Pleosporales - CBS - KNAW
Pleosporales - CBS - KNAW
Pleosporales - CBS - KNAW
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Fungal Diversity<br />
DOI 10.1007/s13225-011-0117-x<br />
<strong>Pleosporales</strong><br />
Ying Zhang & Pedro W. Crous & Conrad L. Schoch &<br />
Kevin D. Hyde<br />
Received: 31 March 2011 /Accepted: 13 June 2011<br />
# The Author(s) 2011. This article is published with open access at Springerlink.com<br />
Abstract One hundred and five generic types of <strong>Pleosporales</strong><br />
are described and illustrated. A brief introduction and detailed<br />
history with short notes on morphology, molecular phylogeny<br />
as well as a general conclusion of each genus are provided.<br />
For those genera where the type or a representative specimen<br />
is unavailable, a brief note is given. Altogether 174 genera of<br />
<strong>Pleosporales</strong> are treated. Phaeotrichaceae as well as Kriegeriella,<br />
Zeuctomorpha and Muroia are excluded from<br />
<strong>Pleosporales</strong>. Based on the multigene phylogenetic analysis,<br />
the suborder Massarineae is emended to accommodate five<br />
families, viz. Lentitheciaceae, Massarinaceae, Montagnulaceae,<br />
Morosphaeriaceae and Trematosphaeriaceae.<br />
Y. Zhang<br />
Division of Microbiology, School of Biological Sciences,<br />
The University of Hong Kong,<br />
Pokfulam Road,<br />
Hong Kong, SAR, People’s Republic of China<br />
P. W. Crous<br />
<strong>CBS</strong>-<strong>KNAW</strong> Fungal Biodiversity Centre,<br />
P.O. Box 85167, 3508 AD Utrecht, The Netherlands<br />
C. L. Schoch<br />
National Center for Biotechnology Information,<br />
National Library of Medicine, National Institutes of Health,<br />
45 Center Drive, MSC 6510,<br />
Bethesda, MD 20892-6510, USA<br />
K. D. Hyde<br />
School of Science, Mae Fah Luang University,<br />
Tasud, Muang,<br />
Chiang Rai 57100, Thailand<br />
K. D. Hyde (*)<br />
Botany and Microbiology Department, College of Science,<br />
King Saud University,<br />
Riyadh 11442, Saudi Arabia<br />
e-mail: kdhyde3@gmail.com<br />
Keywords Generic type . Massarineae . Molecular<br />
phylogeny. Morphology . <strong>Pleosporales</strong> . Taxonomy<br />
Introduction<br />
Historic overview of <strong>Pleosporales</strong><br />
<strong>Pleosporales</strong> is the largest order in the Dothideomycetes,<br />
comprising a quarter of all dothideomycetous species (Kirk<br />
et al. 2008). Species in this order occur in various habitats,<br />
and can be epiphytes, endophytes or parasites of living<br />
leaves or stems, hyperparasites on fungi or insects, lichenized,<br />
or are saprobes of dead plant stems, leaves or bark<br />
(Kruys et al. 2006; Ramesh 2003).<br />
The Pleosporaceae was introduced by Nitschke (1869),<br />
and was assigned to Sphaeriales based on immersed<br />
ascomata and presence of pseudoparaphyses (Ellis and<br />
Everhart 1892; Lindau 1897; Wehmeyer 1975; Winter<br />
1887). Taxa in this family were then assigned to Pseudosphaeriaceae<br />
(Theissen and Sydow 1918; Wehmeyer 1975).<br />
Pseudosphaeriales, represented by Pseudosphaeriaceae,<br />
was introduced by Theissen and Sydow (1918), and was<br />
distinguished from Dothideales by its uniloculate, perithecioid<br />
ascostromata. Subsequently, the uni- or pluri-loculate<br />
ascostromata was reported to be an invalid character to<br />
separate members of Dothideomycetes into different orders<br />
(Luttrell 1955). In addition, the familial type of Pseudosphaeriales<br />
together with its type genus, Pseudosphaeria,was<br />
transferred to Dothideales, thusPseudosphaeriales became a<br />
synonym of Dothideales. The name “Pseudosphaeriales”<br />
has been applied in different senses, thus <strong>Pleosporales</strong> (as an<br />
invalid name due to the absence of a Latin diagnosis) was<br />
proposed by Luttrell (1955) to replace the confusing name,<br />
Pseudosphaeriales, which included seven families, i.e.
Fungal Diversity<br />
Botryosphaeriaceae, Didymosphaeriaceae, Herpotrichiellaceae,<br />
Lophiostomataceae, Mesnieraceae, Pleosporaceae<br />
and Venturiaceae. Müller and von Arx (1962) however,<br />
reused Pseudosphaeriales with 12 families included, viz.<br />
Capnodiaceae, Chaetothyriaceae, Dimeriaceae, Lophiostomataceae,<br />
Mesnieraceae, Micropeltaceae, Microthyriaceae,<br />
Mycosphaerellaceae, Pleosporaceae, Sporormiaceae, Trichothyriaceae<br />
and Venturiaceae.<br />
Familial circumscriptions of the <strong>Pleosporales</strong> were based<br />
on characters of ascomata, morphology of asci and their<br />
arrangement in locules, presence and type of hamathecium,<br />
shape of papilla or ostioles, morphology of ascospores and<br />
type of habitats (Luttrell 1973) (Table1). Based on these<br />
characters, Luttrell (1973) included eight families, i.e.<br />
Botryosphaeriaceae, Dimeriaceae, Lophiostomataceae, Mesnieraceae,<br />
Mycoporaceae, Pleosporaceae, Sporormiaceae<br />
and Venturiaceae in <strong>Pleosporales</strong>. In their review of bitunicate<br />
ascomycetes, von Arx and Müller (1975) accepted only a<br />
single order, Dothideales, with two suborders, i.e. Dothideineae<br />
(including Atichiales, Dothiorales, Hysteriales and<br />
Myriangiales) and Pseudosphaeriineae (including Capnodiales,<br />
Chaetothyriales, Hemisphaeriales, Lophiostomatales,<br />
Microthyriales, Perisporiales, <strong>Pleosporales</strong>, Pseudosphaeriales<br />
and Trichothyriales). This proposal has however, rarely<br />
been followed. Three existing families, i.e. Lophiostomataceae,<br />
Pleosporaceae and Venturiaceae plus 11 other families<br />
were accepted in <strong>Pleosporales</strong> as arranged by Barr (1979a)<br />
(largely using Luttrell’s concepts, Table 1), and she assigned<br />
these families to six suborders. The morphology of pseudoparaphyses<br />
was given much prominence at the ordinal level in<br />
this classification (Barr 1983). In particular the Melanommatales<br />
was introduced to accommodate taxa with trabeculate<br />
pseudoparaphyses (Sporormia-type centrum development)<br />
(Barr 1983), distinguished from cellular pseudoparaphyses<br />
(Pleospora-type centrum development) possessed by members<br />
of <strong>Pleosporales</strong> sensu Barr. The order Melanommatales<br />
included Didymosphaeriaceae, Fenestellaceae, Massariaceae,<br />
Melanommataceae, Microthyriaceae, Mytilinidiaceae,<br />
Platystomaceae and Requienellaceae (Barr 1990a).<br />
<strong>Pleosporales</strong> was formally established by Luttrell and Barr<br />
(in Barr 1987b), characterised by perithecioid ascomata,<br />
usually with a papillate apex, ostioles with or without<br />
periphyses, presence of cellular pseudoparaphyses, bitunicate<br />
asci, and ascospores of various shapes, pigmentation and<br />
septation (Table 1). Eighteen families were included, i.e.<br />
Arthopyreniaceae, Botryosphaeriaceae, Cucurbitariaceae,<br />
Dacampiaceae, Dimeriaceae, Hysteriaceae, Leptosphaeriaceae,<br />
Lophiostomataceae, Parodiellaceae, Phaeosphaeriaceae,<br />
Phaeotrichaceae, Pleomassariaceae, Pleosporaceae,<br />
Polystomellaceae, Pyrenophoraceae, Micropeltidaceae,<br />
Tubeufiaceae and Venturiaceae. Recent phylogenetic analysis<br />
based on DNA sequence comparisons, however, indicated<br />
that separation of the orders (<strong>Pleosporales</strong> and Melanommatales)<br />
based on the Pleospora or Sporormia centrum type, is<br />
not a natural grouping, and Melanommatales has therefore<br />
been combined under <strong>Pleosporales</strong> (Liew et al. 2000;<br />
Lumbsch and Lindemuth 2001; Reynolds 1991). Six more<br />
families, i.e. Cucurbitariaceae, Diademaceae, Didymosphaeriaceae,<br />
Mytilinidiaceae, Testudinaceae and Zopfiaceae,<br />
were subsequently added to <strong>Pleosporales</strong> (Lumbsch and<br />
Huhndorf 2007). After intensive sampling and multigene<br />
phylogenetic studies, 20 families were accepted in <strong>Pleosporales</strong>,<br />
namely Aigialaceae, Amniculicolaceae, Delitschiaceae,<br />
Didymellaceae, Didymosphaeriaceae, Hypsostromataceae,<br />
Lentitheciaceae, Leptosphaeriaceae, Lindgomycetaceae,<br />
Lophiostomataceae, Massarinaceae, Melanommataceae,<br />
Montagnulaceae, Morosphaeriaceae, Phaeosphaeriaceae,<br />
Pleosporaceae, Pleomassariaceae, Sporormiaceae, Tetraplosphaeriaceae<br />
