μm broad, hyaline, branching and anastomosing, septate, embedded in mucilage (Fig. 54e). Two
types of asci and ascospores exist in the same ascoma:
TYPE 1: asci 185–320×40–100 μm (x ¼ 210 78mm, n=
50), 8-spored, cylindro-clavate, bitunicate, fissitunicate,
short-pedicellate, with an ocular chamber (to 13 μm
wide×5 μm high) (Fig. 54c and d). Ascospores 66–84×
20–38 μm (x ¼ 78 25mm, n=50), 2-4-seriate, hyaline,
ellipsoidal, constricted at the central septum, with pad-like
mucilaginous appendage at each end and with some
mucilage associated around the spore, and TYPE 2: asci
158–242×8–15 μm (x ¼ 182 11mm, n=50), 8-spored,
cylindrical, bitunicate, fissitunicate, pedicellate, with an
ocular chamber and faint apical ring, ascospores 29–42×
6–9 μm (x ¼ 35 7mm, n = 50), 1-2-seriate, brown,
ellipsoidal-fusoid, surrounded by a thin mucilaginous
sheath (Fig. 54f, g, h, i and j).
Anamorph: none reported.
Material examined: BRUNEI, on submerged wood,
Aug. 1997, leg. K.D. Hyde (HKU(M) 7425).
Notes
Morphology
Mamillisphaeria was established as a monotypic genus
according to its bitunicate, fissitunicate asci, trabeculate
pseudoparaphyses and dimorphic ascospores, which is
typified by the widely distributed freshwater fungus, M.
dimorphospora (Hyde et al. 1996a, b). The most striking
character of this fungus is its dimorphic ascospores, i.e.
one type is large and hyaline, and the other is comparatively smaller and brown. Only a few ascomycetes have
been reported having dimorphic ascospores, such as
Aquasphaeria dimorphospora and Nectria heterospora
Speg. (Hyde 1995b; Spegazzini 1889). Dimorphic ascospores appear to have evolutionary benefits, for example
the large ascospores with mucilaginous sheaths may
facilitate nutrient storage for germination and enhanced
collision and attachment to substrates. The smaller brown
ascospores may help resist desiccation and damage by UV
light and contribute to aerial dispersal, which might
explain the worldwide distribution of M. dimorphospora
(Hyde et al. 1996a, b).
Phylogenetic study
None.
Concluding remarks
Although in the key by Barr (1990a), M. dimorphospora can be referred to Massariaceae, it is temporarily assigned to Melanommataceae here based on its
trabeculate pseudoparaphyses embedded in mucilage
(Hyde et al. 1996a, b).
Fungal Diversity
Fig. 54 Mamillisphaeria dimorphospora (from HKU(M) 7425,
paratype?). a Ascomata scattered on the host surface. Note the small
papilla. b Section of an ascoma. c, d Asci (TYPE 1). e Trabeculate
pseudoparaphyses in a gelatinous matrix. f–j Ascospores (TYPE 1).
Scale bars: a=0.5 mm, b–d=100 μm, e=10 μm, f–j=20 μm
Fungal Diversity
Massarina Sacc., Syll. fung. (Abellini) 2: 153 (1883).
emend. (Massarinaceae)
Generic description
Habitat terrestrial, saprobic. Ascomata immersed or superficial, scattered or clustered, globose, conical globose to
lenticular, papillate or epapillate, ostiolate. Hamathecium of
dense, cellular pseudoparaphyses. Asci clavate to cylindrical, with short pedicels. Ascospores ellipsoid to fusoid,
hyaline, 1- to 3-septate, with or without mucilaginous
sheath.
Anamorphs reported for genus: Ceratophoma (Sivanesan
1984).
Literature: Aptroot 1998; Barr 1990a; Bose 1961; Eriksson and Yue 1986; Hyde 1995a; Hyde and Aptroot 1998;
Liew et al. 2002; Saccardo 1883; Sivanesan 1984; Tanaka
and Harada 2003d; Zhang et al. 2009a, b.
Type species
Massarina eburnea (Tul. & C. Tul.) Sacc., Syll. fung.
(Abellini) 2: 153 (1883). (Fig. 55)
≡ Massaria eburnea Tul. & C. Tul., Sel. Fung. Carp. 2:
239 (1863).
Ascomata to 250 μm high×500–700 μm diam., solitary or
in small clusters, forming under raised dome-shaped areas,
with blackened centres, with a central ostiole, immersed
within the cortex of thin dead branches, ellipsoidal,
rounded from above, clypeate, neck central, short and
barely noticeable on host surface (Fig. 55a). Clypeus ca.
250 μm diam., 60 μm thick, brown, comprising compact
brown-walled cells of textura angularis to globulosa
beneath host epidermal cells (Fig. 55b). Peridium ca.
20 μm thick comprising 3–5 layers of hyaline compressed
cells, fusing at the outside with the host (Fig. 55e).
Hamathecium filamentous, cellular pseudoparaphyses, ca.
2 μm broad, septate, embedded in mucilage, without
anastomosing (Fig. 55d). Asci 108–170 × 18–22 μm
(x ¼ 144:5 18:8mm, n=10), 8-spored, cylindro-clavate,
pedunculate, bitunicate, fissitunicate, (1-)2-seriate, apically
rounded, with an ocular chamber and faint ring (J-) (Fig. 55c
and f). Ascospores 30–38×8–12 μm (x ¼ 32:4 8:6mm, n=
10), fusoid to ellipsoid, 4-celled, constricted at the septa,
hyaline, with acute rounded ends and surrounded by (5–
8 μm diam.) mucilaginous sheath (Fig. 55g).
Anamorph: Ceratophoma sp. (Sivanesan 1984).
Material examined: FRANCE, on twig of Fagus sp.,
(Desmazières 1764. P, holotype of Sphaeria pupula var
minor), (Mycotheca universalis no. 1951 lectotype). AUSTRIA, Silesia, Karlsbrunn, on dead twigs of Fagus sylvatica
L., Aug. and Sept. 1890, Niessl., De Thümen, sub. Massarina
eburnea, ETH. Saxonia, Königsbrunn, on twigs of Fagus
sylvatica, Apr. 1882, W. Krieger, Rabenhorst & Winter, Fungi
europaei no. 2767, ETH; FRANCE, on a dead twig of Fagus
sylvatica, Deux Sèvres, Villiers en Bois, Forêt de Chizé,
Rimbaud, 14 Apr. 2008, leg. det. Paul Leroy (IFRD 2006).
Notes
Morphology
Massarina was introduced by Saccardo (1883) for
species of pyrenocarpous ascomycetes that had previously
been placed in Massaria, but typically had hyaline
ascospores (Bose 1961). The family Massarinaceae was
described by Munk (1956) to accommodate Massarina.
This family was not commonly used and Massarina was
later placed within the Lophiostomataceae in the Pleosporales (Barr 1990a; Bose 1961; Eriksson and Yue 1986). Of
the 160 epithets listed in his monograph, Aptroot
accepted only 43 species (Aptroot 1998). The concept
of Massarina was widely accepted as having single or
aggregated, immersed to erumpent, spherical to hemispherical, pseudothecioid ascomata; cellular pseudoparaphyses; bitunicate, cylindrical to clavate or obpyriform
asci; and hyaline, 1–3(−7)-septate, fusoid to long ellipsoid
ascospores that mostly have a mucilaginous sheath or
appendages (Aptroot 1998; Hyde and Aptroot 1998;
Tanaka and Harada 2003d).
In the holotype of Sphaeria pupula var. minor (P) and
lectotype of Massarina eburnea (ETH), ascospores are
reported as “not constricted at the septa” (Hyde 1995a).
However, in one of our recent collections, ascospores that
are constricted at their septa were observed (Fig. 55g),
which was consistent with the description by Fallah and
Shearer (2001). This might be because this character is not
clear in the old (over 100 years) and dry herbarium
specimens, or it may be variable between collections.
Phylogenetic study
Recent morphological, molecular and anamorphic results
indicate, however, that Massarina is polyphyletic (Hyde
1995a; Kirk et al. 2001; Liew et al. 2002). Based on the
rDNA dataset, Massarina cisti and the type of Massarina
(M. eburnea) forms a robust clade representing Massarina
sensu stricto (Zhang et al. 2009a, b).
Concluding remarks
Massarina sensu stricto should be accepted, which
seems to only include some terrestrial and saprobic species.
Massariosphaeria (E. Müll.) Crivelli, Diss. Eidgenöss. Techn. Hochschule Zürich 7318: 141 (1983).
(?Amniculicolaceae)
≡ Leptosphaeria subgen. Massariosphaeria E. Müll.,
Sydowia 4: 206 (1950).
Fungal Diversity
Fig. 55 Massarina eburnea
(from IFRD 2006). a Ascomata
on the host surface. b Section
of an ascoma. c Ascus with a
short pedicel. d Cellular pseudoparaphyses. e Section of the
peridium comprising a few
layers of compressed cells. f
Asci in pseudoparaphyses. g
Three-septate ascospores. Scale
bars: a=0.5 mm, b=100 μm,
c–g=20 μm
Generic description
Habitat terrestrial or freshwater, saprobic. Ascomata
medium-sized, scattered, or in small groups, immersed,
erumpent to superficial, subglobose, black; apex with a
wide and usually somewhat compressed papilla. Peridium thick or thin, usually thicker near the apex,
composed of 2–3 layers of thick walled scleroparenchymatous cells. Hamathecium of dense, trabeculate
pseudoparaphyses. Asci 8-spored, bitunicate, cylindrical
to cylindro-clavate, with a short, thick, furcate pedicel.
Ascospores fusoid to narrowly ellipsoid, brown or dark
brown, multi-septate.
Anamorphs reported for genus: none.
Literature: Barr 1989c; Huhndorf et al. 1990; Kohlmeyer et al.
1996; Müller 1950; Tanaka and Harada 2004; Tanaka et al. 2005.
Type species
Massariosphaeria phaeospora (E. Müll.) Crivelli, Ueber
die Heterogene Ascomycetengattung Pleospora Rabh.;
Vorschlag für eine Aufteilung (Diss. Eid genössischen Tech
Hochsch Zürich 7318): 141 (1983). (Fig. 56)
≡ Leptosphaeria phaeospora E. Müll., Sydowia 4: 208
(1950).
Ascomata 400–550 μm high × 300–500 μm diam.,
scattered, or in small groups, immersed, semi-immersed,
subglobose, black, apex wide papilla, sometimes slightly
compressed, 40–70(−100) μm broad (Fig. 56a). Peridium
Fungal Diversity
10–20 μm wide at sides, comprising one cell type of
2–3 layers of thick walled scleroparenchymatous cells,
cell wall 2–5 μm thick, peridium thicker near
the apex (Fig. 56b). Hamathecium of dense, trabeculate
pseudoparaphyses, 1–2 μm broad, septate, branching
and anastomosing. Asci 120–173 × 18–25 μm
(x ¼ 133:2 20:5mm, n=10), 8-spored, bitunicate, fissitunicate dehiscence not observed, broadly cylindrical to
cylindro-clavate, with a short, thick, furcate pedicel, up to
1 5 μ m l o n g . A s c o s p o re s 3 2 . 5 – 4 2 × 1 0 – 1 3 μ m
(x ¼ 36 11:2mm, n = 10), narrowly ellipsoid, usually
slightly curved, dark brown, 7–9 septa, slightly constricted
at the median septum (Fig. 56c and d).
Anamorph: none reported.
Material examined: SWITZERLAND, Kt. Wallis,
Findelen, Artemisiae campestris L., 10 Sept. 1895, H.
Wegelin (ZT, holotype).
Fig. 56 Massariosphaeria
phaeospora (ZT, holotype). a
Ascomata scattering on the host
surface. Note the immersed to
erumpent ascomata. b Section of
a partial peridium. Note the
peridium structure. c, d Released ascospores. Scale bars:
a=1 mm, b–d=20 μm
Notes
Morphology
Massariosphaeria was established by Müller (1950) as
a section of Leptosphaeria based on its large, thick-walled
ascospores with a mucilaginous sheath as well as its
ascomata with a thick apex. Massariosphaeria was
introduced as a separate genus by Crivelli (1983),
characterized by its wide peridial apex comprising
thick-walled cells, compressed to round papilla, and
relatively large, thick-walled, reddish brown to brown,
multi-septate to dictyosporous ascospores, usually surrounded by a sheath (Crivelli 1983; Huhndorf et al. 1990;
Leuchtmann 1984). In particular, Crivelli (1983) emphasized that species of Massariosphaeria often stain the woody
substrate (or culture) purple, and this was accepted by
Leuchtmann (1984). Barr (1989c) had treated Massarios-
Fungal Diversity
phaeria as a synonym of Chaetomastia, but this viewpoint
was rarely followed.
Phylogenetic study
The polyphyletic nature of Massariosphaeria is detected
by analyzing SSU and LSU rDNA sequences (Wang et al.
2007). The purple staining character has shown phylogenetic
significance in Amniculicolaceae, a freshwater family from
France (Zhang et al. 2009a). A single isolate of M.
phaeospora was shown to be unrelated to Amniculicolaceae
and clustered with a single isolate of Thyridaria rubronotata
(Schoch et al. 2009; Zhang et al. 2009a).
Concluding remarks
Based on phylogenetic analysis, staining the substrate purple
may have more phylogenetic significance than morphological
characters (Zhang et al. 2009a). Thus, the generic circumscription of Massariosphaeria should be re-evaluated by further
phylogenetic study with more relevant taxa included.
Mauritiana Poonyth, K.D. Hyde, Aptroot & Peerally,
Fungal Divers. 4: 102 (2000). (?Zopfiaceae)
Generic description
Habitat terrestrial, saprobic. Ascomata medium-sized, gregarious, ovoid, immersed, ostiolate, ostiole rounded. Peridium
thin, thicker near the apex. Hamathecium of dense, cellular
pseudoparaphyses, branching. Asci 8-spored, bitunicate, cylindrical to cylindro-clavate, with a short pedicel and a small
ocular chamber. Ascospores 2-3-seriate, fusoid with rounded
ends, dark brown with paler apical cells, multi-septate,
distoseptate, slightly constricted at the primary septum.
Anamorphs reported for genus: none.
Literature: Hawksworth et al. 1995; Poonyth et al. 2000;
Suetrong et al. 2009.
Type species
Mauritiana rhizophorae Poonyth, K.D. Hyde, Aptroot &
Peerally, Fungal Divers. 4: 102 (2000). (Fig. 57)
Ascomata 390–410 μm high × 310–325 μm diam.,
gregarious, ovoid, immersed, ostiolate, ostiole rounded
(Fig. 57a). Peridium 40–60 μm thick laterally, thicker near
the apex (Fig. 57a and b). Hamathecium of dense, long
cellular pseudoparaphyses, 1.5–2 μm broad, branching. Asci
130–180×20–25 μm (x ¼ 156 21:8mm, n=10), 8-spored,
bitunicate, cylindrical to cylindro-clavate, with a short pedicel,
with a small ocular chamber (Fig. 57c, d and e). Ascospores
29–40×9–13 μm (x ¼ 35:4 11mm, n=10), 2-3-seriate,
fusoid with rounded ends, dark brown with paler apical cells,
9–13-distoseptate, slightly constricted at the primary septum,
smooth (Fig. 57f, g, h and i).
Anamorph: none reported.
Material examined: MAURITIUS, Grand Gaube,
Melville mangrove, on dead decorticated Rhizophora
mucronata Lam. wood still attached to living tree, Jan.
1995, A.D. Poonyth (HKU(M)10219, holotype).
Notes
Morphology
Mauritiana was introduced to accommodate the mangrove fungus, M. rhizophorae, which is characterized by
immersed ostiolate, periphysate ascomata, thin peridium,
bitunicate, 8-spored, cylindrical to cylindro-clavate asci,
fusoid, smooth, hyaline to pale brown, multi-septate and
distoseptate ascospores (Poonyth et al. 2000). But after
carefully studying the type of M. rhizophorae, no typical
distoseptate ascospores observed. The pigmented curved
septum of the ascospore gives a “thickened” appearance.
Based on its immersed ascomata, presence of cellular
pseudoparaphyses, thick-walled, fissitunicate asci and
brown, phragmosporous ascospores constricted at the primary septum, Mauritiana was assigned to the Pyrenulales sensu
stricto (Melanommatales sensu lato, Dothideales sensu lato)
(Hawksworth et al. 1995; Poonyth et al. 2000).
Phylogenetic study
Based on a multigene phylogenetic analysis, Mauritiana
rhizophorae resided within a paraphyletic clade (Suetrong et
al. 2009) sister to marine fungi Halotthia posidonia and
Pontoporeia biturbinata. In this study, the dendrogram shows
it to be closely related to the Sporormiaceae and Lophiostomataceae, which may indicate an uncircumscribed familial clade
(Plate 1). Thus, its familial placement remains undetermined.
Concluding remarks
The “thickened” septa of ascospores of Mauritiana
rhizophorae is quite unique in Pleosporales, which makes
it easily distinguishable from other genera..
Melanomma Nitschke ex Fuckel, Jb. nassau. Ver. Naturk.
23–24: 159 (1870). (Melanommataceae)
Generic description
Habitat terrestrial, saprobic. Ascomata immersed, erumpent
to nearly superficial, medium- to large-sized, globose to
subglobose, coriaceous, gregarious, short papillate. Peridium pseudoparenchymatous cells outside with pale compressed cells inside. Asci cylindric to clavate with short
pedicels. Hamathecium of dense, filamentous, branching,
rarely anastomosing, septate pseudoparaphyses. Ascospores
pale brown, reddish brown to olive-brown, ellipsoid to
fusoid, 2 to multi-septate, constricted at the main septum.
Fungal Diversity
Fig. 57 Mauritiana rhizophorae (from HKU(M)10219,
holotype). a Vertical section of
an ascoma. Note the thin layer of
fungal tissue (pseudostroma?) on
the host surface. b Section of a
partial peridium. c Pseudoparaphyses and immature ascus. d
Fissitunicate asci. e Asci showing thickening of the apical wall.
f–i Ascospores with transverse
septa and paler polar cells. Scale
bars: a=40 μm, b, d–i=10 μm,
c=20 μm
Anamorphs reported for genus: Aposphaeria, Nigrolentilocus, Phoma-like and Pseudospiropes (Chesters 1938;
Sivanesan 1984).
Literature: Barr 1990a; Chesters 1938; Fuckel 1870;
Saccardo 1878; Zhang et al. 2008a.
Type species
Melanomma pulvis-pyrius (Pers.) Fuckel, Jb. nassau. Ver.
Naturk. 23–24: 160 (1870). (Fig. 58)
≡ Sphaeria pulvis-pyrius Pers., Syn. meth. fung.
(Göttingen) 1: 86 (1801).
Ascomata 215–471 μm high × 260–440 μm diam.,
gregarious, substrate surface covered with a thin layer of
brown psueodstroma, superficial, globose, subglobose,
broadly or narrowly conical, often laterally flattened, black,
roughened and irregular, often bearing remnants of wood
fibers; apex short papillate, often somewhat puckered or
sulcate (Fig. 58a). Peridium 70–90 μm wide, to 180 μm
wide at the base, coriaceous, comprising two types of cells,
outer cells small heavily pigmented thick-walled cells of
textura angularis, apical cells smaller and walls thicker,
individual cell walls to 6 μm thick, inner cells lightly
pigmented to hyaline thin-walled cells of textura angularis,
Fungal Diversity
5–8 μm diam., individual cell wall to 1.5–2 μm thick, in
places with columns of textura prismatica, and larger, paler
cells of textura prismatica towards the interior and at the
base (Fig. 58b). Hamathecium of dense, filamentous, 1–2
(−2.5) μm broad, branching, rarely anastomosing, septate
pseudoparaphyses. Asci 98–123 × 6.5–7.5(−9) μm
(x ¼ 109 7:5mm, n=10), 8-spored, bitunicate, fissitunicate,
cylindrical to fusoid, with a short, furcate pedicel, to 25 μm
long, with an ocular chamber (Fig. 58c, d, e, f and g).
Ascospores 14–17.5(−19)×4.5–6.5 μm (x ¼ 15:8 5:2mm,
n=10), obliquely uniseriate and partially overlapping, broadly
fusoid to fusoid with broadly rounded ends, straight or slightly
curved, smooth, olive-brown, 4-celled, slightly constricted at
the septa, the second cell from the top slightly wider than the
others, no sheath (Fig. 58h, i, j, k and l).
Anamorph: Aposphaeria agminalis Sacc. or Phoma
agminalis Sacc. (Sivanesan 1984).
Colonies (of epitype) reaching 4 cm diam. after 20 days
growth on PDA at 25°C, depressed to raised, cottony to woolly,
with rhizoidal margin, grey, reverse darkened. Phoma-like
anamorph has been reported by Chesters (1938) and
Sivanesan (1984), but no anamorphic stage was observed
in the cultures of IFRDCC 2044, CBS 109.77 and CBS
371.75 after culturing 3 months on PDA.
Material examined: on decaying wood (UPS, Scler. suec.
n. 120, holotype, as Sphaeria pulvis-pyrius Pers.); FRANCE,
Ariège, Rimont, Saurine, on bark of Salix caprea, 10 Apr.
2008, Jacques Fournier (IFRD 2001, epitype).
Notes
Morphology
Melanomma, the familial type of Melanommataceae, was
formally established by Fuckel (1870, p 159) based on its small,
carbonaceous ascomata, having: “sporen meist 2–3 mal septirt,
selten ohne Scheidewand, braun oder wasscrhell.” (Chesters
1938; Fuckel 1870). Saccardo (1878, p. 344) emended this
genus as “Spores ovate or oblong, multi-septate, coloured.”
Subsequently, Saccardo (1883, p. 98) extended the description
of Melanomma as “Perithecia gregarious, seldom scattered,
somewhat superficial, sphaerical, papillate or blunt, carbonaceous, smooth or somewhat hairy. Asci elongate, for the most
part accompanied by paraphyses, 8-spored. Spores oblong or
somewhat spindle-shaped, two to many septate, olive or dark
brown. Species of Sphaeria belong here for the most part.”
Melanomma pulvis-pyrius was erected as the lectotype
species (Barr 1990a; Chesters 1938). Barr (1990a) gave a
detailed circumscription for Melanomma, under which
Melanomma contains about 20 species (Kirk et al. 2001).
Melanomma pulvis-pyrius is characterized by its gregarious,
superficial ascomata with short papillate, cylindrical asci with a
short pedicel and fusoid, olive-brown, 3-septate ascospores
(Chesters 1938; Zhang et al. 2008a). One of the diagnostic
characters of Melanommataceae is the trabeculate pseudoparaphyses, although no typical trabeculate pseudoparaphyses could
be found in the neotype (Scler. suec. n. 120, UPS) and epitype
(IFRD 2001) of M. pulvis-pyrius (Zhang et al. 2008a).
Phylogenetic study
Phylogenetic analysis based on five genes (LSU, SSU,
RPB1, RPB2 and EF1) indicates that Melanomma pulvispyrius forms a robust clade with Byssosphaeria, Herpotrichia and Pleomassaria siparia (Pleomassariaceae) and
likely represents a separate family (or families comprising
Melanommataceae) (Zhang et al. 2008a; Mugambi and
Huhndorf 2009b). A more recent phylogenetic analysis
included a group of coelomycete species with stellate
conidia, isolated from Fagales trees clustered in Melanommataceae (Tanaka et al. 2010; Plate 1).
Concluding remarks
The Melanomma concept based on ascospore morphology appears polyphyletic.
Metameris Theiss. & Syd., Annls mycol. 13: 342 (1915).
(Phaeosphaeriaceae)
Generic description
Habitat terrestrial, saprobic or parasitic. Ascostromata erumpent
through the host surface in linear rows parallel to the host fibers.
Ascomata small, globose to subglobose, black, coriaceous.
Peridium composed of large lightly pigmented cells of textura
angularis. Hamathecium of rare, broad pseudoparaphyses,
septate, constricted at the septa. Asci bitunicate, fissitunicate,
broadly cylindrical to slightly obclavate, with a short, thick,
knob-like pedicel. Ascospores hyaline, 1- (rarely 2-) septate.
Anamorphs reported for genus: none.
Literature: von Arx and Müller 1975; Barr 1972; Clements
and Shear 1931; Eriksson 2006; Lumbsch and Huhndorf
2007; Theissen and Sydow 1915.
Type species
Metameris japonica (Syd.) Syd., Annls mycol., 13(3–4):
342 (1915). (Fig. 59)
≡ Monographus japonicus Syd. Annls mycol. 10: 408
(1912).
Ascostromata erumpent through the host surface in linear
rows parallel to the host fibers, 500–750 μm long and 140–
200 μm wide, with three to ten ascomata arranged in a line
(Fig. 59a). Ascomata 115–160 μm diam., semi-immersed in
substrate to erumpent, globose, subglobose, black, coriaceous
(Fig. 59b). Cells of ascostromata heavily pigmented and
thick-walled, cells of peridium composed of large lightly
pigmented cells of textura angularis, cells 5–15 μm diam.,
Fungal Diversity
Fig. 58 Melanomma pulvis-pyrius (a–b, d–e, h–j from UPS, holotype; c, g, k, l from epitype). a Ascomata gregarious on the host surface. b
Vertical section of an ascoma. c–f Asci with pedicels. g Dehiscent ascus. h–l Ascospores. Scale bars: a=0.5 mm, b=200 μm, c–l=10 μm
Fungal Diversity
cell wall <1 μm thick, peridium thicker at the base, up to
50 μm (Fig. 59b). Hamathecium of rare, pseudoparaphyses
3–4 μm broad, septate, constricted at the septa, anastomosing
or branching not observed. Asci (65-)80–90×12–15 μm
(x ¼ 82:8 13:3mm, n=10), 8-spored, bitunicate, fissitunicate,
broadly cylindrical to slightly obclavate, with a short, thick,
knob-like pedicel, lacking an ocular chamber (Fig. 59c).
Ascospores 25–30×5–6 μm (x ¼ 27:4 5:6mm, n=10),
biseriate, oblong, hyaline, 1-2-septate, the secondary septum
exclusively occurring in the upper cells, slightly constricted at
the primary septum which is slightly below the centre of the
ascospore, the upper cells usually swollen near the main
septum (Fig. 59d and e).
Anamorph: none reported.
Material examined: JAPAN, Province Mino. on Osmunda regalis L. var. japonica Milde., 10 May 1912, R. Hale (S,
F7166, type, as Monographos japonicus Syd.).
Fig. 59 Metameris japonica
(from S, F7166, type). a
Ascostroma arrangement on the
host surface. b Section of two
ascomata from one ascostroma.
c Immature asci within pseudoparaphyses. d, e Hyaline ascospores. Scale bars: a=0.5 mm.
b=100 μm, c–e=20 μm
Notes
Morphology
Metameris was formally established by Theissen and
Sydow (1915) to accommodate Monographus japonicus
Syd., which is characterized by the erumpent ascomata
arranged in linear ascostromata, the presence of pseudoparaphyses and hyaline 2-septate ascospores.
Clements and Shear (1931) assigned it to Dothideaceae
(subfamily Dothideae), and von Arx and Müller (1975)
assigned it to Pleosporaceae. Currently, it is considered
as a member of Phaeosphaeriaceae (Pleosporales)
(Eriksson 2006; Lumbsch and Huhndorf 2007). Scirrhodothis and Scirrhophragma are considered synonyms of
Metameris (von Arx and Müller 1975), and all three
closely related to Scirrhia (Barr 1972; Müller and von
Arx 1962).
Fungal Diversity
Phylogenetic study
None.
Concluding remarks
Its small-sized ascomata, broadly cylindrical to slightly
obclavate asci with a short, thick, knob-like pedicel, as well
as its monocotyledonous host preference point Metameris
to the Phaeosphaeriaceae. But DNA comparisons are
needed for confirmation.
Mixtura O.E. Erikss. & J.Z. Yue, Mycotaxon 38: 203
(1990). (Phaeosphaeriaceae)
Generic description
Habitat terrestrial, parasitic. Ascomata small-sized, scattered or clustered on the leaf spots, immersed, erumpent,
minutely papillate, ostiolate. Papilla slightly raised. Peridium thin, comprising one cell type of lightly pigmented
thin-walled cells of textura angularis. Hamathecium of
dense, filliform, septate, cellular pseudoparaphyses, 4–
6.3 μm broad, embedded in mucilage. Asci bitunicate,
ovoid, with a very short stumpy pedicel. Ascospores fusoid
to narrowly fusoid with broadly to narrowly rounded ends,
curved, dark brown, multi-septate, distoseptate, with a germ
pore at the lower end.
Anamorphs reported for genus: none.
Literature: Eriksson and Yue 1990.
Type species
Mixtura saginata (Syd.) O.E. Erikss. & J.Z. Yue, Mycotaxon
38: 203 (1990). (Fig. 60)
≡ Leptosphaeria saginata Syd., Annls mycol. 37: 376 (1939).
Producing elongated yellow spots with brownish margins,
leaf spots up to 45×3–5 mm, opposite side visible as a
brownish spots (Fig. 60a). Ascomata 170–200 μm high×210–
280 μm diam., scattered on the lower side of the leaf,
immersed, erumpent, breaking through the epidermis, minutely papillate. Papilla central, slightly raised, ostiolate,
ostiole surrounded by a white margin (Fig. 60b). Peridium
22–34 μm wide, thicker at the apex, thinner at the base,
comprising one cell type of lightly pigmented thin-walled
cells of textura angularis, cells up to 6×8 μm diam., cell wall
0.5–1.2 μm thick, apex cells smaller and walls thicker
(Fig. 60c). Hamathecium of dense, filliform, septate, cellular
pseudoparaphyses, 4–6.3 μm broad, embedded in mucilage.
Asci 80–128×41–53(−69) μm (x ¼ 100:9 52:8mm, n =10),
8-spored, bitunicate, fissitunicate dehiscence not observed,
sac-like, with a very short stumpy pedicel and a small ocular
chamber (Fig. 60d). Ascospores 86–94(−106)×20.5–23.5 μm
(x ¼ 92:7 21:7mm, n=10), fasciculate, fusoid to narrowly
fusoid, slightly curved, dark brown, 7-septate, distoseptate,
with or without constriction at the primary septum, smoothwalled, with a germ pore at the lower end (Fig. 60e and f).
Anamorph: none reported.
Material examined: ECUADOR, Tungurahua, Hacienda
San Antonio pr. Baños, Province, on the leaves of Chusqueae
serrulatae Pilger., 9 Jan. 1938, H. Sydow. (S reg. nr F8934
type, F8935 isolectotype, as Leptosphaeria saginata).
Notes
Morphology
Mixtura was formally established by Eriksson and
Yue (1990) as a monotypic genus represented by M.
saginata based on its immersed and thin-walled ascomata,
sparse, broad pseudoparaphyses, sac-like asci with a short
pedicel and thick apex. Mixtura has a “mixture” of
characters found in other pleosporalean genera. The
peridium structure is comparable with Phaeosphaeria,
the ascospores with Trematosphaeria and asci with
Wettsteinina (Eriksson and Yue 1990). According to the
structure of ascomata and hamathecium, Mixtura was
provisionally assigned to Phaeosphaeriaceae (Eriksson
and Yue 1990).
Phylogenetic study
None.
Concluding remarks
Morphologically, the sparse broad pseudoparaphyses and
sac-like asci with a thick apical structure in Mixtura seem
more comparable with the generic type of Teratosphaeria
(T. fibrillose Syd. & P. Syd., Teratosphaeriaceae, Capnodiales, Dothideomycetidae) than that of Phaeosphaeria (P.
oryzae). The heavily pigmented, multi-septate ascospores
and the persistent pseudoparaphyses of Mixtura however,
differ from those of Teratosphaeria. Thus, here we assign
Mixtura under Teratosphaeriaceae as a distinct genus until
phylogenetic work is carried out.
Montagnula Berl., Icon. fung. (Abellini) 2: 68 (1896).
(Montagnulaceae)
Generic description
Habitat terrestrial, saprobic. Ascomata small- to mediumsized, immersed to erumpent, gregarious or grouped,
globose to subglobose, black. Hamathecium of dense,
narrowly cellular, septate pseudoparaphyses. Asci bitunicate, fissitunicate, usually cylindro-clavate to clavate with a
long pedicel. Ascospores oblong to narrowly oblong,
straight or somewhat curved, reddish brown to dark
yellowish brown, muriform or phragmosporous.
Anamorphs reported for genus: Aschersonia (Hyde et al. 2011).
Fungal Diversity
Fig. 60 Mixtura saginata
(from S reg. nr F8934, type). a,
b Leaf spots in leaves of Chusquea serrulatae. Note the erumpent ascomata surrounded by
white material in (b). c Section
of an ascoma. Note the peridium
structure which comprises cells
of textura angularis. The arrangement of the asci and pseudoparaphyses can also be seen.
d Immature asci in pseudoparaphyses. Note the stumpy pedicel and thickened apex with
flattened ocular chamber. e, f
Mature ascospores. Note the
hyaline ends and distosepta.
Scale bars: a=10 mm, b, c=
100 μm, d=50 μm, e–f=20 μm
Literature: Aptroot 1995; Barr 2001; Berlese 1896; Clements
and Shear 1931; Crivelli 1983; Leuchtmann 1984; Ramaley
and Barr 1995; Schoch et al. 2006; Wehmeyer 1957, 1961;
Zhang et al. 2009a.
Type species
Montagnula infernalis (Niessl) Berl., Icon. fung. (Abellini).
2: 68 (1896). (Fig. 61)
≡ Leptosphaeria infernalis Niessl, Inst. Coimbra 31: 13
(1883).
Ascomata 220–280 μm high×250–310 μm diam., immersed to erumpent, gregarious or clustered, globose to
subglobose, sometimes triangular in dried material, short
ostiole always filled with hyaline closely adhering cells, black
(Fig. 61a and b). Peridium 40–55 μm thick at sides, up to
80 μm thick near the apex, 3-layered, outer layer composed of
heavily pigmented thick-walled small cells of textura angularis, cells 3–8 μm diam., wall 1.5–3 μm thick, apex thicker
with smaller cells and thicker cell wall, thinner near the base;
mid layer less pigmented, cells 4–13 μm diam.; innermost
layer of narrow compressed rows of cells, merging with
Fungal Diversity
pseudoparaphyses (Fig. 61c). Hamathecium of dense, narrow
cellular pseudoparaphyses, 2–4.5 μm broad, septate (Fig. 61f).
Asci 153–170(−200)×17.5–21.5 μm (including pedicel),
bitunicate, fissitunicate, cylindro-clavate to clavate, pedicel
28–60(−85) μm long, 8-spored, biseriate, with an ocular
chamber best seen in immature ascus (to 3 μm wide×3 μm
high) (Fig. 61d and e). Ascospores 24–29×9–11 μm, oblong to
narrowly oblong, straight or somewhat curved, reddish brown
to dark yellowish brown, verruculose, with five transverse
septa and one vertical septum in each middle cells, constricted
at the primary and secondary primary septa (Fig. 61g).
Anamorph: none reported.
Material examined: PORTUGAL, Coimbra Lusitania,
on leaves of Fourcroya longava pr., Feb., 1881, leg. Moller.
(M 1183, holotype).
However, this character appears to have evolved more than
once and can be found in Kirschsteiniothelia elaterascus
Shearer which clusters with Helicascus (Shearer et al. 2009).
The same ascus character is also found in Xenolophium and
Ostropella in the Platystomaceae (Mugambi and Huhndorf
2009b). Montagnula opulenta is a didymosporous species,
but phylogenetically closely related to those dictyosporous
(Karstenula rhodostoma) and phragmosporous (Paraphaeosphaeria michotii) members of Montagnulaceae (Zhang et al.
2009a). This might indicate that compared to other morphological characters, ascospore type is not a valid character at
family level classification.
Notes
Generic description
Morphology
Montagnula was introduced to accommodate two Pleospora species, i.e. P. infernalis (Niessl) Wehm. and P.
gigantea Mont. by Berlese (1896), based on the presence of
hyphal stromatic tissues over the ascomata and asci with
relatively long pedicels (Barr 2001). Montagnula infernalis
was selected as the lectotype species (Clements and Shear
1931). Subsequently, Wehmeyer (1957, 1961) treated Montagnula as a subgenus of Pleospora. Crivelli (1983) accepted
Montagnula as a separate genus, and divided it into two
subgenera, i.e. Montagnula and Rubiginospora. Montagnula
was characterized by having dark brown ascospores and
exclusively occurring on Agavaceae, while Rubiginospora
has reddish brown ascospores and occurs on Poaceae. This
proposal was not accepted by many workers (Barr 2001).
Subsequently, more species with various ascospores (such as
phragmosporous species by Leuchtmann (1984) and didymosporous species by Aptroot (1995) were added in this
genus), which has obviously become heterogenic. Barr
(2001) assigned species of Montagnula into different genera,
i.e. Kalmusia and Didymosphaerella, respectively and introduced Montagnulaceae to accommodate all of these genera.
Habitat terrestrial, saprobic. Ascomata medium-sized, solitary,
scattered, or in small groups, superficial, cushion-like, circular
in outline, wall black, roughened, containing numerous
locules. Peridium thin, 1-layered. Hamathecium of dense, long
filliform pseudoparaphyses, 2–3 μm broad, septate, branching.
Asci polysporous, with a short, laterally displaced, sometimes
papillate knob-shaped pedicel, apex very thick walled,
bitunicate, fissitunicate, obclavate, ocular chamber not observed. Polyspores oblong to cylindrical, hyaline, non-septate.
Anamorphs reported for genus: none.
Literature: Eriksson 2006; Romero and Samuels 1991.
Phylogenetic study
Montagnula opulenta forms a robust phylogenetic clade
with species of Bimuria, Curreya, Didymocrea, Letendraea, Paraphaeosphaeria, Phaeodothis and Karstenula,
which might represent a familial group (Schoch et al. 2006;
Zhang et al. 2009a). A more convincing conclusion can
only be obtained following sequence data from more
verified fungi being added to the phylogenetic tree.
Concluding remarks
One striking character of Montagnula infernalis is the very
long ascal pedicel once it is released from the ascomata.
Moristroma A.I. Romero & Samuels, Sydowia 43: 246
(1991). (Pleosporales, genera incertae sedis)
Type species
Moristroma polysporum A.I. Romero & Samuels, Sydowia
43: 246 (1991). (Fig. 62)
Ascomata 100–210 μm high×340–600 μm diam., solitary,
scattered, or in small groups of 2–3, superficial, with basal
wall remaining immersed in host tissue, cushion-like,
circular in outline, wall black, roughened, containing
numerous locules, each locule 120–240 μm diam., ostiolate
(Fig. 62a and b). Peridium 14–30 μm thick, 1-layered,
composed of small heavily pigmented thick-walled cells of
textura angularis, cells 2–4 μm diam., cell wall 1.5–3 μm
thick, peridium between the locules hyaline (Fig. 62b and c).
Hamathecium of dense, long filliform pseudoparaphyses, 2–
3 μm broad, septate, branching. Asci 44–60×12–14 μm
(x ¼ 54:3 13mm, n=10), polysporous, with a short, papillate knob-shaped pedicel, apex very thick-walled, bitunicate,
fissitunicate, obclavate, ocular chamber not observed
(Fig. 62d and e). Polyspores 3–4(−5)×0.6–1.2 μm, oblong
to cylindrical, hyaline, non-septate, smooth (Fig. 62f).
Anamorph: none reported.
Material examined: ARGENTINA, Buenos Aires,
Ramallo, on Eucalyptus viminalis Labill., May 1982, Romero
Fungal Diversity
Fig. 61 Montagnula infernalis
(from M 1183, holotype). a
Appearance of ascomata immersed in host tissue. b Section
of an immersed ascoma. Note
the hyaline closely adhering
cells in the ostiole region. c
Section of the peridium comprising a few layers of cells. d
An immature ascus with a long
pedicel. e, g Mature muriform
ascospores in asci. f Cellular
pseudoparaphyses. Scale bars:
a=0.5 mm, b, c=100 μm,
d–g=20 μm
27/4-13 (BAFC 32036, holotype); Nov. 1982, on decorticated
wood, Romero 35/4-13 (BAFC 32037, paratype).
Notes
Morphology
Moristroma was formally established by Romero and
Samuels (1991) based on its “cushion-shaped ascomata
containing lots of locules with numerous asci inside, asci
obclavate, polysporous, with a knob-shaped pedicel”. The
bitunicate asci and numerous cellular pseudoparaphyses
undoubtedly point it to Pleosporales, while the familial
placement of Moristroma is uncertain, and it was
temporarily assigned to Dacampiaceae by Romero and
Samuels (1991), but no 3-layered peridium is found.
Eriksson (2006) assigned it to Teichosporaceae.
Phylogenetic study
None.
Concluding remarks
The familial status of Moristroma cannot be determined yet.
Fungal Diversity
Fig. 62 Moristroma polysporum (from BAFC 32036, holotype). a Two multiculate
ascostroma on the host surface.
b Section of an ascostroma.
Note the multilocula. c Section
of the peridium. Note the thick
walled cells. d, e Broadly cylindrical to fusoid asci containing
numerous part spores. f Released part spores. Scale bars:
a=0.5 mm, b=200 μm, c=
50 μm, d–f=10 μm
Morosphaeria Suetrong, Sakay., E.B.G. Jones & C.L.