and Trematosphaeriaceae (Boehm et al.<br />
2009a, b; Mugambi and Huhndorf 2009b;Schochetal.2009;<br />
Shearer et al. 2009; Suetrongetal.2009; Tanakaetal.2009;<br />
Zhang et al. 2009a) (Table 1). In addition, another five<br />
families, i.e. Arthopyreniaceae, Cucurbitariaceae, Diademaceae,<br />
Teichosporaceae and Zopfiaceae are tentatively included<br />
(Kruys et al. 2006; Plate 1). In the most recent issue of<br />
Myconet, 28 families were included in <strong>Pleosporales</strong><br />
(Lumbsch and Huhndorf 2010).<br />
Species included in <strong>Pleosporales</strong> have different ecological<br />
or morphological characters. For instance, members of<br />
the Leptosphaeriaceae have saprobic or parasitic lifestyles<br />
and lightly pigmented, multi-septate ascospores. Members<br />
of the Lophiostomataceae are mostly saprobic with ascomata<br />
that usually possess a compressed apex. Members of<br />
Sporormiaceae are coprophilous, and are characterized by<br />
heavily pigmented, multi-septate ascospores with germ<br />
slits, and with or without non-periphysate ostioles. The<br />
lack of DNA sequence data for representatives of numerous<br />
families means that their inter-relationships are unclear and<br />
many genera or species are artificially placed based on<br />
morphological classification. The most recent study on<br />
Venturiaceae indicated that this group had a set of unique<br />
morphological and ecological characters, which is distinct<br />
and distantly related to other members of <strong>Pleosporales</strong><br />
(Kruys et al. 2006; Zhang et al. unpublished). Molecular<br />
phylogenetic results indicated that members of Venturiaceae<br />
form a robust clade separate from the core members of<br />
<strong>Pleosporales</strong>, and the clade of Venturiaceae was uncertainly<br />
placed but outside of the two currently designated<br />
dothideomycetous subclasses, i.e. Pleosporomycetidae and<br />
Dothideomycetidae (Schoch et al. 2009). In addition,<br />
phylogenetic analysis of rDNA sequence data indicates<br />
that members of Zopfiaceae (as Testudinaceae) seem to<br />
lack affinity with <strong>Pleosporales</strong> (Kodsueb et al. 2006b).<br />
Thus, 26 families are temporarily accepted in <strong>Pleosporales</strong><br />
in this study, although some such as Zopfiaceae, still<br />
require extensive DNA sequence sampling (Table 4).
Fungal Diversity<br />
Table 1 Major circumscription changes of <strong>Pleosporales</strong> from 1955 to 2011<br />
References<br />
Luttrell 1955<br />
Müller and von Arx 1962<br />
Luttrell 1973<br />
Barr 1979a<br />
Barr 1987b<br />
Kirk et al. 2001, 2008<br />
Boehm et al. 2009a, b; Mugambi<br />
and Huhndorf 2009b; Schoch et<br />
al. 2009; Shearer et al. 2009;<br />
Suetrong et al. 2009; Tanaka et<br />
al. 2009; Zhang et al. 2009a<br />
Circumscriptions distinguishing <strong>Pleosporales</strong> from other orders of Dothideomycetes<br />
Pleospora-type centrum development.<br />
Ascomata perithecoid, with rounded or slit-like ostiole; asci produced within a locule, arranged regularly<br />
in a single layer or irregularly scattered, surrounded with filiform pseudoparaphyses, cylindrical,<br />
ellipsoidal or sac-like.<br />
Ascocarps perithecioid, immersed, erumpent to superficial on various substrates, asci ovoid to mostly<br />
clavate or cylindrical, interspersed with pseudoparaphyses (sometimes form an epithecium) in mostly<br />
medium- to large-sized locules.<br />
Saprobic, parasitic, lichenized or hypersaprobic. Ascomata perithecioid, rarely cleistothecioid or<br />
hysterothecioid, peridium pseudoparenchymatous, pseudoparaphyses cellular, narrow or broad,<br />
deliquescing early at times, not forming an epithecium, asci oblong, clavate or cylindrical, interspersed<br />
with pseudoparaphyses, ascospores mostly asymmetric.<br />
Saprobic, biotrophic or hemibiotrophic. Ascomata globose, subglobose or conical, asci bitunicate, oblong,<br />
clavate or cylindrical, cellular pseudoparaphyses, ascospores hyaline or pigmented, asymmetric or<br />
symmetric, with or without septa.<br />
Ascomata perithecioid or rarely cleistothecioid, sometimes clypeate, mostly globose, thick-walled,<br />
immersed or erumpent, black, sometimes setose, peridium composed of pseudoparenchymatous cells,<br />
pseudoparaphyses trabeculate or cellular, asci cylindrical, fissitunicate, with a well-developed ocular<br />
chamber, rarely with a poorly defined ring (J-), ascospores hyaline to brown, septate, thin or thickwalled,<br />
sometimes muriform, usually with sheath, anamorphs hyphomycetous or coelomycetous.<br />
Hemibiotrophic, saprobic, hypersaprobic, or lichenized. Habitats in freshwater, marine or terrestrial<br />
environment. Ascomata perithecioid, rarely cleistothecioid, immersed, erumpent to superficial, globose<br />
to subglobose, or lenticular to irregular, with or without conspicuous papilla or ostioles. Ostioles with or<br />
without periphyses. Peridium usually composed of a few layers of cells with various shapes and<br />
structures. Hamathecium persistent, filamentous, very rarely decomposing. Asci bitunicate, fissitunicate,<br />
cylindrical, clavate to obclavate, with or without pedicel. Ascospores hyaline or pigmented, ellipsoidal,<br />
broadly to narrowly fusoid or filiform, mostly septate.<br />
Morpho-characters used in taxonomy of <strong>Pleosporales</strong><br />
Sexual characters<br />
According to the Linnean classification system, reproductive<br />
structures are the most important criteria in plant<br />
taxonomy, and this proposal is widely applied in fungal<br />
taxonomy (Gäumann 1952). In the classification of Dothideomycetes,<br />
reproductive characters such as the uni- or<br />
multilocular nature and shape of ascomata, presence and<br />
shape of ostioles/papillae, shape and apical structures of<br />
asci and shape, pigmentation and septation of ascospores<br />
play important roles at different ranks (Clements and Shear<br />
1931; Luttrell 1951, 1955, 1973). Besides the common<br />
morphological characters possessed by Dothideomycetes<br />
(bitunicate and fissitunicate asci as well as the perithecioidlike<br />
ascostromata), most pleosporalean fungi also have<br />
pseudoparaphyses among their well-arranged asci (Zhang et<br />
al. 2009a). Currently, classification of <strong>Pleosporales</strong> at the<br />
family level focuses mostly on morphological characters of<br />
ascomata (such as size, shape of ostiole or papilla), presence<br />
or absence of periphyses, characters of centrum (such as asci,<br />
pseudoparaphyses and ascospores) as well as on lifestyle or<br />
habitat (Barr 1990a; Shearer et al. 2009; Suetrong et al.<br />
2009; Tanaka et al. 2009; Zhang et al. 2009a), whilst relying<br />
extensively on DNA sequence comparisons.<br />
Ascomata<br />
Most species of <strong>Pleosporales</strong> have uniloculate ascomata.<br />
The presence (or absence) and forms of papilla and ostiole<br />
are the pitoval character of ascomata, which serve as<br />
important characteristics in generic or higher rank classification<br />
(Clements and Shear 1931). The vertically flattened papilla has<br />
recently been shown as an effective criterion for familial level<br />
classification, e.g. in the Amniculicolaceae and the Lophiostomataceae<br />
(Zhang et al. 2009a). Papillae and ostioles are<br />
present in most species of <strong>Pleosporales</strong>, except in the<br />
Diademaceae and Sporormiaceae. Members of Diademaceae<br />
have apothecial ascomata, and some genera of Sporormiaceae<br />
have cleistothecioid ascomata. Another coprophilous pleosporalean<br />
family, Delitschiaceae, can be distinguished from<br />
Sporormiaceae by the presence of periphysate ostioles.<br />
Pseudoparaphyses<br />
Presence of pseudoparaphyses is a characteristic of<br />
<strong>Pleosporales</strong> (Kirk et al. 2008; Liew et al. 2000). Although<br />
pseudoparaphyses may be deliquescing in some families<br />
when the ascomata mature (e.g. in Didymellaceae), they are<br />
persistent in most of other pleosporalean members. According<br />
to the thickness, with or without branching and density<br />
of septa, pseudoparaphyses were roughly divided into two<br />
types: trabeculate and cellular, and their taxonomic significance<br />
need to be re-evaluated (Liew et al. 2000).
Fungal Diversity<br />
Asci<br />
The asci of <strong>Pleosporales</strong> are bitunicate, usually fissitunicate,<br />
mostly cylindrical, clavate or cylindro-clavate, and<br />
rarely somewhat obclavate or sphaerical (e.g. Macroventuria<br />
anomochaeta Aa and Westerdykella dispersa). There are<br />
ocular chambers in some genera (e.g. Amniculicola and<br />
Asteromassaria), or sometimes with a large apical ring (J-)<br />
(e.g. Massaria).<br />
Ascospores<br />
Ascospores of <strong>Pleosporales</strong> canbehyalineorcolored<br />
to varying degrees. They may be amerosporous (e.g.<br />
species of Semidelitschia), phragmosporous (e.g. Phaeosphaeria<br />
and Massariosphaeria), dictyosporous (e.g. most<br />
species of Pleospora and Bimuria), or scolecosporous<br />
(e.g. type species of Cochliobolus, Entodesmium or<br />
Lophionema). Although ascospore morphology had been<br />
regarded as a key factor in differentiating genera under<br />
some families, e.g. Arthopyreniaceae (Watson 1929) and<br />
Testudinaceae (Hawksworth 1979), it has been proven<br />
variable even within a single species. For instance, two<br />
types of ascospores are produced by Mamillisphaeria<br />
dimorphospora, i.e. one type is large and hyaline, and the<br />
other is comparatively smaller and brown. Numerous<br />
studies have shown the unreliability of ascospore characters<br />
above genus level classification (e.g. Phillips et al.<br />
2008; Zhang et al. 2009a).<br />
Asexual states of <strong>Pleosporales</strong><br />
Anamorphs of pleosporalean families<br />
Anamorphs of <strong>Pleosporales</strong> are mostly coelomycetous,<br />
but may also be hyphomycetous. Phoma or Phoma-like<br />
anamorphic stages and its relatives are most common<br />
anamorphs of <strong>Pleosporales</strong> (Aveskamp et al. 2010; de<br />
Gruyter et al. 2009, 2010; Hyde et al. 2011). Some of the<br />
reported teleomorph and anamorph connections (including<br />
some listed below) are, however, based on the association<br />
rather than single ascospore isolation followed by induction<br />
of the other stage in culture (Hyde et al. 2011).<br />
<strong>Pleosporales</strong> suborder Pleosporineae<br />
Pleosporineae is a phylogenetically well supported<br />
suborder of <strong>Pleosporales</strong>, which temporarily includes<br />
seven families, namely Cucurbitariaceae, Didymellaceae,<br />
Didymosphaeriaceae, Dothidotthiaceae, Leptosphaeriaceae,<br />
Phaeosphaeriaceae and Pleosporaceae, and<br />
contains many important plant pathogens (de Gruyter et<br />
al. 2010; Zhangetal.2009a). De Gruyter et al. (2009,<br />
2010) systematically analyzed the phylogeny of Phoma<br />
and its closely related genera, and indicated that their<br />
representative species cluster in different subclades of<br />
Pleosporineae.<br />
Plate 1 The best scoring likelihood tree of representative <strong>Pleosporales</strong><br />
obtained with RAxML v. 7.2.7 for a concatenated set of<br />
nucleotides from LSU, SSU, RPB2 and TEF1. Family and suborder<br />
names are indicated where possible. The percentages of nodes present<br />
in 250 bootstrap pseudo replicates are shown above branches. Culture<br />
and voucher numbers are indicated after species names and the<br />
presence of the genes used in the analysis are indicated by pluses in<br />
this order: LSU, SSU, RPB2, TEF1<br />
Cucurbitariaceae<br />
Based on the molecular phylogenetic analysis, some<br />
species of Coniothyrium, Pyrenochaeta, Phoma, Phialophorophoma<br />
and Pleurophoma belong to Cucurbitariaceae (de<br />
Gruyter et al. 2010; Hyde et al. 2011). Other reported<br />
anamorphs of Cucurbitaria are Camarosporium, Diplodialike<br />
and Pleurostromella (Hyde et al. 2011; Sivanesan 1984).<br />
The generic type of Cucurbitaria (C. berberidis Fuckel) is<br />
linked to Pyrenochaeta berberidis (Farr et al. 1989). Curreya<br />
has a Coniothyrium-like anamorphic stage (von Arx and van<br />
der Aa 1983; Marincowitz et al. 2008). The generic type of<br />
Curreya is C. conorum (Fuckel) Sacc., which is reported to<br />
be linked with Coniothyrium glomerulatum Sacc. (von Arx<br />
and van der Aa 1983). The generic type of Rhytidiella (R.<br />
moriformis, Cucurbitariaceae) can cause rough-bark of<br />
Populus balsamifera, andhasaPhaeoseptoria anamorphic<br />
stage (Zalasky 1968). Rhytidiella baranyayi Funk & Zalasky,<br />
another species of Rhytidiella associated with the cork-bark<br />
disease of aspen is linked with Pseudosporella-like anamorphs<br />
(Funk and Zalasky 1975; Sivanesan 1984).<br />
Didymellaceae, Didymosphaeriaceae and Dothidotthiaceae<br />
As has been mentioned before, Phoma sensu lato species<br />
have been proved to be highly polyphyletic, and they cluster<br />
in six distinct familial clades within the <strong>Pleosporales</strong><br />
(Aveskamp et al. 2010). Most Phoma species, including the<br />
generic type (P. herbarum), clustered in Didymellaceae<br />
(Aveskamp et al. 2010). The clade of Didymellaceae also<br />
comprises other sections, such as Ampelomyces, Boeremia,<br />
Chaetasbolisia, Dactuliochaeta, Epicoccum, Peyronellaea,<br />
Phoma-like, Piggotia, Pithoascus, as well as the type species<br />
of Ascochyta and Microsphaeropsis (Aveskamp et al. 2010;<br />
de Gruyter et al. 2009; Kirketal.2008; Sivanesan 1984).<br />
Leptosphaerulina is another genus of Didymellaceae, which<br />
has hyphomycetous anamorphs with pigmented and<br />
muriform conidia, such as Pithomyces (Roux 1986).<br />
The other reported anamorphs of Didymosphaeria are<br />
Fusicladiella-like, Dendrophoma, Phoma-like (Hyde et al.<br />
2011). Hyphomycetous Thyrostroma links to Dothidotthiaceae<br />
(Phillips et al. 2008).<br />
Some important plant pathogens are included within<br />