Schoch, Stud. Mycol. 64: 161 (2009). (Morosphaeriaceae)
Generic description
Habitat marine, saprobic. Ascomata large, solitary or
gregarious, immersed to erumpent, subglobose or depressed
with a flatted base, ostiolate, papillate, brown to black,
coriaceous. Peridium thick. Hamathecium of dense, long
cellular pseudoparaphyses, septate. Asci 8-spored, bitunicate, cylindrical, with short pedicels. Ascospores uniseriate
to partially overlapping, ellipsoidal, hyaline, 1-3-septate,
constricted at the septa, central cells larger, apical cells if
present small and elongated, surrounded with mucilaginous
sheath.
Anamorphs reported for genus: none.
Literature: Hyde and Borse 1986; Hyde 1991a, b;
Suetrong et al. 2009; Zhang et al. 2009a.
Type species
Morosphaeria velataspora (K.D. Hyde & Borse) Suetrong,
Sakay., E.B.G. Jones & C.L. Schoch, Stud. Mycol. 64: 161
(2009). (Fig. 63)
≡ Massarina velataspora K.D. Hyde & Borse, Mycotaxon
27: 163 (1986).
Ascomata 0.7–1.2 mm diam., solitary or gregarious,
immersed to erumpent, subglobose or depressed, with a
flattened base not easily removed from the substrate, ostiolate,
epapillate or papillate, brown to black, coriaceous (Fig. 63a).
Peridium thick, the upper part of the peridium composed of
brown thick-walled cells of textura angularis, cells are
smaller and wall thicker near the apex, at the rim is
composed of vertical, parallel, brown, elongate cells,
wedge-shape in section (Fig. 63a). Hamathecium of dense,
long cellular pseudoparaphyses, 1.1–1.7 μm broad, septate.
Asci 220–320×23–34 μm (x ¼ 251 28:2mm, n=10), 8-
Fungal Diversity
spored, bitunicate, cylindrical, with short pedicels (Fig. 63b).
Ascospores 45–56×14–19 μm (x ¼ 49:5 15:9mm, n=10),
uniseriate to partially overlapping, ellipsoidal, hyaline, 1-3septate, constricted at the septa, central cells larger, apical
cells if present small and elongated, surrounded with
mucilaginous sheath, 5–22 μm wide (Fig. 63c, d and e).
Anamorph: none reported.
Material examined: Jan. 1984, Herb. IMI 297770,
slides 1–10 (holotype) and dried wood (isotype).
two Helicascus species, they belong to Morosphaeriaceae
(another marine family) (Suetrong et al. 2009). Morphologically, Morosphaeria is characterized by solitary to
gregarious, subglobose to lenticular, immersed to superficial ascomata which are ostiolate and papillate, numerous,
filliform pseudoparaphyses, 8-spored, clavate to cylindrical,
bitunicate, fissitunicate asci, and hyaline, 1-3-septate,
fusoid to ellipsoidal ascospores which are surrounded with
mucilaginous sheath.
Notes
Phylogenetic study
Species of Morosphaeria form a sister group with
Helicascus and both of these genera were assigned to a
new family, i.e. Morosphaeriaceae (Suetrong et al. 2009).
In this study, a strain of Asteromassaria pulchra, occuring
on dead twigs of Prunus spinosa, is basal to other species
of Morosphaeriaceae, and gets well support. Thus here we
tentatively assign Asteromassaria in Morosphaeriaceae.
Morphology
Two Massarina sensu lato species described from the
marine environment, viz. M. ramunculicola (Sacc.) O.E.
Erikss. & J.Z. Yue and M. velataspora K.D. Hyde & Borse,
form a robust clade, and a new genus Morosphaeria was
established for them (Suetrong et al. 2009). Together with
Fig. 63 Morosphaeria velataspora (from IMI 297770, type). a Section of an ascoma. b Cylindrical asci embedded in pseudoparaphyses. c–e
Hyaline, 1-3-septate, ascospores with mucilaginous sheath. Scale bars: a=100 μm, b=50 μm, c–e=20 μm
Fungal Diversity
Concluding remarks
The only morphological difference between M. velataspora and M. ramunculicola are their morphology of
ascomata and size of ascospores (Hyde 1991b). But M.
velataspora was reported staining the woody substrate (or
agar in culture) purple (Hyde and Borse 1986; Hyde
1991b). Although this character could not be verified in
the strain used by Suetrong et al. (2009), purple staining has
been reported to have phylogenetic significance at familial
rank in freshwater fungi (Zhang et al. 2009a).
versally septate, constricted at the septa, with one, rarely
two longitudinal septa in all cells except end cells which
are often slightly paler, all cells filled with a large
refractive guttule, smooth to finely verruculose, surrounded by a wide mucilaginous sheath (Fig. 64d).
Anamorph: Phoma sp. (Webster 1957).
Material examined: FRANCE, Haute Garonne,
Avignonet, Lac de Rosel, 16 Jan. 2007, on submerged
dead herbaceous stem (Dipsacus?), leg. Michel Delpont,
det. Jacques Fournier (IFRD 2017).
Murispora Yin. Zhang, C.L. Schoch, J. Fourn., Crous & K.
D. Hyde, Stud. Mycol. 64: 95 (2009b). (Amniculicolaceae)
Notes
Generic description
Habitat freshwater, saprobic. Ascomata medium-sized,
scattered to gregarious, immersed, lenticular, apex slightly
protruding, opening through a small rounded pore,
substrate stained purple. Peridium thin, composed of a
few layers cells of textura angularis, thicker at the apex
with pseudoparenchymatous cells. Hamathecium of narrowly cellular pseudoparaphyses, embedded in mucilage.
Asci 8-spored, bitunicate, fissitunicate, biseriate, cylindroclavate with short pedicels. Ascospores curved- fusoid
with narrowly rounded ends, golden yellow turning brown
when senescent, multi-septate, constricted at the septa,
with one, rarely two longitudinal septa in all cells except
end cells, smooth or finely verruculose, surrounded by a
wide mucilaginous sheath.
Anamorphs reported for genus: Phoma (Webster 1957).
Literature: Zhang et al. 2009a, b.
Type species
Murispora rubicunda (Niessl) Yin. Zhang, J. Fourn. & K.
D. Hyde, Stud. Mycol. 64: 96 (2009a). (Fig. 64)
≡ Pleospora rubicunda Niessl, Notiz. Pyr.: 31 (1876).
Ascomata 170–200 μm high × 380–410 μm diam.,
scattered to gregarious, immersed, lenticular, apex
laterally flattened, black, slightly protruding, opening
through a small rounded pore, substrate stained purple
(Fig. 64a). Peridium 15–18 μm thick at sides, composed
of 3–4 layers cells of textura angularis, up to 28–30 μm
thick at the apex with very thick-walled cells, pseudoparenchymatous, nearly absent at the base (Fig. 64b).
Hamathecium of narrowly cellular pseudoparaphyses,
1–1.7 μm broad, embedded in mucilage. Asci 124–
142 × 19–21 μm, 8-spored, bitunicate, fissitunicate,
biseriate, cylindro-clavate with a small ocular chamber,
with short pedicels (Fig. 64c). Ascospores 30–38× 10–
12 μm, curved-fusoid with narrowly rounded ends,
golden yellow turning brown when senescent, 7–9 trans-
Morphology
Murispora was introduced based on Pleospora rubicunda which is characterized by immersed, erumpent or
nearly superficial, globose to subglobose, elongated weakly
papillate ascomata which stain the woody substrate purple,
trabeculate pseudoparaphyses, 8-spored, bitunicate, fissitunicate, oblong to clavate asci, fusoid, pale or reddish
brown, muriform ascospores (Zhang et al. 2009a). A
phylogenetic study indicated that Murispora forms a robust
clade with species of Amniculicola, and Amniculicolaceae
was introduced to accommodate them (Zhang et al. 2009a).
Phylogenetic study
Murispora rubicunda forms a robust clade with species of
Amniculicola and Neophaeosphaeria (Zhang et al. 2009a).
Concluding remarks
As has mentioned by Eriksson (1981, P. 135), the purplestaining species of Pleospora, treated by Webster (1957),
should not belong to the Pleosporaceae. Both Pleospora
straminis and P. rubelloides should be closely related to
Murispora.
Neomassariosphaeria Yin. Zhang, J. Fourn. & K.D. Hyde,
Stud. Mycol. 64: 96 (2009a). (Amniculicolaceae)
Generic description
Habitat freshwater, saprobic. Ascomata medium-sized,
scattered or in small groups, immersed, with a slightly
protruding elongated papilla, ostiolate, lenticular, stain the
substrate purple. Peridium thin. Hamathecium of dense,
long cellular pseudoparaphyses, septate. Asci 8-spored,
bitunicate, fissitunicate, cylindro-clavate, with short furcate
pedicels. Ascospores 2-3-seriate, narrowly fusoid, somewhat curved, reddish brown, multi-septate, slightly constricted at the primary septum.
Anamorphs reported for genus: none.
Literature: Leuchtmann 1984; Zhang et al. 2009a, b.
Fungal Diversity
Fig. 64 Murispora rubicunda
(from IFRD 2017). a Habitat
section of the immersed ascomata. b Section of an ascoma.
Note the thin peridium and cells
of textura angularis. c Mature
and immature asci. d Muriform
ascospores. Scale bars: a, b=
100 μm, c, d=20 μm
Type species
Notes
Neomassariosphaeria typhicola (P. Karst.) Yin. Zhang, J.
Fourn. & K.D. Hyde, Stud. Mycol. 64: 96 (2009a). (Fig. 65)
≡ Leptosphaeria typhicola P. Karst., Bidr. Känn. Finl.
Nat. Folk 23: 100 (1873).
Ascomata 150–280 μm high × 200–400 μm diam.,
scattered or in small groups, immersed, lenticular, with a
slightly protruding elongated papilla, ostiolate, stain the
substrate purple (Fig. 65a). Peridium 15–30 μm thick.
Hamathecium of dense, long cellular pseudoparaphyses,
1.5–2.5 μm thick, septate. Asci 110–160×13–15 μm, 8spored, bitunicate, fissitunicate, cylindro-clavate, with short
furcate pedicels (Fig. 65b, c and d). Ascospores 30–48×7–
11 μm, 2-3-seriate, narrowly fusoid, somewhat curved,
reddish brown, 7-septate, slightly constricted at the primary
septum, verruculose (Fig. 65c and d).
Anamorph: none reported.
Material examined: DENMARK, Sjaeland, Frederikskilde, Suserup Skove, Tystrup Lake, 25 May 2007, on
submerged culm of Phragmites, leg. & det. Jacques Fournier
(IFRD 2018).
Morphology
Neomassariosphaeria is most comparable with Murispora, and is distinguished from Murispora by its phragmosporous ascospores. Both genera were assigned to
Amniculicolaceae (Zhang et al. 2009a).
Phylogenetic study
Both Neomassariosphaeria grandispora and N. typhicola
clustered with species of Murispora and Amniculicola in
Amniculicolaceae (Zhang et al. 2009a,c).
Concluding remarks
Similar with those purple-staining species of
Pleospora assigned to Murispora, the purple-staining
species of Phaeosphaeria mentioned by Crivelli
(1983) and Leuchtmann (1984) might be assigned to
Neomassariosphaeria.
Neophaeosphaeria M.P.S. Câmara, M.E. Palm & A.W.
Ramaley, Mycol. Res. 107: 519 (2003). (Leptosphaeriaceae)
Fungal Diversity
Fig. 65 Neomassariosphaeria
typhicola (from IFRD 2018). a
Immersed ascomata gregarious
in the host substrate. b–d
Cylindro-clavate asci embedded
in pseudoparaphyses. Note the
phragmosporous ascospores.
Scale bars: a=200 μm,
b–d=20 μm
Generic description
Habitat terrestrial, parasitic or saprobic. Ascomata small,
forming in leaf spots, scattered or clustered, immersed,
depressed globose, under clypeus, coriaceous. Peridium thin.
Hamathecium of dense, cellular pseudoparaphyses, septate,
embedded in mucilage. Asci 8-spored, bitunicate, fissituni-
cate dehiscence not observed, broadly cylindrical to oblong,
with a short furcate pedicel. Ascospores obliquely uniseriate
and partially overlapping, oblong, pale brown, 1-3-septate.
Anamorphs reported for genus: Coniothyrium-like
(Câmara et al. 2003).
Literature: Câmara et al. 2001, 2003; Checa et al. 2002;
Ellis and Everhart 1892.
Fungal Diversity
Type species
Neophaeosphaeria filamentosa (Ellis & Everh.) M.P.S.
Câmara, M.E. Palm & A.W. Ramaley, Mycol. Res. 107:
519 (2003). (Fig. 66)
≡ Leptosphaeria filamentosa Ellis & Everh., J. Mycol. 4:
64 (1888).
Ascomata 115–157 μm high×115–186 μm diam., forming
in leaf spots, scattered or clustered in circular areas, immersed,
depressed globose, with a small ostiolar pore slightly
penetrating above the surface, under clypeus, coriaceous,
papilla not conspicuous (Fig. 66a). Peridium 18–30 μm thick,
composed of large pigmented thin-walled cells of textura
angularis, cells up to 10 μm diam. (Fig. 66c). Hamathecium
of dense, cellular pseudoparaphyses 1.5–2.5 μm broad,
septate, embedded in mucilage (Fig. 66b). Asci 70–105×8–
10 μm (x ¼ 85:3 9:7mm, n=10), 8-spored, bitunicate,
fissitunicate dehiscence not observed, broadly cylindrical to
oblong, with a short, broad, furcate pedicel, 6–13 μm long,
with a small ocular chamber, best seen in immature asci, up to
1.5 μm wide×1 μm high (Fig. 66d, e and f). Ascospores 12–
15×4–5 μm (x ¼ 13:8 5mm, n=10), obliquely uniseriate
and partially overlapping, oblong, yellowish brown, (1-2-)3septate, constricted at the primary septum, the upper second
cell often broader than others, verruculose, containing four
refractive globules (Fig. 66g).
Anamorph: Ellis and Everhart (1892) noted that the
“spermogonial stage is a Coniothyrium (C. concentricum)
with small (4 μm), globose, brown sporidia.”
Material examined: USA, New Jersey, Newfield, on
dead parts in living leaves of Yucca filamentosa L., Jul.
1888, Ellis & Everhart (NY, holotype).
sequences (Câmara et al. 2001; Checa et al. 2002), and
they fall in the group comprising members of Phaeosphaeriaceae and Leptosphaeriaceae (Câmara et al.
2003). Neophaeosphaeria filamentosa, the generic type
of Neophaeosphaeria, nested in Leptosphaeriaceae with
low to moderate bootstrap values (Schoch et al. 2009;
Zhang et al. 2009a).
Concluding remarks
The familial status of Neophaeosphaeria under Leptosphaeriaceae is confirmed, although this family remains
poorly supported in phylogenetic studies.
Nodulosphaeria Rabenh., Klotzschii Herb. Viv. Mycol.,
Edn 2: no. 725 (in sched.) (1858). (Phaeosphaeriaceae)
Generic description
Habitat terrestrial, saprobic or hemibiotrophic. Ascomata
small, immersed to erumpent, globose or subglobose,
black, papillate, ostiolate. Papilla with numerous setae in
the pore-like ostiole. Peridium thin, composed of thickor thin-walled large cells. Hamathecium of cellular
pseudoparaphyses, septate and branching. Asci 8spored, bitunicate, fissitunicate, clavate to cylindroclavate, with a very short, furcate pedicel and a small
ocular chamber. Ascospores filamentous, hyaline or pale
brown, multi-septate, one of the upper cells swollen.
Anamorphs reported for genus: none.
Literature: Barr 1992a; Holm 1957, 1961; Shoemaker
1984b; Shoemaker and Babcock 1987.
Type species
Notes
Morphology
Neophaeosphaeria was formally established by
Câmara et al. (2003) by segregating Paraphaeosphaeria
species with 3-4-septate ascospores and anamorphs of
ovoid to ellipsoid, non-septate, brown, verrucose to
punctuate conidia forming from percurrently proliferating
conidiogenous cells. Neophaeosphaeria filamentosa was
selected as the generic type. Currently, four species are
included under Neophaeosphaeria, i.e. N. barrii, N.
conglomerate (M.E. Barr) M.P.S. Câmara, M.E. Palm &
A.W. Ramaley, N. filamentosa and N. quadriseptata (M.
E. Barr) M.P.S. Câmara, M.E. Palm & A.W. Ramaley
(Câmara et al. 2003). At present all species in Neophaeosphaeria occur on Yucca (Agavaceae).
Phylogenetic study
The four Neophaeosphaeria species form a monophyletic clade based on both ITS and SSU rDNA
Nodulosphaeria hirta Rabenh., Klotzschii Herb. Viv.
Mycol., Edn 2: no. 725 (in sched.) (1858). (Fig. 67)
Ascomata 260–330 μm high × 260–330 μm diam.,
scattered, or in small groups, immersed to erumpent,
globose or subglobose, black, papillate, ostiolate. Papilla
50–80 μm high, numerous setae occur in the pore-like
ostiole (Fig. 67a and b). Peridium 15–30 μm wide at the
sides, thinner at the base, coriaceous, comprising two types
of cells, outer cells of 1–2 layers of heavily pigmented cells
of textura angularis, cells 6–8 μm diam., cell wall 1.5–
3 μm thick, inner of compressed cells, 5×13–3×8 μm
diam., wall 2–3 μm thick (Fig. 67c). Hamathecium of long
cellular pseudoparaphyses 2–3 μm broad, septate and
branching, mucilage not observed. Asci 100–123×12.5–15
(−17.5) μm (x ¼ 110:8 14:3mm, n=10), 8-spored, bitunicate, fissitunicate, clavate to cylindro-clavate, with a very
short, furcate pedicel, with a small ocular chamber (to 2 μm
wide×1 μm high) (Fig. 67d). Ascospores 48–63×5–6.5 μm
(x ¼ 55:3 5:6mm, n=10), 4-seriate, filamentous, pale
Fungal Diversity
Fig. 66 Neophaeosphaeria filamentosa (from NY, holotype). a
Ascomata as a circular cluster on the host surface. b Hamathecium of
wide psuedoparaphyses. c Section of peridium comprising cells of textura
angularis. d–f Cylindrical asci with thickened apex. Note the short furcate
pedicel. g Pale brown, 3-septate ascospores. Note the verruculose
ornamentation. Scale bars: a=200 μm, b, c=20 μm, d–g=10 μm
Fungal Diversity
brown, 8-septate, the 4th upper cell broader than the others,
smooth-walled, without sheath (Fig. 67e and f).
Anamorph: none reported.
Material examined: GERMANY, Dresdae, in herbarum
caulibus emortuis perrara, exeunte majo, 1858 (BR
101945–95, holotype, as Nodulosphaeria hirta).
Notes
Morphology
The name Nodulosphaeria was first used by Rabenhorst
(1858) but was considered as a synonym of Leptosphaeria
for many years (Clements and Shear 1931). The name was
reinstated by Holm (1957) and was represented by N. hirta,
which was concurrently treated as a synonym of N. derasa
(Berk. & Broome) L. Holm. The most outstanding morphological characters of Nodulosphaeria were considered to be
apex of ascomata often covered with setae, ascospore with
three or more transverse septa with a supramedian enlarged
cell or elongated to a scolecospore, mostly with terminal
appendages (Barr 1992a; Holm 1961; Shoemaker 1984b).
The ascomata are usually immersed and the peridium
comprises a few layers of brown, relatively thin-walled cells
of textura angularis and textura prismatica similar to those
of Phaeosphaeria. Thus, Nodulosphaeria is likely to be a
member of Phaeosphaeriaceae. However, this needs to be
confirmed by molecular analysis. The boundary between
Nodulosphaeria and Ophiobolus is not clear-cut, and the
circumscriptions of them usually depend on the viewpoint of
different mycologists. For instance, Shoemaker (1976) has
assigned some Nodulosphaeria species such as N. erythrospora, N. fruticum, N. mathieui and N. megalosporus to
Ophiobolus. Subsequently, more species were added to
Nodulosphaeria (Barr 1992a; Shoemaker 1984b; Shoemaker
and Babcock 1987). Currently, more than 60 names are
included in Nodulosphaeria (http://www.mycobank.org/, 06/
2010).
Phylogenetic study
None.
Concluding remarks
All species included in Nodulosphaeria have an inflated
ascospore cell as mentioned above. However, it is likely
that this character would have evolved more than once as it
is probably an adaption for ascospore ejection from the
ascus (Shoemaker 1976). It occurs in Ophiobolus species
and the ascomata of these species are quite dissimilar to
Nodulosphaeria species and their exclusion from Nodulosphaeria seems warranted. When considering whether a
species belongs in Nodulosphaeria, one must also consider
the ascomata and peridium structure until DNA sequences
are available.
Ohleria Fuckel, Fungi rhenani exsic.: no. 2173 (1868).
(Melanommataceae)
Generic description
Habitat terrestrial, saprobic. Ascomata small to medium size,
solitary, scattered, or in small groups, erumpent to nearly
superficial, papillate, ostiolate. Peridium thin, thicker at the
apex, 1-layered. Hamathecium of dense, long trabeculate
pseudoparaphyses. Asci 8-spored, bitunicate, fissitunicate,
cylindrical, with a short pedicel. Ascospore brown to reddish
brown, broadly to narrowly fusoid, 3-septate, easily separating into two parts at the primary septum.
Anamorphs reported for genus: Monodictys (Samuels 1980).
Literature: Barr 1990b; Clements and Shear 1931; Patel et
al. 1997; Samuels 1980.
Type species
Ohleria modesta Fuckel, Fungi rhenani exsic. (1868) (Fig. 68)
Ascomata 214–357 μm high × 285–400 μm diam.,
solitary, scattered, or in small groups of 2–3, erumpent to
nearly superficial, coriaceous, with basal wall remaining
immersed in host tissue, broadly or narrowly conical, with a
flattened base not easily removed from the substrate, black;
apex with a conical protruding papilla and an often porelike ostiole (Fig. 68a). Peridium 22–53 μm thick laterally,
thicker at the apex, 1-layered, composed of heavily pigmented
thick-walled cells of textura angularis, cells to 7 μm diam.,
cell wall 1.5–3 μm thick, apex cells smaller and walls
thicker, base cells walls thinner (Fig. 68b). Hamathecium of
dense, long trabeculate pseudoparaphyses 1–2 μm broad,
septate, branching and anastomosing (Fig. 68c). Asci 90–
130×(5.5-)7–10 μm (x ¼ 107:3 8mm, n=10), 8-spored,
with a short pedicel up to 20 μm long, bitunicate,
fissitunicate, cylindrical, with a small ocular chamber (to
1.5 μm wide×1.5 μm high) (Fig. 68c, d and e). Ascospore
15–22×4–5 μm (x ¼ 20 4:4mm, n=10), biseriate near the
top and uniseriate at the base, broadly fusoid to fusoid with
broadly to narrowly rounded ends, brown to reddish brown,
3-septum, deeply constricted at the median septum and
breaking into two conical partspores, no constriction at the
secondary septum, smooth (Fig. 68d and e).
Anamorph: none reported.
Material examined: GERMANY, on decorticated,
decaying roots of Fagus sylvatica, very rare, collected in
autumn (G: F. rh. 2173, isotype).
Notes
Morphology
Ohleria is characterized by its subglobose to conic
ascomata, produced on decorticated woody substrates, as
Fungal Diversity
Fig. 67 Nodulosphaeria hirta (from BR 101945–95, holotype). a
Appearance of ascomata on the host surface. b Vertical section of an
ascoma. Note the setae at the apex and in the ostiole. c Section of a
partial peridium. Note the outer layer cells of textura angularis and
inner layer compressed cells. d Squash mount showing asci in
pseudoparaphyses. e, f. The light brown filiform ascospores. Scale
bars: a=0.5 mm, b=100 μm, c=50 μm, d=20 μm, e, f=10 μm
well as its brown and phragmosporous ascospores which
break into two parts at the median septum (Samuels 1980).
Some species of Ohleria are widespread. For instance, O.
brasiliensis is reported from New Zealand, Brazil as well as
United States (Samuels 1980). Ohleria has been considered
closely related to Sporormia and Preussia based on the
ascosporic characters, and several species of Ohleria, such as
O. aemulans Rehm, O. haloxyli Kravtzev, O. silicata
Kravtzev and O. kravtzevii Schwarzman, have been transferred to these genera. Clements and Shear (1931) treated
Ohleria as a synonym of Ohleriella, despite the fact that
Ohleriella is a coprophilous fungus. When the ascomata and
habitats are considered, Ohleria seems closely related to
Melanomma and Trematosphaeria (Samuels 1980).
Phylogenetic study
None.
Concluding remarks
To some degree, habitats show phylogenetic significance
(Zhang et al. 2009a). Thus, Ohleria seems less likely related
to Sporormia and Preussia. But its relationship with
Melanomma is uncertain, because of their differences in
hamathecium and ascospores.
Fungal Diversity
Fig. 68 Ohleria modesta (from g: f. rh. 2173, isotype). a Ascomata scattering on host surface. b Section of a partial peridium. c Asci embedded
in pseudoparaphyses. d, e Cylindrical asci with short pedicels. Scale bars: a=1 mm, b, c=50 μm, d, e=20 μm
Ohleriella Earle, Bull N Y Bot Gard 2: 349 (1902).
(Delitschiaceae)
Generic description
Habitat terrestrial, saprobic. Ascomata medium to large,
immersed, erumpent to nearly superficial, scattered or in
small groups, usually with a wide papilla, ostiolate,
coriaceous. Peridium composed of small pigmented cells
of textura angularis. Asci 8-spored or fewer, cylindroclavate, with a furcate pedicel. Hamathecium of trabeculate
pseudoparaphyses. Ascospores brown to dark brown,
cylindrical to nearly clavate with broadly to narrowly round
ends, multi-septate, easily broken into partspores, smooth,
with elongated germ slit in each cell.
Anamorphs reported for genus: none.
Literature: Ahmed and Cain 1972; von Arx and Müller
1975; Barr 1990a; Clements and Shear 1931.
Type species
Ohleriella neomexicana Earle, Bull N Y Bot Gard 2: 349
(1902). (Fig. 69)
Ascomata 330–420 μm high × 400–575 μm diam.,
solitary, scattered, or in small groups, immersed to
erumpent, to nearly superficial, with basal wall remaining
Fungal Diversity
immersed in host tissue, coriaceous, globose or subglobose, usually a somewhat thick, short papilla, up to
100 μm high, with a pore-like ostiole (Fig. 69a). Peridium
27–35 μm thick laterally, up to 55 μm thick at the apex, 1layered, composed of small pigmented cells of textura
angularis, cells up to 5×8 μm diam., cell wall 1.5–2 μm
thick, apex cells smaller and walls thicker (Fig. 69b).
Hamathecium of dense, long trabeculate pseudoparaphyses, 1–1.5 μm broad, anastomosing and branching between and above the asci. Asci 150–208×17.5–25 μm
(x ¼ 182:5 22mm, n=10), 8-spored, bitunicate, fissitunicate, cylindrical to cylindro-clavate, with a narrowed, furcate,
thin pedicel, 15–55 μm long, 2–3.5 μm broad, with a large
truncate ocular chamber best seen in immature asci (to 4 μm
wide×3 μm high) (Fig. 69e, f and g). Ascospores 55–72.5×
10–12 μm (x ¼ 63 10:4mm, n=10), 3–4 seriate to uniseriate near the base, cylindrical to clavate, with broadly to
narrowly rounded ends, brown, 6–7 transverse septa, easily
separating into partspores, with germ slits, central partspores
of the ascospores shorter than broad, rectangular in vertical
section, round in transverse section, 7–8×6–10 μm diam.,
apical cells usually longer than broad, 11–17.5×6–7 μm
diam. (Fig. 69c and d).
Anamorph: none reported.
Material examined: USA, Albuquerque, Bernalillo Co.,
New Mexico, dry gravelly hill, on wood, 29 Nov. 1901, T.
S.A. Cockerell (NY, holotype).
Notes
Morphology
Ohleriella was formally established by Earle (1902) based
on its “medium to large ascomata with a wide papilla,
relatively wide peridium, cylindro-clavate asci, brown to
deep brown multi-septate ascospore, with elongated germ slit
on each cell”, and was monotypified by O. neomexicana
(Barr 1990a). Ohleriella subsequently has been treated as a
synonym of Ohleria, Sporormiella or Preussia (Ahmed and
Cain 1972; von Arx and Müller 1975; Clements and Shear
1931). Spororminula tenerifae, the generic type of Spororminula, was assigned to Ohleriella, thus Spororminula was
treated as a synonym of Ohleriella (Barr 1990a). Two new
species were introduced by Barr (1990a) from North America.
Currently, three species are included in this genus, i.e. O.
herculean (Ellis & Everh.) M.E. Barr, O. neomexicana and O.
nudilignae M.E. Barr & Malloch (http://www.index
fungorum.org; http://www.mycobank.org, 01/03/2009).
The generic type, O. neomexicana, is morphologically
similar to the coprophilous genus Sporormiella, but is
saprobic on grass stems.
Phylogenetic study
None.
Concluding remarks
Although we maintain Ohleriella as a separate genus
here, its saprobic habitat on grasses and similarity to the
coprophilous Sporormiella may indicate a close evolutionary relationship, with the grass saprobic possibly
being an early relative of the coprophilous Sporormiella.
Alternatively, the species/genera may simply occupy
different ecological niches (i.e. dead grass vs dead grass
in dung). Molecular studies are needed to resolve this
issue.
Ophiobolus Reiss, Hedwigia 1:27 (1854).
(Phaeosphaeriaceae)
Generic description
Habitat terrestrial, saprobic or hemibiotrophic. Ascomata
medium-sized, solitary, scattered, or in groups, globose or
pyriform, coriaceous, black, papillate, ostiolate, periphysate.
Peridium thin, thicker near the apex, thinner at the base.
Hamathecium of long cellular pseudoparaphyses, septate,
anastomosing or branching not observed. Asci 8-spored,
bitunicate, fissitunicate dehiscence not observed, cylindrical,
with a short, furcate pedicel. Ascospores filamentous,
narrower toward the lower end, pale brown, multiseptate, separating into two partspores from the middle
septum, from the breaking point, the second cell of each
partspore enlarged.
Anamorphs reported for genus: Coniothyrium-like,
Rhabdospora, Phoma-like and Scolecosporiella (Hyde et
al. 2011; Shoemaker 1976; Sivanesan 1984).
Literature: Holm 1948, 1957; Müller 1952; Reiss 1854;
Shoemaker 1976; Sivanesan 1984.
Type species
Ophiobolus disseminans Reiss, Hedwigia 1:27 (1854)
(Fig. 70).
Ascomata 220–380 μm high×290–430 μm diam., solitary,
scattered, or in groups often arranged in a row, immersed with
a protruding papilla, globose, pyriform, coriaceous, black,
periphysate. Papilla 40–90 μm high, with a pore-like ostiole
(Fig. 70a and b). Peridium 40–55 μm wide at the sides, up to
70 μm thick at the apex, thinner at the base, comprising two
cell types, outer layer composed of small heavily pigmented
thick-walled cells of textura angularis, cells 2–5 μm diam.,
cell wall 2–3 μm thick, apex cells smaller and walls thicker,
inner layer composed of lightly pigmented or hyaline thinwalled cells of textura angularis, 5–7 μm diam., wall 1.5–
2 μm thick, merging with pseudoparaphyses (Fig. 70c).
Hamathecium of long cellular pseudoparaphyses, 2–3 μm
broad, septate, anastomosing or branching not observed
(Fig. 70e). Asci 150–195×8–12.5 μm (x ¼ 169:5 10:7mm,
Fungal Diversity
Fig. 69 Ohleriella neomexicana (NY, holotype). a Ascoma scattering on the host surface. b Section of a partial peridium. Note the small
cells of textura angularis. c Ascospore in ascus. d Ascospore breaking
into part spores. Note the sigmoid germ slit. e Dehiscent ascus. f, g
Asci with short pedicels. Scale bars: a=100 μm, b=50 μm, c–g=
10 μm
Fungal Diversity
n=10), 8-spored, bitunicate, fissitunicate dehiscence not
observed, cylindrical but narrowing towards the base, with a
short, furcate pedicel which is 10–25 μm long, ocular
chamber not observed (Fig. 70d and e). Ascospores 110–
160×2.5–4 μm (x ¼ 135:3 3mm, n=10), filamentous,
narrower toward the lower end, pale brown, 22–30septate, separating into two partspores from the middle
septum, from the breaking point the second cell of each
partspore enlarged.
Anamorph: none reported.
Material examined: GERMANY, near Kassel, on dead
stem of Cirsium arvense (L.) Scop., Spring 1853 (BPI629021, type).
Notes
Morphology
Ophiobolus was established by Reiss (1854) as a
monotypic genus represented by O. disseminans based on
its “Perithecia discreta, ostiolis prominentibus: sporae ascis
inclusae, binatae, filliformes, multiseptatae”.
A broad generic concept was adopted for the genus by
Holm (1948) and Müller (1952). Shoemaker (1976) surveyed
Canadian species of Ophiobolus using the broad concept of
Holm (1948) and Müller (1952). A narrower generic concept
was used by Holm (1957), which only included species with
ascospores separating into two halves. Holm (1957) assigned
species with enlarged ascospore cells to Nodulosphaeria, and
those with long spirally coiled ascospores to Leptospora
(Shoemaker 1976). This left only three species accepted
under Ophiobolus (Holm 1957), although this concept has
rarely been followed with new species recently being
described (Raja and Shearer 2008).
Walker (1980) provided a detailed description from the
type material and dealt with many species of scolecospored
fungi that had been placed in Ophiobolus by Saccardo
(1883). Thus, currently several Ophiobolus sensu lato
species are separated into Acanthophiobolus, Entodesmium,
Leptosphaeria and Leptospora. Ophiobolus sensu lato
contains about 300 species names (Sivanesan 1984; http://
www.mycobank.org/, 04/02/2009).
Phylogenetic study
Ophiobolus fulgidus (Cooke & Peck) Sacc. (as Leptosphaeria fulgida (Cooke & Peck) M. E. Barr in Dong et al.
1998) lacks support in the clade of Leptosphaeriaceae
(Dong et al. 1998). We expect it may closely related to
Phaeosphaeriaceae.
Concluding remarks
We agree from morphological data that Ophiobolus
should comprise species that have filamentous spores that
break easily into two halves at the central septum, with the
second cell on either side being swollen (Walker 1980) and
that the genus presently comprises three species (i.e. O.
anthrisci (L. Holm) L. Holm, O. ophioboloides (Sacc.) L.
Holm and O. acuminatus). All other Ophiobolus species
need to be re-examined and should be placed in other
genera such as Nodulosphaeria and Leptospora. The genus
is in need of revision and molecular phylogenetic study.
Ophiosphaerella Speg., Anal. Mus. nac. Hist. nat. B. Aires
19: 401–402 (1909). (Phaeosphaeriaceae)
Generic description
Habitat terrestrial, saprobic or hemibiotrophic. Ascomata
small- to medium-sized, solitary or scattered, immersed,
globose or subglobose, papillate, ostiolate. Peridium thin.
Hamathecium of dense, filliform, septate pseudoparaphyses.
Asci bitunicate, fissitunicate dehiscence not observed, cylindrical often narrower near the base, with a short furcate
pedicel. Ascospores filamentous, pale brown, multi-septate.
Anamorphs reported for genus: Scolecosporiella (Farr et
al. 1989).
Literature: von Arx and Müller 1975; Schoch et al. 2006,
2009; Spegazzini 1909; Walker 1980; Wetzel et al. 1999;
Zhang et al. 2009a.
Type species
Ophiosphaerella graminicola Speg., Anal. Mus. nac. Hist.
nat. B. Aires 19: 401 (1909). (Fig. 71)
Ascomata 280–325 μm high×250–300 μm diam., solitary
or scattered, immersed with a short papilla protruding out of
the substrate, globose or subglobose, often laterally flattened,
dark brown to black, papillate, papilla ca. 100 μm high, 140–
180 μm broad, disk-like in appearance from above, periphysate (Fig. 71a and b). Peridium 11–25 μm wide, thicker near
the apex, comprising two cell types of small cells, outer wall
composed 6–10 layers of lightly brown flattened cells of
textura angularis, inner layer composed of paler and thinwalled cells, both layers thicker near the apex (Fig. 71b).
Hamathecium of dense, long pseudoparaphyses 0.8–1.5 μm
broad near the apex, septate, 2–3 μm broad between the asci.
Asci 105–135×5.5–10 μm (x ¼ 118:5 7mm, n=10), 8spored, bitunicate, cylindrical and narrower near the
base, with a short, furcate pedicel, up to 30 μm long,
small inconspicuous ocular chamber (to 1.5 μm wide×
1 μm high) (Fig. 71c, d, e and f). Ascospores 100–125×
1.8–2.2 μm (x ¼ 118 2mm, n=10), filamentous, pale
brown, 12–20 septa, smooth-walled.
Anamorph: none reported.
Material examined: ARGENTINA, Tucumán, on leaf
sheath of Leptochloa virgata (L.) P. Beauv., 14 Apr. 1906,
C. Spegazzini (LPS 858, holotype).
Fungal Diversity
Fig. 70 Ophiobolus
disseminans (from BPI-629021,
type). a Immersed ascomata
scattered on the host surface.
Note the erumpent papilla. b
Section of an ascoma. c. Section
of a partial peridium. Note the
thick-walled outer layer and
thin-walled inner layer (orange
colour due to DIC). d Ascus
with a short furcate pedicel. e
Squash mount showing asci in
pseudoparaphyses. Scale bars:
a=0.5 mm, b=100 μm,
c=50 μm, d, e=20 μm
Notes
Morphology
Ophiosphaerella was introduced by Spegazzini (1909)
who described and illustrated a single new species, O.
graminicola, and thus the genus was validly published
(Walker 1980, p. 70). After checking the type specimen,
Petrak and Sydow (1936) transferred the generic type to
Ophiobolus graminicolus (Speg.) Petrak & Syd, and
assigned Ophiosphaerella as a synonym of Ophiobolus.
This was followed by von Arx and Müller (1975).
Ophiosphaerella differs from Phaeosphaeria by its scolecospores without swollen cells or appendages, and from
Ophiobolus by its ascospores without swollen cells or
separating into partspores, thus was kept as a separating
genus (Eriksson 1967a; Walker 1980).
Phylogenetic study
Ophiosphaerella forms a monophyletic group as a sister
group of Phaeosphaeria located in Phaeosphaeriaceae (Schoch
et al. 2006, 2009; Wetzel et al. 1999; Zhang et al. 2009a).
Concluding remarks
Numerous Ophiobolus species are likely to belong in
Ophiosphaerella. The two genera are distinguished as
Ophiobolus sensu Shoemaker (1976) has swollen central
cells or breaking into partspores or with long spirally
coiled ascospores, and Ophiosphaerella (sensu Walker
Fungal Diversity
1980) has scolecospores without swollen central cells or
breaking into partspores.The recent introduction of Ophiobolus shoemakeri Raja & Shearer (Raja and Shearer 2008)
is probably incorrect since the ascospores do not split up
into partspores and there is no swelling above septum
either. In particular, its freshwater habitat also distinguishes it from other species of Ophiobolus. Like
Ophiobolus, Ophiosphaerella is in need of phylogenetic
analysis but appears to be closely related to Phaeosphaeriaceae (Schoch et al. 2006).
Ostropella (Sacc.) Höhn., Annls mycol. 16: 144 (1918).
(Pleosporales, genera incertae sedis)
≡ Ostropa subgen. Ostropella Sacc., Syll. fung. (Abellini) 2: 805 (1883).
Fig. 71 Ophiosphaerella
graminicola (from LPS 858,
holotype). a Ascomata on the
host surface. Note the protruding disk-like papilla. b Section
of an ascoma. c Asci in
pseudoparaphyses with short
pedicels. d–f Cylindrical asci
with short pedicels. Scale bars:
a=0.5 mm,
b=100 μm, c–f =10 μm
Generic description
Habitat terrestrial, saprobic. Ascomata large, erumpent
to superficial, solitary or gregarious, globose to subglobose, with broad and compressed papilla and slit-like
ostiole. Peridium carbonaceous. Hamathecium of dense,
long trabeculate pseudoparaphyses, anastomosing and
branching, rarely septate, embedded in mucilage. Asci
clavate with very long and thin and furcate pedicels.
Ascospores pale brown, ellipsoid to fusoid, 1-septate,
constricted.
Anamorphs reported for genus: none.
Literature: Barr 1990a; Chesters and Bell 1970; Holm and
Yue 1987; Huhndorf 1993; Müller and von Arx 1962;
Müller and Dennis 1965; Saccardo 1883.