Didymellaceae, such as Phoma medicaginis Malbr. &<br />
Roum., which is a necrotrophic pathogen on Medicago<br />
truncatula (Ellwood et al. 2006). Phoma herbarum is<br />
another plant pathogen, which has potential as a
Fungal Diversity
Plate 1 (continued)<br />
Fungal Diversity
Fungal Diversity<br />
biocontrol agent of weeds (Neumann and Boland 2002).<br />
Ascochyta rabiei is a devastating disease of chickpea in<br />
most of the chickpea producing countries (Saxena and<br />
Singh 1987).<br />
Leptosphaeriaceae<br />
The anamorphic stages of Leptosphaeriaceae can be<br />
Coniothyrium, Phoma, Plenodomus and Pyrenochaeta. All<br />
are coelomycetous anamorphs, and they may have phialidic<br />
or annellidic conidiogenous cells. Phoma heteromorphospora<br />
Aa & Kesteren, the type species of Phoma sect.<br />
Heterospora and Coniothyrium palmarum, the generic type<br />
of Coniothyrium, reside in Leptosphaeriaceae (de Gruyter<br />
et al. 2009).<br />
Pleosporaceae<br />
Various anamorphic types can occur in Pleosporaceae,<br />
which can be coelomycetous or hyphomycetous, and the<br />
ontogeny of conidiogenous cells can be phialidic, annellidic<br />
or sympodial blastic. Both Ascochyta caulina and Phoma<br />
betae belong to Pleosporaceae (de Gruyter et al. 2009).<br />
Some species of Bipolaris and Curvularia are anamorphs<br />
of Cochliobolus. Many species of these two<br />
genera cause plant disease or even infect human beings<br />
(Khan et al. 2000). They are hyphomycetous anamorphs<br />
with sympodial proliferating conidiogenous cells, and<br />
pigmented phragmosporous poroconidia. The generic type<br />
of Lewia (L. scrophulariae) is linked with Alternaria<br />
conjuncta E.G. Simmons (Simmons 1986), and the<br />
generic type of Pleospora (P. herbarum) is linked with<br />
Stemphylium botryosum Sacc. (Sivanesan 1984). Both<br />
Alternaria and Stemphylium are hyphomycetous anamorphs<br />
characterized by pigmented, muriform conidia that develop<br />
at a very restricted site in the apex of distinctive conidiophores<br />
(Simmons 2007).<br />
The generic type of Pleoseptum (P. yuccaesedum) is<br />
linked with Camarosporium yuccaesedum (Ramaley and<br />
Barr 1995), the generic type of Macrospora (M. scirpicola)<br />
with Nimbya scirpicola (Fuckel) E.G. Simmons (Simmons<br />
1989), and the generic type of Setosphaeria (S. turcica)<br />
with Drechslera turcica (Pass.) Subram. & B.L. Jain<br />
(Sivanesan 1984). Pyrenophora has the anamorphic stages<br />
of Drechslera, and the anamorphic stage of Wettsteinina<br />
can be species of Stagonospora (Farr et al. 1989).<br />
Most common anamorphs in Pleosporaceae are Alternaria,<br />
Bipolaris, Phoma-like and Stemphylium, and they<br />
can be saprobic or parasitic on various hosts. Phoma betae<br />
A.B. Frank is a notorious pathogen on sugar beet, which<br />
causes zonate leaf spot or Phomopsis of sugar beet.<br />
Alternaria porri (Ellis) Cif., Stemphylium solani G.F.<br />
Weber, S. botryosum and S. vesicarium (Wallr.) E.G.<br />
Simmons can cause leaf blight of garlic (Zheng et al.<br />
2009). Phoma incompta Sacc. & Martelli is a pathogen on<br />
olive, and Stemphylium botryosum, the anamorph of<br />
Pleospora herbarum, causes leaf disease of olive trees<br />
(Malathrakis 1979).<br />
Phaeosphaeriaceae<br />
The type species of Phoma sect. Paraphoma (Phoma<br />
radicina (McAlpine) Boerema) as well as several pathogens<br />
on Gramineae, i.e. Stagonospora foliicola (Bres.) Bubák, S.<br />
neglecta var. colorata and Wojnowicia hirta Sacc.belongto<br />
Phaeosphaeriaceae (de Gruyter et al. 2009). Other anamorphs<br />
reported for Phaeosphaeriaceae are Amarenographium,<br />
Ampelomyces, Chaetosphaeronema, Coniothyrium,<br />
Hendersonia, Neosetophoma, ?Parahendersonia, Paraphoma,<br />
Phaeoseptoria, Rhabdospora, Scolecosporiella, Setophoma,<br />
Sphaerellopsis and Tiarospora.<br />
These anamorphic fungi can be saprobic, but mostly<br />
pathogenic on herbaceous plants. For instance, Stagonospora<br />
foliicola and Coniothyrium concentricum (Desm.)<br />
Sacc. can cause leaf spots on herbaceous plants (Zeiders<br />
1975), and Ampelomyces quisqualis Ces. is a hyperparasite<br />
of powdery mildews.<br />
<strong>Pleosporales</strong> suborder Massarineae<br />
Massarineae species are mostly saprobic in terrestrial or<br />
aquatic environments. Five families are currently included<br />
within Massarineae, viz. Lentitheciaceae, Massarinaceae,<br />
Montagnulaceae, Morosphaeriaceae and Trematosphaeriaceae.<br />
Anamorphs of the five families are summarized as<br />
follows.<br />
Lentitheciaceae<br />
Stagonospora macropycnidia Cunnell nests within the<br />
clade of Lentitheciaceae (Plate 1). A relatively broad genus<br />
concept of Stagonospora is currently accepted, which<br />
comprises parasitic or saprobic taxa. Keissleriella cladophila<br />
(Niessl) Corbaz is another species nesting within<br />
Lentitheciaceae (Zhang et al. 2009a), and is linked with<br />
Dendrophoma sp., which has branching conidiogenous<br />
cells, and 1-celled, hyaline conidia (Bose 1961; Sivanesan<br />
1984).<br />
Massarinaceae<br />
A relatively narrow concept tends to be accepted for<br />
Massarinaceae, which seems only to comprise limited<br />
species such as Byssothecium circinans, Massarina eburnea,<br />
M. cisti S.K. Bose, M. igniaria (C. Booth) Aptroot<br />
(anamorph: Periconia igniaria E.W. Mason & M.B. Ellis)<br />
and Neottiosporina paspali (G.F. Atk.) B. Sutton & Alcorn<br />
(Zhang et al. 2009a; Plate 1). Similarly, a relatively narrow<br />
generic concept of Massarina was accepted, containing<br />
only M. eburnea and M. cisti (Zhang et al. 2009b), and both<br />
species have been linked with species of Ceratophoma<br />
(Sivanesan 1984).
Fungal Diversity<br />
Montagnulaceae<br />
Montagnula has an Aschersonia anamorph, and Kalmusia<br />
and Paraphaeosphaeria have Coniothyrium-like, Cytoplea,<br />
Microsphaeropsis and Paraconiothyrium anamorphs.<br />
The generic type of Paraphaeosphaeria (P. michotii) is<br />
linked with Coniothyrium scirpi Trail (Webster 1955). The<br />
Coniothyrium complex is highly polyphyletic, and was<br />
subdivided into four groups by Sutton (1980), viz.<br />
Coniothyrium, Microsphaeropsis, Cyclothyrium and Cytoplea.<br />
Paraconiothyrium was introduced to accommodate<br />
Coniothyrium minitans W.A. Campb. and C. sporulosum<br />
(W. Gams & Domsch) Aa, which are closely related to<br />
Paraphaeosphaeria based on 18S rDNA sequences phylogeny<br />
(Verkley et al. 2004).<br />
Morosphaeriaceae<br />
Based on the multigene phylogenetic analysis in this<br />
study, Asteromassaria is tentatively included in Morosphaeriaceae.<br />
Asteromassaria macrospora is linked with<br />
Scolicosporium macrosporium (Berk.) B. Sutton, which is<br />
hyphomycetous. No anamorphic stages have been reported<br />
for other species of Morosphaeriaceae.<br />
Trematosphaeriaceae<br />
Three species from three different genera were included<br />
in Trematosphaeriaceae, i.e. Falciformispora lignatilis,<br />
Halomassarina thalassiae and Trematosphaeria pertusa<br />
(Suetrong et al. data unpublished; Plate 1). Of these, only<br />
Trematosphaeria pertusa, the generic type of Trematosphaeria,<br />
produces hyphopodia-like structures on agar<br />
(Zhang et al. 2008a).<br />
Other families of <strong>Pleosporales</strong><br />
Amniculicolaceae<br />
Three anamorphic species nested within the clade of<br />
Amniculicolaceae, i.e. Anguillospora longissima (Sacc. &<br />
P. Syd.) Ingold, Repetophragma ontariense (Matsush.) W.<br />
P. Wu and Spirosphaera cupreorufescens Voglmayr<br />
(Zhang et al. 2009a). Sivanesan (1984, p.500)described<br />
the teleomorphic stage of Anguillospora longissima as<br />
Massarina sp. II, which fits the diagnostic characters of<br />
Amniculicola well. Thus this taxon may be another species of<br />
Amniculicola.<br />
Hypsostromataceae<br />
A Pleurophomopsis-like anamorph is reported in the<br />
subiculum of the generic type of Hypsostroma (H. saxicola<br />
Huhndorf) (Huhndorf 1992).<br />
Lophiostomataceae<br />
The concept of Lophiostomataceae was also narrowed,<br />
and presently contains only Lophiostoma (Zhang et al.<br />
2009a). Leuchtmann (1985) studied cultures of some<br />
Lophiostoma species, and noticed that L. caulium (Fr.)<br />
Ces. & De Not., L. macrostomum, L. semiliberum (Desm.)<br />
Ces. & De Not., Lophiostoma sp. and Lophiotrema nucula<br />
produced Pleurophomopsis anamorphic stages, which are<br />
similar to those now in Melanomma (Chesters 1938), but<br />
Lophiostoma and Melanomma has no proven phylogenetic<br />
relationship (Zhang et al. 2009a, b; Plate 1). Species of<br />
Aposphaeria have also been reported in Massariosphaeria<br />
(Farr et al. 1989; Leuchtmann 1984), but the polyphyletic<br />
nature of Massariosphaeria is well documented (Wang et<br />
al. 2007).<br />
Melanommataceae<br />
The anamorphs of the Melanommataceae are mostly<br />
coelomycetous and rarely hyphomycetous with various<br />
ontogenic structures, such as annellidic or sympodial for<br />
hyphomycetes (Exosporiella and Pseudospiropes) and<br />
coelomycetes (Aposphaeria-like and Pyrenochaeta).<br />
Herpotrichia is reported as having a Pyrenochaeta<br />
anamorphic stage with or without seta on the surface of<br />
pycnidia (Sivanesan 1984). Aposphaeria and Phoma-like<br />
have been reported in Melanomma species (Chesters<br />
1938; Sivanesan 1984). Similarly, the anamorphs of<br />
Karstenula are reported as coelomycetous, i.e. Microdiplodia<br />
(Constantinescu 1993). The anamorphic stage of<br />
Anomalemma is Exosporiella (Sivanesan 1983), and that<br />
of Byssosphaeria is Pyrenochaeta (Barr 1984). Ohleria<br />
brasiliensis Starbäck has been linked with Monodictys<br />
putredinis (Wallr.) S. Hughes (Samuels 1980). Astrosphaeriella<br />
is a contentious genus as its familial status is<br />
not determined yet. Here we temporarily assigned it under<br />
Melanommataceae, which is linked with the anamorph<br />
genus Pleurophomopsis.<br />
Pleomassariaceae<br />
Shearia and Prosthemium are all anamorphs of Pleomassaria,<br />
andProsthemium betulinum is linked with the generic<br />
type of Pleomassaria (P. siparia) (Barr1982b; Sivanesan<br />
1984; Sutton 1980; Tanaka et al. 2010). Splanchnonema is a<br />
genus of Pleomassariaceae, the teleomorphic morphology of<br />
which is difficult to distinguish from two other genera, i.e.<br />
Asteromassaria and Pleomassaria, and the reported anamorphs<br />
of Splanchnonema are Ceuthodiplospora, Myxocyclus<br />
and Stegonsporium, which are comparable with those of<br />
Asteromassaria and Pleomassaria.<br />
Tetraplosphaeriaceae<br />
Tetraplosphaeriaceae was introduced to accommodate<br />
the Massarina-like bambusicolous fungi that produce<br />
Tetraploa sensu stricto anamorphs (Tanaka et al. 2009).<br />
Tetraploa aristata Berk. & Broome, the generic type of<br />
Tetraploa is widely distributed, associated with various
Fungal Diversity<br />
substrates and many occur in freshwater or has been<br />
isolated from air. The polyphyletic nature of T. aristata<br />
has been well documented (Tanaka et al. 2009). Anamorphic<br />
stages can serve as a diagnostic character for this<br />
family.<br />
Diademaceae, Massariaceae, Sporormiaceae<br />
and Teichosporaceae<br />
The Sporormiaceae is coprophilous having Phoma or<br />
Phoma-related anamorphic states (Cannon and Kirk 2007).<br />
Comoclathris (Diademaceae) is linked with Alternaria-like<br />
anamorphs (Simmons 1952). Myxocyclus links to Massaria<br />
(Massariaceae) (Hyde et al. 2011). The anamorphic stage of<br />
Chaetomastia (Teichosporaceae) isAposphaeria- orConiothyrium-like<br />
(Barr 1989c).<br />
Generally speaking, the morphologically simple conidiophores<br />
are usually considered phylogenetically uninformative<br />
(Seifert and Samuels 2000). Phoma-like anamorphs commonly<br />
occur in <strong>Pleosporales</strong>, while their colorless and unicellular<br />
conidia are also not phylogenetically informative (Seifert and<br />
Samuels 2000).<br />
All of the above mentioned anamorphic taxa of<br />
<strong>Pleosporales</strong> have phialidic, annellidic or sympodial conidiogenous<br />
cells, representing apical wall-building type<br />
(compared to ring wall-building and diffused wallbuilding)<br />
(Nag Raj 1993), which may indicate that the<br />
wall-building type probably has phylogenetic significance.<br />
Molecular phylogeny of <strong>Pleosporales</strong><br />
Numerous genes have been applied in phylogenetic<br />
studies of <strong>Pleosporales</strong>, mostly including LSU, SSU,<br />
mtSSU and ITS as well as the protein genes, such as<br />
RPB1, RPB2, TEF1, β-tubulin (TUB1) and actin (ACT1).<br />
A single gene such as ITS or LSU, has been used to study<br />
phylogenetic relationships between Leptosphaeria and<br />
Phaeosphaeria (Câmara et al. 2002) or Pleosporaceae<br />
and Tubeufiaceae (Kodsueb et al. 2006a, b) (Table2). The<br />
use of these phylogenetic markers, although making<br />
important contributions, has not been successful in<br />
resolving numerous relationships in single gene dendrograms.<br />
One exception is the use of SSU sequences to<br />