Fungal Diversity
Type species
Ostropella albocincta (Berk. & M.A. Curtis) Höhn., Annls
mycol. 16: 144 (1918). (Fig. 72)
≡ Ostropa albocincta Berk & M.A. Curtis, in Berkeley,
J. Linn. Soc., Bot. 10: 372 (1868).
Ascomata 1000–1730 μm high×1050–1450 μm diam.,
scattered to gregarious, erumpent to superficial, globose to
subglobose, roughened, often covered with white crustose
covering, with subiculum, with a broad compressed papilla
and long and slit-like ostiole (Fig. 72a). Peridium 100–
250 μm thick, not of uniform thickness throughout entire wall
area, composed of two cell types, one is of lightly pigmented
thin-walled cells of textura prismatica, cells up to 17×3 μm
diam., cell wall <1 μm thick, intermingled with small heavily
pigmented thick-walled cells of textura globosa, cells up to
5 μm diam., cell wall 2–3 μm thick (Fig. 72b). Hamathecium
of dense, long trabeculate pseudoparaphyses, 1.2–1.8 μm
broad, anastomosing and branching, rarely septate, embedded
in mucilage (Fig. 72c). Asci 90–150(−180)×8–13(−17) μm
(x ¼ 120:5 11:5mm, n=10), 8-spored, bitunicate, fissitunicate dehiscence not observed, cylindro-clavate, with a long,
narrowed, furcate pedicel which is up to 75 μm long, and with
a small ocular chamber best seen in immature asci (up to
2 μm wide×1 μm high) (Fig. 72d and e). Ascospores 18–26×
5–6 μm (x ¼ 22:4 5:6mm, n=10) biseriate in upper part
and uniseriate in lower part, fusoid, pale brown, 1-septate,
deeply constricted at the septum, smooth or rarely verrucose
(Fig. 72f, g and h).
Anamorph: none reported.
Material examined: Wright s.n., Herb. G.E. Massee,
(NY 921990, possible isotype); CUBA, as Ostropa
albocincta, C. Wright 345, 1879 (K(M): 143941, syntype).
differences in ascomatal vestiture and the rather thick
wall comprising two cell types of Ostropella albocincta
differ from those of Schizostoma pachythele. Chesters and
Bell (1970) suggested that S. pachythele, Xenolophium
leve and X. verrucosum Syd. are three varieties under
Lophiostoma pachythele (Berk. & Broome) Chesters &
A.E. Bell. The conspecific status of the three taxa was
supported by Holm and Yue (1987). Although no
combination was made, Holm and Yue (1987) assigned
these taxa to Xenolophium instead of Lophiostoma. Barr
(1990a) suggested that either Ostropella or Xenolophium
could accommodate these closely related taxa, i.e. O.
fusispora (Seaver) E. Müll., S. pachythele, X. leve, and X.
verrucosum. Huhndorf (1993) formally transferred S. applanata Petch and S. pachythele to Xenolophium.
Phylogenetic study
Phylogenetic analysis based on LSU sequences indicated
that Ostropella albocincta clusters together with Xenolophium applanatum as well as species of Platystomum, but
they receive poor support (Mugambi and Huhndorf 2009b).
They all were temporarily assigned under Platystomaceae
(Mugambi and Huhndorf 2009b).
Concluding remarks
Although the placement of Ostropella albocincta under
Platystomaceae lacks support, Ostropella should be excluded from Melanommataceae despite its trabeculate
pseudoparaphyses.
Paraliomyces Kohlm., Nova Hedwigia 1: 81 (1959).
(Pleosporales, genera incertae sedis)
Generic description
Notes
Morphology
Ostropella was established by Saccardo (1883) as a
subgenus of Ostropa and was monotypic being represented
by O. albocincta. The genus was formally established (as
Ostropella) and redescribed by von Höhnel (1918b) and
later the description was modified by several workers (Barr
1990a; Huhndorf 1993; Müller and von Arx 1962; Müller
and Dennis 1965). Ostropella is characterized by having
large ascomata, a conspicuous ridged compressed papilla
with an elongated slit-like ostiole, and 1-septate lightly
pigmented ascospores.
The affinity of Ostropella to Schizostoma sensu Sacc. was
first recognized by von Höhnel (1918b) and this was accepted
by Müller and von Arx (1962) and they transferred
Schizostoma pachythele (Berk. & Broome) Sacc. and Ostreionella fusispora Seaver to Ostropella. Holm and Yue (1987),
however, disagreed with this transfer because of the
Habitat marine, saprobic. Ascostromata immersed, penetrating into the substrate with dark brown hyphae. Ascomata
medium-sized, solitary, immersed or erumpent, subglobose
to pyriform, subiculate or nonsubiculate, papillate or
epapillate, ostiolate, periphysate, carbonaceous. Peridium
thick. Hamathecium of long trabeculate pseudoparaphyses.
Asci 8-spored, bitunicate, fissitunicate, cylindrical, with a
short furcate pedicel, without apical apparatus, uniseriate.
Ascospores ellipsoid to broadly fusoid with broadly rounded
ends, 1-septate, constricted at the septum, hyaline, smoothwalled, surrounded by a gelatinous sheath.
Anamorphs reported for genus: none.
Literature: Kohlmeyer 1959; Tam et al. 2003.
Type species
Paraliomyces lentifer Kohlm. [as ‘lentiferus’], Nova
Hedwigia 1: 81 (1959). (Fig. 73)
Fungal Diversity
Ascostromata black, immersed, penetrating into the
substrate with dark brown hyphae. Ascomata up to
680 μm high × 540 μm diam., solitary, immersed or
erumpent, subglobose to pyriform, subiculate or nonsubiculate, papillate or epapillate, ostiolate, periphysate, carbonaceous (Fig. 73a). Peridium thick. Hamathecium of long
trabeculate pseudoparaphyses, 1–1.5 μm broad. Asci 90–
130×12–17 μm (x ¼ 116 15mm, n=10), bitunicate, fissitunicate, cylindrical, 8-spored, uniseriate, with a short
furcate pedicel, without apical apparatus (Fig. 73b, c and
d). Ascospores 17.5–25×10–12.5 μm (x ¼ 21 11mm, n=
10), ellipsoid to broadly fusoid with broadly rounded ends,
1-septate, constricted at the septum, hyaline, smooth-walled,
surrounded by a gelatinous sheath that contracts to form a
lateral, lentiform, viscous appendage over the septum, 7.5–
12.5 μm diam., 1–3 μm thick (Fig. 73e, f, g and h).
Anamorph: none reported.
Material examined: USA, Florida, Charlotte Harbor in
Punta Garda, 10 Jan. 1964, leg., det. J. J. Kohlmeyer (Herb.
J. Kohlmeyer No. 1720).
Notes
Morphology
Paraliomyces was introduced to accommodate the
marine fungus P. lentifer, which is characterized by
immersed ascomata produced within the ascostroma,
trabeculate pseudoparaphyses, cylindrical, 8-spored asci,
ellipsoidal, hyaline, 1-septate ascospores surrounded by a
gelatinous sheath, which forms a lentiform, viscous
appendage over the septum (Kohlmeyer 1959).
Phylogenetic study
Based on analysis of SSU sequences, Paraliomyces
lentifer nested within Pleosporales, but its familial status
was left undetermined (Tam et al. 2003).
Concluding remarks
None.
Phaeosphaeria I. Miyake, Bot. Mag., Tokyo 23: 93 (1909).
(Phaeosphaeriaceae)
Fig. 72 Ostropella albocincta (K(M): 143941, syntype). a Ascomatab
gregarious on host surface. b Section of the partial peridium. Note the
peridium comprising two cell types and the whitening tissue
(arrowed). c Pseudoparaphyses. d, e Asci with long pedicel. f–
h Ascospores, which are strongly constricted at the central septum.
Scale bars: a=1 mm, b=100 μm, d, e, h=20 μm, c, g, f=10 μm
Anamorphs reported for genus: Amarenographium,
Hendersonia-like, Phaeoseptoria, Scolecosporiella and
Stagonospora (Hyde et al. 2011; Leuchtmann 1984;
Shoemaker and Babcock 1989b).
Literature: von Arx and Müller 1975; Câmara et al.
2002; Eriksson 1967a, 1981; Holm 1957; Khashnobish and
Shearer 1996; Leuchtmann 1984; Miyake 1909; Shoemaker
and Babcock 1989b.
Type species
Phaeosphaeria oryzae I. Miyake, Bot. Mag., Tokyo 23:
136 (1909). (Fig. 74)
Ascomata 120–140 μm high × 100–140 μm diam.,
solitary, scattered, or in small groups, immersed, globose,
subglobose, wall black, forming black spots on the leaves
of hosts (Fig. 74a). Apex with a pore-like ostiole. Peridium
4–8 μm wide at the sides, composed of heavily pigmented
thin-walled cells of textura angularis, cells 2–2.5×3–5 μm
diam., cell wall less than 1 μm thick (Fig. 74b). Hamathecium of dense, long cellular pseudoparaphyses 2–2.5 μm
broad, embedded in mucilage, rarely branched, septate. Asci
53–80(−90) × 7–10 μm (x ¼ 65:3 8:3mm, n = 10), 8spored, bitunicate, fissitunicate, broadly cylindrical to
narrowly fusoid, with a short pedicel which is ca. 8 μm
long, with a small ocular chamber and an inconspicuous
apical apparatus (to 2 μm wide × 1 μm high) (Fig. 74c, d
a n d e ) . Ascospores 17–22(−28) × 4–5 μm
(x ¼ 20:5 4:6mm, n=10), obliquely uniseriate, partially
overlapping or biseriate, narrowly fusoid with rounded ends,
pale brown, 3-septate, slightly constricted at primary septum,
granulate (Fig. 74f and g).
Anamorph: none reported.
Material examined: JAPAN, Suruya, Shizuoka, on the
leaves of Oryza sativa, Sept. 1907 (S nr F9572, F9573,
lectotype).
Generic description
Notes
Habitat terrestrial, saprobic or hemibiotrophic. Ascomata
small, solitary, scattered, or in small groups, immersed,
globose, subglobose, wall black. Apex with a pore-like
ostiole. Peridium thin. Hamathecium of dense, filliform,
septate pseudoparaphyses. Asci 8-spored, bitunicate, fissitunicate, broadly cylindrical to narrowly fusoid, with a
short pedicel. Ascospores fusoid to narrowly fusoid, pale
brown to brown, 3-septate.
Morphology
Phaeosphaeria was introduced by Miyake (1909), but
was regarded as a synonym of Leptosphaeria for a long
time. Holm (1957), however, reinstated Phaeosphaeria,
assigning some Leptosphaeria sensu lato species with
relatively small ascomata and which occurred on monocotyledons to Phaeosphaeria. Although this division based
Fungal Diversity
Fungal Diversity
Fig. 73 Paraliomyces lentifer (from Herb. J. Kohlmeyer No. 1720). a
Section of an immersed ascoma. b Eight-spored cylindrical asci
embedded in pseudoparaphyses. c, d Cylindrical asci with short
pedicels. e–h One-septate hyaline ascospores. Scale bars: a=100 μm,
b–d=20 μm, e–h=10 μm
on host range is considered unnatural by some workers
(Dennis 1978; Sivanesan 1984), it has been widely
accepted (von Arx and Müller 1975; Eriksson 1967a;
Hedjaroude 1969; Shoemaker and Babcock 1989b).
Eriksson (1981) further revised the generic concept of
Phaeosphaeria by including dictyosporous taxa as well as
some perisporium taxa. Phaeosphaeria was further divided into six subgenera, i.e. Ovispora, Fusispora, Phaeosphaeria, Spathispora, Vagispora and Sicispora, based on
differences in ascospore shape and the number of septa
Fungal Diversity
(Shoemaker and Babcock 1989b). Phaeosphaeria species
are usually associated or parasitic on annual monocots,
such as Cyperaceae, Juncaceae or Poaceae but have also
been recorded as saprobes and on dicotyledons (e.g. P.
viridella and P. vagans).
Phylogenetic study
The separation of Phaeosphaeria from Leptosphaeria
sensu stricto was supported by phylogenetic studies based
on ITS sequences. The peridium structure, pseudoparenchymatous cells in Phaeosphaeria versus scleroplectenchymatous cells in Leptosphaeria had phylogenetic
significance in the distinction between these two genera,
while the subgenus division was not supported by the
phylogenetic results (Câmara et al. 2002; Morales et al.
1995). The familial status of both Phaeosphaeriaceae and
Leptosphaeriaceae was verified by multigene phylogenetic
analysis (Schoch et al. 2009; Zhang et al. 2009a).
Concluding remarks
Phaeosphaeria was originally thought to be a synonym of
Leptosphaeria (Müller 1950; Munk 1957), however, molecular analysis has shown these two genera differ with
Phaeosphaeria having pseudoparenchymatous peridium, Stagonospora-like anamorph and mostly monocotyledonous
hosts and Leptosphaeria having scleroplectenchymatous
peridium, Phoma-like anamorph and mostly dicotyledonous
hosts (Câmara et al. 2002; Schoch et al. 2009; Shoemaker and
Babcock 1989b; Zhang et al. 2009a). It is now recognized
that Phaeosphaeria is the type genus of Phaeosphaeriaceae and related genera include Entodesmium and
Setomelanomma and probably Ophiosphaerella (Schoch
et al. 2009; Zhang et al. 2009a). Paraphaeosphaeria was
introduced as an off-shoot of Phaeosphaeria and differs
in ascospore shape and septation as well as anamorphic
stages (Eriksson 1967a, b). Similarly, Nodulosphaeria
was recently reinstated and differs from Phaeosphaeria
because of setae over the apex as well as its ascospores
with swelling supramedian cells and terminal appendages
(Holm 1957, 1961). While the newly reinstated Phaeosphaeria was confined to monocotyledons and particularly
grasses, there are now many species that have been
described from dicotyledons (Farr et al. 1989). Whether
these taxa form a monophyletic group needs to be
investigated with fresh collections and molecular data.
Phaeosphaeriopsis M.P.S. Câmara, M.E. Palm & A.W.
Ramaley, Mycol. Res. 107: 519 (2003). (Phaeosphaeriaceae)
Generic description
Habitat terrestrial, saprobic or hemibiotrophic? Ascomata small, scattered or in small groups, immersed,
globose, subglobose. Peridium thin, comprising one cell
type of textura angularis. Hamathecium of dense, wide
cellular pseudoparaphyses. Asci 8-spored, bitunicate,
cylindrical to broadly fusoid, with a short pedicel and a
small ocular chamber. Ascospores obliquely uniseriate
and partially overlapping to biseriate even triseriate,
cylindrical, pale brown, multi-septate, primary septum
submedian, with or without constriction, verrucose or
baculate.
Anamorphs reported for genus: Coniothyrium-like,
Phaeostagonospora (Câmara et al. 2003).
Literature: Câmara et al. 2003.
Type species
Phaeosphaeriopsis glaucopunctata (Grev.) M.P.S. Câmara,
M.E. Palm & A.W. Ramaley, Mycol. Res. 107: 519 (2003).
(Fig. 75)
≡ Cryptosphaeria glaucopunctata Grev. Fl. Edin.: 362
(1824).
Ascomata 120–150 μm high × 140–200 μm diam.,
scattered, or in small groups, immersed, globose, subglobose (Fig. 75a). Peridium 10–25 μm wide, comprising one
type of cells, composed of thick-walled cells of textura
angularis, cells 4–9 μm diam., cell wall 2–3 μm thick,
almost equal in thickness. Hamathecium of dense, wide
cellular pseudoparaphyses, 3–5 μm broad. Asci (50-)60–
110 × 10–15 μm (x ¼ 82:3 12mm, n = 10), 8-spored,
bitunicate, fissitunicate dehiscence not observe, cylindrical to broadly fusoid, with a short pedicel, with a small
ocular chamber (to 0.8 μm wide×1 μm high) (Fig. 75b).
Ascospores 18–28×5–7.5 μm (x ¼ 23:5 6:2mm, n=10),
obliquely uniseriate and partially overlapping to biseriate even triseriate, cylindrical, pale brown, 4(−5)septate, without constriction or slightly constricted at
the basal septum, the forth cell from the apex usually
slightly inflated, the basal cell often longer, baculate
(Fig. 75c, d, e and f).
Anamorph: none reported.
Material examined: UK, Epping, Sept. 1863 (E, M.C.
Cooke 166, barcode: E00074286).
Notes
Morphology
Phaeosphaeriopsis was introduced to accommodate
some species of Paraphaeosphaeria based on both morphological characters and results of SSU rDNA sequence
analyses (Câmara et al. 2003). Most of the Phaeosphaeriopsis species occur on the Agavaceae, although P.
glaucopunctata occurs on Liliaceae (Ruscus). Phaeosphaeriopsis is characterized by having uni- or multioculate
stromata and 4- or 5-septate ascospores. Although the
Fungal Diversity
Fig. 74 Phaeosphaeria oryzae
(from S nr F9572, F9573, lectotype). a Appearance of ascomata on the host surface. b
Section of an ascoma. c Squash
mount showing asci in pseudoparaphyses. Note that asci with
short pedicels. d, e Asci with
short pedicels. F, G. Light
brown 3-septate ascospores.
Scale bars: a=100 μm, b–g=
10 μm
morphological characters of Phaeosphaeriopsis species is
more diverse than those of Paraphaeosphaeria sensu
stricto or Neophaeosphaeria, the ITS sequences are more
similar to each other than those of the other two genera
(Câmara et al. 2003). Currently, Phaeosphaeriopsis comprises seven species, namely P. agavensis (A.W. Ramaley,
M.E. Palm & M.E. Barr) M.P.S. Câmara, M.E. Palm & A.
W. Ramaley, P. amblyospora A.W. Ramaley, P. glaucopunctata, P. musae Arzanlou & Crous, P. nolinae (A.W.
Ramaley) M.P.S. Câmara, M.E. Palm & A.W. Ramaley, P.
obtusispora (Speg.) M.P.S. Câmara, M.E. Palm & A.W.
Ramaley and P. phacidiomorpha (Ces.) D.F. Farr & M.E.
Palm (http://www.mycobank.org/, 06/2010).
is also shown to belong to Phaeosphaeriaceae in recent
phylogenetic studies (Schoch et al. 2009; Plate 1).
Phylogenetic study
The generic type of Phaeosphaeriopsis, P. glaucopunctata, located in Phaeosphaeriaceae based on SSU rDNA
sequences (Câmara et al. 2003). Phaeosphaeriopsis musae
Habitat terrestrial, saprobic? Ascomata small, scattered,
immersed, semi-immersed to nearly superficial, globose,
subglobose, black, smooth; apex with a protruding
papilla and pore-like ostiole, without periphyses. Perid-
Concluding remarks
None.
Platysporoides (Wehm.) Shoemaker & C.E. Babc., Can. J.
Bot. 70: 1648 (1992). (Pleosporaceae)
≡ Pleospora subgenus Platysporoides Wehmeyer, A
World Monograph of the genus Pleospora and its Segregates,
p. 236. 1961.
Generic description
Fungal Diversity
ium thin, composed of a few layers of textura
angularis. Hamathecium of numerous, cellular pseudoparaphyses, anastomosing, septate. Asci bitunicate, fissitunicate, cylindrical to cylindro-clavate, with a short,
furcate pedicel. Ascospores broadly ellipsoid, reddish
brown, muriform.
Anamorphs reported for genus: none.
Literature: Shoemaker and Babcock 1992; Wehmeyer
1961.
Type species
Platysporoides chartarum (Fuckel) Shoemaker & C.E.
Babc., Can. J. Bot. 70: 1650 (1992) (Fig. 76)
≡ Pleospora chartarum Fuckel, Jb. nassau. Ver. Naturk.
23–24: 133–134 (1870).
Fig. 75 Phaeosphaeriopsis
glauco-punctata (from M.C.
Cooke 166). a Ascomata immersed in the substrate. b Eightspored cylindrical asci. c–f. Pale
brown baculate ascospores
which are released from asci.
Scale bars: a=200 μm,
b=20 μm, c, d–f=10 μm
Ascomata 150–230 μm high × 180–260 μm diam.,
scattered, immersed, semi-immersed to rarely superficial,
globose, subglobose, black, smooth; apex with a protruding papilla, 50–85 μm long, 60–85 μm broad, ostiolate
(Fig. 76a and b). 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, without periphyses.
Hamathecium of dense, long cellular pseudoparaphyses,
2–3 μm broad, anastomosing, septate (Fig. 76c). Asci
110–140 × 12.5–16.5 μm (x ¼ 121:5 14:7mm, n = 10),
(6-)8-spored, bitunicate, fissitunicate, cylindrical to
cylindro-clavate, with a short, furcate pedicel, 8–17 μm
long, ocular chamber not observed (Fig. 76c, d and e).
Ascospores 20–26× 8–11 μm (x ¼ 23:7 9mm, n =10),
obliquely uniseriate and partially overlapping, flattened,
broadly ellipsoid in front view, reddish brown, 3
Fungal Diversity
transverse septa, 1 longitudinal septum in each central
cell, 1 oblique septum in each end cell, constricted at all
septa, granulate, with a sheath 2–3 μm wide (as reported
in Shoemaker and Babcock 1992) (Fig. 76f, g and h).
Anamorph: none reported.
Material examined: GERMANY, Budenheim, Leopold
Fuckel, Nassau’s Flora, on old paper (G NASSAU: 210558
(a), as Sphaeria chartarum Wallr., type).
Notes
Morphology
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 “applanodictyospore” and “terete 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 “an intraepidermal discoid lid” (Shoemaker and
Babcock 1992).
Phylogenetic study
None.
Concluding remarks
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 are of little phylogenetic
significance.
Pleomassaria Speg., Anal. Soc. cient. argent. 9: 192
(1880). (Pleomassariaceae)
Shoemaker and LeClair 1975; Sivanesan 1984; Tanaka
et al. 2005.
Type species
Pleomassaria siparia (Berk. & Broome) Sacc., Syll. fung.
2: 239 (1883) (Fig. 77)
≡ Sphaeria siparia Berk. & Broome, Ann. Mag. nat.
Hist., Ser. 2 9: 321 (1852).
Ascomata 150–410 μm high × 440–740 μm diam.,
solitary, scattered, or in small groups, immersed, erumpent
by a minute slit or a small conical swelling in the bark,
depressed globose, papillalte, ostiolate (Fig. 77a). Peridium 45–60 μm wide, thicker at the apex, thinner at the
base, 1-layered, composed of small pigmented thickwalled compressed cells, cells ca. 15×3 μm diam., cell
wall 2–3.5 μm thick, apex cells larger, base composed of
small pigmented thick-walled cells of textura angularis,
ca. 5 μm diam. (Fig. 77b). Hamathecium of dense, cellular
pseudoparaphyses, 1–2 μm broad, embedded in mucilage,
anastomosing or branching not observed. Asci 180–250×
28–42 μm (x¼206:3 36:8mm, n=10), 8-spored, bitunicate, fissitunicate, broadly cylindrical to broadly cylindroclavate, with a short, thick pedicel, 15–45 μm long, with
inconspicuous ocular chamber (Fig. 77c and d). Ascospores 45–58×12.5–17.5 μm (x ¼ 50:5 14:8mm, n=10),
biseriate, narrowly oblong with broadly to narrowly
rounded ends, brown, muriform with 5–8 transverse septa
and 1–2 vertical septa in some cells, smooth to verrucose,
constricted at the septa, surrounded by a mucilaginous
sheath (Fig. 77e, f and g).
Anamorph: Prosthemium betulinum Kunze (Sivanesan
1984).
Conidia to 120 μm diam., with 3–5 arms, each arm 3–5septate, 40–55×13–16 μm, connected to a central cell
(Fig. 77h, i and j).
Material examined: UK, Wiltshire, Spye Park, on
branch of Betulina with Hendersonia polycystis Berk., et
Br. leg. C.E. Broome, 1850? (BR, type).
Generic description
Notes
Habitat terrestrial, saprobic. Ascomata medium to large,
solitary, scattered, or in small groups, immersed, erumpent
by a minute slit or a small conical swelling in the bark,
flattened, papillate, ostiolate. Hamathecium of dense,
cellular pseudoparaphyses, embedded in mucilage. Asci
bitunicate, fissitunicate, broadly cylindrical to broadly
cylindro-clavate, with a short, thick pedicel. Ascospores
muriform, brown, constricted at the septa.
Anamorphs reported for genus: Prosthemium and Shearia
(Barr 1982b; Sivanesan 1984).
Literature: Barr 1982b, 1990b, 1993a; Clements and Shear
1931; Eriksson 2006; Lumbsch and Huhndorf 2007;
Morphology
Pleomassaria as characterized by Barr (1982b) has
medium- to large-sized, immersed ascomata, cellular
pseudoparaphyses, clavate to oblong asci and large,
muriform ascospores (Barr 1982b; Sivanesan 1984). The
muriform and somewhat asymmetrical ascospores with a
submedian primary septum distinguish Pleomassaria from
Asteromassaria in the family Pleomassariaceae, while in
Splanchnonema ascospores have distinct bipolar asymmetry. Barr (1982b) included five North American species in
the genus, while Kirk et al. (2008) listed four species.
Fungal Diversity
Fig. 76 Platysporoides chartarum (from G NASSAU:
210558, type). a, b Ascomata
scattered among fibers. Note the
central ostioles. c Asci in numerous cellular pseudoparaphyses. d, e Cylindro-clavate asci
with short pedicels. f–h. Muriform ascospores. Scale bars:
a, b=200 μm, c–e=20 μm,
f–h=10 μm
Barr (1993a) treated Pleomassaria as a synonym of
Splanchnonema based on a morphological cladistic analysis, but this proposal was not followed by later workers
(Eriksson 2006; Lumbsch and Huhndorf 2007; Tanaka et
al. 2005).
Phylogenetic study
Pleomassaria siparia forms a robust phylogenetic clade
with Melanomma pulvis-pyrius (generic type) (Schoch et al.
2009; Zhang et al. 2009a), which might represent a
phylogenetic family (or suborder?).
Fungal Diversity
Concluding remarks
The genera Asteromassaria, Pleomassaria and Splanchnonema of Pleomassariaceae are considered to be closely
related and difficult to separate (Barr 1982b; Crivelli 1983).
They all have ascomata which are immersed in bark and are
visible as slightly raised pustules with small ostioles, but
may eventually become erumpent (e.g. Asteromassaria
macrospora). Pseudoparaphyses are cellular, asci are bitunicate, while ascospores vary from 1-septate and pale brown
(e.g. Asteromassaria macrospora) to muriform (e.g. Pleomassaria siparia) and may be symmetrical (e.g. Asteromassaria macrospora) or highly asymmetrical (e.g.
Splanchnonema pustulatum). The peridium ranges from
thick-walled textura angularis (e.g. Asteromassaria macrospora) to thin-walled compressed cells (e.g. Splanchnonema
pustulatum) and medium textura prismatica (e.g. Pleomassaria siparia). Anamorphs also vary distinctly, Prosthemium
in Pleomassaria siparia, Scolicosporium in Asteromassaria
macrospora but no anamorphic stage reported for Splanchnonema pustulatum. Furthermore, Asteromassaria pulchra
clusters in Morosphaeriaceae in this study, thus here we
tentatively assign Asteromassaria in Morosphaeriaceae
(Plate 1). There seems to be considerable confusion in this
family, especially when Pleomassaria siparia forms a robust
phylogenetic clade with Melanomma pulvis-pyrius (Melannomataceae). Thus in this study, Pleomassariaceae is
restated as a separate family from Melannomataceae.
Therefore, fresh collections of the types of these genera are
needed for molecular analysis and to establish which
characters are important for classification.
Pleophragmia Fuckel, Jb. nassau. Ver. Naturk. 23–24: 243
(1870). (Sporormiaceae)
Generic description
Habitat terrestrial, saprobic (coprophilous). Ascomata smallto medium-sized, gregarious, immersed to erumpent, globose
to subglobose, black, coriaceous; apex with a short papilla,
or sometimes forming an ostiolar pore. Peridium thin,
composed of several layers of thin-walled cells of textura
angularis. Hamathecium of dense, delicate pseudoparaphyses. Asci 8-spored, bitunicate, fissitunicate, clavate to
cylindro-clavate, with a relatively long pedicel and an ocular
chamber. Ascospores muriform, narrow oblong to cylindrical
with rounded ends, dark brown, constricted at each septum.
Anamorphs reported for genus: none.
Literature: von Arx and Müller 1975; Cain 1934.
Ascomata 330–480 μm high × 320–430 μm diam.,
gregarious, immersed to slightly erumpent, globose to
subglobose, black; apex with a short papilla, sometimes
forming a ostiolar pore (Fig. 78a). Peridium 25–35 μm
thick at the sides, composed of one cell type of lightly
pigmented thin-walled cells of textura angularis, cells 6–
10 μm diam., cell wall 1.5–2 μm thick (Fig. 78b).
Hamathecium of numerous, long pseudoparaphyses, 1–
2 μm broad, anastomosing not observed. Asci 160–250×
22.5–27.5 μm (x ¼ 203:6 25mm, n=10), 8-spored, bitunicate, fissitunicate, clavate to cylindro-clavate, with a 20–
50 μm long pedicel and an ocular chamber (to 5 μm wide×
2 μm high) (Fig. 78e and f). Ascospores 42–50×8–10 μm
(x ¼ 46 10mm, n=10), biseriate to uniseriate and partially
overlapping, narrowly oblong to cylindrical with rounded
ends, dark brown, often slightly curved, with 9 transverse
septa with two crossing longitudinal septa in the centre,
constricted at each septum, smooth-walled (Fig. 78c, d, g
and h).
Anamorph: none reported.
Material examined: GERMANY, between Königstein
and Glashütten, on the same dung with Delitschia minuta.
s.d. (G, Fungi rhenani n2272, type).
Notes
Morphology
Pleophragmia was formally established by Fuckel
(1870) and monotypified by Pleophragmia leporum. The
most comparable genus to Pleophragmia is Sporormia, as
ascospores of both have no germ slits and the inner layer of
wall is considerably thinner than the outer layer (Barr
1990a, b). But the muriform ascospores of Pleophragmia
can be readily distinguished from the phragmosporous
ascospores of Sporormia. Currently, only four species are
accommodated under this genus (http://www.mycobank.
org, 28-02-2009).
Phylogenetic study
None.
Concluding remarks
The presence of both transverse and crossing longitudinal
septa is the most striking character of Pleophragmia, although
the phylogenetic significance of this character is unclear.
Pleoseptum A.W. Ramaley & M.E. Barr, Mycotaxon 54: 76
(1995). (Phaeosphaeriaceae)
Type species
Generic description
Pleophragmia leporum Fuckel, Jb. nassau. Ver. Naturk.
23–24 (1870) [1869–70]. (Fig. 78)
Habitat terrestrial, saprobic? Ascomata medium-sized,
scattered, or in small groups, immersed, globose to conoid,
Fungal Diversity
Fig. 77 1 Pleomassaria siparia
(from BR, type). a Ascomata on
the host surface (after removing
the cortices). b Section of a
partial peridium. c, d Asci with
short pedicels. e–g Ascospores
with thin sheath. Scale bars:
a=0.5 mm, b–d=50 μm, e–g=
20 μm. 2 Prosthemium betulinum (from BR, type). h–j Conidia with arms. Scale bars:
h–j=20 μm
black, papillate, ostiolate. Peridium 1-layered. Hamathecium of dense, long cellular pseudoparaphyses, septate,
branching. Asci 8-spored, bitunicate, fissitunicate, cylindrical to cylindro-clavate, with furcate pedicel. Ascospores
obliquely uniseriate and partially overlapping, muriform,
ellipsoid, ovoid to fusoid, yellowish to dark brown.
Anamorphs reported for genus: Camarosporium (Ramaley
and Barr 1995).
Fungal Diversity
Fig. 77 (continued)
Literature: Ramaley and Barr 1995.
Type species
Pleoseptum yuccaesedum A.W. Ramaley & M.E. Barr,
Mycotaxon 54: 76 (1995). (Fig. 79)
Ascomata 300–500 μm diam., scattered, or in small
groups of 2–3, immersed with a flattened top, globose to
conoid, black, papillate, ostiolate (Fig. 79a). Papilla
small, slightly protruding from the host surface. Peridium
30–50 μm thick at sides, up to 100 μm thick at the apex,
1-layered, composed of 5–8 layers of heavily pigmented
purplish-brown cells of textura angularis, cells 5–12 μm
diam., cell wall 1–2 μm thick, apex cells smaller and
walls thicker (Fig. 79c). Hamathecium of dense, long
cellular pseudoparaphyses 1–2 μm broad, septate,
branching (Fig. 79b). Asci 125–170(−195) × 15–22 μm
(x ¼ 153:8 19:3mm, n = 10), 8-spored, bitunicate, fissitunicate, cylindrical to cylindro-clavate, with a short,
narrowed, furcate pedicel which is 10–20 μm long, with
an ocular chamber best seen in immature asci (to 5 μm
broad × 3 μm high) (Fig. 79d and e). Ascospores 22–30 ×
11–14 μm (x ¼ 27:1 12:6mm, n = 10) obliquely uniseriate and partially overlapping, ellipsoid, ovoid to fusoid,
yellowish to yellowish brown, becoming reddish brown
to dark brown, muriform, with 3-(4) transverse septa,
constricted at the primary septum, part above central
septum wider, vertical septa exist in each cell, ornamentation of foveolae in linear rows (Fig. 79f and g).
Anamorph: Camarosporium yuccaesedum Fairm.
(Ramaley and Barr 1995).
Conidiomata 200–450 μm diam., pycnidial, immersed,
scattered, subglobose to conoid, ostiolate. Macroconidiogenous cells determinate or indeterminate, enteroblastic,
hyaline, smooth. Macroconidia holoblastic, 20–36×10–
15 μm diam., ellipsoid to narrowly ovoid, muriform,
yellowish brown, 3–7 transverse septa, constricted at the
septa. Microconidiogenous cells produced near or in the
ostiole, hyaline, smooth. Microconidia 5–10×5–7 μm
diam., globose to ovoid, aseptate, hyaline, smooth.
Material examined: USA, Colorado, Montezuma County,
hillside near entrance to Mesa Verde National Park, on dead
leaves of Yucca baccata, 11 Oct. 1992, Ramaley Annette
(9237A) (BPI 802381, holotype).
Notes
Morphology
Pleoseptum is a monotypic genus established by
Ramaley and Barr (1995) and represented by P. yuccaesedum based on its “immersed ascomata, thick peridium,
muriform ascospores, anamorphic stage and the linoeate
ornamentation of the ascospores and conidia”. The shape
of ascomata of Pleoseptum is comparable with that of
Chaetoplea, but the peridium structure easily distinguishes
them. Some species of Curreya, Leptosphaeria and
Heptameria are comparable with Pleoseptum, but their
anamorphic stages differ.
Pleoseptum yuccaesedum and its Camarosporium yuccaesedum anamorph both formed in the leaves of Yucca
baccata and the ascomata and conidiomata were indistinguishable. Camarosporium is the anamorph of diverse
teleomorph genera included in Botryosphaeriales and
Cucurbitariaceae (Kirk et al. 2008). The genus is in need
of revision (Sutton 1980) and is no doubt polyphyletic.
Phylogenetic study
None.
Concluding remarks
The placement of Pleoseptum under Phaeosphaeriaceae
is still tentative.
Pleospora Rabenh. ex Ces. & De Not., Comm. Soc. crittog.
Ital. 1: 217 (1863). (Pleosporaceae)
Generic description
Habitat terrestrial, saprobic or parasitic. Ascomata smallto medium-sized, immersed, erumpent to superficial,
Fungal Diversity
Fig. 78 Pleophragmia leporum (from G. Fungi rhenani n2272,
type). a Appearance of ascomata on the substrate surface. Note the
ostiolar pore. b Section of a partial peridium. c, h Apical part of an
ascus. Note the apical apparatus in (c). d Released ascospores. e, f
Clavate Asci with pedicels. g Part of a broken ascospore. Note the
crossing septa. Scale bars: a=0.5 mm, B=50 μm, c–f=20 μm, g,
h=10 μm
Fungal Diversity
Fig. 79 Pleoseptum yuccaesedum (from BPI 802381, holotype). a
Appearance of ascomata scattered on the host surface. Only the
upper region is visible. b Squash mount of asci in pseudoparaphyses.
c Section of an ascoma. Note the peridium comprising cells of
textura angularis. d, e Asci with short furcate pedicels. f, g
Muriform dark-brown ascospores. Scale bars: a = 0.5 mm, b =
40 μm, c=100 μm, d, e=20 μm, f, g=10 μm
Fungal Diversity
papillate, ostiolate. Peridium thin. Hamathecium of dense,
cellular pseudoparaphyses. Asci 8-spored, bitunicate,
fissitunicate, cylindrical to clavate, with furcate pedicel
and small inconspicuous ocular chamber. Ascospores
muriform, brown or pale brown, with or without
sheath.
Anamorphs reported for genus: Stemphylium (Simmons
1985).
Literature: Barr 1981; Frisullo and Braun 1996; Kodsueb
et al. 2006a; Luttrell 1951; Wehmeyer 1946, 1961, 1975;
Zhang et al. 2009a.
Type species
Pleospora herbarum (Pers.) Rabenh., Klotzschii Herb. Viv.
Mycol. 2: no. 547 (1854). (Fig. 80)
≡ Sphaeria herbarum Pers., Syn. meth. fung. (Göttingen)
1: 78 (1801).
Ascomata 130–220 μm high × 250–420 μm diam.,
scattered, or in small groups of 2–3, immersed, semiimmersed to erumpent, broadly to narrowly oblong and
flattened, with flattened base not easily removed from the
substrate, wall black, papillate, ostiolate (Fig. 80a and b).
Peridium 30–50 μm thick on sides, thinner at the base,
coriaceous, 2-layered, outer layer composed of one or two
layers of heavily pigmented thick-walled cells of textura
angularis, cells 5–10 μm diam., cell wall 2–4 μm thick,
apex cells smaller and walls thicker, inner layer composed
of hyaline thin-walled cells of textura angularis, 8–12 μm
diam., wall hyaline, 0.5–1.5 μm thick (Fig. 80c). Hamathecium of dense, cellular pseudoparaphyses, 2–3 μm
broad, filling the gaps between the asci. Asci 100–210×
27.5–30 μm (x ¼ 142:2 28:3mm, n=10), 8-spored, bitunicate, fissitunicate, broadly cylindrical to clavate, with a
short, thick, furcate pedicel, 8-12(−20) μm long, with
small inconspicuous ocular chamber (ca. 3 μm wide×
1 μm high) (Fig. 80d and e). Ascospores 28–38×12.5–
15 μm (x ¼ 33 14:5mm, n=10), ellipsoidal, straight or
sometimes curved, with broadly rounded ends and upper
hemispore slightly shorter and broader; spores usually
divided by 3 A-transsepta, all 4 segments by longisepta
and then by one stratum of B-transsepta (mature spores as
a rule with 7 transsepta, 3A + 4B), yellowish brown,
smooth; each hemispore with thick gelatinous sheath, the
lower one with umbilicus (sheaths fused in mature spores)
(Fig. 80f, g, h, i, j and k).
Anamorph: Stemphyllium herbarum E. Simmons
(Simmons 1985).
Material examined: GERMANY, on stalks of Melilotusalla? at the bank of the Elbe in Konigstein, 1882 (E,
Krieger 683); as Sphaeria herbarum Persoon Syn. fung. p.
78 (E, 81); as Sphaeria herbarum Fries, Scleromyceti
Sueciae 38 (E, lectotype).
Notes
Morphology
Pleospora was originally assigned within Sphaeriales.
Subsequently, it was assigned within Pseudosphaeriales and
Pleosporales (Wehmeyer 1961). Pleospora is a large group,
which is widely distributed and associated with a wide range
of species of monocotyledons as well as dicotyledons
(Wehmeyer 1975). All species of Pleospora have muriform
ascospores (Wehmeyer 1961, 1975). Pleospora has downward growing pseudoparaphyses within the ascomata of
“Pleospora-type” development (Luttrell Univ. Mo. Stud.
1951), which subsequently served as a diagnostic character.
However, only a limited number of species had detailed
studies on this character (Wehmeyer 1961). The heterogeneous nature of Pleospora has been noted, and several
subgenera have been erected, such as Scleroplea to include
all “sclerotioid” species of Pleospora, Teichosporoides to
accommodate species of Pleospora with immersed ascomata,
Pleosphaeria for those having superficial and setose ascomata
(Wehmeyer 1961). Similarly, Cucurbitaria, Fenestella and
Montagnula are also separated as a section from Pleospora.
Most of these subgenera are currently at genus rank.
Phylogenetic study
The polyphyletic nature of Pleospora is clear (Kodsueb et
al. 2006a), and those that stain the woody substrate purple
should be assigned to Amniculicolaceae (Zhang et al. 2009a).
Concluding remarks
As some Pleospora species have a wide range of host
spectrum, especially on both monocotyledons and dicotyledons, it is highly possible they are cryptic species.
Preussia Fuckel, Hedwigia 6: 175 (1867) [1869–70].
(Sporormiaceae)
Generic description
Habitat terrestrial, saprobic (on decaying fibers or coprophilous). Ascomata small- to medium-sized, cleistothecial or
perithecial, solitary or scattered on substrate surface,
globose, membraneous, black. Peridium thin, composed
of thick-walled, poly-angular cells from the surface view.
Pseudoparaphyses not observed. Asci (4-) 8-spored, bitunicate, clavate to broadly clavate, with a long and thin and
furcate pedicel. Ascospores 3–6 seriate to uniseriate near
the base, cylindrical with rounded ends, brown, septate,
easily breaking into partspores, with germ slits in each cell.
Anamorphs reported for genus: Phoma (von Arx 1973;
Cain 1961; Malloch and Cain 1972).