demonstrate the phylogenetic significance of pseudoparaphyses<br />
(Liew et al. 2000) whilst rejecting the phylogenetic<br />
utility of pseudoparaphyses morphology (cellular or<br />
trabeculate). Analyses with combined genes have had<br />
more success. For instance combined analyses with LSU<br />
and SSU sequence data could be used to define family<br />
level classification in a few cases (Dong et al. 1998; de<br />
Gruyter et al. 2009; Lumbsch and Lindemuth 2001; Pinnoi<br />
et al. 2007; Zhang et al. 2009b) (Table2). The addition of<br />
more than two genes has been used to determine relationships<br />
between orders. For instance, genes such as LSU,<br />
SSU and mtSSU have been used to analyze ordinal<br />
relationships in Loculoascomycetes (Lindemuth et al.<br />
2001), and to analyze phylogenetic relationships of<br />
coprophilous families in <strong>Pleosporales</strong> (Kruys et al.<br />
2006). Phaeocryptopus gaeumannii (T. Rohde) Petr. was<br />
showntobelonginDothideales based on LSU, SSU and<br />
ITS sequence analysis (Winton et al. 2007), while Schoch<br />
et al. (2006) used four genes, i.e. LSU, SSU, RPB2 and<br />
TEF1 to evaluate the phylogenetic relationships among<br />
different orders of the Dothideomycetes. Fivegenes,viz.<br />
LSU, SSU, TEF1, RPB1 andRPB2, were used to study the<br />
phylogenetic relationships of different orders within<br />
Dothideomycetes (Schoch et al. 2009) and of different<br />
families within <strong>Pleosporales</strong> (Zhang et al. 2009a)<br />
(Table 2). It is clear that even more genes will be required<br />
to address the remaining issues and the promise of genome<br />
analyses is within reach (www.jgi.doe.gov/sequencing/<br />
why/dothideomycetes.html) forDothideomycetes.<br />
The importance of generic type specimens<br />
The type specimen (collection type) is a fundamental<br />
element in the current Code of Botanical Nomenclature at<br />
familial or lower ranks (Moore 1998). A type specimen<br />
fixes the name to an exact specimen at family, genera,<br />
species and variety/subspecies rank and is ultimately based<br />
on this single specimen, i.e. a family name is based on a<br />
genus, the genus name is based on a species, and the<br />
species name is based on a specimen (Kirk et al. 2008).<br />
The generic type is of great importance in defining<br />
generic circumscriptions in fungal taxonomy. The generic<br />
types of <strong>Pleosporales</strong> have been studied previously by<br />
many mycologists. For instance, Müller and von Arx<br />
(1962) studied the generic types of “Pyrenomycetes”, and<br />
described and illustrated them in detail. Sivanesan (1984)<br />
described and illustrated the generic representatives of<br />
Loculoascomycetes for both their teleomorphs and anamorphs,<br />
and their links were emphasized. A large number<br />
of pleosporalean genera have been studied by Barr (1990a,<br />
b). Almost all of the previous work was conducted more<br />
than 20 years ago, when no molecular phylogenetic studies<br />
could be carried out and thus had been carried out in a<br />
systematic fashion.<br />
Aim and outline of present study<br />
The present study had two principal objectives:<br />
1. To explore genera under <strong>Pleosporales</strong> based on the<br />
generic types and provide a detailed description and<br />
illustration for the type species of selected genera,<br />
discuss the study history of those genera, and explore<br />
their ordinal, familial, and generic relationships;
Fungal Diversity<br />
Table 2 List of phylogenetic studies on <strong>Pleosporales</strong><br />
Year Author(s) Loci used Target fungi General conclusion<br />
1998 Dong et al. LSU, SSU Leptosphaeriaceae, Pleosporaceae<br />
and three other families<br />
Leptosphaeriaceae is paraphyletic and<br />
Pleosporaceae is monophyletic.<br />
2000 Liew et al. SSU <strong>Pleosporales</strong> and Melanommatales <strong>Pleosporales</strong> and Melanommatales are not<br />
naturial groups.<br />
2001 Lindemuth et al. LSU, SSU, mtSSU loculoascomycetes Loculoascomycetes are not monophyletic.<br />
2001 Lumbsch and<br />
Lindemuth<br />
LSU, SSU Dothideomycetes Presence of pseudoparaphyses is a major<br />
character at order level classification<br />
2002 Câmara et al. ITS Leptosphaeria and Phaeosphaeria Accepted Leptosphaeria sensu stricto.<br />
2006 Kodsueb et al. LSU Pleosporaceae Wettsteinina should be excluded from the<br />
Pleosporaceae.<br />
2006 Kodsueb et al. LSU Tubeufiaceae Tubeufiaceae is more closely related to the<br />
Venturiaceae.<br />
2006 Kruys et al. LSU, SSU, mtSSU coprophilous familes of<br />
<strong>Pleosporales</strong><br />
2006 Schoch et al. LSU, SSU, TEF1,<br />
RPB2<br />
Dothideomycetes<br />
coprophilous familes of <strong>Pleosporales</strong> form<br />
phylogenetic monophyletic groups<br />
respectively<br />
Proposed the subclasses<br />
Pleosporomycetidae<br />
2007 Pinnoi et al. LSU, SSU <strong>Pleosporales</strong> phylogenetic relationships of different<br />
families of <strong>Pleosporales</strong>, introduced a<br />
new fungus–– Berkleasmium crunisia<br />
2007 Wang et al. LSU, SSU, RPB2 Massariosphaeria Massariosphaeria is not monophyletic<br />
2007 Winton et al. LSU, SSU, ITS Phaeocryptopus gaeumannii Phaeocryptopus gaeumannii nested in<br />
Dothideales.<br />
2008a Zhang et al. LSU, SSU Melanomma and Trematosphaeria Melanomma and Trematosphaeria belong<br />
to different families<br />
2009 de Gruyter et al. LSU, SSU; Phoma and related genera They are closely related with<br />
Didymellaceae, Leptosphaeriaceae,<br />
Phaeosphaeriaceae and Pleosporaceae<br />
2009a Zhang et al. LSU, SSU, TEF1,<br />
RPB1, RPB2<br />
2009 Mugambi and<br />
Huhndorf<br />
LSU, TEF1<br />
<strong>Pleosporales</strong><br />
Melanommataceae,<br />
Lophiostomataceae<br />
Amniculicolaceae and Lentitheciaceae were<br />
introduced, and Pleosporineae<br />
recircumscribed.<br />
Recircumscribed Melanommataceae and<br />
Lophiostomataceae, and reinstated<br />
Hypsostromataceae.<br />
2009 Nelsen et al. LSU and mtSSU lichenized Dothideomycetes Pyrenocarpous lichens with bitunicate asci<br />
are not monophyletic, but belong to at<br />
least two classes (Dothideomycetes and<br />
Erotiomycetes).<br />
2009 Suetrong et al. LSU, SSU, TEF1, RPB1 marine Dothideomycetes Two new families are introduced<br />
Aigialaceae and Morosphaeriaceae.<br />
2009 Shearer et al. LSU, SSU freshwater Dothideomycetes Freshwater Dothideomycetes are related to<br />
terrestrial taxa and have adapted to<br />
freshwater habitats numerous times.<br />
2009 Tanaka et al. LSU, SSU, TEF1, ITS, BT bambusicolous <strong>Pleosporales</strong> Introduced Tetraplosphaeriaceae with<br />
Tetraploa-like anamorphs.<br />
2009 Kruys and Wedin ITS-nLSU, mtSSU rDNA<br />
and β-tubulin<br />
Sporormiaceae<br />
Analyzed the inter-generic relationships as<br />
well as evaluated the morphological significance<br />
used in this family.<br />
2010 Hirayama et al. LSU, SSU Massarina ingoldiana sensu lato Massarina ingoldiana sensu lato is<br />
polyphyletic, and separated into two<br />
clades within <strong>Pleosporales</strong>.<br />
2010 Aveskamp et al. LSU, SSU, ITS and β-tubulin Phoma and related genera within Rejected current Boeremaean subdivision.<br />
Didymellaceae<br />
2010 de Gruyter et al. LSU, SSU Phoma and related genera within<br />
Pleosporineae<br />
Introduced Pyrenochaetopsis, Setophoma<br />
and Neosetophoma and reinstated<br />
Cucurbitariaceae within Pleosporineae
Fungal Diversity<br />