Literature: Ahmed and Cain 1972; Arenal et al. 2005; von
Arx 1973; von Arx and van der Aa 1987; Auerswald 1866;
Fungal Diversity
Fig. 80 Pleospora herbarium (from E, Krieger 683). a Immersed
ascomata scattering on host surface. b Ascomata in small groups. Note:
the surface layer of the host is removed. c Section of an ascoma. Note
the peridium cells of textura angularis. d, e. Asci with short pedicels f–j
Ascospores. Scale bars: a, b=0.5 mm, c=100 μm, d, e=30 μm, f–j=
20 μm
Barr 1987b, 1990a; Boylan 1970; Cain 1961; Eriksson 1992;
Fuckel 1866; Guarro et al. 1981, 1997a, b; Khan and Cain
1979a, b; Kruys and Wedin 2009; Lodha 1971; Lorenzo
1994; Luck-Allen and Cain 1975; Maciejowska and
Fungal Diversity
Williams 1963; Malloch and Cain 1972; Munk 1957;
Narendra and Rao 1976; Rai and Tewari 1963; Sultana
and Malik 1980.
Type species
Preussia funiculata (Preuss) Fuckel, Jb. nassau. Ver.
Naturk. 23–24: 91 (1870) [1869–70]. (Fig. 81)
≡ Perisporium funiculatum Preuss, Fung. Hoyersw.: no.
145 (1851).
Ascomata 240–500 μm diam., cleistothecial, solitary,
scattered on substrate, superficial, globose, membraneous,
black (Fig. 81a). Peridium thin, composed of thick-walled,
poly-angular cells in front view (Fig. 81b). Pseudoparaphyses
not observed. Asci 42–65×20–25 μm (x ¼ 55:8 21:8mm, n=
10), (4-)8-spored, bitunicate, broadly clavate, with a long and
thin and furcate pedicel, up to 115 μm long, ocular chamber
not observed (Fig. 81c and d). Ascospores 30–40×6.3–
7.5 μm (x ¼ 35:6 6:9mm, n=10), 3–6 seriate to uniseriate
near the base, cylindrical with rounded ends, brown, with
3 transverse septa, easily breaking into partspores, central
cells round in transverse section but rectangular in
vertical section, with a germ slit in each cell, 6.5–8.5×
4–7.5 μm broad, apical cells 8.8–10×5–7 μm broad,
sheath not observed.
Anamorph: none reported.
Material examined: USA, Ontario, York Co., Nashville,
on old jute sack on ground, 1 Jul. 1960, leg. & det. R.F.
Cain (in part Preussia typharum) (TRTC 46985).
Notes
Morphology
Preussia was introduced by Fuckel (1866) to accommodate species having cleistothecioid ascomata, bitunicate asci, multi-septate ascospores with a germ slit in
each cell and with a gelatinous sheath, and occurring in
soil or plant debris. Preussia, Sporormia and Sporormiella are regarded as closely related genera, which share
numerous morphological characters. Sporormia can be
distinguished from Preussia by its perithecioid ascomata
and cylindrical asci. The only distinguishing morphological character for Preussia from Sporormiella are the
cleistothecioid ascomata in Preussia (Barr 2000; Cain
1961), but this has been shown to have little phylogenetic
significance (von Arx 1973; Zhang et al. 2009a).
Substrate preference has been used to distinguish species
of Sporormiella and Preussia, with Sporormiella being
restricted to a coprophilous habitat, while Preussia grows
in plant debris, wood or soil (von Arx and van der Aa
1987). This proposal was rejected, as P. intermedia
(Clum) Cain can be isolated from either soil or dung
(Guarro et al. 1997b). In a review of Preussia, Cain
(1961) accepted 12 species, and some of them are coprophilous. Subsequently, numerous additional new species have
been published (Arenal et al. 2005; Barr 1987b, 1990a;
Boylan 1970; Eriksson 1992; Guarro et al. 1981, 1997a, b;
Khan and Cain 1979a; Lodha 1971; Lorenzo 1994; LuckAllen and Cain 1975; Maciejowska and Williams 1963;
Malloch and Cain 1972; Narendra and Rao 1976; Rai and
Tewari 1963; Sultana and Malik 1980). Currently, 84 species
are listed under Preussia (http://www.mycobank.org/
mycotaxo.aspx, 10/2010) and Kirk et al. (2008) estimates
there are 51 species.
Phylogenetic study
In phylogenetic analysis based on ITS, nLSU, mtSSU
and β-tubulin gene fragments, Preussia, Sporormiella and
Spororminula clustered together. Thus, Sporormiella together with Spororminula are treated as synonyms of
Preussia (Kruys and Wedin 2009).
Concluding remarks
Preussia sensu lato (including Sporormiella and Spororminula) based on both morphology and molecular data
should be accepted pending further research.
Quintaria Kohlm. & Volkm.-Kohlm., Bot. Mar. 34: 34
(1991). (Pleosporales, genera incertae sedis)
Habitat marine, saprobic. Ascomata medium-sized, scattered or loosely gregarious, immersed, mostly subglobose,
rarely globose, with a protruding papilla, ostiolate. Peridium thin, 2-layered, coriaceous, thicker near the apex.
Hamathecium of dense, filamentous, trabeculate pseudoparaphyses, branching and anastomosing between and
above asci. Asci 8-spored, bitunicate, fissitunicate,
cylindro-clavate, with a short furcate pedicel. Ascospores
biseriate, broadly fusoid to fusoid, hyaline, mostly 5septate, rarely up to 7-septate.
Anamorphs reported for genus: none.
Literature: Hyde and Goh 1999; Kohlmeyer and
Volkmann-Kohlmeyer 1991; Suetrong et al. 2009; Zhang
et al. 2008b.
Type species
Quintaria lignatilis (Kohlm.) Kohlm. & Volkm.-Kohlm.,
Bot. Mar. 34: 35 (1991). (Fig. 82)
≡ Trematosphaeria lignatilis Kohlm., Marine Ecology,
[Pubblicazioni della Stazione Zoologica Napoli I] 5(4): 365
(1984).
Ascomata 240–500 μm diam., scattered or loosely
gregarious, immersed, globose to subglobose, coriaceous,
ostiolate, ostiole is encrusted with thick-walled black cells,
papilla up to 400 μm long (Fig. 82a). Peridium thin, 20–
30 μm wide, thinner at the base, thicker near the apex, up to
Fungal Diversity
Fig. 81 Preussia funiculata
(from TRTC 46985). a Superficial cleistothecoid ascomata. b
Part of peridium from front
view. c Squash mounts showing
a large number of asci. d A
clavate ascus with a long and
thin pedicel. Scale bars: a=
0.5 mm, b=20 μm, c, d=50 μm
300 μm, 2-layered, outer layer composed of hyphoid cells,
inner layer composed of compressed cells of textura
angularis (Fig. 82b). Hamathecium of dense, filamentous,
trabeculate pseudoparaphyses, 0.8–1.5 μm broad, branching
and anastomosing between and above asci (Fig. 82e).
Asci 175–250×25–35 μm (x ¼ 220 28mm, n=10), 8spored, bitunicate, fissitunicate, cylindro-clavate, with a
short, furcate pedicel, to 20 μm long (Fig. 82c and e).
Ascospores 55–73×12.5–15 μm (x ¼ 63:3 13:1mm, n=
10), biseriate, broadly fusoid to fusoid, usually slightly
curved, smooth, hyaline, mostly 5-septate, rarely up to 7septate, smooth-walled, lacking a sheath.
Anamorph: none reported.
Material examined: BELIZE, Twin Cays, on attached
dead tip of prop root of Rhizophora mangle, with shipworms, 3 Apr. 1983, leg. & det. J.K. Kohlmeyer (J.
Kohlmeyer No. 4365a, holotype).
Notes
Morphology
Quintaria was introduced to accommodate the marine
fungus, Trematosphaeria lignatilis, based on its immersed
ascomata with rounded bases, black incrustations surround-
ing the sides of the ostiolar canal as well as its hyaline
ascospores (Kohlmeyer and Volkmann-Kohlmeyer 1991).
Subsequently, three more species were introduced to this
genus, viz. Q. aquatica K.D. Hyde & Goh, Q. microsporum
Yin. Zhang, K.D. Hyde & J. Fourn. and Q. submerse K.D.
Hyde & Goh, which are all from freshwater (Hyde and Goh
1999; Zhang et al. 2008b).
Phylogenetic study
Multigene phylogenetic study indicated that Quintaria
lignatilis forms a separate sister clade to other families of
Pleosporales, which may represent a new familial linage
(Suetrong et al. 2009). This was supported by phylogenetic
studies which place the freshwater Q. submersa separate from
Q. lignatilis (Schoch et al. 2009; Suetrong et al. 2009; Plate 1).
Concluding remarks
The freshwater members of Quintaria should likely be
excluded from this genus, and only the generic type, Q.
lignatilis retained, but this needs confirmation.
Roussoëlla Sacc., in Saccardo & Paoletti, Atti Inst. Veneto
Sci. lett., ed Arti, Sér. 3 6: 410 (1888). (Arthopyreniaceae
(or Massariaceae))
Fungal Diversity
Generic description
Habitat terrestrial, saprobic. Ascomata medium-sized, clustered, immersed in host tissue, forming under darkened,
slightly raised, somewhat liner or dome-shaped stroma on the
host, with a flush intra-epidermal papilla; immersed under
clypeus, papillate, ostiolate. Peridium thin, comprising
several layers of compressed cells. Hamathecium of dense,
long trabeculate pseudoparaphyses, embedded in mucilage,
hyaline, anastomosing and septate. Asci 8-spored, bitunicate,
cylindrical, with furcate pedicel, and a conspicuous ocular
chamber. Ascospores uniseriate to partially overlapping, fusoid
or ellipsoidal, brown, 1-septate, constricted at the septum.
Anamorphs reported for genus: Cytoplea (Hyde et al.
1996a).
Literature: Hyde et al. 1996a; Hyde 1997; Ju et al. 1996;
Tanaka et al. 2009.
Type species
Roussoëlla nitidula Sacc. & Paol., Atti Ist. Veneto Sci., Ser.
6, 6:410. (1888). (Fig. 83)
Ascomata 160–200 μm high×400–500 μm diam., clustered, immersed in host tissue, forming under darkened,
slightly raised, somewhat liner or dome-shaped stroma on the
host, with a flush intra-epidermal papilla; in vertical section
subglobose with a flattened base, immersed under clypeus,
subglobose with a flattened base, papillate, ostiolate (Fig. 83a).
Peridium up to 20 μm thick, comprising several layers of
compressed cells. Hamathecium of dense, long trabeculate
pseudoparaphyses, 1–1.5 μm broad, embedded in mucilage,
anastomosing and septate. Asci 123–220×7–11 μm, 8-spored,
bitunicate, cylindrical, with furcate pedicels, and a conspicuous ocular chamber (Fig. 83b, c and d). Ascospores 17.5–
22×5.5–8 μm, uniseriate to partially overlapping, fusoid or
ellipsoidal, brown, 1-septate, constricted at the septum,
ornamented with longitudinal wall striations and surrounded
by a wide mucilaginous sheath (Fig. 83e, f, g and h).
Anamorph: Cytoplea hysterioides K.D. Hyde (Hyde et
al. 1996a).
Material examined: MALAYSIA, Malacca, on culms of
Bambusa Bar & Grill, 1885, B. Scortechini 15 (PAD,
Roussoëlla nitidula Sacc. Paol. 2484, holotype, on a loose
label Roussoëlla nitidula S. & P. Est Phyllachora phaeodidym./15 prob. original material from Malacca Peninsula).
Notes
Morphology
Roussoëlla was introduced by Saccardo for the single
species R. nitidula Sacc. & Paol. (Saccardo and Paoletti
1888). It was redescribed by Hyde et al. (1996a) and the
anamorph of Roussoëlla hysterioides (Ces.) Höhn., Cyto-
plea hysterioides K.D. Hyde was determined and described.
Roussoëlla was then reviewed by Hyde (1997) and a
modified key for Roussoëlla species was provided based on
the one proposed by Ju et al. (1996). Roussoëlla is
characterized as having immersed ascomata containing
long cylindrical asci and brown 1-septate ornamented
ascospores. In this study, we have checked the type species
and it matches Hyde et al. (1996a). The asci are bitunicate,
but we could not observe the fissitunicate dehiscence.
Phylogenetic study
Species of Roussoëlla, Roussoellopsis as well as Arthopyrenia salicis form a robust phylogenetic clade, which form
a sister group with pleosporalean families, but the generic
type of Roussoëlla (R. nitidula) was not included in the
phylogenetic study (Tanaka et al. 2009).
Concluding remarks
The bambusicolous habitat of Roussoëlla is a striking
character at generic rank classification but its relationship to
the lichenized Arthopyrenia is unexpected and will require
more analysis.
Saccharicola D. Hawksw. & O.E. Erikss., in Eriksson &
Hawksworth, Mycologia 95: 431 (2003). (Massarinaceae)
Generic description
Habitat terrestrial, parasitic. Ascomata medium-sized, solitary,
scattered, immersed, globose to subglobose, carbonaceous,
papillate, ostiolate. Peridium relatively thin, composed of one
cell type of pale brown to hyaline pseudoparenchymatous cells.
Hamathecium of trabeculate pseudoparaphyses. Asci bitunicate,
8-spored, cylindro-clavate to clavate. Ascospores biseriate and
sometimes laterally uniseriate, fusoid with narrowly rounded
ends, septate, constricted at the septa, the upper second cell
becoming pigmented when mature, smooth or verruculose.
Anamorphs reported for genus: Stagonospora (Eriksson
and Hawksworth 2003; Kaiser et al. 1979; Leuchtmann
1984).
Literature: Eriksson and Hawksworth 2003.
Type species
Saccharicola bicolor (D. Hawksw., W.J. Kaiser & Ndimande)
D. Hawksw. & O.E. Erikss., Mycologia 95: 431 (2003).
(Fig. 84)
≡ Leptosphaeria bicolor D. Hawksw., W.J. Kaiser &
Ndimande, Mycologia 71: 483 (1979).
Ascomata 125–175 μm high × 175–220 μm diam.,
solitary, scattered, immersed, globose to subglobose, wall
black, carbonaceous, with a protruding papilla, with a
central ostiole (Fig. 84a). Peridium 15–20 μm thick
Fungal Diversity
Fig. 82 Quintaria lignitalis
(from J. Kohlmeyer No. 4365a,
holotype). a Ascomata immersed in substrate. b Section of
an ascoma. Note the thin peridium and elongated papilla. c, e
Asci embedded in pseudoparaphyses. d Five septate fusoid
hyaline ascospores. Scale bars:
a=0.5 mm, b=200 μm, c, e=
50 μm, d =20 μm
Fungal Diversity
composed of one cell type of pale brown to hyaline
pseudoparenchymatous cells, becoming thicker near the apex
(Fig. 84a). Hamathecium of 1–2 μm broad, filliform, hyaline,
septate pseudoparaphyses, branching and anastomosing in
mucilage. Asci (90-)125–150×(20-)25–30 μm, 8-spored, with
a short pedicel, bitunicate, cylindro-clavate to clavate, with a
small ocular chamber at the apex (Fig. 84c). Ascospores 29–
Fig. 83 Roussoëlla nitidula
(from PAD Paol. 2484, holotype). a Appearance of the
stroma on host surface. b Asci
and pseudoparaphyses. c, d
Long cylindrical furcate asci.
e–h. Ascospores. Note the striate ornamentation. Scale bars:
a=0.5 mm, b–d=20 μm,
e–h=10 μm
42×8–11 μm, biseriate and sometimes laterally uniseriate,
fusoid with narrowly rounded ends, (2-)3-septate, deeply
constricted at the septa, the upper second cell subhyaline to
pale brown when young and becoming dark brown to almost
black at maturity, smooth or verruculose (Fig. 84d). (data from
the original description by Kaiser et al. (1979) because of the
bad condition of the type material).
Fungal Diversity
Anamorph: Pycnidia typical of Stagonospora (Sphaeropsidales), “scattered, arising singly both on the host and in pure
culture, in culture generally surrounded by an envelope of
mycelial hyphae, numerous, immersed on the host, but nearly
superficial in culture, subglobose to slightly applanate, black,
150–250 μm diam., with a central slightly papillate ostiole,
lacking a distinct neck; walls mainly 15–20 μm thick,
composed of three to six layers of pseudoparenchymatous
cells, the outermost layers dark brown and inner pale brown to
hyaline cells somewhat compressed radially, very variable in
size, cells of the outer layers mainly 7–12 μm long×4–6 μm
wide in vertically section and 10–12 μm diam. in surface
view, wall not or only slightly thicked near the ostiole.
Conidiogenous cells lining the inner surface of the pycnidial
cavity, holoblastic, minute and difficult to distinguish from the
pseudoparenchymatous cells with which they are mixed,
mammiform with a flattened apex, hyaline, smooth walled,
about 4–6 μm tall and 4–6 μm wide. Conidia copiously
produced, ellipsoid, with somewhat truncated ends, hyaline,
smooth walled, (2-)3 septate, not or slightly constricted at the
septa, often guttulate, rather thin walled, (21-)24–28(−34)
μm×7–8.5(−11.5) μm” (from Kaiser et al. 1979).
Material examined: KENYA, near Nairobi, on leaves of
Saccharum officinarum L.; 24 Aug. 1977; leg. W.J. Kaiser
(IMI 215888, holotype).
Notes
Morphology
Saccharicola was separated from Leptosphaeria as a new
genus based on its Stagonospora anamorph and its biotrophic habitat in leaves of sugar cane, and two species were
included, i.e. Saccharicola bicolor and S. taiwanensis (J.M.
Yen & C.C. Chi) O.E. Erikss. & D. Hawksw. (Eriksson and
Hawksworth 2003). Saccharicola is characterized by its
parasitic habitat on monocots, small ascomata, bitunicate
asci, presence of pseudoparaphyses as well as its 3-septate
ascospores (Eriksson and Hawksworth 2003).
Phylogenetic study
Based on the limited phylogenetic analysis of SSU
sequences, Saccharicola is considered to be closely related
to Massarina eburnea, the generic type of Massarina
(Eriksson and Hawksworth 2003). Thus, Saccharicola was
assigned to Massarinaceae, which includes Keissleriella,
Massarina and Saccharicola (Eriksson and Hawksworth
2003).
similar to Pleosporineae. Further molecular study is needed
for confirmation.
Salsuginea K.D. Hyde, Bot. Mar. 34: 315 (1991). (Pleosporales, genera incertae sedis)
Generic description
Habitat marine, saprobic. Ascomata large, solitary, fusoid,
conical or subglobose, with or without a flattened base,
immersed under a darkened clypeus, papillate, ostiolate.
Peridium thin, composed of round cells (in cross section) at
sides, fusing at the top with the clypeus, thin at the base.
Hamathecium of dense, long trabeculate pseudoparaphyses,
anastomosing, embedded in mucilage. Asci 8-spored, bitunicate, fissitunicate, clavate to cylindro-clavate, pedunculate,
with a large ocular chamber and conspicuous apical ring.
Ascospores uniseriate, obovoid, brown to black, with hyaline
apical germ pores, 1-septate, constricted at the septum, dark
brown with paler apical cells, lacking sheath, smooth.
Anamorphs reported for genus: none.
Literature: Hyde 1991a; Suetrong et al. 2009.
Type species
Salsuginea ramicola K.D. Hyde, Bot. Mar. 34: 316 (1991).
(Fig. 85)
Ascomata 1040–2600 μm high×455–1430 μm diam.,
solitary, fusoid, conical or subglobose, with or without a
flattened base, immersed under a darkened clypeus, papillate,
ostiolate, ostiole rounded (Fig. 85a). Peridium up to 39 μm
thick, composed of round cells (in cross section) at sides,
fusing at the top with the clypeus, thin at the base (Fig. 85b).
Hamathecium of dense, long trabeculate pseudoparaphyses,
1–2 μm broad, anastomosing, embedded in mucilage. Asci
440–512×29–34 μm, 8-spored, bitunicate, fissitunicate,
clavate to cylindro-clavate, pedunculate, with a large ocular
chamber and conspicuous apical ring (Fig. 85c and e).
Ascospores 59–72×24–30 μm, uniseriate, obovoid, brown to
black, with hyaline apical germ pores, 1-septate, constricted
at the septum, dark brown with paler apical cells, lacking
sheath, smooth (Fig. 85d and f).
Anamorph: none reported.
Material examined: THAILAND, Ranong mangrove,
Aegiceras corniculatum (L.) Blanco., Oct. 1988, leg. & det.
K.D. Hyde (BRIP 17102, holotype).
Notes
Concluding remarks
Based on the parasitic habitat on monocots and its small
ascomata and Stagonospora (or Cercospora? for S. taiwanensis, see Eriksson and Hawksworth 2003; Shoemaker
and Babcock 1989b) anamorph, Saccharicola seems more
Morphology
Salsuginea was introduced to accommodate the mangrove
fungus, S. ramicola, which is characterized by large,
immersed, ostiolate and papillate ascomata under a clypeus,
Fungal Diversity
Fig. 84 Saccharicola bicolor
(from IMI 215888, holotype). a
Section of an ascomata immersed in the host tissue. b
Section of a partial pycnidia.
Note the phragmosporous conidia. c Clavate ascus with ocular chamber and short pedicel. d
Ascospores. Note the pigmented
central cell(s). Scale bars: a, b=
50 μm, c=20 μm, d=10 μm
dense, trabeculate pseudoparaphyses embedded in gel matrix,
fissitunicate, 8-spored, cylindrical asci with short pedicel and
conspicuous apical apparatus, 1-septate, dark brown ascospores with paler apical cells (Hyde 1991a). Salsuginea is
considered closely related to Helicascus and Caryospora, and
they are all proposed to Melanommataceae (Hyde 1991a).
Phylogenetic study
Based on a multigene phylogenetic analysis, Salsuginea
ramicola nested in a paraphyletic clade within Pleosporales;
its familial status is undetermined (Suetrong et al. 2009).
Concluding remarks
It has been shown that trabeculate pseudoparaphyses has
no phylogenetic significance at familial rank, so a well
resolved phylogeny based on DNA comparisons will be
necessary to categorize this genus.
Semidelitschia Cain & Luck-Allen, Mycologia 61: 581
(1969). (Delitschiaceae)
Generic description
Habitat terrestrial, saprobic (coprophilous). Ascomata
immersed to slightly erumpent, scattered, coriaceous,
papillate, ostiolate. Hamathecium of non-typical trabeculate pseudoparaphyses, thin, septate, rarely branching.
Asci cylindrical, pedicellate, each with a conspicuous
large apical ring. Ascospores non-septate, dark brown to
nearly black, each with an elongated germ slit.
Anamorphs reported for genus: none.
Literature: Barr 2000; Cain and Luck-Allen 1969.
Type species
Semidelitschia agasmatica Cain & Luck-Allen, Mycologia
61: 581 (1969). (Fig. 86)
Ascomata 550–900 μm diam., solitary, immersed to
erumpent, globose to subglobose, black, semicoriaceous,
smooth-walled, with a protruding papilla and a conspicuous
ostiole (Fig. 86a). Peridium thin, comprising multi-angular
Fungal Diversity
cells from front view. Hamathecium of non-typical trabeculate pseudoparaphyses, 1–2 μm broad, septate, rarely
branching, anastomosing not observed. Asci 410–505×(38-)
43–50 μm (x ¼ 470:6 46:4mm, n=10), 8-spored, bitunicate, fissitunicate dehiscence not observed, cylindrical, with
a thick pedicel which is up to 90 μm long, and with a large
Fig. 85 Salsuginea ramicola
(from BRIP 17102, holotype).
a Habitat section of an ascoma.
b Section of the partial peridium. c Clavate mature and immature asci. d Ascospores
within ascus. e Apical part of
immature asci. f Ascospores
with an apical chamber at each
end. Scale bars: a=0.5 mm,
b–e=50 μm, f=10 μm
and conspicuous dome-shaped ocular chamber surrounded by
apical ring (to 18 μm wide×4 μm high) (Fig. 86b and e).
Ascospores 53–65×30–38 μm (x ¼ 61:3 34:1mm, n=10),
uniseriate to obliquely uniseriate and partially overlapping, broad fusoid to subglobose, hyaline when
young, then becoming yellowish brown, reddish brown
Fungal Diversity
and nearly black and opaque when mature, non-septate,
smooth-walled, with a full length germ slit, surrounded
by a broad gelatinous sheath (Fig. 86c and d).
Anamorph: none reported.
Material examined: CANADA, Alberta, North of
Beaver Mines, on sheep dung, 28 Jul. 1962, E.R. LuckAllen, (TRTC 41607, paratype); USA, Montana: Gallatin
County, 60 min S of Bozeman, on sheep dung, 2 Sept.
1957, Cain (TRTC 42032, paratype); Stillwater County
Columbus, on cow dung, 3 Sept. 1957, Cain (TRTC 42031,
paratype); South Dakota, Meade Co.: South of Wall, on
cow dung, 3 Sept. 1962, Cain (TRTC 40697, holotype).
Notes
Morphology
Semidelitschia was formally established by Cain and LuckAllen (1969) and was assigned to Sporormiaceae. Although it
is similar to Delitschia, it differs as the ascospores are 1celled, as opposed to 2-celled. Subsequently, Semidelitschia
was transferred to Delitschiaceae together with Delitschia
(Barr 2000). Currently, three species are listed under this
genus, i.e. S. agasmatica Cain & Luck-Allen, S. nanostellata
A.E. Bell & Mahoney and S. tetraspora J.H. Mirza & S.M.
Khan (Index Fungorum) although the number of species in
the genus are given as only two in Kirk et al. (2008).
Phylogenetic study
None.
Concluding remarks
This is a clearly defined genus that differs from
Delitschia in having 1-celled ascospores. Cultures of S.
agasmatica are needed for sequencing and for establishing
the placement and uniqueness of the genus.
Setomelanomma M. Morelet, Bull. Soc. Sci. nat. Arch.
Toulon et du Var 227:15 (1980). (Phaeosphaeriaceae)
Generic description
Habitat terrestrial, hemibiotrophic or biotrophic. Ascomata
small, solitary, scattered, immersed, erumpent to superficial,
globose to subglobose, black; with or without a small
papilla, apex covered with setae and a periphysate ostiole.
Peridium thin, 1-layered, composed of several layers of
cells of textura angularis. Hamathecium of dense, 1–2 μm
broad pseudoparaphyses, septate, anastomosing. Asci 8spored, bitunicate, broadly cylindrical. Ascospores fusoid to
broadly clavate, pale brown to brown, 3-septate.
Anamorphs reported for genus: none.
Literature: Leonard and Suggs 1974; Morelet 1980;
Rossman et al. 2002; Schoch et al. 2009; Zhang et al. 2009a.
Type species
Setomelanomma holmii M. Morelet, Bulletin de la Société
des Sciences naturelles et d’Archéologie de Toulon et du
Var 36 (no. 227): 15 (1980). (Fig. 87)
(Some information in the following description is from
Rossman et al. (2002))
Ascomata 80–250 μm diam., solitary, scattered, immersed, erumpent to superficial, globose to subglobose,
black, with setae; with or without a small papilla, apex
covered with setae and a periphysate ostiole. Peridium 15–
25 μm thick, 1-layered, composed of several layers of cells
of textura angularis, cell wall thinner and more lightly
pigmented towards centrum, cell wall thicker near the apex.
Hamathecium of dense, 1–2 μm broad pseudoparaphyses,
thicker near the base, septate, anastomosing (Fig. 87a and
d). Asci 70–100×11–14 μm, 8-spored, bitunicate, broadly
cylindrical with a short, thick, furcate pedicel, with a small
ocular chamber (Fig. 87a, b and c). Ascospores 16–21×5–
6.5 μm, obliquely uniseriate and partially overlapping to
biseriate, fusoid to broadly clavate with broadly to narrowly
rounded ends, pale brown to brown, 3-septate, slightly
constricted at the median septum, smooth (Fig. 87e).
Anamorph: none reported.
Material examined: FRANCE, Leuglay, on dying twigs
of Picea pungens. 8 May 1987, leg. M. Morelet (UPS F117969 (slide), isotype).
Notes
Morphology
Setomelanomma was formally established by Morelet
(1980) as a monotypic genus represented by S. holmii,
which was collected in France. The description, however, is
not detailed and lacks illustrations. Rossman et al. (2002)
collected this species in North America and detailed studies
were conducted including both morphology and phylogeny.
The bitunicate, broadly cylindrical asci, cellular pseudoparaphyses as well as the pale brown, septate ascospores with
a median primary septum point Setomelanomma to
Phaeosphaeriaceae as defined by Barr (1992a) and
Eriksson et al. (2002) (Rossman et al. 2002). However, its
setose ascomata, brown and 3-septate ascospores together
with its residence in conifers distinguish it from all other
genera under Phaeosphaeriaceae (Rossman et al. 2002).
Setomelanomma is mostly comparable with Kalmusia and
Phaeosphaeria. Setomelanomma can be distinguished from
Kalmusia by its erumpent to superficial ascomata with
almost no papilla, and Phaeosphaeria differs from Setomelanomma by its host spectrum and reported anamorphic
stages (Rossman et al. 2002). Currently, five species are
included in Setomelanomma, namely S. holmii, S. monoceras,
S. prolata K.J. Leonard & Suggs, S. rostrata (K.J. Leonard) K.
Fungal Diversity
Fig. 86 Semidelitschia agasmatica (from TRTC 40697, holotype). a Immersed ascomata
scattered on the surface of the
substrate. b Squash of ascoma.
Note the numerous released asci. c Apical ring of cylindrical
asci. d One-celled ascospores.
Note the germ slits (see arrow).
e Cylindrical ascus. Note the
tapering pedicel. Scale bars:
a=0.5 mm, b–e=100 μm
J. Leonard & Suggs and S. turcica (Luttr.) K.J. Leonard &
Suggs (http://www.mycobank.org/, 06/2010).
Shiraia Henn., Bot. Jb. 28: 274 (1900). (Pleosporales,
genera incertae sedis)
Phylogenetic study
Setomelanomma forms a well supported phylogenetic
clade with other members of Phaeosphaeriaceae (Schoch
et al. 2009; Zhang et al. 2009a).
Generic description
Concluding remarks
None.
Habitat terrestrial, parasitic. Ascostroma warty-like or
tuber-like. Ascomata medium to large, subglobose, gregarious on the surface layer of ascostroma, immersed,
ostiolate, with a small black opening seen on the surface
of the ascostroma, ostiole rounded. Hamathecium of
Fungal Diversity
dense, long trabeculate pseudoparaphyses, anastomosing
and branching between the asci. Asci bitunicate, fissitunicate, cylindrical to cylindro-clavate, with a short furcate
pedicel, with a big and truncate ocular chamber. Ascospores obliquely uniseriate and partially overlapping,
narrowly fusoid to fusoid or broadly fusoid with tapering
or narrowly rounded ends, hyaline to pale brown or
brown, muriform.
Anamorphs reported for genus: coelomycetous with
muriform conidia (see Liu 2009).
Literature: Cheng et al. 2004; Hino 1961; Kishi et al.
1991; Liu 2009; Morakotkarn et al. 2008.
Type species
Shiraia bambusicola Henn., Bot. Jb. 28: 274 (1900).
(Fig. 88)
Ascostroma 1–1.5 cm high×1–2.5 cm diam., subglobose, oblong to irregular, slightly pink with cracking
surface. Ascomata 350–800 μm high×300–700 μm diam.,
subglobose, gregarious on the surface layer of ascostroma,
immersed, ostiolate, with a small black opening seen on
the surface of the ascostroma, ostiole rounded, the inner
tissue of ascostroma carnation red (Fig. 88a and b).
Hamathecium of dense, long trabeculate pseudoparaphyses, 0.8–1.5 μm broad, anastomosing and branching
between the asci. Asci 300–425 × 20–35 μm
(x ¼ 360:5 28mm, n=10), 6-spored, bitunicate, fissitunicate, cylindrical to cylindro-clavate, with a short furcate
pedicel, up to 50 μm long, with a big and truncate ocular
chamber (Fig. 88c and d). Ascospores 62.5–80×17.5–
22.5 μm (x ¼ 72:3 19:3mm, n=10), obliquely uniseriate
and partially overlapping, narrowly fusoid to fusoid with
tapering or narrowly rounded ends, hyaline turning pale
brown when mature, muriform, with 9–13 transversal
septa, 1–3 longitudinal septa in central cells, slightly
constricted at the septa, usually with a gelatinous cap at
each end (Fig. 88e, f and g).
Anamorph: coelomycetous with muriform conidia (see
Liu 2009).
Material examined: CHINA, Zhejiang, Hangzhou,
Panan, on bamboom, 15 Jun. 2009, leg. Liu Yongxiang
(IFRD 2040).
Notes
Morphology
Shiraia is reported as a parasite on branches of several
genera of bamboo distributed mainly in southern regions of
China and Japan (Hino 1961; Kishi et al. 1991; Liu 2009).
Shiraia is characterized by its bambusicolous habitat, large
ascostroma and muriform ascospores. Asci comprise 6
ascospores in this study and some previous studies (Hino
1961; Liu 2009). Shiraia bambusicola is well studied
because of its medical effect in anticancer treatment (Kishi
et al. 1991).
Phylogenetic study
Based on the SSU and ITS rDNA sequences analysis, its
pleosporalean status was verified, and Shiraia was suggested to be closely related to Leptosphaeriaceae and/or
Phaeosphaeriaceae (Pleosporineae) (Cheng et al. 2004).
Based on the molecular phylogenetic analysis, another
Shiraia-like fungus was reported which produced distinctive prawn-shaped conidioma-like structures (Morakotkarn
et al. 2008), and differed from conidiomata in the anamorph
of S. bambusicola described by Liu (2009).
Concluding remarks A relatively broad species concept of
Shiraia bambusicola is currently used, which could
comprise several species.
Sinodidymella J.Z. Yue & O.E. Erikss., Mycotaxon 24: 295
(1985). (Teichosporaceae)
Generic description
Habitat terrestrial, saprobic? Ascomata medium to large,
scattered, or in small groups, immersed, erumpent, to
superficial, globose, subglobose, coriaceous, apex flattened, with radial ridges arranged around the central
region. Peridium thick, 2-layered. Hamathecium of
dense, broadly trabeculate pseudoparaphyses, anastomosing and branching between the asci. Asci 8-spored, with a
short, furcate pedicel, bitunicate, cylindrical. Ascospores
broadly ellipsoid, hyaline, becoming pale brown when
mature, 1-septate, constricted at the median septum.
Anamorphs reported for genus: none.
Literature: Yue and Eriksson 1985.
Type species
Sinodidymella verrucosa (Petr.) J.Z. Yue & O.E. Erikss.,
Mycotaxon 24: 295 (1985). (Fig. 89)
≡ Amphididymella verrucosa Petr., Meddn Göteb. Bot.
17: 129 (1947).
Ascomata 620–930 μm high×800–1250 μm diam., scattered, or in small groups, immersed, becoming erumpent, to
nearly superficial, globose, subglobose, coriaceous, apex
flattened, with 3–6 radial ridges arranged around the central
region, with a flattened base not easily removed from the
substrate, wall black, roughened (Fig. 89a and b). Peridium
100–150 μm thick, thinner at the base, 2-layered, outer layer
thin, up to 40 μm thick, composed of small heavily pigmented
thick-walled cells of textura globulosa, cells up to 5 μm
diam., cell wall 3–6 μm thick, inner layer thick, composed of
Fungal Diversity
hyaline small cells of textura epidermoidea, 2–4 μm diam.,
cell wall 1–3 μm thick, interspersed with interwoven
mycelium in places (Fig. 89b and c). Hamathecium of dense,
broadly trabeculate pseudoparaphyses 1–2 μm broad, anastomosing between and above the asci (Fig. 89d). Asci 140–190
(−205)×12.5–15(−17.5) μm (x ¼ 164 14:3mm, n=10), 8spored, bitunicate, cylindrical, with a short, furcate pedicel,
20–45 μm long, and an inconspicuous ocular chamber (to
2 μm wide×1 μm high) (Fig. 89d and e). Ascospores 20–25×
10–12 μm (x ¼ 22:1 10:3mm, n=10), obliquely uniseriate
and partially overlapping, broadly ellipsoid with rounded
ends, hyaline, becoming pale brown when mature, 1-septate,
constricted at the median septum, smooth (Fig. 89f).
Anamorph: none reported.
Material examined: CHINA, Kansu Prov., between
Scharakuto and Kweite, on rotten stems of Salsola gemmascens Pall., 25 Jul. 1935, G. Fenzel 2400 (W 16366, type).
Notes
Morphology
Sinodidymella was formally established by Yue and
Eriksson (1985) as they noticed that Amphididymella
verrucosa Petr. was not congeneric with the generic type,
A. adeana Petr., which is a pyrenolichen. Thus a new
monotypic genus, Sinodidymella was introduced to accommodate it. The most outstanding morphological character of
Sinodidymella is its radial ridges, which are somewhat
comparable with that of Lophiostoma rugulosum Yin.
Zhang, J. Fourn. & K.D. Hyde, although their pseudoparaphyses are dissimilar. Lophiostoma rugulosum has “tightly
aggregated cellular pseudoparaphyses” and “apically ending
into bunches of clavate cells” (Zhang et al. 2009b).
Phylogenetic study
None.
Concluding remarks
The radial ridges have little phylogenetic significance in
genus level classification (Zhang et al. 2009b), but the
broadly trabeculate pseudoparaphyses of Sinodidymella
may fit Melanommataceae.
Splanchnonema Corda, in Sturm, Deutschl. Fl., 3 Abt.
(Pilze Deutschl.) 2(9), Tome 3: 115 (1829). (?Pleomassariaceae)
Generic description
Habitat terrestrial, saprobic. Ascomata medium to large,
solitary or scattered, immersed in cortex with a pseudostromal covering, with a small ostiole appearing on the host
surface, flattened subglobose. Peridium thin. Hamathecium
Fig. 87 Setomelanomma holmii (from UPS F-117969 (slide), isotype). b
a, b Asci with short pedicels in pseudoparaphyses. c Partial view of
ascus. d Branching and septate pseudoparaphyses. a Three-septate
lightly pigmented ascospores in ascus. Scale bars: a–e=10 μm
of dense, cellular pseudoparaphyses, embedded in mucilage, anastomosing and branching. Asci bitunicate, fissitunicate, clavate to broadly cylindrical, with a short,
narrowed, furcate pedicel. Ascospores clavate with a
rounded apex and acute base, reddish brown, constricted
at the septa.
Anamorphs reported for genus: Myxocyclus, Steganosporium (Barr 1982b).
Literature: Barr 1982b, 1993a; Boise 1985; Corda 1829;
Eriksson 1981; Ramaley and Barr 1995; Shoemaker and
LeClair 1975; Sivanesan 1984; Tanaka et al. 2005.
Type species
Splanchnonema pustulatum Corda, in Sturm, Deutschl. Fl.,
3 Abt. (Pilze Deutschl.) 2(9), Tome 3: 115 (1829). (Fig. 90)
Ascomata 400–600 μm high×550–1000 μm diam.,
solitary or scattered, immersed in cortex with a pseudostromal
covering, with a small ostiole appearing on the host surface,
flattened subglobose (Fig. 90a). Peridium 15–25 μm thick,
composed of small lightly pigmented thin-walled compressed
cells (Fig. 90b). Hamathecium of dense, long cellular
pseudoparaphyses 2–3 μm broad, embedded in mucilage,
anastomosing and branching. Asci 200–250 × 30–
45 μm (x ¼ 219:6 38:2mm, n =10), 8-spored, bitunicate,
fissitunicate, clavate to broadly cylindrical, with a short,
narrowed, furcate pedicel up to 35 μm long, without
conspicuous ocular chamber (Fig. 90c and d). Ascospores
45–53×20–24 μm (x ¼ 48:5 22:3mm, n=10), obliquely
uniseriate and partially overlapping to biseriate, clavate
with a rounded apex and acute base, reddish brown, 2septate, apical cell largest, broader than the lower cells,
basal cell smallest, constricted at the septa, smoothwalled, surrounded by a regular hyaline gelatinous
sheath, 3–6 μm thick (Fig. 90e and f).
Anamorph: none reported.
Material examined: UK, Avon, nr Bath, Batheaston, on
branch of Ulmus, C.E. Broome (L, No. 910.251-352, No.
910.251-371).
Notes
Morphology
A confusing outline of the history of Splanchnonema
was provided by Shoemaker and LeClair (1975), which at
the time was a valid, but little used name. Eriksson (1981)
and Sivanesan (1984) stated (without comment) that the
lectotype of Splanchnonema is S. pupula (Fr.) O. Kuntze.
However, S. pustulatum is listed as the generic type in the
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Fig. 88 Shiraia bambusium
(from IFRD 2040). a Ascostroma form a nubby structures
on the twigs of host. b Vertical
section of an ascostroma. Note
the reddish staining of the inner
tissue. c, d Cylindrical asci with
a short pedicel. e–g Muriform
fusoid hyaline ascospores. Scale
bars: a=1 cm, b=1 mm, c, d=
50 μm, e–g=20 μm
online databases MycoBank and Index Fungorum. We
assume Eriksson (1981) gained his data from Shoemaker
and LeClair (1973), who considered S. pustulatum to be a
synonym of S. pupula. Since we were unable to locate
material of Corda or Fries we used a later collection of C.E.
Broome.
Splanchnonema can be distinguished from the morphologically comparable genera, i.e. Pleomassaria or
Splanchospora by its depressed ascomata, and obovoid
and asymmetrical ascospores (Barr 1982b). Currently,
about 40 species are included in this genus. Barr
(1993a) provided a key to 27 North American species,
however, the inclusion of species with a range of
ascospore types and immersed to superficial ascomata
suggests the genus to be polyphyletic. Tanaka et al.
(2005) suspected that the genus might include species of
Pleomassaria, thus this genus needs further study.
Phylogenetic study
Splanchnonema platani (= Massaria platani) is poorly
supported to be related to Lentitheciaceae (Schoch et al.
2009).
Fig. 89 Sinodidymella verrucosa (from W 16366, type). a Ascomata
on the host surface. Note the radial ridges around the pseudostiolar
region. b Section of an ascoma. c Section of peridium. Note the
hyaline small cells and interwoven hyphae. d Cylindrical asci in
pseudoparaphyses. e Eight-spored ascus with short pedicel. f Hyaline,
1-septate ascospores which turn pale brown when mature. Scale bars:
a=1 mm, b=100 μm, c=50 μm, d–f=20 μm
b
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Concluding remarks
Splanchnonema pustulatum has unique ascospores formed
in immersed ascomata with thin walls, indicating that Splanchnonema sensu stricto should be confined to a few similar
species. The type needs recollecting, sequencing and epitypifying in order to establish the phylogenetic relationships of
this genus and to study what may be important defining
characters. Also see entry under Pleomassaria.
Sporormia De Not., Micromyc. Ital. Novi 5: 10 (1845).
(Sporormiaceae)
Fig. 90 Splanchnonema pustulatum (from L, No. 910.251–352, No.
910.251–371). a Appearnce of ascomata on the host surface beneath a
slightly raised area with minute ostiolar opening. b Section of the
partial peridium. Note the compressed cells. c Dehiscent ascus. d
Generic description
Habitat terrestrial, saprobic (coprophilous). Ascomata small,
solitary, scattered, immersed to erumpent, globose, subglobose, wall black; apex without obvious papilla, ostiolate.
Peridium thin. Hamathecium of rare, broad, septate pseudoparaphyses. Asci 8-spored, bitunicate, fissitunicate dehiscence not observed, short cylindrical, with a short, narrowed,
furcate pedicel. Ascospores fasciculate, broadly filliform,
reddish brown, multi-septate, easily separating into partspores, without visible germ-slits or pores.
Cluster of three asci joined in hymenium and pseudoparaphyses. e, f
Asymmetric ascospores. Note the conspicuous sheath. Scale bars: a=
1 mm, b–d=50 μm, e, f=20 μm
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Anamorphs reported for genus: none.
Literature: Ahmed and Asad 1968; Ahmed and Cain 1972;
Kirschstein 1944; de Notaris 1849.
Type species
Sporormia fimetaria De Not., Micromyc. Ital. Novi 5: 10
(1845). (Fig. 91)
Ascomata 100–150 μm diam., solitary, scattered,
immersed to erumpent, globose, subglobose, wall
black; apex without obvious papilla, ostiolate
(Fig. 91a). Peridium thin (other characters unknown).
Hamathecium of rare, 2–3 μm wide, septate pseudoparaphyses. Asci 70–100 × 13–18 μm (x ¼ 86:4 14:9mm,
n = 10), 8-spored, bitunicate, fissitunicate dehiscence not
observed, shortly cylindrical, with a short, narrowed,
furcate pedicel up to 20 μm long, no apical apparatus
could be observed (Fig. 91b, c and d). Ascospores 50–
58 × 4–5 μm (x ¼ 54:7 4:8mm, n = 10), fasciculate,
broadly filliform, reddish brown, with 16 cells, easily
separating into partspores, central cells of the ascospores
shorter than broad, rectangular in vertical section, round
in transverse section, 4–5 × 2.5–3.5 μm, without visible
germ-slits or pores, apical cells usually longer than
broad, 5–6.5 μm long, also without apertures (sheath is
reported (Ahmed and Cain 1972), but not observed in
this study) (Fig. 91e).
Anamorph: none reported.
Material examined: 1832, (RO, type, as Hormospora
fimetaria De Not.).
Sporormia with other genera of Sporormiaceae is unclear
and not resolved yet.
Concluding remarks
Several coprophilous taxa (e.g. Chaetopreussia and
Pleophragmia as well as Sporormiella nigropurpurea) in
the Pleosporales were not included in the study by Kruys
and Wedin (2009). Strains of these genera need to be
collected and analyzed and their relationship with Sporormia established.
Trematosphaeria Fuckel, Jb. nassau. Ver. Naturk. 23–24:
161 (1870). (Trematosphaeriaceae)
Generic description
Habitat terrestrial or freshwater, saprobic. Ascomata
subglobose, unilocular, erumpent to superficial, with
papillate ostiole. Peridium thin, comprising several cell
types. Hamathecium of dense, delicate, filliform, septate
pseudoparaphyses. Asci bitunicate, fissitunicate,
cylindro-clavate, normally 8-spored. Ascospores
ellipsoid-fusoid to biconic, septate, smooth to finely
verruculose, brown.
Anamorphs reported for genus: hyphopodia-like (Zhang
et al. 2008a).
Literature: von Arx and Müller 1975; Barr 1979a;
Boise 1985; Clements and Shear 1931; Zhang et al.
2008a.
Type species
Notes
Morphology
Sporormia was formally established by de Notaris (1849),
and only one species was described, i.e. S. fimetaria, which
subsequently was selected as the generic type. Sporormia
sensu stricto was accepted by several workers, and only
includes members with a fasciculate ascospore arrangement,
parallel to the ascus, and the part cells of the ascospores
lacking germ-slits (Ahmed and Asad 1968; Ahmed and Cain
1972; Kirschstein 1944). Species whose ascospores are not
fasciculate and have partspores with germ-slits were assigned
to Sporormiopsis by Kirschstein (1944) and to Sporormiella
by Ahmed and Cain (1972).
Phylogenetic study
The generic status of Sporormia in Pleosporales was
verified based on a phylogenetic analysis of ITS-nLSU
rDNA, mtSSU rDNA and ß-tubulin sequences (Kruys and
Wedin 2009). Sporormia clustered together with species of
Westerdykella (including Eremodothis and Pycnidiophora),
but lacks clear statistical support. Thus, the relationship of
Trematosphaeria pertusa (Pers.) Fuckel, Jb. nassau. Ver.
Naturk. 23–24: 161 (1870). (Fig. 92)
≡ Sphaeria pertusa Pers., Syn. meth. fung. (Göttingen)
1: 83 (1801).
Ascomata 350–550 μm high × 320–480 μm diam.,
solitary, scattered, or in groups, initially immersed, becoming erumpent, to semi-immersed, subglobose, black; apex
with a short ostiole usually slightly conical and widely
porate, to 100 μm high (Fig. 92a and b). Peridium 48–
55 μm wide laterally, to 80 μm at the apex, thinner at the
base, 30–40 μm thick, coriaceous, 3-layered, comprising
several cell types, one is of small heavily pigmented thickwalled cells of textura angularis, cells 4–8 μm diam., cell
wall 1.5–3 μm thick in places with columns of textura
prismatica orientated perpendicular to the ascomatal surface, apex cells smaller and walls thicker, forming thickwalled cells of textura pseudoparenchymata, and larger,
paler cells of mixture of textura epidermoidea and textura
angularis at the base (Fig. 92b, c and h). Hamathecium of
dense, filamentous, 1.5–2.5 μm broad, septate pseudoparaphyses, embedded in mucilage, branching and anastomos-
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ing between and above the asci (Fig. 92d, e and f). Asci
100–145×15–17 μm (x ¼ 118 15:5mm, n=10), 8-spored,
bitunicate, fissitunicate, cylindro-clavate, with a short,
thick, furcate pedicel which is 12–30 μm long, with a
truncate ocular chamber (Fig. 92d, e, f, g and i). Ascospores
27.5–32.5×7.5–8.5 μm (x ¼ 29:5 8mm, n=10), biseriate
to uniseriate near the base, fusoid with broadly to narrowly
rounded ends, dark brown, 1–3-septate, secondary septum
forming late or often absent, constricted at the median
septum, the upper cell often shorter and broader than the
lower one, smooth to finely verruculose, containing
refractive globules (Fig. 92j and k).
Anamorph: Only hyphopodia-like structures (or conidia?)
observed (Zhang et al. 2008a).
Colonies (of epitype) reaching 5 cm diam. after 20 days
growth on MEA at 25°C, raised, woolly, deep grey, with
irregular to rhizoidal margin, reverse darkened. Hyphopodialike structures (or conidia?) produced after 6 months, hyaline
to pale brown, lobed, 4–4.5(−5) μm long and 3–3.5 μm diam.
Material examined: EUROPE, Upsala, on decaying
wood, designated by Boise (1985), (L-Pers 910269–172,
as Sphaeria pertusa Pers., neotype); FRANCE, Deux
Sèvres, Sansais, Le Vanneau, Les Grandes Mottines,
swamp, on bark of a dead stump of Fraxinus excelsior,
25 Apr. 2004, J. Fournier (IFRD 2002, epitype); Haute
Garonne, Avignonet, Canal du Midi, on submerged wood
of Platanus in a canal, 23 Nov. 2006, Michel Delpont, det.
J. Fournier (IFRD2003).
Notes
Morphology
Trematosphaeria was formally established in ‘Rhenish
fungi’ by Fuckel (1870) based on the broadly pertuse
ascomata, and Fries (1823) assigned it under Ascomycetes,
Pyrenomycetes, Lophiostomataceae. Subsequently, Winter
(1885) placed Trematosphaeria in Amphisphaeriaceae.
Berlese (1890), however, treated Trematosphaeria as a
synonym of Melanomma (Melanommataceae). After establishment of Loculoascomycetes (Luttrell 1955), Trematosphaeria was assigned to Pleosporaceae (Loculoascomycetes,
Pleosporales) (Holm 1957), and this was followed by von
Arx and Müller (1975). Trematosphaeria was assigned to
Melanommataceae by Barr (1979a), and this has been
widely followed (Eriksson 2006; Kirk et al. 2001; Lumbsch
and Huhndorf 2007).
Trematosphaeria pertusa, the lectotype species of Trematosphaeria (Clements and Shear 1931), is characterized by
having semi-immersed to erumpent ascomata, filamentous
pseudoparaphyses, cylindro-clavate asci, fusoid, 1-septate
reddish brown to dark brown ascospores (Zhang et al.
2008a). All of these characters are quite different from those
of Melanomma, the familial type of Melanommataceae.
Fig. 91 Sporormia fimetaria (from RO, type). a Appearance of
ascomata on the host surface. Note the scattered distribution. b–d
Broad cylindrical asci with a short and thick pedicel. e Released
filiform ascospores which may break up into part spores. Scale bars:
a=0.5 mm, b–d=20 μm, e=10 μm
Phylogenetic study
Trematosphaeria pertusa forms a robust phylogenetic
clade with Falciformispora lignatilis and Halomassarina
thalassiae, and they are all assigned to Trematosphaeriaceae (Suetrong et al. 2009; Zhang et al. 2009a; Plate 1).
Concluding remarks
Trematosphaeria pertusa is a terrestrial species which
can also survive in a freshwater environment. However,
both Falciformispora lignatilis and Halomassarina thalassiae are marine fungi. Their habitat difference may indicate
their distant relationship, at least above genus level.
Verruculina Kohlm. & Volkm.-Kohlm., Mycol. Res. 94:
689 (1990). (Testudinaceae)
Generic description
Habitat marine, saprobic. Ascomata medium-sized, solitary
under clypeate, immersed to semi-immersed, subglobose to
depressed ellipsoidal, papillate, ostiolate, periphysate,
black, carbonaceous. Peridium thin, comprising a few
layers of cells of textura angularis. Hamathecium of long
cellular pseudoparaphyses, embedded in mucilage, hyaline,
septate and sparsely branching. Asci 8-spored, bitunicate,
fissitunicate, cylindrical, with short pedicels, ocular chamber not observed. Ascospores biseriate, ovoid or ellipsoidal,
dark brown, 1-septate, constricted at the septum, verrucose
or verruculose, with or without germ pore.
Anamorphs reported for genus: none.
Literature: Kohlmeyer and Volkmann-Kohlmeyer 1990;
Suetrong et al. 2009.
Type species
Verruculina enalia (Kohlm.) Kohlm. & Volkm.-Kohlm.,
Mycol. Res. 94: 689 (1990). (Fig. 93)
≡ Didymosphaeria enalia Kohlm., Ber. dt. bot. Ges. 79:
28 (1966).
Ascomata 295–480 μm high×140–520 μm diam., solitary
under clypeate, immersed to semi-immersed, subglobose to
depressed ellipsoidal, ostiolate, papillate, periphysate, black,
carbonaceous. Peridium thin, comprising a few layers of cells
of textura angularis. Hamathecium of long cellular pseudoparaphyses, 1.5–2 μm broad, embedded in mucilage, hyaline,
septate and sparsely branching. Asci 177–135×12.5–15.5 μm,
8-spored, bitunicate, fissitunicate, cylindrical, with short
furcate pedicels, ocular chamber not observed (Fig. 93a).
b
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Ascospores 16.5–23×7.5–10 μm, biseriate, ovoid or ellipsoidal, dark brown, 1-septate, constricted at the septum, verrucose
or verruculose, with or without germ pore (Fig. 93b).
Anamorph: none reported.
Material examined: SEYCHELLES, Victoria, on submerged branch of Rhizophora mangle L., Mar. 2004, K.D.
Hyde (KDH 2137, slide).
Fig. 92 Trematosphaeria pertusa (a, d, f–i from epitype, b, c, e, j
from neotype). a Ascomata on the host surface. b Section of an ascoma.
c, h Section of the peridium. c shows the peridium structure at sides,
and h indicates the basal peridium structure. Note the hyaline and thinwalled cells in (h). d Asci amongst pseudoparaphyses. e Ascus with
pedicle. f, g Dehiscent ascus. i Upper part of the ascus, showing the
ocular chamber and the mucilage covering the apex. j, k Ascospores.
Scale bars: a=0.5 mm, b, c=100 μm, d–h=20 μm, i–k=10 μm
Notes
Type species
Morphology
Verruculina was introduced to accommodate an obligate marine species, i.e. Verruculina enalia (Kohlmeyer
and Volkmann-Kohlmeyer 1990). Verruculina is characterized by immersed, clypeate, carbonaceous, ostiolate and
papillate ascomata. The peridium is composed of cells of
textura angularis. Pseudoparaphyses are trabeculate and
embedded in mucilage. Asci are 8-spored, cylindrical with
short pedicels and ocular chamber, and ascospores are
ellipsoidal, 1-septate, dark brown, verrucose or verruculose. The partly or completely immersed clypeate ascomata of V. enalia is comparable with those of
Didymosphaeria futilis, but it differs from the later by
the dark peridium, gelatinous matrix around the pseudoparaphyses, stipitate asci with an ocular chamber, and the
verruculose ascospores (Kohlmeyer and VolkmannKohlmeyer 1990).
Westerdykella ornata Stolk, Trans. Br. Mycol. Soc. 38: 422
(1955). (Fig. 94)
Ascomata 100–300 μm diam., cleistothecoid, scattered
on the upper layer of the culture medium, wall black
(Fig. 94a). Peridium composed of one layer of cells of
polygonal in front view, dark brown, thick-walled cells,
ca. 5 μm diam. Hamathecium not observed. Asci 25–32×
16–22 μm, 32-spored, bitunicate nature undetermined,
fissitunicate dehiscence not observed, subglobose to
ellipsoid, arranged in the centre of the ascomata, with
a short furcate pedicel best seen in immature asci
(Fig. 94b, c, d and f). Ascospores 6.2–7× 6–6.8 μm,
globose, brown, 1-celled, ornamented with irregular spiral
bands, which occur in four to five coils, without germ
pore (Fig. 94e).
Anamorph: none reported.
On MEA colonies spreading, but somewhat erumpent,
with moderate aerial mycelium and even, lobate margins;
surface dirty white with luteous to orange patches; reverse
orange to sienna. On PDA similar but with sparse aerial
mycelium; surface with patches of orange to luteous and
dirty white; reverse luteous with cream margins. On OA
flat, spreading with sparse aerial mycelium; surface with
luteous and dirty white patches and transparent margins;
sporulating on OA, visible as black masses of aggregated
ascomata; colonies reaching 4 cm diam. on all media (based
on CBS 379.55).
Material examined: MOZAMBIQUE, Inhaca, leg. H.J.
Swart, mangrove mud (CBS 379.55, holotype).
Phylogenetic study
Based on multigene phylogenetic analysis, Verruculina
enalia nested within Testudinaceae (Suetrong et al. 2009).
Thus, its familial placement seems clarified.
Concluding remarks
None.
Westerdykella Stolk, Trans. Br. Mycol. Soc. 38: 422
(1955). (Sporormiaceae)
Generic description
Notes
Habitat terrestrial, saprobic (coprophilous). Ascomata
small, scattered on the upper layer of the culture medium,
wall black. Peridium thin, composed of one layer of cells of
polygonal, dark brown, thick-walled cells. Hamathecium
not observed. Asci 32-spored, bitunicate nature undetermined, fissitunicate dehiscence not observed, subglobose to
ellipsoid, arranged in the centre of the ascomata, with or
without a short pedicel. Ascospores globose, brown, 1-celled,
without germ pore.
Anamorphs reported for genus: Phoma-like (von Arx 1974).
Literature: von Arx 1973, 1981; Kruys et al. 2006; Kruys
and Wedin 2009; Stolk 1955a.
Morphology
Westerdykella was introduced to accommodate a
coprophilous fungus, which is characterized by cleistothecioid and membraneous ascomata (Stolk 1955a). Asci
are subglobose to ellipsoid, stalked, many-spored and
evanescent. Ascospores are globose to subglobose,
brown, ornamented with spiral bands, without germ pores
(Stolk 1955a). Westerdykella was assigned under Phaeosporeae of the Eurotiaceae (Stolk 1955a), and was
assigned to Sporormiaceae by von Arx and Müller
(1975). Based on the spore ornamentation, von Arx and
b
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Fig. 93 Verruculina enalia (from KDH 2137, slide). a Cylindrical asci with short pedicels. b One-septate verruculose ascospores. Scale bars:
a=20 μm, b=10 μm
van der Aa (1987) and Barr (2000) accepted Westerdykella as
a separate genus, but this is not supported by molecular
phylogenetic analysis (Kruys and Wedin 2009).
Phylogenetic study
Phylogenetic reconstructions indicated that both Pycnidiophora and Eremodothis should be treated as synonyms
of Westerdykella (Kruys and Wedin 2009).
Concluding remarks
Westerdykella is another example where ascospore ornamentation can be phylogenetically uninformative. Westerdykella
is proved a good genus of Sporormiaceae (Kruys and Wedin
2009).
Wettsteinina Höhn., Sber. Akad. Wiss. Wien, Math.-naturw.
Kl., Abt. I 116: 126 (1907). (?Lentitheciaceae)
Generic description
Habitat terrestrial or freshwater? hemibiotrophic or
saprobic. Ascomata generally small, scattered, immersed
with a protruding broad papilla. Peridium very thin,
composed of few layers of thin-walled large polygonal
cells in surface view. Hamathecium deliquescing at
maturity. Asci bitunicate, fissitunicate, subglobose to
obpyriform, without a pedicel, with small truncate ocular
chamber. Ascospores hyaline and turning pale brown
when mature, septate, upper second cell enlarged, slightly
constricted at the second septum, smooth, surrounded by a
hyaline gelatinous sheath.
Anamorph reported for genus: Stagonospora (Farr et al.
1989).
Literature: Barr 1972; Müller 1950; Shoemaker and
Babcock 1987, 1989b.
Type species
Wettsteinina gigaspora Höhn., Sber. Akad. Wiss. Wien,
Math.-naturw. Kl., Abt. 1 116: 126 (1907). (Fig. 95)
Ascomata 150–250 μm diam., scattered, immersed
with protruding broad papillae, 50–90 μm diam.
Peridium thin, composed of few layers of thin-walled
large polygonal cells in surface view, 6–15 μm diam.
(Fig. 95a). Hamathecium deliquescing at maturity. Asci
140–200×75–120 μm, 8-spored, bitunicate, fissitunicate,
subglobose to obpyriform, lacking a pedicel, with a small
truncate ocular chamber (to 8 μm wide×5 μm high)
(Fig. 95b and c). Ascospores 90–110×25–30 μm, 2–4seriate, hyaline and turning pale brown when mature,
broadly clavate, 4-septate, primary septum distinct and
constricted forming 1/3rd from the apex of the ascospore,
complete, secondary septa less distinct and slightly
constricted, incomplete, with one forming above and two
forming below the primary septum, largest cell the second
cell from apex, smooth, surrounded by a hyaline gelatinous sheath 5–8 μm thick (Fig. 95d and e).
Anamorph: none reported.
Material examined: SLOVENIA, Postojna, on Genista
sagittalis leg. Stapf. det. H. Rehm. (S, holotype of
Massarina gigantospora).
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Fig. 94 Westerdykella ornata (from CBS 379.55, holotype). a Appearance of the ascomata on culture substrate surface. b–f Mature and
immature asci as well as the released ascospores. Note the spiral bands around the ascospores. Scale bars: a=1 mm, b–f=10 μm
Notes
Morphology
Confusion exists in the generic type of Wettsteinina.
Höhnel (1907) described W. gigaspora when introducing
Wettsteinina, and listed it as the first species of Wettsteinina. Clements and Shear (1931) accepted W. gigaspora as
the generic type of Wettsteinina, which is followed by
Shoemaker and Babcock (1987). But according to http://
www.indexfungorum.org (June 2011), W. gigantospora is
the generic type of Wettsteinina. Both W. gigantospora
and W. gigaspora were treated as the synonyms of W.
mirabilis (Niessl) Höhn. http://www.indexfungorum.org
(June, 2011, Synonymy Contributor: CBS (2010)). We
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tentatively described the generic type of W. gigantospora
as a representing of the type of W. gigaspora here.
New family names, i.e. Pseudosphaeriaceae and Wettsteininaceae (as Wettsteiniaceae) and a new order, Pseudosphaeriales had been introduced to accommodate Wettsteinina and
its synonym Pseudosphaeria (Höhnel 1907; Locquin 1972).
After a systematic study, Wettsteinina was included in
Pleosporaceae based on its “Pleospora-type” centrum, and
Pseudosphaeriaceae and Wettsteininaceae are treated as
synonyms of Pleosporaceae (Shoemaker and Babcock 1987).
Phylogenetic study
Wettsteinina macrotheca (Rostr.) E. Müll., W. pachyasca
(Niessl) Petr. and W. dryadis (Rostr.) Petr. were reported to be
closely related to Pleomassaria siparia (Melanommataceae)
(Kodsueb et al. 2006a), and W. lacustris (Fuckel) Shoemaker
& C.E. Babc. nested within Lentitheciaceae (Schoch et al.
2009). The generic type has not been sequenced.
Concluding remarks
The most striking character for Wettsteinina is its
asymmetrical ascospores, thick-walled obpyriform asci
and lack of pseudoparaphyses at maturity. These
characters are comparable with genera in the Capnodiales and Venturiales. The phylogenetic significance of
these characters are not fully understood, while the
hemibiotrophic or saprobic life style may indicate its
polyphyletic nature (Shoemaker and Babcock 1987).
Strains from the genus, in particular the generic type
require DNA sequence data so that the phylogenetic
placement can be investigated.
Wilmia Dianese, Inácio & Dorn. -Silva, Mycologia 93:
1014 (2001). (Phaeosphaeriaceae)
Generic description
Habitat terrestrial, hemibiotrophic or biotrophic. Ascomata
small, scattered, immersed, globose to subglobose, papillate.
Peridium thin, composed of a few layers of brown, thickwalled cells of textura angularis to prismatica. Hamathecium
comprising filliform, septate, rarely branching, evanescent,
cellular pseudoparaphyses embedded in mucilage. Asci
bitunicate, fissitunicate, cylindrical to clavate, with a short,
furcate pedicel and ocular chamber. Ascospores fusoid, pale
brown, 1-septate.
Anamorphs reported for genus: see below.
Literature: Dianese et al. 2001.
Ascomata 175–240 μm high×95–145 μm diam., scattered, immersed, globose to subglobose; apex with a short
papilla, 40–80 μm long, ostiolate, periphysate, periphyses
up to 90 μm long (Fig. 96a and b). Peridium 6–15 μm
wide, 1-layered, composed of 3–7 layers of brown, thickwalled cells of textura angularis to prismatica, cells 4–
9 μm diam., cell wall 2–4 μm thick (Fig. 96a and b).
Hamathecium of long cellular pseudoparaphyses 2–3 μm
broad, septate, rarely branching, embedded in mucilage,
evanescent. Asci 65–95×9.5–14 μm (x ¼ 78:5 11:5mm,
n=10), 8-spored, bitunicate, fissitunicate, cylindrical to
clavate, with a short, furcate pedicel and a small ocular
chamber (Fig. 96c, d and f). Ascospores 22.5–28×5–
8.5 μm (x ¼ 26:5 6:8mm, n=10), biseriate, fusoid with
narrowly rounded ends, pale brown, 1-septate, constricted
at the septum, the upper cell often shorter and broader than
the lower one, smooth, with or without sheath (Fig. 96d and
e).
Anamorph: Conidiomata 170–200 μm high×85–130 μm
diam., eustromatic, immersed, subglobose to irregular,
ostiolate, brown. Peridium thin, 1–2 wall layers, 6–8 μm
thick, thicker near the apex. Ostiole 50–63 μm high×30–35
broad. Conidiogenous cells ampulliform or lageniform,
phialidic, aseptate. Conidia 13–20×4–7 μm, ellipsoid,
oblong, ovoid, hyaline (Dianese et al. 2001).
Material examined: BRAZIL, Distrito Federal, Vargem
Bonita, Fazenda Agua Limpa, on leaves of Memora
pedunculata (Vell.) Miers, 18 May 1995, Carlos A. Inácio
(UB Col. Microl 8438 holotype).
Notes
Morphology
Wilmia was formally established by Dianese et al. (2001)
as a monotypic genus represented by W. brasiliensis, which
causes leaf spots on Memora pedunculata. The peridium of
W. brasiliensis comprises a few layers of brown, thickwalled textura angularis to prismatica cells, and it also has
cellular pseudoparaphyses, clavate asci, 1-septate pale
brown ascospores (Dianese et al. 2001).
Phylogenetic study
None.
Concluding remarks
The dicotyledonous host habit of Wilmia brasiliensis seems
in agreement with Leptosphaeriaceae rather than Phaeosphaeriaceae. But a verified conclusion can only be reached by
further molecular phylogenetic study.
Type species
Wilmia brasiliensis Dianese, Inácio & Dorn.-Silva, Mycologia 93: 1014 (2001). (Fig. 96)
Xenolophium Syd., Bulletin of the Bernice P. Bishop
Museum, Honolulu, Hawaii 19: 96 (1925). (Pleosporales,
genera incertae sedis)
Fungal Diversity
Fig. 95 Wettsteinina gigantospora (from S, holotype of
Massarina gigantospora). a
Ascomata with protruding papilla scattered on the host surface. b Obpyriform thick-walled
ascus with small apical apparatus. c Fissitunicate ascus. d
Released hyaline ascospores.
Note the distinct primary septum
and less distinct secondary septa. e Ascospore with sheath.
Scale bars: a=0.5 mm, b–d=
100 μm, e=50 μm
Generic description
Habitat terrestrial, saprobic on wood. Ascomata nearly
superficial, scattered to gregarious, globose, large, with a
conspicuous compressed papilla and large slit-like ostiole.
Peridium carbonaceous. Hamathecium of dense, long
trabeculate pseudoparaphyses, branching and anastomosing
between and among asci. Asci 8-spored, clavate, with very
Fungal Diversity
long furcate pedicels. Ascospores fusoid to narrowly fusoid,
light to dark brown, 1-septate, constricted at the septum.
Anamorphs reported for genus: none.
Literature: Chesters and Bell 1970; Huhndorf 1993;
Mugambi and Huhndorf 2009b; Müller and von Arx
1962; Stevens 1925.
Type species
Xenolophium leve Syd., Bulletin of the Bernice P. Bishop
Museum, Honolulu, Hawaii 19: 97 (1925) (Fig. 97)
Current name: Xenolophium applanatum (Petch)
Huhndorf, Mycologia 85: 493 (1993).
≡ Schizostoma applanatum Petch, Ann. Roy. Bot. Gard.
(Peradeniya) 6: 231 (1916).
Ascomata 1–1.5 mm diam., scattered to clustered, erumpent
to superficial, globose with base immersed in host tissue, wall
black, carbonaceous, roughened with ridges, papillate. Apex
with a conspicuous hysteriform papilla extending on the sides,
1–1.4 mm long, 0.4–0.5 mm wide, 0.2–0.3 mm high, smooth,
ostiole slit-like, nearly as long as papilla length (Fig. 97a).
Peridium 140–160 μm thick, pseudoparenchymatous, composed of two distinct layers: outer crust 16–45 μm thick,
blackish, of heavily melanized, nearly opaque thick-walled
angular cells, of uneven thickness forming irregular strands
extending into the inner layer; inner layer subhyaline,
composed of thick-walled prismatic to angular cells, with
columns or patches of darker thick-walled cells extending
inwardly from the outer layer; papilla wall 200–220 μm thick,
of heavily melanized angular thick-walled cells (Fig. 97b and
c). Hamathecium of dense, very long trabeculate pseudoparaphyses 0.8–1.5 μm broad, embedded in mucilage, anastomosing and branching between and above the asci. Asci 104–152×
9–12 μm (excluding pedicel) (x ¼ 149 10:2mm, n=10), 8spored, bitunicate, fissitunicate dehiscence not observed,
clavate, with a long, narrowed, furcate pedicel which is 50–
75 μm long (Fig. 97d). Ascospores 17–26×4–5.5 μm
(x ¼ 22:5 4:8mm, n=10), upper biseriate and lower uniseriate,
fusoid, straight to slightly curved, equally 1-septate, constricted
at the septum, the upper cell slightly wider, with one or rarely
two additional septa appearing on a small number of senescent
ascospores, pale brown, median septum darker, constricted,
smooth, without sheath or appendages (Fig. 97e, f and g).
Anamorph: none reported.
Material examined: MARTINIQUE, Morne Rouge, on
rotten wood, leg C. Lécuru, det Jacques Fournier, 29 Aug.
2007, IFRD 2038.
X. verrucosum, of which X. leve is selected as the generic type
(Huhndorf 1993). Because of its morphological similarity with
some genera, such as Ostropella and Schizostoma, Xenolophium has been treated as a synonym of Ostropella (Müller
and von Arx 1962) or even of Lophiostoma (Chesters and Bell
1970). Huhndorf (1993) clarified the circumscription of
Xenolophium and treated X. leve as a synonym of Schizostoma
applanata. Xenolophium mainly differs from Ostropella in
lack of “organized cell composition and triangular pattern of
melanization” in the peridium (Huhndorf 1993).
Phylogenetic study
The polyphyletic nature of Xenolophium has been
demonstrated (Mugambi and Huhndorf 2009b). The generic
type of Xenolophium (X. leve, current name X. applanatum)
clustered together with Ostropella albocincta (generic type
of Ostropella), and both locate in Platystomaceae
(Mugambi and Huhndorf 2009b).
Concluding remarks
The large ascomata with slit-like ostioles, hamathecium of
numerous and trabeculate pseudoparaphyses, clavate asci with
long pedicels, and the pale brown, 1-septate ascospores of
Xenolophium leve are all comparable with those of Ostropella
albocincta. However, the phylogenetic results do not support
them being congeneric (Mugambi and Huhndorf 2009b).
Synonyms
Javaria Boise, J.R., Acta Amazonica 14(Supl.): 50 (1984).
(Melanommataceae)
Current name: Astrosphaeriella Syd. & P. Syd., Annls
mycol. 11: 260 (1913).
Generic description
Habitat terrestrial, saprobic. Ascomata medium-sized, scattered, erumpent to nearly superficial, reflexed pieces of the
ruptured host tissue usually persisting around the surface of
the ascomata; ascomata broadly conical, with a flattened
base not easily removed from the substrate, wall black,
papillate. Peridium carbonaceous. Hamathecium of trabeculate pseudoparaphyses. Asci 8-spored, bitunicate, fissitunicate, cylindro-clavate to narrowly fusoid, with a short,
narrowed, furcate pedicel. Ascospores elongate-fusoid,
hyaline, 1-septate, constricted at the septum.
Anamorphs reported for genus: none.
Literature: Barr 1990a; Boise 1984.
Notes
Type species
Morphology
Xenolophium was formally established by Sydow (in
Stevens 1925) to accommodate two species, i.e. X. leve and
Javaria samuelsii Boise, J.R., Acta Amazonica 14(Supl.):
50 (1984) (Fig. 98)
Fungal Diversity
Fig. 96 Wilmia brasiliensis (from UB Col. Microl 8438, holotype). a
Section of an ascoma. Note the setae in the ostiole. b Conidioma of
the coelomycetous anamorphic stage. c, d Clavate asci with short
furcate pedicels. e, f Released 1-septate pale brown ascospores. Scale
bars: a, b=100 μm, c, d=20 μm, e, f=10 μm
Fungal Diversity
Fig. 97 Xenolophium applanatum (from IFRD 2038). a Gregarious
ascomata on the host surface. Note protruding papilla and slit-like
ostiole. b Vertical section of the papilla and ostiole. c Section of the
partial peridium. Note the two layers of the peridium. d Eight-spored
asci in trabeculate pseudoparaphyses. Note the long pedicels. e–g Pale
brown ascospores. Scale bars: a=2 mm, b=200 μm, c=50 μm, d=
20 μm, e–g=10 μm
Fungal Diversity
Current name: Astrosphaeriella samuelsii Boise, Acta
Amazon., Supl. 14(1–2, Suppl.): 50 (1986) [1984].
Ascomata 300–380 μm diam., scattered, erumpent
through the outer layers of the host tissues, to nearly
superficial, reflexed pieces of the ruptured host tissue
usually persisting around the surface of the ascomata;
ascomata broadly conical, with a flattened base not easily
removed from the substrate, wall black, papillate (Fig. 98a).
Peridium 50–80 μm thick, carbonaceous and crisp, 1layered. Hamathecium of dense, long trabeculate pseudoparaphyses, 0.8–1.5 μm broad, embedded in mucilage,
anastomosing between and above the asci. Asci 140–185×
17.5–20 μm (x ¼ 158 19:4mm, n=10), 8-spored, bitunicate, fissitunicate, cylindro-clavate to narrowly fusoid, with
a short, narrowed, furcate pedicel up to 20 μm long
(Fig. 98b and c). Ascospores 48–55(−60)×6–7.5(−10) μm
(x ¼ 52:2 7:7mm, n= 10), biseriate, elongate- fusoid,
gradually tapering towards the ends, hyaline, surrounded
with sheath, 2–5 μm thick, 1-septate, constricted at the
septum (Fig. 98d).
Anamorph: none reported.
Material examined: Serra Araca, 60 m, terra firme,
open forest, deep litter. Dry. 10–13 Mar. 1984, det. Jean R.
Boise, G.J. Samuels (isotype).
Notes
Morphology
Javaria was introduced by Boise (1984) based on
seven Amazonian collections on decaying palm petioles; it
is comparable with Astrosphaeriella in numerous characters. But Javaria differs from Astrosphaeriella by its
hyaline ascospores with sheath, and its apical ring can be
stained with Congo Red, as well as its small ascomata.
Barr (1990a) introduced a second species J. shimekii
which occurs on woody substrate. Some mycologists treat
Javaria as a synonym of Astrosphaeriella (Hyde and
Fröhlich 1998).
Phylogenetic study
None.
Concluding remarks
The size of ascomata and pigmentation of ascospores has
little significance at generic level classification (Zhang et al.
2009a). Likewise, the staining of endotunica with Congo
Red has not been shown to have great significance. Thus,
we accept Javaria as a synonym of Astrosphaeriella.
Pycnidiophora Clum, Mycologia 47: 899 (1955).
(Sporormiaceae)
Current name: Westerdykella Stolk, Trans. Br. Mycol.
Soc. 38(4): 422 (1955).
Generic description
Habitat terrestrial, saprobic (coprophilous). Ascomata
small, cleistothecial, scattered on surface of agar media,
semi-immersed, globose to subglobose, black. Peridium
thin, composed of thin-walled, polyangular cells from front
view. Hamathecium not apparent. Asci numerous, irregularly arranged, bitunicate nature undetermined, fissitunicate
nature undetermined, globose, without pedicel. Ascospores
gathering in the globose asci, smooth.
Anamorphs reported for genus: Phoma-like.
Literature: Cain 1961; Clum 1955; Stolk 1955b; Thompson
and Backus 1966.
Type species
Pycnidiophora dispersa Clum, Mycologia 47: 900 (1955)
[1955]. (Fig. 99)
Current name: Westerdykella dispersa (Clum) Cejp &
Milko.
Ascomata 200–290 μm diam., cleistothecial, scattered
on surface of agar media, semi-immersed, globose to
subglobose, black (Fig. 99a). Peridium thin, composed of
thin-walled, poly-angular cells from front view (Fig. 99b).
Hamathecium not apparent. Asci numerous, 11–14 μm
diam. (x ¼ 12:3mm, n=10), irregularly arranged, 32-spored
when mature, bitunicate nature undetermined, fissitunicate
nature undetermined, globose, without pedicel (Fig. 99b
and c). Ascospores 4–5.5×2.5–3 μm (x ¼ 4:7 2:8mm, n=
10), in the globose asci, olivaceous, oblong, 1-celled,
smooth (Fig. 99d).
Anamorph: Phoma-like coelomycetes.
On MEA colonies spreading, flat with sparse aerial
mycelium, covering the dish after 1 month; surface smokegrey with dirty white margins; reverse olivaceous-grey with
luteous patches. On PDA spreading without aerial mycelium,
colonies transparent, sporulating profusely with black, globose ascomata and pycnidia of a Phoma-like anamorph. On
OA similar, lacking aerial mycelium, sporulating profusely
with black, globose ascomata (based on CBS 297.56).
Material examined: USA, Michigan, East Lansing,
Science Greenhouse, isolated from damped off Phlox seedling, Dec. 1952, F.M. Clum (No. 27) (MSC 133.118, type).
Notes
Morphology
Pycnidiophora was formally established by Clum (1955)
based on its “imperfect stage of pycnidium”, which was
subsequently confirmed as the sexual stage (Cain 1961;
Thompson and Backus 1966). Clum (1955) has described
and tentatively assigned P. dispersa (Clum) Cain to
Fungal Diversity
Fig. 98 Javaria samuelsii (from isotype). a Ascoma on the host surface. Note reflexed pieces of the ruptured host tissue. b, c Cylindro-clavate
asci within narrow pseudoparaphyses in gelatinous matrix. d Released ascospore with sheath. Scale bars: a=1 mm, b=50 μm, c, d=20 μm
Aspergillaceae (= Eurotiaceae), and Stolk (1955b) has
proposed to assign the morphologically comparable species
P. multispora Saito & Minoura ex Cain to Eurotiaceae as
well. Cain (1961), however, suspected that the 32 asco-
Fungal Diversity
spores are actually the disarticulated segments of eight 4celled ascospores, thus assigned it under Preussia (Sporormiaceae). After detailed study, Thompson and Backus
(1966) confirmed that the so-called “eight 4-celled ascospores” do not exist in the development of the asci in both
P. dispersa and P. multisporum. Thus, Pycnidiophora was
assigned to Eurotiaceae (Eurotiales) (Thompson and
Backus 1966).
Phylogenetic study
Phylogenetic study based on the ITS-nLSU rDNA
sequences indicated that Pycnidiophora dispersa nested
within clade of Westerdykella (including the generic type,
W. ornata) (Kruys and Wedin 2009). Morphologically, both
genera have cleistothecioid ascomata, asci with short or
without pedicels and ascospores 1-celled and no germ slits.
Thus, Pycnidiophora is treated as a synonym of Westerdykella (Kruys and Wedin 2009).
Concluding remarks
Although the pleosporalean status of Pycnidiophora is
verified, morphological characters such as the cleistothecioid ascomata and irregularly arranged asci, which do not
show typical bitunicate or fissitunicate characters, absence
of pseudoparaphyses as well as the ascospores separating
into partspores very early all challenge the traditional
concept of Pleosporales (Zhang et al. 2009a). Obviously,
most of these morphological characters overlap with those
of the Eurotiales.
Type species
Sporormiella nigropurpurea Ellis & Everh., N. Amer.
Pyren.: 136 (1892). (Fig. 100)
Current name: Preussia nigropurpurea (Ellis & Everh.)
Kruys, Syst. Biod. 7: 476.