2. To investigate the phylogeny of <strong>Pleosporales</strong>, its interfamilial<br />
relationships, and the morphological circumscription<br />
of each family;<br />
In order to clarify morphological characters, the generic<br />
types of the majority of teleomorphic pleosporalean genera<br />
(> 60%) were studied. Most of them are from the “core<br />
families” of <strong>Pleosporales</strong>, i.e.Delitschiaceae, Lophiostomataceae,<br />
Massariaceae, Massarinaceae, Melanommataceae,<br />
Montagnulaceae, Phaeosphaeriaceae, Phaeotrichaceae, Pleomassariaceae,<br />
Pleosporaceae, Sporormiaceae and Teichosporaceae.<br />
Notes are given for those where type<br />
specimens could not be obtained during the timeframe<br />
of this study. A detailed description and illustration of<br />
each generic type is provided. Comments, notes and<br />
problems that need to be addressed are provided for each<br />
genus. Phylogenetic investigation based on five nuclear loci,<br />
viz. LSU, SSU, RPB1, RPB2 andTEF1 was carried out using<br />
available strains from numerous genera in <strong>Pleosporales</strong>. In<br />
total, 278 pleosporalean taxa are included in the phylogenetic<br />
analysis, which form 25 familial clades on the dendrogram<br />
(Plate 1). The suborder, Massarineae, is emended to<br />
accommodate Lentitheciaceae, Massarinaceae, Montagnulaceae,<br />
Morosphaeriaceae and Trematosphaeriaceae.<br />
Materials and methods<br />
Molecular phylogeny<br />
Four genes were used in this analysis, the large and small<br />
subunits of the nuclear ribosomal RNA genes (LSU, SSU)<br />
and two protein coding genes, namely the second largest<br />
subunit of RNA polymerase II (RPB2) and translation<br />
elongation factor-1 alpha (TEF1). All sequences were<br />
downloaded from GenBank as listed in Table 3. Eachof<br />
the individual ribosomal genes was aligned in SATé under<br />
default settings with at least 20 iterations. The protein coding<br />
genes were aligned in BioEdit (Hall 2004) and completed by<br />
manual adjustment. Introns were removed and all genes were<br />
concatenated in a single nucleotide alignment with 43%<br />
missing and gap characters out of a total set of 5081. The<br />
alignment had 100% representation for LSU, 75% for SSU,<br />
48% for RPB2 and65%forTEF1. The final data matrix had<br />
280 taxa including outgroups (Table 3).<br />
Previous results indicated no clear conflict amongst the<br />
majority of the data used (Schoch et al. 2009). A phylogenetic<br />
analysis of the concatenated alignment was performed on<br />
CIPRES webportal (Miller et al. 2009) using RAxML v. 7.2.7<br />
(Stamatakis 2006; Stamatakis et al. 2008) applying unique<br />
model parameters for each gene and codon (8 partitions). A<br />
general time reversible model (GTR) was applied with a<br />
discrete gamma distribution and four rate classes. Fifty<br />
thorough maximum likelihood (ML) tree searches were done<br />
in RAxML v. 7.2.7 under the same model, each one starting<br />
from a separate randomized tree and the best scoring tree<br />
selected with a final likelihood value of −95238.628839. Two<br />
isolates of Hysterium angustatum (Hysteriales, Pleosporomycetidae)<br />
were used as outgroups based on earlier work<br />
(Boehm et al. 2009a). Bootstrap pseudo-replicates were run<br />
with the GTRCAT model approximation, allowing the<br />
program to halt bootstraps automatically under the majority<br />
rule criterion (Pattengale et al. 2010). The resulting 250<br />
replicates were plotted on to the best scoring tree obtained<br />
previously. The phylogram with bootstrap values on the<br />
branches is presented in Plate 1 by using graphical options<br />
available in TreeDyn v. 198.3 (Chevenet et al. 2006).<br />
Morphology<br />
Type specimens as well as some other specimens were loaned<br />
from the following herbaria: BAFC, BISH, BPI, BR, BRIP,<br />
<strong>CBS</strong>, E, ETH, FFE, FH, G, H, Herb. J. Kohlmeyer, HHUF,<br />
IFRD, ILLS, IMI, K(M), L, LPS, M, MA, NY, PAD, PC, PH,<br />
RO, S, TNS, TRTC, UB, UBC, UPS and ZT. Attempts were<br />
made to trace and borrow all the type specimens from herbaria<br />
worldwide, but only some of them could be obtained. Some of<br />
the type specimens are in such bad condition that little<br />
information could be obtained. In order to obtain the location<br />
of specimens, original publications were searched.<br />
Ascostroma and ascomata were examined under an<br />
Olympus SZ H10 dissecting microscope. Section of the<br />
fruiting structures was carried out by cryotome or by handcutting.<br />
Measurements and descriptions of sections of the<br />
ascomata, hamathecium, asci and ascospores were carried out<br />
by immersing ascomata in water or in 10% lactic acid.<br />
Microphotography was taken with material mounted in water,<br />
cotton blue, Melzer’s reagent or 10–100% lactic acid.<br />
Terminologies are as in Ulloa and Hanlin (2000). In<br />
addition, ascomata size is defined as: small-sized: < 300 μm<br />
diam., medium-sized: from 300 μm to 600 μm diam., largesized:<br />
> 600 μm diam.<br />
Question mark (“?”) before family (or genus) name<br />
means its familial (or generic) status within <strong>Pleosporales</strong><br />
(or some particular family) is uncertain. Other question<br />
marks after habitats, latin names or other substantives mean<br />
the correctness of their usages need verification.<br />
Results<br />
Molecular phylogeny<br />
In total, 278 pleosporalean taxa are included in the phylogenetic<br />
analysis. These form 25 familial clades in the dendrogram,<br />
i.e. Aigialaceae, Amniculicolaceae, Arthopyreniaceae,
Fungal Diversity<br />
Table 3 Taxa used in the phylogenetic analysis and their corresponding GenBank numbers. Culture and voucher abbreviations are indicated were<br />
available<br />
Species Culture/voucher 1 LSU SSU RPB2 TEF1<br />
Acrocordiopsis patilii BCC 28166 GU479772 GU479736 GU479811<br />
Acrocordiopsis patilii BCC 28167 GU479773 GU479737 GU479812<br />
Aigialus grandis BCC 18419 GU479774 GU479738 GU479813 GU479838<br />
Aigialus grandis JK 5244A GU301793 GU296131 GU371762<br />
Aigialus mangrovis BCC 33563 GU479776 GU479741 GU479815 GU479840<br />
Aigialus mangrovis BCC 33564 GU479777 GU479742 GU479816 GU479841<br />
Aigialus parvus A6 GU301795 GU296133 GU371771 GU349064<br />
Aigialus parvus BCC 32558 GU479779 GU479743 GU479818 GU479843<br />
Aigialus rhizophorae BCC 33572 GU479780 GU479745 GU479819 GU479844<br />
Aigialus rhizophorae BCC 33573 GU479781 GU479746 GU479820 GU479845<br />
Alternaria alternata <strong>CBS</strong> 916.96 DQ678082 DQ678031 DQ677980 DQ677927<br />
Amniculicola immersa <strong>CBS</strong> 123083 FJ795498 GU456295 GU456358 GU456273<br />
Amniculicola parva <strong>CBS</strong> 123092 FJ795497 GU296134 GU349065<br />
Anteaglonium abbreviatum ANM 925.1 GQ221877 GQ221924<br />
Anteaglonium abbreviatum GKM 1029 GQ221878 GQ221915<br />
Anteaglonium globosum ANM 925.2 GQ221879 GQ221925<br />
Anteaglonium latirostrum L100N 2 GQ221876 GQ221938<br />
Arthopyrenia salicis 1994 Coppins AY607730<br />
Arthopyrenia salicis <strong>CBS</strong> 368.94 AY538339 AY538333<br />
Ascochyta pisi <strong>CBS</strong> 126.54 DQ678070 DQ678018 DQ677967 DQ677913<br />
Ascocratera manglicola BCC 09270 GU479782 GU479747 GU479821 GU479846<br />
Ascocratera manglicola JK 5262 C GU301799 GU296136 GU371763<br />
Asteromassaria pulchra <strong>CBS</strong> 124082 GU301800 GU296137 GU371772 GU349066<br />