Ascomata 314–528 μm high×(250-)357–500 μm diam.,
solitary, scattered, or in small groups, immersed, semiimmersed to nearly superficial, globose, subglobose, wall
black, coriaceous, smooth, papillate, papilla 43–115 μm long,
72–157 μm broad, ostiolate, ostiole filled with periphyses
(Fig. 100a and b). Peridium 20–28 μm thick laterally, up to
40 μm thick at the apex, composed of small heavily pigmented
cells of textura angularis, cells 5–8 μm diam., cell wall
1–3 μm thick, apex cells smaller and walls thicker
(Fig. 100c). Hamathecium of dense, long, septate, cellular
pseudoparaphyses, 1.5–2 μm broad, embedded in mucilage.
Asci (70-)110–158×9–12.5(−15) μm (x ¼ 114:3 11:1mm, n=
10), 8-spored, bitunicate, fissitunicate, cylindrical to cylindroclavate, with a narrowed, furcate pedicel, 13–38 μm long,
ocular chamber apparent (Fig. 100d and e). Ascospores 15–
20×4–5.5 μm (x ¼ 17:3 4:9mm, n=10), obliquely uniseriate and partially overlapping to biseriate, shortly cylindrical
with rounded ends, brown, 3-septate, deeply constricted at
each septum, with sigmoid germ slit in each cell, smoothwalled (Fig. 100f and g).
Anamorph: none reported.
Material examined: USA, New field, New Jersey:
Gloucester Co., on cow dung, Mar. 1891 (NY, holotype).
Notes
Sporormiella Ellis & Everh., N. Amer. Pyren.: 136 (1892).
(Sporormiaceae)
Current name: Preussia Fuckel, Hedwigia 6: 175
(1867) [1869–70].
Generic description
Habitat terrestrial, saprobic (coprophilous). Ascomata
medium-sized, solitary, scattered, or in small groups,
semi-immersed to nearly superficial, globose, subglobose,
black, coriaceous, ostiolate, periphysate. Peridium thin,
composed of small heavily pigmented cells of textura
angularis, apex cells smaller and walls thicker. Hamathecium of dense, septate, cellular pseudoparaphyses, embedded in mucilage. Asci 8-spored, bitunicate, fissitunicate,
cylindro-clavate, with a narrowed, furcate pedicel. Ascospores cylindrical with rounded ends, brown, 3-septate,
deeply constricted at each septa, with sigmoid germ slit in
each cell.
Anamorphs reported for genus: none.
Literature: Ahmed and Cain 1972; Ellis and Everhart
1892; Khan and Cain 1979a, b; Luck-Allen and Cain 1975.
Morphology
Sporormiella was formally established by Ellis and
Everhart (1892) based on the single species, Sporormiella
nigropurpurea, which is characterized by its “immersed to
semi-immersed, papillate ascomata, cylindrical to cylindroclavate asci with a pedicel, three to multi-septate ascospores
with elongated germ slits through the whole cell” (Ahmed
and Cain 1972; Khan and Cain 1979a, b). Barr (1990a) has
indicated that Sporormiella might be a synonym of
Ohleriella, while Sporormiella is assigned to Sporormiaceae as a separate genus (Eriksson 2006; Lumbsch and
Huhndorf 2007). Currently, about 90 species are included
in this genus (http://www.mycobank.org).
Phylogenetic study
The phylogenetic analysis based on ITS-nLSU rDNA,
mtSSU rDNA and ß-tubulin sequences indicated that Sporormiella nested in Preussia, and a Sporormiella–Preussia complex
is formed (Kruys and Wedin 2009). Thus, Sporormiella was
assigned under Preussia (Kruys and Wedin 2009).
Fungal Diversity
Fig. 99 Pycnidiophora dispersa (a from CBS 297.56; b-d from MSC
133.118, type). a Ascomata scattering on the surface of the substrate.
b Crashed ascoma. Note the numerous released asci. c Globose asci
and released ascospores. d One-celled ascospores. Scale bars: a=
200 μm, b–d=20 μm
Concluding remarks
several layers of heavily pigmented, elongate cells of
textura angularis. Hamathecium of dense trabeculate,
aseptate, decomposing pseudoparaphyses. Asci bitunicate,
broadly cylindro-clavate with a narrow furcated pedicel.
Ascospores cylindrical to cylindro-clavate, with round ends,
brown, multi-septate, easily breaking into partspores.
Anamorphs reported for genus: none.
Literature: von Arx and van der Aa 1987.
It is clear that the presence or absence of an ostiole cannot
distinguish Sporormiella from Preussia according to the
findings of Guarro et al. (1997a, b) and Kruys and Wedin
(2009). Thus, Sporormiella should be treated as a synonym
of Preussia (Kruys and Wedin 2009).
Spororminula Arx & Aa, Trans. Br. Mycol. Soc. 89: 117
(1987). (Sporormiaceae)
Current name: Preussia Fuckel, Hedwigia 6: 175
(1867) [1869–70].
Generic description
Habitat terrestrial, saprobic (coprophilous). Ascomata small
to medium, solitary, scattered, immersed to erumpent,
globose, subglobose, to ovate, black, membraneous, papillate, ostiolate. Peridium thin, membraneous, composed of
Type species
Spororminula tenerifae Arx & Aa, Trans. Br. Mycol. Soc.
89: 117 (1987).(Fig. 101)
Current name: Preussia tenerifae (Arx & Aa) Kruys,
Syst. Biod. 7: 476.
Ascomata 290–400 μm diam., solitary, scattered, initially immersed, becoming erumpent when mature, globose,
subglobose to ovate, black, membraneous, with a cylindrical
or somewhat conical beak, 90–150(−230) μm broad and 110–
Fungal Diversity
Fig. 100 Sporormiella nigropurpurea (from NY, holotype). a
Section of an ascoma. b Section of the papilla. Note the dense
pseudoparaphyses. c Section of a partial peridium. d, e Eight-spored
cylindro-clavate asci with furcate pedicels. f, g Four-celled, brown
ascospores. Note the sigmoid germ slit in each cell. Scale bars: a=
200 μm, b, c=50 μm, d, e=20 μm, f, g=10 μm
Fungal Diversity
190 μm high (Fig. 101a). Peridium 20–33 μm thick, 1layered, composed of several layers of heavily pigmented,
elongate cells of textura angularis, cells up to 6.3×5 μm
diam., cell wall 1–1.5 μm thick (Fig. 101b and c).
Hamathecium of dense, long trabeculate pseudoparaphyses
1–2 μm broad, hyaline, aseptate, decomposing when mature.
Asci 165–220×33–42.5 μm, 8-spored, bitunicate, broadly
clavate, with a small, thin and furcate pedicel, 35–50 μm long,
3–5 μm broad, ocular chamber not observed (Fig. 101d and
e). Ascospores 68–93×12.5–16 μm, 3–4 seriate to uniseriate
near the base, cylindrical to cylindro-clavate with rounded
ends, brown, (6-)7 transverse septa, easily breaking into
partspores, central cells triangular in transverse section but
rectangular in vertical section, 14–16.5×8–10 μm long, apical
cells 12.5–15×11.5–17.5 μm long (Fig. 101f and g).
Anamorph: none reported.
Material examined: SPAIN, Canary Islands, Tenerifa
Las Canadas, on rabbit? droppings, Mar. 1986, J.A. von
Arx (HCBS 9812, holotype).
Notes
Morphology
Spororminula was formally established by von Arx and van
der Aa (1987) according to its “ostiolate ascomata, elongated
ascospore separated into part cells by transverse septa and
without germ slits”, and was monotypified by S. tenerifae.
Currently, only one species was included in this genus.
Phylogenetic study
Based on a phylogenetic analysis of ITS-nLSU rDNA, mtSSU
rDNA and ß-tubulin sequences, Spororminula tenerifae
nested in the clade of Preussia, thus Spororminula was
treated as a synonym of Preussia (Kruys and Wedin 2009).
Concluding remarks
To clarify its relationship with other genera of Sporormiaceae, further phylogenetic study is needed, which should
include additional related taxa.
Excluded and doubtful genera
Kriegeriella Höhn., Annls mycol. 16: 39 (1918). (Dothideomycetes, families incertae sedis, Microthyriaceae)
Generic description
Habitat terrestrial, saprobic? Ascomata small, solitary,
scattered, superficial, subglobose, black, roughened, apex
Fig. 101 Spororminula tenerifae (from HCBS 9812, holotype). a
Appearance of ascomata on the host surface. b, c Sections of the
partial peridium. Note the elongate cells of textura angularis. d, e
Asci with thin pedicels. f, g Ascospores, which may break into part
spores. Scale bars: a=0.5 mm, b=100 μm, c=50 μm, d–g=20 μm
no obvious opening. Peridium thin, composed of a single
type of lightly pigmented thin-walled cells. Hamathecium
long cellular pseudoparaphyses, septate. Asci 8-spored,
bitunicate, obpyriform. Ascospores hyaline, turning brown
when mature, multi-septate, constricted at each septum.
Anamorphs reported for genus: none.
Literature: von Arx and Müller 1975; Barr 1975, 1987b;
Eriksson 2006; Lumbsch and Huhndorf 2007.
Type species
Kriegeriella mirabilis Höhn., Annls mycol. 16: 39 (1918)
(Fig. 102)
Ascomata 100–120 μm high×150–220 μm diam., solitary,
scattered, superficial, with basal wall flattened on the surface of
the substrate, subglobose, black, roughened, apex no obvious
opening (Fig. 102a and e). Peridium thin, composed of a
single type of lightly pigmented thin-walled cells, cells up to
12×5 μm diam. in front view, cell wall less than 1 μm thick,
apex cells smaller and walls thicker (Fig. 102a and f).
Hamathecium long cellular pseudoparaphyses, 1.5–2 μm
wide, septate. Asci 65–85×31–36 μm (x ¼ 63:1 33mm, n=
10), 8-spored, bitunicate, fissitunicate undetermined, obpyriform, no pedicel, no ocular chamber was seen (Fig. 102c and
g). Ascospores 28–37.5×8–11 μm (x ¼ 32:6 10mm, n=
10), 8-seriate, fusoid with broadly rounded ends, hyaline,
turning brown when mature, 5–6-septate, constricted at each
septum, the upper first-cell longer and broader than the lower
ones with semi-round shape, smooth (Fig. 102d, h, i).
Anamorph: see Fig. b.
Material examined: On the leaves of Faulenden nadeln
von Pinus silvestris, bei Roñigstein, Sept. 1896, W. Rueges.
(S reg. nr F12638, isolectotype).
Notes
Morphology
Kriegeriella was formally established by von Höhnel (1918b)
and was represented by two species, i.e. K. mirabilis and K.
transiens; it was typified by K. mirabilis and assigned to
Microthyriaceae. Subsequently, Kriegeriella was assigned to
the subfamily of Aulographiodeae (Microthyriaceae)
(Batista et al. 1959), Asterinaceae (Hemisphaeriales) (Luttrell 1973) and Pseudosphaeriaceae (Dothideales) (Barr
1975).
After checking the original description and the type
specimens of K. mirabilis and K. transiens, no significant
b
Fungal Diversity
Fungal Diversity
difference could be observed, and both are described from
rotting needles of conifers (Barr 1975; Batista et al. 1959;
Höhnel 1918b). 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 under the genus
Kriegeriella (von Arx and Müller 1975; Barr 1975). The
other section of Kriegeriella, Extrawettsteinina, includes
two previous Extrawettsteinina species, i.e. K. minuta and
K. mediterranea. Barr (1987b) introduced a family, i.e.
Kriegeriellaceae (Dothideales) to accommodate Kriegeriella and Extrawettsteinina. This proposal is rarely
followed, and Kriegeriella is usually assigned to Pleosporaceae (Pleosporales) (Eriksson 2006; Lumbsch and Huhndorf
2007).
Phylogenetic study
None.
Concluding remarks
Kriegeriella might belong to Microthyriaceae, although it
would be unusual in this family in having 5-6-septate
ascospores. Micropeltidaceae better accommodates the
ascospores, however, the parallel arrangement of cells of
the upper peridium are not typical. Asterinaceae may be
most suitable as Luttrell (1973) suggested.
Phaeotrichum Cain & M.E. Barr, Can. J. Bot. 34: 676
(1956). (Dothideomycetes, family incertae sedis,
Phaeotrichaceae)
Type species
Phaeotrichum hystricinum Cain & M.E. Barr, Can. J. Bot.
34: 677 (1956). (Fig. 103)
(Some information for the following description is from
Cain 1956)
Ascomata 170–280 μm diam., cleistothecial, solitary, or
in small groups, superficial, with 15–20 long straight or
slightly flexed, thin, black appendages evenly scattered on
the surface of the ascomata, 0.5–1 mm long, 15–25 μm
wide at base, tapering to less than 5 μm at the blunt apex,
with few or without septa, globose, black, smooth
(Fig. 103a). Peridium thin, carbonaceous-membraneous,
1-layered, composed of dark brown thick-walled cells
of textura angularis, cells 8–16 μm diam., cell wall
0.5–1.5 μm thick (data obtained from Cain 1956)
(Fig. 103b). Hamathecium not observed. Asci 42–48×
14–17 μm, 8-spored, bitunicate form not typical, lacking
fissitunicate dehiscence, broadly clavate, with a relatively
thick pedicel which is about 18 μm (data obtained from
Cain 1956). Ascospores 14–16×4–5 μm, 4-seriate, oblong to ellipsoid, hyaline when young, turning reddish
brown at maturity, 1-septate, deeply constricted at the
septum, each end with a subhyaline and broadly rounded
germ pore, smooth, readily separating into partspores at
the septum at maturity (Fig. 103c, d, e and f).
Anamorph: none reported.
Material examined: CANADA, Ontario, Muskoka,
Stoneleigh, on porcupine dung, 18 Aug. 1932, Cain (TRTC
4361, holotype).
Note: the ascomata of the specimen are fragile and no
asci could be obtained.
Notes
Generic description
Morphology
Habitat terrestrial, saprobic (coprophilous). Ascomata
small, cleistothecial, solitary, or in small groups,
superficial, with long straight or slightly flexed, thin,
black appendages evenly scattered on the surface of the
ascomata, globose, black. Peridium thin, carbonaceousmembraneous, 1-layered, composed of dark brown thickwalled cells of textura angularis. Hamathecium not
observed. Asci bitunicate form not clear, fissitunicate
dehiscence not observed, broadly clavate, with a relatively thick pedicel. Ascospores oblong to ellipsoid,
hyaline when young, turning reddish brown at maturity,
1-septate, deeply constricted at the septum, each end with
a subhyaline and broadly rounded germ pore, readily
forming partspores at the septum at maturity.
Anamorphs reported for genus: none.
Literature: Cain 1956; Malloch and Cain 1972.
Phaeotrichum was formally established by Cain (1956)
to accommodate two new coprophilous fungi, i.e. P.
hystricinum and P. circinatum Cain, and P. hystricinum
was selected as the generic type. Phaeotrichum is
mainly characterized by its coprophilous habitat, superficial cleistothecial ascocarps covered by long hairy
appendages, reddish brown 1-septate ascospore with a
broadly rounded germ pore at each end, readily breaking
into partspores (Cain 1956). According to Cain (1956),
Phaeotrichum possesses untypical bitunicate ascus, and
the ascospore releasing is described as “simply break
down and allow the contents to become free in the
cavity of the ascocarp”. This ascospore releasing
mechanism is considered as evolutionarily developed
compared to those that “discharge the ascospores
through an apical pore” (Cain 1956). Although lacking
Fungal Diversity
Fig. 102 Kriegeriella mirabilis (from S reg. nr F12638, isolectotype). a Section of a superficial ascoma. b Anamorphic stage. c
Obpyriform ascus. Note the pigmented ascospores and hyaline
ascospores coexisted in a single ascus. d Ascospores. Scale bars: a=
50 μm, b–d=10 μm. e Ascomata on the host surface. f, g Crashed
ascoma. Note the peridium structure. h, i Hyaline asymmetric
ascospores. Scale bars: e, f =100 μm, g=50 μm, h, i=10 μm
a typical bitunicate ascus, Phaeotrichum is still
assigned to Pleosporales, because the lack of bitunicate
ascus does not “taken by itself, exclude a fungus from
close relationship” (Cain 1956).
Fungal Diversity
Fig. 102 (continued)
Fungal Diversity
Phylogenetic study
Concluding remarks
A single unverified isolate of Phaeotrichum benjaminii is
placed well outside of Pleosporales in a broad phylogenetic
study (Schoch et al. 2009).
The superficial cleistothecial ascocarps covered by long hairy
appendages, the absence of hamathecium as well as the
nontypical bitunicate ascus are all distinct from members of
Fig. 103 Phaeotrichum hystricinum (from TRTC 4361, holotype). a Superficial ascomata on
host surface. Note the long and
black appendages. b Part of
peridium. Note the large cells in
surface view. c–f Released reddish brown ascospores with hyaline end cells. Note the strongly
constricted middle septum.
Scale bars: a=0.5 mm,
b–f=20 μm
Fungal Diversity
Pleosporales, but definite conclusions could only be obtained
by further molecular phylogenetic analysis. In this study, we
assign it to Dothideomycetes incertae cedis.
Zeuctomorpha Sivan., P.M. Kirk & Govindu, Bitunicate
Ascomycetes and their Anamorphs: 572 (1984). (Venturiaceae)
Generic description
asci, ellipsoid and 1-septate ascospores, and presence of
pseudoparaphyses, and was monotypified by Z. arecae.
Zeuctomorpha arecae is widely distributed in tropical
regions of East South Asia exclusively on the leaves of
Areca catechu (Sivanesan 1984).
Phylogenetic study
None.
Habitat terrestrial, hemibiotrophic. Ascomata small, gregarious, superficial, globose to slightly flattened, ostiolate,
covered with setae. Peridium thin, composed of heavily
pigmented pseudoparenchymatous cells of textura angularis.
Hamathecium of rare, septate, branching and anastomosing
pseudoparaphyses. Asci 8-spored, with a short thick pedicel,
bitunicate, fissitunicate, broadly clavate to obclavate. Ascospores ellipsoid, dark brown, 1-septate, asymmetrical, deeply
constricted at the septum.
Anamorphs reported for genus: Acroconidiellina (Sivanesan 1984).
Literature: Sivanesan 1984.
Concluding remarks
Type species
Generic description
Zeuctomorpha arecae Sivan., P.M. Kirk & Govindu, in
Sivanesan, Bitunicate Ascomycetes and their Anamorphs:
572 (1984). (Fig. 104)
Ascomata 175–300 μm diam., gregarious, superficial,
globose to slightly flattened, collapsed at the apex when
dry, ostiolate, covered with numerous long setae
(Fig. 104a). Peridium up to 25 μm wide, composed of
heavily pigmented pseudoparenchymatous cells of textura
angularis, to 7 μm diam. Hamathecium of rare, 2–5 μm
broad, septate, branching and anastomosing pseudoparaphyses. Asci 83–185×29–40(−50) μm (x ¼ 134 35:3mm, n=
10), 8-spored, bitunicate, fissitunicate, broadly clavate to
obclavate, with a short thick pedicel, up to 40 μm long,
apically rounded, with a small ocular chamber (to 4 μm
wide×7 μm high) (Fig. 104b, c and d). Ascospores 35–43×
12.5–18 μm (x ¼ 36:5 15:4mm, n=10), 2–4 seriate, ellipsoid, dark brown, 1-septate, deeply constricted at the septum,
usually slightly asymmetric, smooth (Fig. 104e, f, g, h and i).
Anamorph: Acroconidiellina arecae (Sivanesan 1984).
Material examined: INDIA, Shimogee, on Areca
catechu L. leaf, 1 Nov. 1979, H.C. Govindu (IMI 246067,
holotype).
Habitat terrestrial, saprobic or parasitic. Ascostromata erumpent through the host surface in linear rows parallel to the
host fibers. Ascomata small- to medium-sized, semiimmersed to erumpent, subglobose to rectangular, black,
coriaceous, cells of ascostromata pseudoparenchymatous,
cells of peridium composed of pigmented cells of textura
angularis. Hamathecium of rare, pseudoparaphyses. Asci
bitunicate, clavate to cylindro-clavate. Ascospores oblong to
elongated oblong, hyaline, 1-celled, usually slightly curved.
Anamorphs reported for genus: none.
Literature: Hino and Katumoto 1958.
Notes
Morphology
Zeuctomorpha was formally established by Sivanesan
(1984) based on its superficial setose ascomata, clavate
This taxon is unusual amongst the Pleosporaceae as it
has hairy superficial ascomata, few pseudoparaphyses,
broadly clavate to obclavate asci and 1-septate pigmented
ascospores. All of these morphological characters are
most comparable with species of Acantharia, which
might be closely related to Venturiaceae (Zhang et al.
data unpublished).
Muroia I. Hino & Katum., J. Jap. Bot. 33: 79 (1958).
(Ascomycota)
Type species
Muroia nipponica I. Hino & Katum., J. Jap. Bot. 33: 79
(1958). (Fig. 105)
Ascostroma 1–6 mm long, 360–470 μm broad, linear
parallel to the host fibers with several linearly arranged
ascomata (Fig. 105a). Ascomata 250–400 μm diam.,
semi-immersed in substrate to erumpent, subglobose to
rectangular with a furrow-shaped ostiole, black, coriaceous, cells of ascostromata pseudoparenchymatous.
Peridium composed of pigmented cells of textura
angularis. Hamathecium of rare, 3–4.5 μm broad
pseudoparaphyses. Asci (120-)150–190 × 30–45 μm, 8spored, bitunicate, fissitunicate dehiscence not observed,
clavate to cylindro-clavate, with a short, thin, knob-like
pedicel, lacking an ocular chamber (Fig. 105b). Ascospores
43–50×13–18 μm (x ¼ 46:6 15:2mm, n=10), biseriate, ob-
Fungal Diversity
Fig. 104 Zeuctomorpha arecae (from IMI 246067, holotype). a
Gregarious ascomata on host surface. Note the numerous setae on the
surface of ascomata. b Asci with ocular chamber and short peduncles.
c, d Ascus with ocular chamber and knob-like pedicel. e–i One septate
ascospores which are slightly asymmetrical. Scale bars: a=0.5 mm,
b–i=20 μm
Fungal Diversity
long to elongated oblong, hyaline, 1-celled, usually slightly
curved (Fig. 105c,d and e).
Anamorph: none reported.
Material examined: JAPAN, Province Ugo. on moribund culm of Sasa kurilensis, 4 Aug. 1957, coll. H. Muroi,
Det. I. Hino & K. Katumoto (TNS-F-230252, isotype).
Notes
Morphology
Muroia was introduced based on M. nipponica, which is a
parasite on the lower part of Sasa kurilensis (Hino and
Katumoto 1958). Muroia is characterized by its 1-celled
ascospores. Considering the perithecial structure and linear
ostiole, it was assigned to the Lophiostomataceae, and was
regarded as closely related to the amerosporous Lophiella
(Hino and Katumoto 1958).
Phylogenetic study
None.
Concluding remarks
The linear ascostroma and 1-celled, hyaline ascospores
make it less likely to fit the concept of Lophiostomataceae.
Because of the condition of the specimen, its bitunicate
nature could not be confirmed.
Genera not studied
Aglaospora De Not., G. bot. ital. 2: 43 (1844).
Type species: Aglaospora profusa (Fr.) De Not., G. bot.
ital. 2: 43 (1844).
Aglaospora, which was introduced by de Notaris
(1844), has 35 species epithets (http://www.mycobank.
org/mycotaxo.aspx) and was considered to be a synonym
of Massaria (Voglmayr and Jaklitsch 2011) or separate
(Barr 1990a). In a recent phylogenetic study, Voglmayr and
Jaklitsch (2011) confirmed that Aglaospora is a synonym of
Massaria and is treated as such here. The immersed
ascomata with short beaks, together with ascostroma under
pseudostromatic tissues, cylindrical asci with a large and
refractive apical ring, trabeculate pseudoparaphyses within a
gel matrix, and distoseptate ascospores, are all similar to
species of Massaria. The large and conspicuous apical ring
of the ascus of Aglaospora has the appearance of being
unitunicate, and thus Shoemaker and Kokko (1977) treated it
as a unitunicate taxon. Currently, its bitunicate status is
widely accepted.
Allewia E.G. Simmons, Mycotaxon 38: 260 (1990).
Type species: Allewia proteae E.G. Simmons, Mycotaxon
38: 262 (1990).
Allewia was introduced by Simmons (1990) to accommodate Lewia-like species but with Embellisia anamorphs.
Embellisia differs from other similar genera by a combination of characters including the percentage of dictyoconidia,
shape of conidia, thickness of septa, umbilicate sites of
conidiophore geniculation, proliferating chlamydospores and
hyphal coils in culture (Simmons 1971). Based on multigene
phylogenetic analysis, A. eureka, which is closely related to
A. proteae, clustered together with species of Alternaria.
Thus, Allewia should be treated as a synonym of Lewia.
Anteaglonium Mugambi & Huhndorf, System. Biodivers.
7: 460 (2009).
Type species: Anteaglonium abbreviatum (Schwein.)
Mugambi & Huhndorf, System. Biodivers. 7: 460 (2009).
≡ Hysterium abbreviatum Schwein., Trans. Am. phil.
Soc., New Series 4: no. 2094 (1832).
Anteaglonium was introduced to accommodate a monophyletic hysterothecial clade within Pleosporales, and four
species (A. abbreviatum, A. globosum Mugambi & Huhndorf, A. parvulum (W.R. Gerard) Mugambi & Huhndorf
and A. latirostrum Mugambi & Huhndorf) are included
(Mugambi and Huhndorf 2009a). Anteaglonium is characterized by erumpent to superficial, globose to subglobose or
elongate, fusoid to oblong ascomata, which are brown to
shiny black, opening by a pronounced or indistinct
longitudinal slit running entire length of fruit body or apex
raised and laterally compressed; asci cylindrical with short
pedicel, 8-spored, uniseriate or biseriate; ascospores fusoid
to oblong, septate, constricted at the primary septum,
hyaline or pigmented. A phylogenetic analysis based on
DNA comparisons indicated that Anteaglonium resides as a
separate clade but related to Tetraplosphaeria, Lophiotrema
and other species without clear resolution. Therefore, the
familial placement of Anteaglonium remains unclear
(Mugambi and Huhndorf 2009a).
Arthopyrenia A. Massal., Ric. auton. lich. crost. (Verona):
165 (1852).
Type species: Arthopyrenia rhyponta (Ach.) A. Massal.,
Ric. auton. lich. crost. (Verona): 166, fig. 329 (1852).
≡ Verrucaria rhyponta Ach., K. Vetensk-Acad. Nya
Handl.: 150 (1809).
Arthopyrenia is a lichen genus with a Trentepohlia
photobiont and is characterized by dimidiate perithecoid
ascomata, which are scattered to irregularly confluent, and
have an upper thick clypeate wall composed of periderm
cells intermixed with dark hyphae. The pseudoparaphyses are
branched and asci are obpyriform, obclavate to subcylindrical
and 8-spored. Ascospores are oblong, ovoid, slipper-shaped,
1-3-septate, hyaline and smooth-walled (Coppins 1988;
Fungal Diversity
Fig. 105 Muroia nipponica (TNS-F-230252, isotype). a Linear ascostroma parallel to the host fibers. b Crashed ascus with ascospores released.
c–e Released hyaline ascospores. Scale bars: a=5 mm, b–e=30 μm
Upreti and Pant 1993). Multigene phylogenetic studies
indicated that Arthopyrenia salicis, a typical species of
Arthopyrenia, is located within Pleosporales in close
proximity to bambusicolous species in the genus Roussoella,
with its familial status remaining undetermined (Del Prado et
al. 2006; Schoch et al. 2009; Zhang et al. 2009a).
Ascocratera Kohlm., Can. J. Bot. 64: 3036 (1986).
Type species: Ascocratera manglicola Kohlm., Can. J. Bot.
64(12): 3036 (1986).
Ascocratera is a monotypic obligate marine fungus and
is characterized by conical, crater-like, erumpent to
superficial and carbonaceous ascomata, a depressed
ostiole, a thick peridium, trabeculate pseudoparaphyses,
bitunicate, fissitunicate and cylindrical asci, and ellipsoi-
dal, hyaline, 1-septate (3-septate when senescent) ascospores surrounded by a sheath (Kohlmeyer 1986).
Ascocratera was reported to be one of the most common
marine fungi of the upper intertidal zone of dead
mangrove roots, trunks and branches (Kohlmeyer 1986).
Based on a multigene phylogenetic analysis, Ascocratera
nested within the clade of Aigialaceae (Schoch et al.
2009; Suetrong et al. 2009).
Atradidymella M.L. Davey & Currah, Am. J. Bot. 96: 1283
(2009).
Type species: Atradidymella muscivora M.L. Davey &
Currah, Am. J. Bot. 96: 1283 (2009).
Atradidymella was introduced as a pleosporalean genus
parasitic on boreal bryophytes, and is characterized by
Fungal Diversity
minute, unilocular, setose pseudothecia with 2–3 wall
layers; brown, fusoid, 1-septate ascospores, and an anamorphic stage (Phoma muscivora M.L. Davey & Currah)
(Davey and Currah 2009). Based on an ITS rDNA
sequences analysis, Atradidymella nested within Didymellaceae (Davey and Currah 2009).
Bertiella (Sacc.) Sacc. & P. Syd., in Saccardo, Syll. fung.
(Abellini) 14: 19 (1899).
≡ Bertia subgen. Bertiella Sacc., Syll. fung. (Abellini) 1:
584 (1882).
Type species: Bertiella macrospora (Sacc.) Sacc. &
Traverso, Syll. fung. (Abellini) 19: 147 (1910).
≡ Bertia macrospora Sacc., Michelia 1(no. 8): 452 (1882).
Bertia subg. Bertiella was raised to generic rank by Saccardo
(1899), and is typified by B. macrospora. After studying the
type specimen of B. macrospora, Eriksson and Yue (1986)
assigned it to Massarina (as M. macrospora (Sacc.) O.E. Erikss.
& J.Z. Yue). Concurrently, Bertiella is treated as a synonym of
Massarina. Hyde et al. (2002) assigned Bertia macrospora to
Lophiostoma as (L. bertiellum Aptroot & K.D. Hyde).
The superficial ascomata, cylindro-clavate asci and
hyaline 1-septate ascospores which may become 3-septate
and pale brown when senescent and, in particular, the
woody habitat indicate that B. macrospora may be related
to Lophiostoma sensu Holm and Holm (1988). A single
isolate of Bertiella macrospora clusters with Byssosphaeria
in the Melanommataceae in a recent DNA based phylogeny
(Mugambi and Huhndorf 2009b). The relationship between
Bertiella and Byssosphaeria needs further study.
Byssothecium Fuckel, Bot. Ztg. 19: 251 (1861).
Type species: Byssothecium circinans Fuckel, Bot. Ztg.
19: 251 (1861).
The isotype of Byssothecium circinans is in FH as
exiccatae (Fungi rhenani 730c); it was described by Boise
(1983) and could not be loaned. Byssothecium circinans is
regarded as a saprobe or weak parasite of Medicago sativa
(Semeniuk 1983), and a Pleospora-type centrum was
observed (Boise 1983). A Chaetophoma-like anamorph
was produced in culture, however, no culture or herbarium
specimen is listed (Boise 1983). Boise (1983) regarded
Byssothecium circinans as closely related to Teichospora,
however, confirmation is required. An isolate of Byssothecium circinans was sequenced and a multigene phylogeny
placed it in close proximity to members of Massarinaceae
(Schoch et al. 2009; Zhang et al. 2009a; Plate 1).
Caryospora De Not., Micr. Ital. Novi 9: 7 (1855).
Type species: Caryospora putaminum (Schwein.) De Not.,
Micr. Ital., Dec. 9: 7 (1855).
After studying the Caryospora species in North America, Barr (1979b) indicated that species of Caryospora may
closely relate to Trematosphaeria. Boise (1985) distinguished Caryospora from Trematosphaeria based on the
structure of ascospores. Currently, 17 taxa, from freshwater,
marine, or terrestrial habitats (Raja and Shearer 2008), are
included within Caryospora and might be polyphyletic.
Celtidia J.D. Janse, Ann. Jard. Bot. Buitenzorg 14: 202 (1897).
Type species: Celtidia duplicispora J.D. Janse, Ann. Jard.
Bot. Buitenzorg 14: 202 (1897).
Celtidia is a monotypic genus, which is characterized by
its echinulate ascospores (Hawksworth 1979). It is only
known from an illustration accompanying the original
description from root nodules of Celtis in Java. A new
collection is needed for further study of this genus.
Chaetopreussia Locq.-Lin., Revue Mycol., Paris 41: 185
(1977).
Type species: Chaetopreussia chadefaudii Locq.-Lin.,
Revue Mycol., Paris 41: 187 (1977).
Chaetopreussia is a monotypic genus characterized by
cleistothecioid ascomata with seta, and 3-septate ascospores
without germ slits. Recent molecular analysis has shown that
cleistothecioid ascomata and the presence of germ slits lack
significance at the generic rank (Kruys and Wedin 2009).
Chaetopreussia is possibly another synonym of Preussia.
Clathrospora Rabenh., Hedwigia 1(18): 116 (1857).
Type species: Clathrospora elynae Rabenh., Hedwigia 1:
116 (1857).
The most striking character of Clathrospora is its
ascomata opening with an intraepidermal discoid lid and
muriform applanate ascospores with more than one row of
longitudinal septa (Shoemaker and Babcock 1992). The
form of opening and applanate ascospores, however, might
have limited significance at generic rank and thus,
Clathrospora may be closely related to Pleosporaceae.
Phylogenetic analysis based on nLSU, nSSU and mtSSU
indicate that C. diplospora (Ellis & Everh.) Sacc. &
Traverso nests in Pleosporaceae (Kruys et al. 2006).
Clathrospora elynae is saprobic on monocots (Shoemaker
and Babcock 1992).
Cochliobolus Drechsler, Phytopathology 24: 973 (1934).
Type species: Cochliobolus heterostrophus (Drechsler)
Drechsler, Phytopathology 24: 973 (1934).
Cochliobolus and its asexual relatives are well studied
taxa in Pleosporales because of their economic importance.
Cochliobolus includes both saprobic and pathogenic species
that are significant monocot pathogens worldwide, which
attack corn, rice, barley, sugarcane, wheat, and oats, all
major cereal crops. Cochliobolus is characterized by globose
or subglobose ascomata with a well defined long ostiolar
papilla or cylindrical neck, a peridium composed of
Fungal Diversity
pseudoparenchymatous cells, filliform, septate and branched
pseudoparaphyses, and thin-walled cylindrical or broadly
clavate asci. Ascospores are distinctively hyaline or pale
brown, filliform, and strongly helicoid to loosely coiled in
the asci (Sivanesan 1984). The anamorphs of Cochliobolus
belong to Bipolaris and Curvularia (Sivanesan 1984).
Bipolaris and Curvularia can be distinguished by characters
of conidial morphology, conidial germination, hilum structure, conidial septum and wall structure, conidial septum
ontogeny (Sivanesan 1987). Multigene phylogenetic analysis
indicated that Cochliobolus heterostrophus and C. sativus (S.
Ito & Kurib.) Drechsler ex Dastur nested within the clade of
Pleosporaceae (Zhang et al. 2009a; Plate 1). Thus, its
familial placement is confirmed.
Comoclathris Clem., Gen. fung. (Minneapolis): 37, 173
(1909).
Type species: Comoclathris lanata Clem. [as ‘Comochlatris’],
Gen. fung. (Minneapolis) (1909).
Comoclathris is temporarily placed in Diademaceae, and
its pivotal characters are the circular lid-like opening and
applanate reddish-brown to dark reddish-brown muriform
ascospores with single longitudinal septa (versus two or
more rows of longitudinal septa of Clathrospora) (Shoemaker
and Babcock 1992). Barr (1990b) treated it as a synonym of
Graphyllium. Comoclathris has been linked with an Alternaria-like anamorphs (Simmons 1952), which may suggest
its close relationship with Pleosporaceae.
Coronopapilla Kohlm. & Volkm.-Kohlm., Mycol. Res. 94:
686 (1990).
Type species: Coronopapilla avellina Kohlm. & Volkm.Kohlm., Mycol. Res. 94: 687 (1990).
Coronopapilla is characterized by immersed ascomata
with a conical papilla, thin peridium, 8-spored and thickwalled, cylindrical and fissitunicate asci. Ascospores are
ellipsoidal, 1-3-septate, brown and distoseptate. Coronopapilla
avellina is an obligate marine species, and was originally
assigned to Didymosphaeriaceae (Kohlmeyer and VolkmannKohlmeyer 1990). The marine habitat of Coronopapilla
makes it readily distinguishable from Didymosphaeria futilis
(the generic type of Didymosphaeria). Thus, the familial
placement of Coronopapilla is yet to be determined.
Cucurbitaria Gray, Nat. Arr. Brit. Pl. (London) 1: 508, 519
(1821).
Type species: Cucurbitaria berberidis (Pers.) Gray, Nat.
Arr. Brit. Pl. (London) 1: 508, 519 (1821).
≡ Sphaeria berberidis Pers., Neues Mag. Bot. 1: 83
(1794).
A narrow generic concept of Cucurbitaria was accepted
by Welch (1926), who restricted Cucurbitaria to five
closely related species, which have turbinate ascomata that
develop cespitosely in a massive subiculum or over
compressed stromatic tissues and have a thick and obconoid
base. A broader generic concept was accepted by Mirza
(1968), who also included species with globose or ovoid to
pyriform ascomata that are gregarious on the substrate with
only sparse subiculum and lack an obconoid region in the
base of the locule. Barr (1990b) accepted an intermediate
concept, and described 11 related species from North
America. Currently, 450 species are accepted in Cucurbitaria (http://www.mycobank.org/mycotaxo.aspx), and the
genus was assigned to Cucurbitariaceae. In this study, an
isolate of C. berberidis clustered with some species of
Pyrenochaeta and Didymosphaeria futilis, and they get
moderate bootstrap support (Plate 1). Cucurbitariaceae
may be another family within Pleosporineae.
Curreya Sacc., Syll. fung. (Abellini) 2: 651 (1883).
Type species: Curreya conorum (Fuckel) Sacc., Syll. fung.
(Abellini) 2: 651 (1883).
Curreya is a contentious genus which had been assigned to
Pleospora (Barr 1981). von Arx and van der Aa (1983),
however, maintained it as distinct, because of its Coniothyrium anamorph, and considered Curreya should be closely
related to Didymosphaeria, Melanomma, Paraphaeosphaeria or Massarina. Because of the small sclerotial cells of its
peridium, the narrower, thinner-walled asci and its Coniothyrium-like anamorph, Barr (1990b) assigned it to the
Leptosphaeriaceae. Previous phylogenetic studies indicated
that a strain of Curreya pityophila (J.C. Schmidt & Kunze)
Petr. nested within Massarineae (Kruys et al. 2006).
Decorospora Inderb., Kohlm. & Volkm.-Kohlm., Mycologia 94: 657 (2002).
Type species: Decorospora gaudefroyi (Pat.) Inderb.,
Kohlm. & Volkm.-Kohlm., Mycologia 94: 657 (2002).
≡ Pleospora gaudefroyi Pat., Tabl. analyt. Fung. France
(Paris) 10: 40 (no. 602) (1886).
Decorospora gaudefroyi (as Pleospora gaudefroyi) had
been considered a synonym of Pleospora herbarum,
despite its striking sheath of ascospores (Wehmeyer
1961). Molecular phylogenetic analysis based on partial
SSU and ITS rDNA sequences indicated that Decorospora
gaudefroyi was a sister taxon in the Pleosporaceae
represented by Alternaria alternata (Fr.) Keissl., Cochliobolus sativus, Pleospora herbarum, Pyrenophora triticirepentis (Died.) Drechsler and Setosphaeria rostrata K.J.
Leonard (Inderbitzin et al. 2002). Decorospora was
introduced as a monotypic genus represented by Decorospora gaudefroyi, which is characterized by black ascomata becoming superficial on the substrate at maturity,
septate and branched pseudoparaphyses, fissitunicate, clavate asci, as well as yellowish brown ascospores with seven
transverse septa and one to three longitudinal septa in each
Fungal Diversity
segment, enclosed in a sheath with 4–5 apical extensions
(Inderbitzin et al. 2002). Decorospora gaudefroyi is an
obligate marine fungus, growing at or above the high water
mark (Inderbitzin et al. 2002).
Diadema Shoemaker & C.E. Babc., Can. J. Bot. 67: 1349
(1989).
Type species: Diadema tetramerum Shoemaker & C.E.
Babc. [as ‘tetramera’], Can. J. Bot. 67: 1354 (1989).
During their study of Leptosphaeria and Phaeosphaeria,
Shoemaker and Babcock (1989c) found some alpine fungi
with typical pleosporalean characters (such as perithecoid
ascomata, bitunicate asci and presence of pseudoparaphyses)
having relatively large, very dark brown ascospores, mostly
with a peculiar disc-like opening (as reported in some
species of Wettsteinina, Shoemaker and Babcock 1987).
Thus, they introduced a new genus Diadema (typified by D.
tetramerum) to accommodate them (Shoemaker and
Babcock 1989c). Currently, Diadema is assigned to Diademaceae, and differs from other genera in the family in having
ascospores which lack longitudinal septa (Shoemaker and
Babcock 1992). The large, dark brown ascospores and the
disc-like opening, however, may be an adaptation to
environmental factors.
Diademosa Shoemaker & C.E. Babc., Can. J. Bot. 70: 1641
(1992).
Type species: Diademosa californiana (M.E. Barr) Shoemaker
& C.E. Babc. [as ‘californianum’], Can. J. Bot. 70: 1641 (1992).
≡ Graphyllium californianum M.E. Barr, Mem. N. Y.
bot. Gdn 62: 40 (1990).
Diademosa is the only genus in Diademaceae that has
terete (cylindrical, circular in cross section) ascospores
(Shoemaker and Babcock 1992).
Didymella Sacc., Michelia 2(no. 6): 57 (1880).
Type species: Didymella exigua (Niessl) Sacc., Syll. fung.
(Abellini) 1: 553 (1882).
≡ Didymosphaeria exigua Niessl, Öst. bot. Z.: 165
(1875).
The type specimen of Didymella (D. exigua) is lost and a
neotype specimen was selected by de Gruyter et al. (2009).
Didymella was characterized by the immersed or erumpent,
globose or flattened and ostiolate ascomata with dense, rare
(or lack?) of pseudoparaphyses. Asci are cylindrical,
clavate or saccate, and 8-spored. Ascospores are hyaline,
1-septate (symmetrical or asymmetrical) and constricted at
the septum. Didymella has been assigned under Mycosphaerellaceae, Pleosporales (Sivanesan 1984), Phaeosphaeriaceae (Barr 1979a; Silva-Hanlin and Hanlin 1999),
Venturiaceae (Reddy et al. 1998) or Pleosporales genera
incertae sedis (Lumbsch and Huhndorf 2007). Based on a
multigene phylogenetic analysis, the Didymella clade forms
a familial rank within Pleosporineae, thus the Didymellaceae was introduced (Aveskamp et al. 2010; de Gruyter et
al. 2009; Zhang et al. 2009a; Plate 1). Anamorphs of
Didymellaceae include Ascochyta, Ampelomyces, Boeremia, Chaetasbolisia, Dactuliochaeta, Epicoccum, Microsphaeropsis, Peyronellaea, Phoma, Piggotia, Pithoascus,
Pithomyces and Stagonosporopsis (Aveskamp et al. 2010;
de Gruyter et al. 2009; Hyde et al. 2011).
Didymocrea Kowalski, Mycologia 57: 405 (1965).
Type species: Didymocrea sadasivanii (T.K.R. Reddy)
Kowalski, Mycologia 57: 405 (1965).
≡ Didymosphaeria sadasivanii T.K.R. Reddy, Mycologia 53: 471 (1962).
Didymocrea is a monotypic genus, and was separated
from Didymosphaeria based on its “unitunicate asci”,
presence of pseudoparaphyses and absence of spermatia,
and assigned under Hypocreales (Kowalski 1965). Following Kowalski (1965), Luttrell (1975) also studied the
centrum development of Didymocrea, and concluded that it
should be a true pleosporalean fungus with functionally
unitunicate asci, and retained it in Didymosphaeria. After
studying the type specimen of Didymocrea sadasivanii,
Aptroot (1995) concluded that it should be closely related to
the loculoascomycetous genus Zopfia. Rossman et al. (1999)
also kept it as a unique genus in Pleosporales. Based on a
multigene phylogenetic analysis, D. sadasivanii nests within
Montagnulaceae (Kruys et al. 2006; Schoch et al. 2009).
Dothivalsaria Petr., Sydowia 19: 283 (1966) [1965].
Type species: Dothivalsaria megalospora (Auersw.) Petr.,
Sydowia 19: 283 (1966) [1965].
≡ Valsaria megalospora Auersw., Leipzig. Bot.
Tauschver. 5. (1866).
Dothivalsaria is monotypic and is represented by D.
megalospora (Petrak 1965). The taxon is characterized by
immersed, medium- to large-sized ascomata which usually
aggregate under blackened stromatic tissues and have
trabeculate pseudoparaphyses. Asci are cylindrical, while
ascospores are brown, ellipsoid, and 1-septate and uniseriate
in the asci (Barr 1990a). The ascostroma of D. megalospora is
comparable with those of Aglaospora profusa as has been
mentioned by Barr (1990a), but their relationships are unclear.
Epiphegia G.H. Otth, Mitt. naturf. Ges. Bern: 104 (1870).
Type species: Epiphegia alni G.H. Otth, Mitt. naturf. Ges.
Bern: 104 (1870).
Epiphegia was reinstated to accommodate a species
which has Phragmoporthe-like ascocarps and Massarinalike asci, pseudoparaphyses and ascospores (Aptroot 1998).
Ascomata are grouped within stromatic tissues, pseudoparaphyses are cellular, asci are bitunicate and ascospores are
hyaline and trans-septate (Aptroot 1998).
Fungal Diversity
Eremodothis Arx, Kavaka 3: 34 (1976) [1975] (IMI 90223=
CBS 610.74 type).
Type species: Eremodothis angulata (A.C. Das) Arx,
Kavaka 3: 34 (1976) [1975].
≡ Thielavia angulata A.C. Das, Trans. Br. Mycol. Soc.
45: 545 (1962).
The type species of Eremodothis (E. angulata) was
originally isolated from rice field soil in Fulta, India and it
was assigned to Sporormiaceae because of the orange
pigmentation of the colony (von Arx 1976). The cleistothecoid ascomata, sphaerical asci and 1-celled ascospores of E.
angulata are comparable with those of Pycnidiophora.
Based on a multigene phylogenetic study, both Eremodothis
and Pycnidiophora were treated as synonyms of Westerdykella (Kruys and Wedin 2009).
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).
Extrawettsteinina 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 (Barr
1987a). Extrawettsteinina is characterized by superficial, conical
ascomata, containing a few saccate bitunicate asci, ellipsoidal,
obovate-clavate, septate, smooth and hyaline ascospores which
turn dull brown at maturity (Barr 1972). The diagnostic
character of Extrawettsteinina is its conic ascocarps which are
superficial on the substrate, and radiating arrangement of wall
cells, which makes it distinguishable from comparable genera
such as Stomatogene and Wettsteinina.
Graphyllium Clem., Botanical Survey of Nebraska 5: 6 (1901).
Type species: Graphyllium chloës Clem., Botanical Survey
of Nebraska 5: 6 (1901).
Graphyllium was first described as a hysteriaceous
fungus with elongate ascomata, but von Höhnel (1918b,
1919) recognized its similarity to Clathrospora. Petrak
(1952) transferred Graphyllium to Pleospora, and noted
that the elongate ascomata and closely grouped rows of
small ascomata are not sufficient to recognize the genus.
Barr (1987b, 1990b) supported this proposal and considered Graphyllium differs from Clathrospora by shape,
septation and pigmentation of ascospores. A narrow
generic concept of Graphyllium was adapted by Shoemaker
and Babcock (1992), which is characterized by hysterothecia,
applanate ascospores that are at least 3-septate in side view
and have some longitudinal septa in front view, and it was
assigned under Hysteriaceae (order Pleosporales, Shoemaker
and Babcock 1992). But subsequent classification systems
tend to assign it to Diademaceae (e.g. Lumbsch and
Huhndorf 2007, 2010). This seems unlikely as pointed out
by Zhang et al. (2011) and the genus could be placed in one
of five families containing hysteriotheciod ascomata. Recollection and molecular studies are needed.
Halomassarina Suetrong, Sakay., E.B.G. Jones, Kohlm.,
Volkm.-Kohlm. & C.L. Schoch, Stud. Mycol. 64: 161 (2009).
Type species: Halomassarina thalassiae (Kohlm. & Volkm.Kohlm.) Suetrong, Sakay., E.B.G. Jones, Kohlm., Volkm.Kohlm. & C.L. Schoch, Stud. Mycol. 64: 161 (2009).
≡ Massarina thalassiae Kohlm. & Volkm.-Kohlm., Can.
J. Bot. 65: 575 (1987).
Halomassarina is another marine genus which morphologically fits Massarina sensu lato, and is typified by H.
thalassiae, which is characterized by subglobose to pyriform, immersed or erumpent, ostiolate, periphysate, papillate
or epapillate, coriaceous ascomata, simple, rarely anastomosing pseudoparaphyses, 8-spored, cylindrical to clavate,
pedunculate, thick-walled, fissitunicate asci, and ellipsoidal,
(1-)3-septate, hyaline ascospores. Based on a multigene
phylogenetic analysis, Halomassarina thalassiae clustered
together with Trematosphaeria pertusa and another marine
fungus Falciformispora lignatilis, and they are all assigned
to Trematosphaeriaceae (Suetrong et al. data unpublished).
Hypsostroma Huhndorf, Mycologia 84: 750 (1992).
Type species: Hypsostroma saxicola Huhndorf, Mycologia
84: 750 (1992).
Hypsostroma was introduced as a tropical woodinhabiting genus by Huhndorf (1992). Hypsostroma has
several striking characters including the large superficial
ascomata which form on a subiculum, pseudoparenchymatous peridial cells, trabeculate pseudoparaphyses, clavate
asci with long pedicels and a conspicuous apical apparatus,
and ascospores that separate into partspores with a germ slit
in each partspore (Huhndorf 1992). Phylogenetic study
indicated that Hypsostroma should be a new genus and the
Hypsostromataceae was reinstated to accommodate Hypsostroma (Mugambi and Huhndorf 2009b; Plate 1).
Julella Fabre, Annls Sci. Nat., Bot., sér. 6 9: 113 (1879) [1878].
Type species: Julella buxi Fabre, Annls Sci. Nat., Bot., sér.
6 9: 113 (1879) [1878].
Julella has been assigned to Thelenellaceae, a family of
Ostropomycetidae (Lumbsch and Huhndorf 2007), and
Arthopyreniaceae (= Xanthopyreniaceae sensu O. Eriksson,
Pleosporales) (Barr 1985). Julella is characterized by its
immersed, medium-sized ascomata with pseudoparenchymatous peridial cells, cellular pseudoparaphyses, and
hyaline and muriform ascospores (Barr 1985). With the
exception of hyaline ascospores, these characters are typical
of Montagnulaceae. The taxonomic affinity of the generic
Fungal Diversity
type of Julella needs confirmation following recollection.
Julella avicenniae (Borse) K.D. Hyde is a marine fungus. A
DNA based phylogeny containing most currently accepted
families placed two isolates of J. avicenniae as sister to the
families in the Pleosporineae with good support, which might
suggest a novel family within Pleosporales (Suetrong et al.
2009). However, J. avicenniae is not the generic type and
therefore this conclusion must be treated with caution as only
J. avicenniae can be considered pleosporalean.
Lautitia S. Schatz, Can. J. Bot. 62: 31 (1984).
Type species: Lautitia danica (Berl.) S. Schatz, Can. J. Bot.
62: 31 (1984).
≡ Leptosphaeria danica Berl., Icon. fung. (Abellini) 1:
87 (1892).
Lautitia is monotypified by L. danica, which is characterized by subglobose, immersed, ostiolate ascomata with a
pseudoclypeus, a thin peridium, broad, cellular pseudoparaphyses, and 8-spored, bitunicate, cylindrical to clavate asci.
Ascospores are hyaline, 1-septate, and obovate and the fungus
is parasitic on algae (Schatz 1984). Marine or maritime fungi
have been reported in Phaeosphaeria, such as P. spartinae
(Ellis & Everh.) Shoemaker & C.E. Babc. and P. ammophilae (Lasch) Kohlm. & E. Kohlm. (Zhang et al. 2009a). In
addition, the prosenchymatous peridium of L. danica agrees
with that of Phaeosphaeriaceae (Schatz 1984).
Lepidosphaeria Parg.-Leduc, C. r. hebd. Séanc. Acad. Sci.,
Paris, Sér. D 270: 2786 (1970).
Type species: Lepidosphaeria nicotiae Parg.-Leduc, Pubbl.
Staz. Zool. Napoli, 1 270: 2786 (1970).
Lepidosphaeria is a genus likely in Testudinaceae, which
is distinguished from other genera of this family by its
smaller ascospores, which lack furrows, and have minute
granulate ornamentation (Hawksworth 1979). In DNA
sequence-based phylogenies, L. nicotiae clustered with
species of Ulospora and Verruculina (Schoch et al. 2009;
Zhang et al. 2009a), but more recent work including
species of Platystomaceae lacks support (Plate 1).
Letendraea Sacc., Michelia 2: 73 (1880).
Type species: Letendraea eurotioides Sacc., Michelia 2: 73
(1880).
Letendraea was introduced for L. eurotioides, which is
characterized by superficial, globose to conical ascomata,
filliform pseudoparaphyses, obclavate to cylindrical, 8spored asci, and fusoid to oblong, 1-septate ascospores
(Saccardo 1880). Because L. helminthicola (Berk. &
Broome) Weese clustered with Karstenula rhodostoma,
Letendraea was assigned to Melanommataceae (Kodsueb
et al. 2006b). But subsequent multigene phylogenetic
analysis indicated that both L. helminthicola and L. padouk
Nicot & Parg.-Leduc nested within Montagnulaceae
(Schoch et al. 2009; Zhang et al. 2009a; Plate 1), and its
familial status seems confirmed.
Lindgomyces K. Hirayama, Kaz. Tanaka & Shearer,
Mycologia 102: 133 (2010).
Type species: Lindgomyces ingoldianus (Shearer & K.D.
Hyde) K. Hirayama, Kaz. Tanaka & Shearer, Mycologia
102: 733 (2010).
≡ Massarina ingoldiana Shearer & K.D. Hyde, Mycologia 89: 114 (1997).
Lindgomyces was introduced to accommodate a freshwater
lineage, which belongs to Massarina ingoldiana sensu lato,
and is characterized by scattered, subglobose to globose,
erumpent, papillate, ostiolate ascomata, cellular pseudoparaphyses, and 8-spored, fissitunicate, cylindrical to clavate
asci. Ascospores are fusoid to narrowly fusoid, hyaline and 1septate but become 3–5-septate when senescent (Hirayama et
al. 2010). A new family, Lindgomycetaceae, was introduced
to accommodate Lindgomyces (Hirayama et al. 2010).
Lophiella Sacc., Michelia 1: 337 (1878).
Type species: Lophiella cristata (Pers.) Sacc., Michelia 1:
337 (1878).
≡ Sphaeria cristata Pers., Syn. meth. fung. (Göttingen)
1: 54 (1801).
The generic type of Lophiella, L. cristata, was treated as
a synonym of Lophiostoma angustilabrum var. crenatum
(Pers.) Chesters & A.E. Bell (see http://www.indexfungorum.
org/names/Names.asp).
Loratospora Kohlm. & Volkm.-Kohlm., Syst. Ascom. 12:
10 (1993).
Type species: Loratospora aestuarii Kohlm. & Volkm.Kohlm., Syst. Ascom. 12: 10 (1993).
Loratospora was introduced as a marine genus and is
monotypified by L. aestuarii (Kohlmeyer and VolkmannKohlmeyer 1993). The generic type is characterized by
ellipsoid, immersed to erumpent, carbonaceous ascomata,
which are ostiolate, and with or without a papilla. Pseudoparaphyses comprise small subglobose cells forming irregular
chains and finally breaking apart, and asci are 8-spored,
clavate to ellipsoidal, and fissitunicate. Ascospores are
hyaline, cylindrical, 3-septate and surrounded by a mucilaginous sheath (Kohlmeyer and Volkmann-Kohlmeyer 1993).
The distinctive pseudoparaphyses of Loratospora aestuarii
makes it readily distinguishable from other taxa. Based on a
multigene phylogenetic analysis, Loratospora aestuarii nested
within the clade of Phaeosphaeriaceae (Schoch et al. 2009;
Suetrong et al. 2009; Plate 1), and ascospores of L. aestuarii
are in agreement with those of Phaeosphaeria as has been
mentioned by Kohlmeyer and Volkmann-Kohlmeyer (1993).
Macrospora Fuckel, Jb. nassau. Ver. Naturk. 23–24: 139
(1870) [1869–70].
Fungal Diversity
Type species: Macrospora scirpicola (DC.) Fuckel, Jb.
nassau. Ver. Naturk. 23–24: 139 (1870) [1869–70].
≡ Sphaeria scirpicola DC., in Lamarck & de Candolle,
Fl. franç., Edn 3 (Paris) 2: 300 (1805).
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 first septum and ascospores 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 in Pleosporaceae, and this proposal was
widely followed (Eriksson 2006; Lumbsch and Huhndorf
2010). Nimbya anamorphs were reported for Macrospora
(Johnson et al. 2002).
Massaria De Not., G. bot. ital. 1: 333 (1844).
Type species: Massaria inquinans (Tode) De Not., G. bot.
ital. 1: 333 (1844).
≡ Sphaeria inquinans Tode, Fung. mecklenb. sel.
(Lüneburg) 1: Fig. 85 (1790).
Colonies on MEA erumpent, not spreading; surface
irregular, folded; margins even, feathery; surface olivaceous
grey, with thin, umber margin; reverse olivaceous-grey. On
PDA similar; surface olivaceous grey, margin dirty white;
reverse smoke-grey to olivaceous grey; colonies reaching
1 cm diam. On OA similar, surface olivaceous grey in
centre, margins wide, dirty white; colonies reaching 12 mm
diam. on all media tested; colonies sterile (based on CBS
125591).
Massaria was formally established by de Notaris (1844),
and is typified by M. inquinans. Numerous fungi with
brown septate ascospores surrounded by gelatinous sheath
were included in the genus (Barr 1979b; Shoemaker and
LeClair 1975). Shoemaker and LeClair (1975) accepted a
narrow concept for Massaria, with only a few species
characterized by large, symmetric, 4-celled ascospores
surrounded by a massive gelatinous sheath. Barr (1979b,
1990a) had considered Aglaospora a separate genus, but this
subsequently proved congeneric with Massaria (Voglmayr
and Jaklitsch 2011). Based on intensive sample collection
and multi-gene phylogenetic analysis, Voglmayr and
Jaklitsch (2011) accepted Massaria as the sole genus within
Massariaceae, which is characterized by a set of well
defined morphological and ecological characters; Europe is
regarded as the centre of diversity.
Misturatosphaeria Mugambi & Huhndorf, Stud. Mycol.
64: 108 (2009).
Type species: Misturatosphaeria aurantonotata Mugambi &
Huhndorf, Stud. Mycol. 64: 108 (2009).
Misturatosphaeria was introduced to accommodate a
group of fungi which are phylogenetically closely related to
Amniculicolaceae, Lophiostomataceae sensu stricto and
Sporormiaceae (Mugambi and Huhndorf 2009b; Zhang et
al. 2009a). Species of Misturatosphaeria are characterized
by erumpent to superficial ascomata which are scattered or
in groups, with or without papilla; asci cylindrical or
clavate, 8-spored; pseudoparaphyses numerous, septate,
ascospores brown or hyaline, phragmosporous or dictyosporous, with or without sheath. The terrestrial saprobic
habitat on wood, as well as its distinct morphological
characters may indicate that this genus belongs to an
undescribed family. A close relationship with the marine
anamorphic species Floricola striata is unexpected and
may suggest that some of the species in this genus could
have marine affinities (Plate 1).
Navicella Fabre, Annls Sci. Nat., Bot., sér. 6 9: 96 (1879)
[1878].
Type species: Navicella julii Fabre, Annls Sci. Nat., Bot.,
sér. 6 9: 96 (1879) [1878].
Navicella is characterized by medium- to large-sized,
immersed to erumpent, globose ascomata, apex elongated or
rarely rounded, asci clavate or cylindrical, pseudoparaphyses
trabeculate, ascospores reddish to dark brown, ellipsoid to
fusoid, multi-septate, the primary septum is euseptate, and
others distoseptate, obliquely uniseriate or biseriate (Barr
1990a). Navicella is saprobic on bark, and was considered
closely related to the Lophiostomataceae (Holm and Holm
1988). Based on the wide endotunica, thin apical ring and
distoseptate ascospores, Barr (1990a) transferred it to the
Massariaceae. The morphological characters of Navicella do
not match the Massariaceae sensu stricto (Voglmayr and
Jaklitsch 2011).
Neotestudina Segretain & Destombes, C. r. hebd. Séanc.
Acad. Sci., Paris 253: 2579 (1961).
Type species: Neotestudina rosatii Segretain &Destombes,
C. r. hebd. Séanc. Acad. Sci., Paris 253: 2579 (1961).
Neotestudina is characterized by medium- to large-sized,
superficial, gregarious, cleistothecioid and globose ascomata
which split on opening. Asci are 4- or 8-spored, and cylindrical
or oblong, pseudoparaphyses are sparse and trabeculate, and
ascospores are dark brown, ellipsoid, 1-septate, with a small
germ pore at each end, and uniseriate or crowded in the asci
(Barr 1990a). Based on the cleistothecioid ascomata, Neotestudina was assigned under Zopfiaceae (von Arx and Müller
1975) or Testudinaceae (Hawksworth 1979). Barr (1990a)
assigned it to Didymosphaeriaceae based on its ascospore
morphology. A DNA based phylogeny showed that sequence
obtained from Neotestudina rosatii resides as sister to
Ulospora bilgramii (D. Hawksw., C. Booth & MorganJones) D. Hawksw., Malloch & Sivan. and other species that
may represent Testudinaceae or Platystomaceae (Kruys et al.
2006; Plate 1).
Fungal Diversity
Paraphaeosphaeria O.E. Erikss., Ark. Bot., Ser. 2 6: 405
(1967).
Type species: Paraphaeosphaeria michotii (Westend.) O.E.
Erikss., Cryptogams of the Himalayas 6: 405 (1967).
≡ Sphaeria michotii Westend., Bull. Acad. R. Sci. Belg.,
Cl. Sci., sér. 2 7: 87 (1859).
Paraphaeosphaeria was separated from Leptosphaeria
(Eriksson 1967a), and it is also quite comparable with
Phaeosphaeria. Paraphaeosphaeria can be distinguished
from Phaeosphaeria by its ascospores. Ascospores of Paraphaeosphaeria michotii have two septa, and they are biseriate,
straight, subcylindrical with broadly rounded ends, rather dark
brown and punctate. The primary septum is laid down closer
to the distal end than to the proximal, and the larger, proximal
hemispore is divided by one transversal septum. There are
more septa in the proximal hemispore of other species such as
Par. castagnei (Durieu & Mont.) O.E. Erikss., Par. obtusispora (Speg.) O.E. Erikss. and Par. vectis (Berk. & Broome)
Hedjar. Anamorphic characters can also distinguish Paraphaeosphaeria and Phaeosphaeria. Paraphaeosphaeria has
Paraconiothyrium or Coniothyrium-related anamorphs, but
Phaeosphaeria has Hendersonia-Phaeoseptoria anamorphs
(Eriksson 1967a). Shoemaker and Babcock (1985) redescribed some Canadian and extralimital species, and excluded
Par. longispora (Wegelin) Crivelli and Par. oblongata
(Niessl) Crivelli from Paraphaeosphaeria based on their
longitudinal septa as well as beak-like papilla and wall
structures. Molecular phylogenetic results based on multigenes indicated that Paraphaeosphaeria should belong to
Montagnulaceae (Zhang et al. 2009a; Plate 1).
Passeriniella Berl., Icon. fung. (Abellini) 1: 51 (1890).
Type species: Passeriniella dichroa (Pass.) Berl., Icon.
fung. (Abellini) 1: 51 (1890).
≡ Leptosphaeria dichroa Pass.
Passeriniella was introduced by Berlese in 1890 based on
the black, ostiolate and papillate ascomata, 8-spored asci, as
well as transverse septate ascospores, with pigmented central
cells and hyaline terminal cells. Two species were included, i.
e. P. dichroa and P. incarcerata (Berk. & M.A. Curtis) Berl.
(Berlese 1890). Subsequently, more species were introduced
including some marine taxa such as P. mangrovei G.L. Maria
& K.R. Sridhar, P. obiones (P. Crouan & H. Crouan) K.D.
Hyde & Mouzouras and P. savoryellopsis K.D. Hyde &
Mouzouras (Hyde and Mouzouras 1988; Maria and Sridhar
2002). Currently, eight species are included (http://www.
mycobank.org, Jan. 2011). Both P. dichroa and P. incarcerata were considered as synonyms of Leptosphaeria obiones
(P. Crouan & H. Crouan) Sacc. (Kohlmeyer and Kohlmeyer
1979). The familial placement of the marine species P.
savoryellopsis could not be resolved in a DNA based
phylogeny but it did suggest a close relationship to
Acrocordiopsis patilii (Suetrong et al. 2009) in Pleosporales.
Peridiothelia D. Hawksw., Bull. Br. Mus. nat. Hist., Bot.
14: 120 (1985).
Type species: Peridiothelia fuliguncta (Norman) D.
Hawksw., Bull. Br. Mus. nat. Hist., Bot. 14: 121 (1985).
≡ Microthelia fuliguncta Norman, Öfvers. kongl.
Svensk. Vetensk.-Akad. Förhandl., Stockholm 41(no. 8):
36 (1884).
When dealing with the names under Microthelia, Peridiothelia was introduced to accommodate species having nonclypeate peridium which composed cells of textura globulosa but sometimes angularis, “dark reddish brown except
below the generative locule where the wall is poorly
developed or almost absent at maturity, colour not changed
significantly in potassium hydroxide, centrum turning blue in
iodine” (Hawksworth 1985a). Three species were included, i.
e. P. grandiuscula (Anzi) D. Hawksw., P. fuliguncta and P.
oleae (Körb.) D. Hawksw., and Peridiothelia was referred to
Phaeosphaeriaceae (Hawksworth 1985a, b). However, its
familial placement is not confirmed yet.
Phaeodothis Syd. & P. Syd., Annls mycol. 2: 166 (1904).
Type species: Phaeodothis tricuspidis Syd. & P. Syd.,
Annls mycol. 2: 166 (1904).
Phaeodothis is characterized by its 1-septate euseptate
ascospores with a sparse hamathecium consisting of thin
pseudoparaphyses and immersed to superficial ascomata
(Aptroot 1995). The genus had been previously assigned to
Didymosphaeria, but Aptroot (1995) considered it to be closely
related to Phaeosphaeriaceae. A strain named Phaeodothis
winteri (a synonym of P. tricuspidis Syd. & P. Syd.) nested
within the clade of Montagnulaceae (Schoch et al. 2009).
Platychora Petr., Annls mycol. 23: 102 (1925).
Type species: Platychora ulmi (Schleich.) Petr., Annls
mycol. 23(1/2): 103 (1925).
Platychora is characterized by immersed to erumpent crustlike ascostroma with globose locules scattered inside (Barr
1968). Asci are oblong to saccate or nearly cylindrical and
bitunicate, and ascospores are hyaline 1-septate apiosporous
and turn olivaceous when old. Platychora had been previously
assigned to Venturiaceae by Barr (1968), but molecular
phylogenetic analysis indicated that a strain named Platychora
ulmi (the generic type of Platychora) belongs to Didymellaceae (Winton et al. 2007; Plate 1). The generic type needs
recollecting and epitypifying to stabilize the generic name.
Platystomum Trevis., Bull. Soc. R. Bot. Belg. 16: 16 (1877).
Type species: Platystomum compressum (Pers.) Trevis.,
Bull. Soc. R. Bot. Belg. 16: 16 (1877).
≡ Sphaeria compressa Pers., Syn. meth. fung. (Göttingen)
1: 56 (1801).
Platystomum was introduced by Trevisan in 1877, and
has been considered a synonym of Lophidium, as the
Fungal Diversity
ascospores of Platystomum have both transverse and
vertical septa (Barr 1990a, b; Chesters and Bell 1970).
However, the boundary between Lophiostoma and Platystomum is not clear (Chesters and Bell 1970). Holm and
Holm (1988) treated Platystomum as a synonym of
Lophiostoma, and concurrently, the Platystomaceae
should be treated as a synonym of Lophiostomataceae.
Based on a phylogenetic analysis, however, the generic
type of Platystomum (P. compressum) separated from
other species of Lophiostoma, and nested with the clade of
Platystomaceae (Mugambi and Huhndorf 2009b) which
may be closely related to species in the Testiduniaceae
(Plate 1).
Polyplosphaeria Kaz. Tanaka & K. Hirayama, Stud.
Mycol. 64: 192 (2009).
Type species: Polyplosphaeria fusca Kaz. Tanaka & K.
Hirayama, Stud. Mycol. 64: 193 (2009).
Polyplosphaeria is characterized by globose ascomata
surrounded by numerous brown hyphae and a reddish
pigment on the host surface around the ascomata (Tanaka et
al. 2009). Asci are cylindro-clavate with fissitunicate
dehiscence and ascospores are narrowly fusoid surrounded
by a sheath. The anamorph is Piricauda-like (Tanaka et al.
2009). The cylindro-clavate asci, narrowly fusoid ascospores
as well as its thin and numerous pseudoparaphyses are
comparable with those of Massarina sensu lato, especially
Lentithecium (Zhang et al. 2009a). The terrestrial and
bambusicolous habitat of Polyplosphaeria and Piricauda
anamorph readily distinguishes the genus from Lentithecium.
Pontoporeia Kohlm., Nova Hedwigia 6: 5 (1963).
Type species: Pontoporeia biturbinata (Durieu & Mont.)
Kohlm., Nova Hedwigia 6: 5 (1963)
≡ Sphaeria biturbinata Durieu & Mont., Flora Algéricae
1: 497 (1849).
Pontoporeia was introduced by Kohlmeyer in 1963,
and is monotypified by P. biturbinata. Pontoporeia was
treated as a synonym of Zopfia (Malloch and Cain 1972),
which is followed by Hawksworth and Booth (1974).
Based on its asci originating at the periphery of the
subglobose locus, filaments occupying the center of the
ascocarps, the irregular peridial structure, the ascospores
having 2-layered walls with a germ pore at each end and
its marine habitat, Pontoporeia was kept as a separate
genus within Pleosporaceae (Kohlmeyer and Kohlmeyer
1979). A DNA based phylogeny placed an isolate on a
long branch in relationship with other marine species,
Halotthia posidoniae and Mauritiana rhizophorae, but a
familial placement awaits further resolution (Suetrong et
al. 2009).
Pseudotrichia Kirschst., Annls mycol. 37: 125 (1939).
Type species: Pseudotrichia stromatophila Kirschst., Annls
mycol. 37: 125 (1939).
Pseudotrichia can be distinguished from Byssosphaeria,
Herpotrichia and Lojkania by its lacking of subiculum,
larger ascomata usually with compressed apices, the
peripheral arrangement of asci and trabeculate pseudoparaphyses (Barr 1984). Phylogenetic study of strains Pseudotrichia mutabilis and some Herpotrichia species indicated
that these species are closely related, and both nested within
Melanommataceae (Mugambi and Huhndorf 2009b). But in
this study, Pseudotrichia guatopoensis nested in the
Testudinaceae (or Platystomaceae) (Plate 1). The types of
both Herpotrichia and Pseudotrichia need recollecting,
redescribing and epitypifying in order to stabiles the use of
these generic names and clarify their familial status.
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).
≡ Leptosphaeria thalictri G. Winter, Hedwigia 10: 40
(1872).
Pseudoyuconia was introduced by Vassiljeva (1983), and
was monotypified by P. thalictri. Currently, Pseudoyuconia
is included in Pleosporaceae (Lumbsch and Huhndorf
2010).
Pyrenophora Fr., Summa veg. Scand., Section Post.
(Stockholm): 397 (1849).
Type species: Pyrenophora phaeocomes (Rebent.) Fr.,
Summa veg. Scand., Section Post. (Stockholm): 397
(1849).
≡ Sphaeria phaeocomes Rebent., Prodr. fl. neomarch.
(Berolini): 338 (1804).
Pyrenophora is characterized by immersed, erumpent to
nearly superficial ascomata, indefinite pseudoparaphyses,
clavate to saccate asci usually with a large apical ring, and
muriform terete ascospores. Morphologically, the terete
ascospores of Pyrenophora can be readily distinguished
from Clathrospora and Platyspora. The indefinite pseudoparaphyses and smaller ascospores of Pyrenophora can be
readily distinguished from those of Pleospora (Sivanesan
1984). Based on both morphology and molecular phylogeny, Pyrenophora is closely related to Pleosporaceae
(Zhang et al. 2009a).
Rechingeriella Petr., in Rechinger et al. Annln naturh. Mus.
Wien 50: 465 (1940).
Type species: Rechingeriella insignis Petr., Annln naturh.
Mus. Wien, Ser. B, Bot. Zool. 50: 465 (1940).
Rechingeriella is characterized by its erumpent to
superficial, cleistothecioid ascomata and thin, branching
pseudoparaphyses (Hawksworth and Booth 1974). Asci are
Fungal Diversity
obovate, thick-walled, bitunicate and evanescent, and
ascospores are globose, simple, dark brown to black (based
on the type specimen of R. insignis) (Hawksworth and
Booth 1974). Based on these characters, R. insignis was
treated as a species of Zopfia (as Z. insignis (Petr.) D.
Hawksw. & C. Booth). Rechingeriella has been assigned to
Botryosphaeriaceae by von Arx and Müller (1975). Further
study should be conducted on the type specimen of R.
insignis in order to clarify its taxonomic status and fresh
collections are needed for epitypification.
Rhytidiella Zalasky, Can. J. Bot. 46: 1383 (1968).
Type species: Rhytidiella moriformis Zalasky, Can. J. Bot.
46: 1383 (1968).
Rhytidiella was introduced based on R. moriformis, which
causes perennial rough-bark of Populus balsamifera
(Zalasky 1968), and produces macroconidia belonging to
Phaeoseptoria. Subsequently, three more species were
introduced, viz. R. baranyayi A. Funk & Zalasky, R. hebes
P.R. Johnst. and R. beloniza (Stirt.) M.B. Aguirre (AguirreHudson 1991; Funk and Zalasky 1975; Johnston 2007), Both
R. baranyayi and R. hebes seem closely related to R.
moriformis on both biology and morphology (Funk and
Zalasky 1975; Johnston 2007), but R. beloniza is saprobic on
Cordyline australis bark (Aguirre-Hudson 1991). Rhytidiella
was temporarily assigned to Cucurbitariaceae (Barr 1987b).
Richonia Boud., Revue mycol., Toulouse 7: 224 (1885).
Type species: Richonia variospora Boud., Revue mycol.,
Toulouse 7: 265 (1885).
Richonia is characterized by its 1-septate, relatively large
ascospores which are broadly rounded at both ends, and
have a thick ornamented undulating sheath giving an
irregularly ridged appearance to mature spores (Hawksworth
1979). Richonia variospora has been isolated from several
localities in France, but it is rare (Hawksworth 1979).
Richonia was assigned under Zopfiaceae (von Arx and
Müller 1975; Hawksworth 1979), and there are presently no
better suggestions for its familial placement. The taxon needs
recollecting and epitypifying.
Rimora Kohlm., Volkm.-Kohlm., Suetrong, Sakay. & E.B.
G. Jones, Stud. Mycol. 64: 166 (2009).
Type species: Rimora mangrovei (Kohlm. & Vittal)
Kohlm., Volkm.-Kohlm., Suetrong, Sakay. & E.B.G. Jones,
Stud. Mycol. 64: 166 (2009).
≡ Lophiostoma mangrovei Kohlm. & Vittal [as ‘mangrovis’], Mycologia 78: 487 (1986).
Rimora was introduced based on a marine fungus R.
mangrovei (syn. Lophiostoma mangrovei), and is characterized by its erumpent ascomata with elongated flat tops,
cellular pseudoparaphyses and cylindrical asci (Suetrong et
al. 2009). Ascospores are fusoid, hyaline, 3-septate and
surrounded with an evanescent sheath (Kohlmeyer and
Vittal 1986; Suetrong et al. 2009). Rimora forms a robust
clade with other marine fungi, such as species of Aigialus
and Ascocratera, and a new family, Aigialaceae was
introduced to accommodate them (Suetrong et al. 2009).
Roussoellopsis I. Hino & Katum., J. Jap. Bot. 40: 86 (1965).
Type species: Roussoellopsis japonica (I. Hino & Katum.)
I. Hino & Katum., J. Jap. Bot. 40: 86 (1965).
≡ Didymosphaeria japonica I. Hino & Katum., Bulletin
of the Faculty of Agriculture, Yamaguchi University 5: 229
(1954).
Roussoellopsis was introduced by Hino and Katumoto
(1965) based on three bambusicolous fungal species, i.e. R.
japonica, R. macrospora (I. Hino & Katum.) I. Hino &
Katum. and R. tosaensis (I. Hino & Katum.) I. Hino &
Katum. These three species have immersed and gregarious
ascomata, clavate to cylindro-clavate asci, numerous and
filliform pseudoparaphyses, and 1-septate, asymmetrical
ascospores (Hino and Katumoto 1965). All these characters
point Roussoellopsis to Pleosporales, but its familial
placement cannot be determined.
Saccothecium Fr., Fl. Scan.: 349 (1836).
Type species: Saccothecium sepincola (Fr.) Fr. [as ‘saepincola’], Summa veg. Scand., Section Post. (Stockholm): 398
(1849).
≡ Sphaeria sepincola Fr. [as ‘saepincola’], Observ.
mycol. (Havniae) 1: 181 (1815).
Saccothecium is characterized by its subglobose, immersed to erumpent ascomata, absence of pseudoparaphyses
and hyaline, muriform to phragmosporous ascospores. It has
been assigned to the Dothioraceae (Barr 1987b; Müller and
von Arx 1950). Molecular phylogenetic analysis indicated
that a strain named S. sepincola nested within Didymellaceae
(Schoch et al. 2009; Plate 1). The generic type needs
recollecting, redescribing and epitypifying.
Setosphaeria K.J. Leonard & Suggs, Mycologia 66: 294
(1974).
Type species: Setosphaeria turcica (Luttr.) K.J. Leonard &
Suggs, Mycologia 66: 295 (1974).
≡ Trichometasphaeria turcica Luttr., Phytopathology 48:
282 (1958).
Setosphaeria was segregated from Keissleriella on the
basis of lacking 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, and four species were included, i.e. S.
prolata, S. holmii, S. pedicellata (R.R. Nelson) K.J.
Leonard & Suggs and S. turcica (Leonard and Suggs
1974). Currently, nine species are included in Setosphaeria (http://www.mycobank.org, Jan/2011). Setosphaeria
Fungal Diversity
monoceras Alcorn nested within Pleosporaceae based on
multigene phylogenetic analysis (Schoch et al. 2009;
Plate 1).
Syncarpella Theiss. & Syd., Annls mycol. 13: 631 (1915).
Type species: Syncarpella tumefaciens (Ellis & Harkn.)
Theiss. & Syd., Annls mycol. 13(5/6): 633 (1915).
≡ Sphaeria tumefaciens Ellis & Harkn., J. Mycol. 2: 41
(1886).
Syncarpella was introduced by Theissen and Sydow
(1915) as a genus of Montagnellaceae within Dothideales.
A detailed description of S. tumefaciens can be seen in Barr
and Boise (1989). Syncarpella was considered closely
related to Leptosphaeria, and was treated as a synonym
(Clements and Shear 1931). Syncarpella is characterized by
its abundant globose, ovoid to turbinate ascomata with
minute papillae which are seated on a common basal stroma
and which are erumpent through fissures in the host tissues
(Barr and Boise 1989). The peridium is thicker at the base,
pseudoparaphyses are cellular, and asci are bitunicate,
clavate to oblong with a furcate pedicel. Ascospores are
pale brown to brown, oblong to narrowly obovoid, ends
obtuse, transversely septate, smooth-walled. All these
characters fit Cucurbitariaceae, where Barr and Boise
(1989) transferred Syncarpella.
Teichospora Fuckel, Jb. nassau. Ver. Naturk. 23–24: 160
(1870) [1869–70].
Type species: Teichospora trabicola Fuckel, Jb. nassau.
Ver. Naturk. 23–24: 161 (1870) [1869–70].
Teichospora was introduced by Fuckel (1870), and was
typified by T. trabicola, with four more species included,
i.e. T. brevirostris Fuckel, T. dura Fuckel, T. morthieri
Fuckel and T. obducens (Schumach.) Fuckel. Only T.
brevirostris and T. trabicola were kept in Teichospora
(Barr 1987b). After studying the type specimens, Barr
(1987b) indicated that Teichospora was different from
Strickeria with Teichospora belonging to Pleosporales,
and Strickeria closely related to Melanomma (Melanommatales). Currently, more than 250 names are included
within Teichospora (http://www.mycobank.org, Jan/2011),
but almost no molecular phylogenetic study has been
conducted on this genus.
Testudina Bizz., Atti Inst. Veneto Sci. lett., ed Arti, Sér. 6
3: 303 (1885).
Type species: Testudina terrestris Bizz., Fungi venet. nov.
vel. Crit. 3: 303 (1885).
Testudina terrestris is characterized by its reticulately
ridged ascospores, which readily distinguish it from other
genera of Zopfiaceae (Hawksworth 1979). The species is
usually associated with other fungi, or on the wood of Abies?
and Pinus or on the fallen leaves of Taxus in Europe
(Hawksworth and Booth 1974; Hawksworth 1979).
Tetraplosphaeria Kaz. Tanaka & K. Hirayama, Stud.
Mycol. 64: 177 (2009).
Type species: Tetraplosphaeria sasicola Kaz. Tanaka & K.
Hirayama, Stud. Mycol. 64: 180 (2009).
Tetraplosphaeria was introduced by Tanaka et al. (2009)
to accommodate bambusicolous fungi with immersed to
erumpent, globose to subglobose and smaller (mostly<
300 μm) ascomata. The peridium is thin, and is composed
of thin-walled cells of textura angularis. The pseudoparaphyses are cellular, and asci are fissitunicate, 8-spored,
cylindrical to clavate with short pedicels. Ascospores are
narrowly fusoid, hyaline and surrounded with a sheath.
Species of Tetraplosphaeria have Tetraploa sensu stricto
anamorphic stage, which is quite unique in Tetraplosphaeriaceae (Tanaka et al. 2009).
Tingoldiago K. Hirayama & Kaz. Tanaka, Mycologia 102:
740 (2010).
Type species: Tingoldiago graminicola K. Hirayama &
Kaz. Tanaka, Mycologia 102(3): 740 (2010).
Tingoldiago is a genus of freshwater ascomycetes characterized by flattened, globose, immersed to erumpent ascomata,
and numerous cellular pseudoparaphyses (Hirayama et al.
2010). Asci are fissitunicate and cylindrical, and ascospores
are 1-septate, which usually turn 3-septate and pale brown
when old, usually with a sheath (Hirayama et al. 2010).
Based on both morphology and multigene phylogenetic
analysis, Tingoldiago should be treated as a synonym of
Lentithecium (Shearer et al. 2009a; Zhang et al. 2009a).
Tremateia Kohlm., Volkm.-Kohlm. & O.E. Erikss., Bot.
Mar. 38: 165 (1995).
Type species: Tremateia halophila Kohlm., Volkm.-Kohlm.
& O.E. Erikss., Bot. Mar. 38: 166 (1995).
Tremateia was introduced as a facultative marine genus
which is characterized by depressed globose, immersed
ascomata, numerous and cellular pseudoparaphyses, fissitunicate and clavate asci, ellipsoid muriform ascospores,
and a Phoma-like anamorph (Kohlmeyer et al. 1995).
These characters point Tremateia to Pleosporaceae
(Kohlmeyer et al. 1995). DNA sequence based phylogenies
placed T. halophila as sister to Bimuria novae-zelandiae
in Montagnulaceae (Schoch et al. 2009; Suetrong et al.
2009).
Triplosphaeria Kaz. Tanaka & K. Hirayama, Stud. Mycol.
64: 186 (2009).
Type species: Triplosphaeria maxima Kaz. Tanaka & K.
Hirayama, Stud. Mycol. 64: 188 (2009).
Triplosphaeria was introduced as a bambusicolous genus
characterized by immersed ascomata, numerous cellular
Fungal Diversity
pseudoparaphyses, bitunicate, cylindrical to clavate asci with
a short pedicel, fusoid, hyaline, 1-septate ascospores surrounded with a sheath, and with a Tetraploa-like anamorph
(Tanaka et al. 2009). Together with Tetraplosphaeria,
Pseudotetraploa, Quadricrura and Polyplosphaeria, Triplosphaeria was assigned to the Tetraplosphaeriaceae (Tanaka
et al. 2009).
Ulospora D. Hawksw., Malloch & Sivan., in Hawksworth,
Can. J. Bot. 57: 96 (1979).
Type species: Ulospora bilgramii (D. Hawksw., C. Booth
& Morgan-Jones) D. Hawksw., Malloch & Sivan., Can. J.
Bot. 57: 96 (1979).
Ulospora was introduced as a monotypic genus to
accommodate taxa of Testudinaceae whose ascospore has
3–6 fissures (Hawksworth 1979). Genera of Testudinaceae
are distinguished based on the morphology of ascospores,
although the validity of this classification needs to be
confirmed by molecular study. DNA sequence based
phylogenies placed sequences from an unverified culture
of U. bilgramii in a clade together with Verruculina enalia,
and Lepidosphaeria nicotiae and it may have a close
relationship to species in Platystomaceae (Mugambi and
Huhndorf 2009b; Schoch et al. 2009; Plate 1).
Zopfia Rabenh., Fungi europ. exsicc.: no. 1734 (1874).
Type species: Zopfia rhizophila Rabenh., Fungi europ.
exsicc.: no. 1734 (1874).
Zopfia was introduced by Rabenhorst (1874) as a
monotypic genus (typified by Z. rhizophila), and it was
assigned to the Perisporiaceae by Saccardo (1882) and
Winter (1884). Arnaud (1913) described the Zopfiaceae to
accommodate Zopfia, and considered that it should be
excluded from the Perisporiaceae. A relatively broad
generic concept was accepted by Hawksworth and Booth
(1974), in which they take the ascospore size and
ornamentation variation as criteria under generic rank
classification, and they treat Celtidia, Lepidosphaeria,
Marchaliella, Neotestudina, Pontoporeia, Pseudophaeotrichum, Rechingeriella, Richonia and Testudina as synonyms
of Zopfia. A narrow generic concept was adopted by
Hawksworth (1979), and Zopfia is characterized by 1septate ascospores, which are apiculate at both ends,
smooth-walled by light microscope, with minute irregular
pitting by SEM, and larger than other species of Zopfia
sensu Hawksworth and Booth (1974). Three species were
accepted, viz. Z. albiziae Farr, Z. biturbinata (Dur. &
Mont.) Malloch & Cain and Z. rhizophila, and they all
occur on roots of plants (Hawksworth 1979). DNA
sequences from an unverified culture of Zopfia rhizophila
placed it in close proximity to species in Delitschiaceae
without strong statistical support (Kruys et al. 2006; Schoch
et al. 2009; Plate 1).
Zopfiofoveola D. Hawksw., Can. J. Bot. 57: 98 (1979).
Type species: Zopfiofoveola punctata (D. Hawksw. & C.
Booth) D. Hawksw., Can. J. Bot. 57: 98 (1979).
≡ Zopfia punctata D. Hawksw. & C. Booth, Mycol. Pap.
153: 23 (1974).
Zopfiofoveola was hesitantly separated from Zopfia as a
monotypic new genus based on its evenly distributed
ornamentation with pale minute pits readily visible under
the light microscope, and the more elongate shape and less
pronounced apical papilla than those of Zopfia (Hawksworth
1979). The type specimen of this species however, cannot be
redescribed, because “the type species is only known from a
microscopic preparation obtained from earthworm excrements
in Sweden” as has been mentioned by Hawksworth (1979).
General discussion
Molecular phylogenetic studies based on four to five genes
indicate that 20 families should be included in Pleosporales
(Schoch et al. 2009; Shearer et al. 2009; Suetrong et al.
2009; Tanaka et al. 2009; Zhang et al. 2009a). Together
with five unverified families (marked with “?”), 26 families
are currently assigned under Pleosporales (Table 4). The
Phaeotrichaceae lacks pseudoparaphyses, has cleistothecial
ascomata with long setae, and conspicuous ascospores with
germ pores at each end. These characters do not agree with
the current concept of Pleosporales (Zhang et al. 2009a),
and therefore Phaeotrichaceae is excluded from Pleosporales (Table 4).
Families in Pleosporales
Based on LSU and SSU rDNA, RPB1, RPB2 and TEF1
sequence analysis, Pleosporineae is emended, and in this
study, seven families are tentatively included, i.e. Cucurbitariaceae, Didymellaceae, Didymosphaeriaceae, Dothidotthiaceae, Leptosphaeriaceae, Phaeosphaeriaceae and
Pleosporaceae (Zhang et al. 2009a; Plate 1). In this study,
Massarineae was emended to accommodate another five
families, viz. Lentitheciaceae, Massarinaceae, Montagnulaceae, Morosphaeriaceae, Trematosphaeriaceae. The subordinal affinity of other families remained undetermined.
Most of the families accepted within Pleosporales received
high bootstrap support (Plate 1). The characters used to
define a family, however, do not appear to have clear cut
boundaries, as the ascomatal and hamathecial characters also
seem to be poorly defined in some families. For example,
both trabeculate and cellular pseudoparaphyses coexist in the
Amniculicolaceae. Pycnidiophora, a genus of Sporormiaceae, has cleistothecial ascomata with spherical asci irregularly arranged in it. Brown phragmosporous ascospores are
reported in Amniculicolaceae, Leptosphaeriaceae, Lophios-
Fungal Diversity
tomataceae, Melanommataceae, Montagnulaceae, Phaeosphaeriaceae and Pleosporaceae. Similarly muriform ascospores occur in Aigialaceae, Amniculicolaceae,
Didymellaceae, Lophiostomataceae, Montagnulaceae, Pleosporaceae and Sporormiaceae. Anamorphs of Pleosporales
are also variable to a large degree at the family level. Both
hyphomycetous and coelomycetous anamorphs co-exist in
Didymellaceae, Melanommataceae or Pleosporaceae.
Phoma and Phoma-like anamorphs exist in Didymellaceae,
Leptosphaeriaceae, Phaeosphaeriaceae, Pleosporaceae and
Melanommataceae (de Gruyter et al. 2009; Zhang et al.
2009a). It is clear that some characters, e.g. cleistothecial or
perithecial ascomata, shape, colour and septation of ascospores, shape or arrangement (regular or irregular) of asci, or
even presence or absence of pseudoparaphyses have evolved
on numerous occasions which make the use of morphological characters in segregating families complicated. It is
therefore unclear with our present state of knowledge which
characters are taxonomically important at the family level or
whether a suit of characters are necessary to define a family.
DNA sequence comparisons are essential in delineating
these taxa in combination with other characters. It is hoped
that additional characters, i.e. biochemical, genomic and
subcellular will be used to further distinguish these groups
into natural taxa. Below we discuss each of the families,
their genera and their considered important characteristics.
Aigialaceae Suetrong, Sakay., E.B.G. Jones, Kohlm.,
Volkm.-Kohlm. & C.L. Schoch 2010
The Aigialaceae was introduced by Suetrong et al.
(2009) based on its carbonaceous ascomata without papilla,
cylindrical asci with apical apparatus, trabeculate pseudoparaphyses and ascospores with a sheath. The type genus
(Aigialus) of the Aigialaceae was previously incorporated
within the Massariaceae (Lumbsch and Huhndorf 2007).
Currently, three genera are assigned under Aigialaceae, viz.
Ascocratera, Aigialus and Rimora (Suetrong et al. 2009).
The genera included in Aigialaceae have a wide range of
morphological variation, with very few shared features as
mentioned above, but all are found in mangrove habitats
(Suetrong et al. 2009). The ascospores, however, vary
widely from having 1 to 3 transverse septa and being hyaline
to muriformly septate and brown (Suetrong et al. 2009). It is
still unclear which characters unify the family and therefore
placement of unsequenced genera is difficult. Further
molecular work is needed to better understand this family.
Table 4 Families currently accepted in Pleosporales (syn. Melanommatales) with included genera
Pleosporales subordo. Pleosporineae
?Cucurbitariaceae
Cucurbitaria Gray
Curreya Sacc.
?Rhytidiella Zalasky
Syncarpella Theiss. & Syd.
Didymellaceae
?Appendispora K.D. Hyde
Didymella Sacc. ex D. Sacc.
Didymosphaerella Cooke
Leptosphaerulina McAlpine
Macroventuria Aa
?Platychora Petr.
Didymosphaeriaceae
Didymosphaeria Fuckel
Phaeodothis Syd. & P. Syd.
Dothidotthiaceae
Dothidotthia Höhn.
Leptosphaeriaceae
Leptosphaeria Ces. & De Not.
Neophaeosphaeria Câmara, M.E. Palm & A.W. Ramaley
Phaeosphaeriaceae
Barria Z.Q. Yuan
?Bricookea M.E. Barr
?Chaetoplea (Sacc.) Clem.
?Eudarluca Speg.
Entodesmium Reiss
?Hadrospora Boise
Lautitia S. Schatz
Loratospora Kohlm. & Volkm.-Kohlm.
Metameris Theiss. & Syd.
Mixtura O.E. Erikss. & J.Z. Yue
Nodulosphaeria Rabenh.
Ophiobolus Reiss
Ophiosphaerella Speg.
Phaeosphaeria I. Miyake
Phaeosphaeriopsis Câmara, M.E. Palm & A.W.
Ramaley
Pleoseptum A.W. Ramaley & M.E. Barr
Setomelanomma M. Morelet
Wilmia Dianese, Inácio & Dornelo-Silva
Pleosporaceae
Cochliobolus Drechsler
Crivellia Shoemaker & Inderbitzin
Decorospora Inderbitzin, Kohlm. & Volkm.-Kohlm.
Extrawettsteinina M.E. Barr
Lewia M.E. Barr & E.G. Simmons
Macrospora Fuckel
Platysporoides (Wehm.) Shoemaker & C.E. Babc.
Pleospora Rabenh. ex Ces. & De Not.
Amniculicolaceae Yin. Zhang, C.L. Schoch, J. Fourn.,
Crous & K.D. Hyde 2009
Members of Amniculicolaceae form a well supported
clade, and all are freshwater fungi which usually stain the
woody substrate purple (Zhang et al. 2009a, c). Genera of
Amniculicolaceae have ascomata with compressed papilla
Pseudoyuconia Lar. N. Vasiljeva
Pyrenophora Fr.
Setosphaeria K.J. Leonard & Suggs
Pleosporales subordo. Massarineae
Lentitheciaceae
Lentithecium K.D. Hyde, J. Fourn. & Yin. Zhang
Fungal Diversity
Table 4 (continued)
Katumotoa Kaz. Tanaka & Y. Harada
Keissleriella Höhn.
?Wettsteinina Höhn.
Massarinaceae
Table 4 (continued)
Lophiostoma Ces. & De Not.
Melanommataceae
?Astrosphaeriella Syd. & P. Syd. (Syn. Javaria)
?Anomalemma Sivan.
Byssothecium Fuckel
?Asymmetricospora J. Fröhl. & K.D. Hyde
Massarina Sacc.
Bertiella (Sacc.) Sacc. & P. Syd.
Saccharicola D. Hawksw. & O.E. Erikss.
?Bicrouania Kohlm. & Volkm.-Kohlm.
Montagnulaceae
Byssosphaeria Cooke
Bimuria D. Hawksw., Chea & Sheridan
Calyptronectria Speg.
?Didymocrea Kowalsky
?Caryosporella Kohlm.
Kalmusia Niessl
Herpotrichia Fuckel
Karstenula Speg.
?Mamillisphaeria K.D. Hyde, S.W. Wong & E.B.G. Jones
Letendraea Sacc.
Melanomma Nitschke ex Fuckel
Montagnula Berl.
Ohleria Fuckel
Paraphaeosphaeria O.E. Erikss.
Tremateia Kohlm., Volkm.-Kohlm. & O.E. Erikss.
Morosphaeriaceae
?Asteromassaria Höhn
Pseudotrichia Kirschst.
Pleomassariaceae
?Lichenopyrenis Calatayud, Sanz & Aptroot
?Splanchnonema Corda
Helicascus Kohlm.
?Peridiothelia D. Hawksw.
Morosphaeria Suetrong, Sakay., E.B.G. Jones & C.L. Schoch
Pleomassaria Speg.
Trematosphaeriaceae
Sporormiaceae
Falciformispora K.D. Hyde
Chaetopreussia Locq.-Lin.
Halomassarina Suetrong, Sakay., E.B.G. Jones, Kohlm., Volkm.-Kohlm. & C.L.
Schoch
Pleophragmia Fuckel
Trematosphaeria Fuckel
Preussia Fuckel
Other families
Aigialaceae
Aigialus Kohlm. & S. Schatz
Eremodothis Arx
Sporormia De Not.
Westerdykella Stolk
?Teichosporaceae
Ascocratera Kohlm.
Chaetomastia (Sacc.) Berl
Rimora Kohlm., Volkm.-Kohlm., Suetrong, Sakay. & E.B.G. Jones
Immotthia M.E. Barr
Amniculicolaceae
Amniculicola Y. Zhang & K.D. Hyde
Murispora Yin. Zhang, C.L. Schoch, J. Fourn., Crous & K.D. Hyde
?Massariosphaeria (E. Müll.) Crivelli
Neomassariosphaeria Yin. Zhang, J. Fourn. & K.D. Hyde
?Arthopyreniaceae (Massariaceae)
Arthopyrenia A. Massal.
Dothivalsaria Petr.
Loculohypoxylon M.E. Barr
Sinodidymella J.Z. Yue & O.E. Erikss.
Teichospora Fuckel
Tetraplosphaeriaceae
Polyplosphaeria Kaz. Tanaka & K. Hirayama
Tetraplosphaeria Kaz. Tanaka & K. Hirayama
Triplosphaeria Kaz. Tanaka & K. Hirayama
?Zopfiaceae (syn Testudinaceae)
?Dubitatio Speg.
Caryospora De Not.
Massaria De Not.
Celtidia J.M. Janse
Navicella Fabre
?Coronopapilla Kohlm. & Volkm.-Kohlm.
Roussoëlla Sacc.
Halotthia Kohlm.
?Roussoellopsis I. Hino & Katum.
Lepidosphaeria Parg.-Leduc
Delitschiaceae
Delitschia Auersw.
?Mauritiana Poonyth, K.D. Hyde, Aptroot & Peerally
Pontoporeia Kohlm.
Ohleriella Earle
?Rechingeriella Petr.
Semidelitschia Cain & Luck-Allen
Richonia Boud.
?Diademaceae
Clathrospora Rabenh.
Testudina Bizz.
Verruculina Kohlm. & Volkm.-Kohlm.
Comoclathris Clem.
Ulospora D. Hawksw., Malloch & Sivan.
Diadema Shoemaker & C.E. Babc.
Zopfia Rabenh.
Diademosa Shoemaker & C.E. Babc.
Graphyllium Clem.
Hypsostromataceae
Hypsostroma Huhndorf
Lindgomycetaceae
Lindgomyces K. Hirayama, Kaz. Tanaka & Shearer 2010
Lophiostomataceae
Zopfiofoveola D. Hawksw.
Pleosporales genera incertae sedis
Acrocordiopsis Borse & K.D. Hyde
Aglaospora De Not.
Anteaglonium Mugambi & Huhndorf
Ascorhombispora L. Cai & K.D. Hyde
Atradidymella Davey & Currah
Fungal Diversity
Table 4 (continued)
Belizeana Kohlm. & Volkm.-Kohlm.
Biatriospora K.D. Hyde & Borse
Byssolophis Clem.
Carinispora K.D. Hyde
Cilioplea Munk
Decaisnella Fabre
Epiphegia Nitschke ex G.H. Otth
Isthmosporella Shearer & Crane
Julella Fabre
Lineolata Kohlm. & Volkm.-Kohlm.
Lophiella Sacc.
Lophionema Sacc.
Lophiotrema Sacc.
Moristroma A.I. Romero & Samuels
Neotestudina Segretain & Destombes
Ostropella (Sacc.) Höhn.
Paraliomyces Kohlm.
Passeriniella Berl.
Quintaria Kohlm. & Volkm.-Kohlm.
Saccothecium Fr.
Salsuginea K.D. Hyde
Shiraia P. Henn.
Xenolophium Syd.
Family excluded
Phaeotrichaceae
Echinoascotheca Matsush.
Phaeotrichum Cain & M.E. Barr
Trichodelitschia Munk
Genera excluded
Kriegeriella Höhn.
Muroia I. Hino & Katum.
Zeuctomorpha Sivan., P.M. Kirk & Govindu
and cylindrical to cylindro-clavate asci. Neomassariosphaeria
typhicola was traditionally assigned to Massariosphaeria (as
M. typhicola), and Massariosphaeria is characterized by
staining the woody substrate purple (Crivelli 1983;
Leuchtmann 1984). Eriksson (1981 p. 135) had pointed out
that “Purple-staining species of Pleospora, treated by Webster
(1957), are not congeneric with P. herbarum (Eriksson 1967b:
13), and certainly do not even belong to the Pleosporaceae”. This is mirrored in Murispora rubicunda, a
previous Pleospora species (as P. rubicunda) staining the
woody substrate purple, closely related to the Amniculicolaceae in a subsequent phylogenetic study (Zhang et al.
2009a). The anamorphs of this family are possibly
Anguillospora longissima, Spirosphaera cupreorufescens
and Repetophragma ontariense (Zhang et al. 2009a).
?Arthopyreniaceae (or Massariaceae) W. Watson 1929
The Arthopyreniaceae was introduced as a lichenized
family of Pyrenocarpales, which comprises Acrocordia,
Arthopyrenia, Athrismidium, Bottaria, Celothelium, Lau-
rera, Leptorhaphis, Microthelia, Microtheliopsis, Polyblastiopsis, Pseudosagedia, Raciborskiella and Tomasellia
(Watson 1929). Subsequently, Arthopyreniaceae was
assigned under Dothideales (suborder Pseudosphaeriineae)
(von Arx and Müller 1975). The generic type of Massaria
(M. inquinans) and Torula herbarum and Arthopyrenia
salicis together with members of Roussoella as well as
Roussoellopsis form a robust clade, which makes their
familial placement uncertain (Massariaceae or Arthopyreniaceae) (Schoch et al. 2009; Zhang et al. 2009a).
?Cucurbitariaceae G. Winter 1885
The Cucurbitariaceae is characterized by its aggregated
ascomata which form from a basal stromatic structure,
ostiolate, fissitunicate and cylindrical asci, and pigmented,
phragmosporous or muriform ascospores (Cannon and Kirk
2007). Currently, no molecular study has been able to
resolve its ordinal status, but some characters are similar to
Leptosphaeriaceae or Phaeosphaeriaceae (Cannon and
Kirk 2007). Cucurbitaria elongata clustered within Pleosporales (Schoch et al. 2006).
Delitschiaceae M.E. Barr 2000
The Delitschiaceae was established to accommodate
some species of the Sporormiaceae, which is characterized
by its ascomata with periphysate ostioles, ocular chamber
surrounded by a dome and usually in having four refractive
rods, ascospores with or without a septum, having a germ
slit in each cell and being surrounded by a mucilaginous
sheath (Barr 2000). Species of the Delitschiaceae are
hypersaprotrophic on old dung or exposed wood (Barr
2000). Based on a molecular phylogenetic studies, Delitschia didyma and D. winteri form a robust clade basal to
other pleosporalean fungi (Schoch et al. 2009; Zhang et al.
2009a). The familial status of two other genera, Ohleriella
and Semidelitschia, remains undetermined.
?Diademaceae Shoemaker & C.E. Babc. 1992
The Diademaceae was introduced by Shoemaker and
Babcock (1992) based on its ascomata opening as a flat
circular lid and bitunicate asci, ascospores are fusiform,
brown, mostly applanate, and having three or more
transverse septate and with or lacking longitudinal septa
and usually having a sheath. Five genera had been included
viz. Clathrospora, Comoclathris, Diadema, Diademosa and
Macrospora (Shoemaker and Babcock 1992).
Didymellaceae Gruyter, Aveskamp & Verkley 2009
The generic type of Didymella (D. exigua) together with
some Phoma or Phoma-related species form a robust
familial clade on the phylogenetic tree, thus the Didymellaceae was introduced to accommodate them (de Gruyter et
al. 2009). Subsequently, Didymellaceae was assigned to
Fungal Diversity
Pleosporineae (suborder of Pleosporales) (Zhang et al.
2009a). A detailed study was conducted on the Didymellaceae based on LSU, SSU rDNA, ITS as well as β-tubulin,
which indicated that many Phoma or Phoma-related
species/fungi reside in this clade of the Didymellaceae
(Aveskamp et al. 2010).
Didymosphaeriaceae Munk 1953
The Didymosphaeriaceae was introduced by Munk (1953),
and was revived by Aptroot (1995) based on its distoseptate
ascospores and trabeculate pseudoparaphyses, mainly anastomosing above the asci. The familial status of the Didymosphaeriaceae is debatable, and Lumbsch and Huhndorf (2007)
assigned it to the Montagnulaceae, while von Arx and Müller
(1975) treated it as a synonym of the Pleosporaceae. In this
study, Didymosphaeria futilis (the generic type of Didymosphaeria) is closely related to the Cucurbitariaceae (Plate 1).
Herein, we accept it as a separate family containing three genera,
namely Appendispora, Didymosphaeria and Phaeodothis. More
information could only be obtained by further molecular work
based on correctly identified strains.
Dothidotthiaceae Crous & A.J.L. Phillips 2008
Dothidotthiaceae was introduced to accommodate the
single genus Dothidotthia, which is characterized by gregarious, erumpent, globose ascomata, hyaline, septate pseudoparaphyses, 8-spored, bitunicate, clavate asci, ellipsoid, 1septate ascospores, and has anamorphic Thyrostroma
(Phillips et al. 2008). In this study, Dothidotthiaceae is
closely related to Didymellaceae, but it is still treated as a
separate family (Plate 1).
Hypsostromataceae Huhndorf 1994
Hypsostromataceae was introduced based on two tropical
genera (i.e. Hypsostroma and Manglicola), which have superficial, large, elongate ascomata with a soft-textured, pseudoparenchymatic wall, trabeculate pseudoparaphyses and stipitate asci
attached in a basal arrangement in the centrum; asci with an
apical chamber and fluorescing ring; and fusiform, septate
ascospores (Huhndorf 1994). Hypsostromataceae was assigned
to Melanommatales sensu Barr (Huhndorf 1994). In a
subsequent phylogenetic study, Hypsostromataceae was recovered as a strongly supported monophyletic group nested within
Pleosporales (Mugambi and Huhndorf 2009b).
Lentitheciaceae Yin. Zhang, C.L. Schoch, J. Fourn., Crous
& K.D. Hyde 2009
Phylogenetic analysis based on multi-genes indicate that
freshwater taxa, e.g. Lentithecium fluviatile, L. arundinaceum, Stagonospora macropycnidia, Wettsteinina lacustris,
Keissleriella cladophila, Katumotoa bambusicola and
Ophiosphaerella sasicola form a well supported clade, which
most likely represent a familial rank (Zhang et al. 2009a).
Their morphology, however, varies widely, e.g. ascomata
small- to medium-sized, ascospores fusoid to filliform,
hyaline to pale yellow, 1- to multi-septate (Zhang et al.
2009a). In particular, they are saprobic on monocotyledons
or dicotyledons. Currently, no conspicuous, unique morphological character has been noted in Lentitheciaceae, which
makes it difficult to recognize based on morphology.
Leptosphaeriaceae M.E. Barr 1987a
The Leptosphaeriaceae was introduced by Barr (1987a)
based on Leptosphaeria. The familial status of the Leptosphaeriaceae is subsequently supported by molecular phylogenetic studies, in which members of the Leptosphaeriaceae
form a paraphyletic clade with moderate bootstrap support
(Dong et al. 1998; de Gruyter et al. 2009; Schoch et al. 2009;
Zhang et al. 2009a). Coniothyrium palmarum, the generic
type of Coniothyrium nested within this family (de Gruyter
et al. 2009). Further molecular phylogenetic study is needed,
in which more related taxa are included.
Lindgomycetaceae K. Hirayama, Kaz. Tanaka & Shearer 2010
Lindgomycetaceae was introduced as a monotypic
family represented by Lindgomyces (Hirayama et al.
2010). Lindgomycetaceae is another freshwater family in
Pleosporales, which is characterized by its subglobose to
globose, ostiolate and papillate ascomata, numerous, septate, branching and anastomosing pseudoparaphyses, fissitunicate, cylindrical to clavate, 8-spored asci, fusiform to
cylindrical, uni- to multiseptate, hyaline to brown ascospores usually covered with an entire sheath and/or bipolar
mucilaginous appendages (Hirayama et al. 2010).
Lophiostomataceae Sacc. 1883
The Lophiostomataceae had been characterized by its
slot-like ostiole on the top of a flattened neck (Holm and
Holm 1988). Based on this, 11 genera were assigned under
the Lophiostomataceae, viz. Byssolophis, Cilioplea, Entodesmium, Herpotrichia, Lophiella, Lophionema, Lophiostoma, Lophiotrema, Massariosphaeria, Muroia and
Quintaria (Holm and Holm 1988). The Lophiostomataceae
was thought to be heterogeneous, as the “papilla form is an
unstable and highly adaptive character” (Holm and Holm
1988). Most recent phylogenetic analysis support the
monophyletic status of the Lophiostomataceae sensu stricto
(which tends to comprise a single genus of Lophiostoma)
(Zhang et al. 2009a, b). The familial placement of other
genera, however, remains unresolved.
Massarinaceae Munk 1956
The Massarinaceae was established based on Keissleriella,
Massarina, Metasphaeria, Pseudotrichia and Trichometasphaeria (Munk 1956). Subsequently, the Massarinaceae is
sometimes treated as a synonym of Lophiostomataceae (Barr
Fungal Diversity
1987b). Based on a multigene phylogenetic study, the generic
type of Massarina (M. eburnea) together with M. cisti,
Neottiosporina paspali and Byssothecium circinans form a
well supported clade (Zhang et al. 2009a, b). It seems that a
relatively narrow familial concept should be accepted.
Melanommataceae G. Winter 1885
The traditional circumscription of the Melanommataceae
was based on its globose or depressed perithecial ascomata,
bitunicate and fissitunicate asci, pigmented phragmosporous ascospores as well as the trabeculate pseudoparaphyses
(Barr 1990a; Sivanesan 1984). However, the family has
recently proved polyphyletic (Liew et al. 2000; Kodsueb et
al. 2006a; Kruys et al. 2006; Wang et al. 2007). Bimuria,
Ostropella, Trematosphaeria and Xenolophium occur outside Melanommataceae (Mugambi and Huhndorf 2009b;
Zhang et al. 2009a). Species of Byssosphaeria, Bertiella,
Herpotrichia, Pseudotrichia, Pleomassaria as well as
Melanomma resided in the clade of Melanommataceae
(Mugambi and Huhndorf 2009b; Schoch et al. 2009; Zhang
et al. 2009a). The familial status of many genera previously
listed under this family remains to be sorted out (Lumbsch
and Huhndorf 2007).
Montagnulaceae M.E. Barr 2001
The Montagnulaceae was introduced to accommodate
some pleosporalean genera with ascomata immersed under
a clypeus, a pseudoparenchymatous peridium with small
cells, cylindric or oblong asci with pedicels and brown
ascospores (Barr 2001). Three genera were included, i.e.
phragmosporous Kalmusia, dictyosporous Montagnula and
didymosporous Didymosphaerella (Barr 2001). Our molecular phylogenetic analysis based on multi-genes indicated
that species from Kalmusia, Phaeosphaeria, Bimuria,
Didymocrea, Paraphaeosphaeria, Karstenula, Letendraea
as well as Montagnula resided in the monophylogenetic
clade of the Montagnulaceae (Schoch et al. 2009; Zhang et
al. 2009a).
Morosphaeriaceae Suetrong, Sakay., E.B.G. Jones & C.L.
Schoch 2009
Four marine species, viz. Massarina ramunculicola
(as Morosphaeria ramunculicola), Massarina velataspora
(Morosphaeria velataspora), Helicascus kanaloanus and
H. nypae together with the freshwater species Kirschsteiniothelia elaterascus form a well supported clade, which
most likely represent a familial rank (Suetrong et al.
2009). Thus, Morosphaeriaceae was introduced to accommodate these taxa (Suetrong et al. 2009). In
this study, Asteromassaria pulchra is basal to other
species of Morosphaeriaceae, and gets well support
(Plate 1). Thus we tentatively assign Asteromassaria under
Morosphaeriaceae.
Phaeosphaeriaceae M.E. Barr 1979a
The Phaeosphaeriaceae was introduced to accommodate
some pleosporalean genera which have saprobic, parasitic or
hyperparasitic lifestyles and have small- to medium-sized,
subglobose or conical ascomata, bitunicate asci and hyaline or
pigmented ascospores with or without septation (Barr 1979a).
Fourteen genera were included, viz. Comoclathris, Didymella,
Eudarluca, Heptameria, Leptosphaeria, Loculohypoxylon,
Metameris, Microthelia, Nodulosphaeria, Ophiobolus, Paraphaeosphaeria, Rhopographus, Scirrhodothis and Teichospora (Barr 1979a), which were subsequently assigned to
various families, such as Loculohypoxylon and Teichospora to
the Teichosporaceae, Paraphaeosphaeria to the Montagnulaceae, Leptosphaeria to the Leptosphaeriaceae, Comoclathris
to the Diademaceae, Didymella to the Didymellaceae and
Heptameria and Rhopographus to genera incertae sedis of
Dothideomycetes (Aveskamp et al. 2010; de Gruyter et al.
2009; Lumbsch and Huhndorf 2007; Zhang et al. 2009a).
Based on multi-gene phylogenetic analysis, a relatively narrow
familial concept is accepted, which is mostly associated with
monocotyledons, with perithecoid, small- to medium-sized
ascomata, and septate ascospores which are fusiform to
filliform (Zhang et al. 2009a). Four genera were accepted,
Ophiosphaerella, Phaeosphaeria, Entodesmium and Setomelanomma (Zhang et al. 2009a). Together with Cucurbitariaceae,
Didymellaceae, Didymosphaeriaceae, Dothidotthiaceae, Leptosphaeriaceae and Pleosporaceae, the Phaeosphaeriaceae is
assigned under Pleosporineae (Zhang et al. 2009a).
Pleomassariaceae M.E. Barr 1979a
Both Asteromassaria and Splanchnonema were designated as representative genera of Pleomassariaceae (Barr
1979a). Currently, four genera are included in Pleomassariaceae, viz. ?Lichenopyrenis, ?Splanchnonema, ?Peridiothelia and Pleomassaria (Table 4). The generic type of
Pleomassaria (P. siparia) clustered with species of Melanommataceae in previous and present studies (Schoch et al.
2009; Zhang et al. 2009a; Plate 1). Zhang et al. (2009a) has
attempted to assign Pleomassariaceae to Melanommataceae
(Zhang et al. 2009a). Based on the distinct morphology and
anamorphic stage of Pleomassaria siparia as well as the
divergence of dendrogram, we hesitantely reinstate Pleomassariaceae as a separate family in this study.
Pleosporaceae Nitschke 1869
The Pleosporaceae is one of the earliest introduced
families in Dothideomycetes. The Pleosporaceae was
originally assigned under Sphaeriales, which accommodated species with paraphyses and immersed perithecia (Ellis
and Everhart 1892; Lindau 1897; Winter 1887). Subsequently, many of the Pleosporaceae species were transferred to the Pseudosphaeriaceae, which was subsequently
elevated to ordinal rank as Pseudosphaeriales (Theissen
Fungal Diversity
and Sydow 1918). Luttrell (1955) introduced the Pleosporales (lacking a Latin description), which is characterized by
its Pleospora-type of centrum development. Based on this,
the Pleosporaceae and the Lophiostomataceae as well as
other five families were placed in Pleosporales (Luttrell
1955). Pleosporaceae is the largest and most typical family
in Pleosporales. Wehmeyer (1975) stated that the Pleosporatype centrum development is verified in a small number of
genera, and centrum development in the majority of genera
is unknown; thus the placement of families or genera is quite
arbitrary. In addition, the circumscription of Pleosporaceae
is not clear-cut, and “……ascostromata of many different
types, which are previously placed in various other families
(Trichosphaeriaceae, Melanommataceae, Cucurbitariaceae,
Amphisphaeriaceae etc.) are to be found here” (Wehmeyer
1975). Thus, the heterogeneous nature of Pleosporales is
obvious (Eriksson 1981), and had been confirmed by
subsequent molecular phylogenetic studies (e.g. Kodsueb et
al. 2006a). Based on the multi-gene phylogenetic analysis,
some species from Lewia, Cochliobolus, Pleospora, Pyrenophora and Setosphaeria resided in the Pleosporaceae
(Zhang et al. 2009a).
Sporormiaceae Munk 1957
The Sporormiaceae is the largest coprophilous family in
Pleosporales, which bears great morphological variation.
Ascomata vary from cleistothecoid to perithecoid, asci are
regularly or irregularly arranged, clavate or spherical,
ascospores with or without germ slits or ornamentations.
Based on phylogenetic analysis, Sporormiaceae is most
likely monophyletic as currently circumscribed (Kruys et al.
2006; Kruys and Wedin 2009).
?Teichosporaceae M.E. Barr 2002
The Teichosporaceae was introduced by segregating some
non-lichenized members of the Dacampiaceae which are
apostrophic on woody stems and periderm or hypersaprotrophic on other ascomycetous fungi (Barr 2002). The Dacampiaceae together with its synonym, Pyrenidiaceae was only
maintained to accommodate its lichenicolous genera (Barr
2002). This proposal does not have any molecular phylogenetic support.
Tetraplosphaeriaceae Kaz. Tanaka & K. Hirayama 2009
The Tetraplosphaeriaceae was introduced to accommodate five genera, i.e. Tetraplosphaeria, Triplosphaeria,
Polyplosphaeria and the anamorphic genera Pseudotetraploa and Quadricrura (Tanaka et al. 2009). The Tetraplosphaeriaceae is characterized by its Massarina-like
teleomorphs and its Tetraploa-like anamorphs with setaelike appendages, and its monophylogenetic status has been
recently confirmed based on DNA phylogenetic studies
(Tanaka et al. 2009).
Trematosphaeriaceae
Three species, viz. Falciformispora lignatilis, Halomassarina thalassiae and Trematosphaeria pertusa form a
robust clade, which forms a sister group with other
pleosporalean families (Schoch et al. 2009; Suetrong et al.
2009). Trematosphaeriaceae is waiting to be formally
proposed (Suetrong et al. data unpublished).
?Zopfiaceae G. Arnaud ex D. Hawksw. 1992
The Zopfiaceae was introduced by Arnaud (1913), but
was invalid due to the lack of a Latin diagnosis (see
comments by Eriksson and Hawksworth 1992). The Zopfiaceae was formally introduced by Eriksson and Hawksworth
(1992), and is characterized by its cleistothecial ascomata,
thick-walled peridium, globose or saccate asci and oneseptate, dark brown ascospores (Cannon and Kirk 2007).
Currently, eleven genera are included, but the family is likely
polyphyletic (Kruys et al. 2006).
Excluded family
Phaeotrichaceae Cain 1956
The cleistothecioid ascomata, ascospores with germ pore
at each end and the absence of pseudoparaphyses indicate
that the Phaeotrichaceae may not be closely related to
Pleosporales. This was confirmed by DNA based phylogenies (Schoch et al. 2009). Thus, we exclude it from
Pleosporales.
Final remarks
Problems and concerns
Recently, many new pleosporalean lineages from freshwater (Shearer et al. 2009; Zhang et al. 2009a), marine
(Suetrong et al. 2009) or from bambusicolous hosts (Tanaka
et al. 2009) have been reported. In particular, large-scale
phylogenetic analysis indicate that numerous unresolved
clades still exist, which may also indicate that a large
number of fungal lineages are not resolved. As has been
estimated, 95% of all fungi are unreported (Hawksworth
1991), and a large portion of them might exist only as
hyphae (or DNA-only fungi, Taylor 1993). Under the
influence of human activities, environmental situations are
changing quickly, which may result in numerous fungal
taxa losing their habitats and/or become endangered. More
field work is urgently needed.
A future polyphasic approach to study Pleosporales
The use of DNA sequence comparisons have proved
invaluable in modern concepts of fungal taxonomy. It is now
clear many fungi do not produce reproductive structures or
Fungal Diversity
only do so under very rare circumstances and many fungi
cannot be cultured (Begerow et al. 2010). More and more
morpho-species have proven to be cryptic taxa, and already a
large percentage of fungal diversity is documented only by
DNA sequences. DNA sequence analysis is an essential way
to resolve these problems. But are they enough for fully
informed fungal taxonomy? Each single morphological
character may be the outcome of the expression of one to
numerous genes, which might be composed of thousands of
base pairs. DNA barcoding methods are “a breakthrough for
identification, but they will not supplant the need to
formulate and rigorously test species hypothesis” (Wheeler
et al. 2004). Thus, integration of classical morphological
approaches and DNA and protein based sequence comparisons are critical to produce a modern taxonomy that reflects
evolutionary similarities and differences (DeSalle et al. 2005;
Godfray 2002). In particular, the advent of comparative
genomics and advances in our understanding of secondary
metabolites and host or habitat spectra allow the possibility
to tie phylogenetic hypotheses derived from DNA and
protein sequence to the biology of the organisms. (Bitzer et
al. 2008; Stajich et al. 2009; Zhang et al. 2009a, b).
Acknowledgement We are grateful to the Directors and Curators of
the following herbaria for loan of specimens in their keeping: BAFC,
BISH, BPI, BR, BRIP, CBS, E, ETH, FFE, FH, G, H, Herb. J.
Kohlmeyer, HHUF, IFRD, ILLS, IMI, K(M), L, LPS, M, MA, NY,
PAD, PC, PH, RO, S, TNS, TRTC, UB, UBC, UPS and ZT; to Dr. L.
Cai, Dr. A.J.L. Phillips, Dr. C. Shearer and some other mycologists for
their permission to use or refer to their published figures, to J.K. Liu,
H. Zhang, Y.L. Yang and J. Fournier for helping me loan or collect
specimens, to H. Leung for technical help. The third coauthor
acknowledges the Intramural Research Program of the NIH, National
Library of Medicine. The Global Research Network and King Saud
University are also thanked for support.
Open Access This article is distributed under the terms of the
Creative Commons Attribution Noncommercial License which permits any noncommercial use, distribution, and reproduction in any
medium, provided the original author(s) and source are credited.
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