Astrosphaeriella aggregata MAFF 239485 AB524590 AB524449<br />
Astrosphaeriella aggregata MAFF 239486 AB524591 AB524450 AB539105 AB539092<br />
Astrosphaeriella bakeriana <strong>CBS</strong> 115556 GU301801 GU349015<br />
Astrosphaeriella stellata MAFF 239487 AB524592 AB524451<br />
Beverwykella pulmonaria <strong>CBS</strong> 283.53 GU301804 GU371768<br />
Biatriospora marina CY 1228 GQ925848 GQ925835 GU479823 GU479848<br />
Bimuria novae-zelandiae <strong>CBS</strong> 107.79 AY016356 AY016338 DQ470917 DQ471087<br />
Byssolophis sphaerioides IFRDCC2053 GU301805 GU296140 GU456348 GU456263<br />
Byssosphaeria jamaicana SMH1403 GU385152 GU327746<br />
Byssosphaeria rhodomphala GKM L153N GU385157 GU327747<br />
Byssosphaeria salebrosa SMH2387 GU385162 GU327748<br />
Byssosphaeria schiedermayeriana GKM1197 GU385161 GU327750<br />
Byssosphaeria schiedermayeriana GKM152N GU385168 GU327749<br />
Byssosphaeria villosa GKM204N GU385151 GU327751<br />
Byssothecium circinans <strong>CBS</strong> 675.92 AY016357 AY016339 DQ767646 GU349061<br />
Chaetosphaeronema hispidulum <strong>CBS</strong> 216.75 EU754144 EU754045 GU371777<br />
Cochliobolus heterostrophus <strong>CBS</strong> 134.39 AY544645 AY544727 DQ247790 DQ497603<br />
Cochliobolus sativus DAOM 226212 DQ678045 DQ677995 DQ677939<br />
Corynespora cassiicola <strong>CBS</strong> 100822 GU301808 GU296144 GU371742 GU349052<br />
Corynespora olivacea <strong>CBS</strong> 114450 GU301809 GU349014<br />
Corynespora smithii CABI 5649b GU323201 GU371783 GU349018<br />
Cucurbitaria berberidis <strong>CBS</strong> 394.84 GQ387605 GQ387544<br />
Decaisnella formosa BCC 25616 GQ925846 GQ925833 GU479825 GU479851<br />
Decaisnella formosa BCC 25617 GQ925847 GQ925834 GU479824 GU479850<br />
Decorospora gaudefroyi <strong>CBS</strong> 332.63 EF177849 AF394542
Fungal Diversity<br />
Table 3 (continued)<br />
Species Culture/voucher 1 LSU SSU RPB2 TEF1<br />
Delitschia cf. chaetomioides GKM 1283 GU385172<br />
Delitschia cf. chaetomioides GKM 3253.2 GU390656<br />
Delitschia chaetomioides GKM1283 GU385172 GU327752<br />
Delitschia chaetomioides SMH3253.2 GU390656 GU327753<br />
Delitschia winteri <strong>CBS</strong> 225.62 DQ678077 DQ678026 DQ677975 DQ677922<br />
Didymella exigua <strong>CBS</strong> 183.55 EU754155 EU754056<br />
Didymocrea sadasivanii <strong>CBS</strong> 438 65 DQ384103 DQ384066<br />
Didymosphaeria futilis CMW 22186 EU552123<br />
Didymosphaeria futilis HKUCC 5834 GU205219 GU205236<br />
Dothidotthia aspera CPC 12933 EU673276 EU673228<br />
Dothidotthia symphoricarpi <strong>CBS</strong>119687 EU673273 EU673224<br />
Entodesmium rude <strong>CBS</strong> 650.86 GU301812 GU349012<br />
Falciformispora lignatilis BCC 21117 GU371826 GU371834 GU371819<br />
Falciformispora lignatilis BCC 21118 GU371827 GU371835 GU371820<br />
Floricola striata JK 5603 K GU479785 GU479751<br />
Floricola striata JK 5678I GU301813 GU296149 GU371758<br />
Halomassarina thalassiae BCC 17055 GQ925850 GQ925843<br />
Halomassarina thalassiae JK 5262D GU301816 GU349011<br />
Halotthia posidoniae BBH 22481 GU479786 GU479752<br />
Helicascus nypae BCC 36751 GU479788 GU479754 GU479826 GU479854<br />
Helicascus nypae BCC 36752 GU479789 GU479755 GU479827 GU479855<br />
Herpotrichia diffusa <strong>CBS</strong> 250.62 DQ678071 DQ678019 DQ677968 DQ677915<br />
Herpotrichia juniperi <strong>CBS</strong> 200.31 DQ678080 DQ678029 DQ677978 DQ677925<br />
Herpotrichia macrotricha GKM196N GU385176 GU327755<br />
Herpotrichia macrotricha SMH269 GU385177 GU327756<br />
Hypsostroma caimitalense GKM 1165 GU385180<br />
Hypsostroma saxicola SMH 5005 GU385181<br />
Hysterium angustatum <strong>CBS</strong> 123334 FJ161207 FJ161167 FJ161129 FJ161111<br />
Hysterium angustatum <strong>CBS</strong> 236.34 FJ161180 GU397359 FJ161117 FJ161096<br />
Julella avicenniae BCC 18422 GU371823 GU371831 GU371787 GU371816<br />
Julella avicenniae BCC 20173 GU371822 GU371830 GU371786 GU371815<br />
Julella avicenniae JK 5326A GU479790 GU479756<br />
Kalmusia scabrispora MAFF 239517 AB524593 AB524452 AB539093 AB539106<br />
Kalmusia scabrispora NBRC 106237 AB524594 AB524453 AB539094 AB539107<br />
Karstenula rhodostoma <strong>CBS</strong> 690.94 GU301821 GU296154 GU371788 GU349067<br />
Katumotoa bambusicola MAFF 239641 AB524595 AB524454 AB539095 AB539108<br />
Keissleriella cladophila <strong>CBS</strong> 104.55 GU301822 GU296155 GU371735 GU349043<br />
Keissleriella rara <strong>CBS</strong> 118429 GU479791 GU479757<br />
Kirschsteiniothelia elaterascus A22-5A/HKUCC7769 AY787934 AF053727<br />
Lentithecium aquaticum <strong>CBS</strong> 123099 GU301823 GU296156 GU371789 GU349068<br />
Lentithecium arundinaceum <strong>CBS</strong> 123131 GU456320 GU456298 GU456281<br />
Lentithecium arundinaceum <strong>CBS</strong> 619.86 GU301824 GU296157 FJ795473<br />
Lentithecium fluviatile <strong>CBS</strong> 122367 GU301825 GU296158 GU349074<br />
Lepidosphaeria nicotiae <strong>CBS</strong> 101341 DQ678067 DQ677963 DQ677910<br />
Leptosphaeria biglobosa <strong>CBS</strong> 298.36 GU237980 GU238207<br />
Leptosphaeria biglobosa <strong>CBS</strong> 303.51 GU301826 GU349010<br />
Leptosphaeria doliolum <strong>CBS</strong> 505.75 GU301827 GU296159 GU349069<br />
Leptosphaeria dryadis <strong>CBS</strong> 643.86 GU301828 GU371733 GU349009<br />
Leptosphaerulina argentinensis <strong>CBS</strong> 569.94 GU301829 GU349008
Fungal Diversity<br />
Table 3 (continued)<br />
Species Culture/voucher 1 LSU SSU RPB2 TEF1<br />
Leptosphaerulina australis <strong>CBS</strong> 311.51-T FJ795500 GU456357 GU456272<br />
Leptosphaerulina australis <strong>CBS</strong> 317.83 GU301830 GU296160 GU371790 GU349070<br />
Leptosphearia maculans DAOM 229267 DQ470946 DQ470993 DQ470894 DQ471062<br />
Letendraea helminthicola <strong>CBS</strong> 884.85 AY016362 AY016345<br />
Letendraea padouk <strong>CBS</strong> 485.70 AY849951 GU296162<br />
Lindgomyces breviappendiculatus KT 1399 AB521749 AB521734<br />
Lindgomyces cinctosporae R56-1 AB522431 AB522430<br />
Lindgomyces cinctosporae R56-3 GU266245 GU266238<br />
Lindgomyces ingoldianus KH 100 JCM 16479 AB521737 AB521720<br />
Lindgomyces rotundatus KH 114 JCM 16484 AB521742 AB521725<br />
Lophiostoma alpigenum GKM 1091b GU385193<br />
Lophiostoma arundinis <strong>CBS</strong> 621.86 DQ782384 DQ782383 DQ782386 DQ782387<br />
Lophiostoma caulium <strong>CBS</strong> 623.86 GU301833 GU296163 GU371791<br />
Lophiostoma compressum IFRD 2014 GU301834 GU296164 FJ795457<br />
Lophiostoma crenatum <strong>CBS</strong> 629.86 DQ678069 DQ678017 DQ677965 DQ677912<br />
Lophiostoma fuckelii <strong>CBS</strong> 101952 DQ399531<br />
Lophiostoma fuckelii <strong>CBS</strong> 113432 EU552139<br />
Lophiostoma fuckelii GKM 1063 GU385192<br />
Lophiostoma macrostomum <strong>CBS</strong> 122681 EU552141<br />
Lophiostoma macrostomum HHUF 27293 AB433274<br />
Lophiostoma macrostomum KT 635 AB433273 AB521731<br />
Lophiostoma quadrinucleatum GKM1233 GU385184 GU327760<br />
Lophiostoma sagittiforme HHUF 29754 AB369267<br />
Lophiotrema brunneosporum <strong>CBS</strong> 123095 GU301835 GU296165 GU349071<br />
Lophiotrema lignicola <strong>CBS</strong> 122364 GU301836 GU296166 GU349072<br />
Massarina arundinariae MAFF 239461 AB524596 AB524455 AB539096 AB524817<br />
Massarina arundinariae NBRC 106238 AB524597 AB524456 AB539097 AB524818<br />
Lophiotrema nucula <strong>CBS</strong> 627.86 GU301837 GU296167 GU371792 GU349073<br />
Loratospora aestuarii JK 5535B GU301838 GU296168 GU371760<br />
Macroventuria anomochaeta <strong>CBS</strong> 525.71 GU456315 GU456346 GU456262<br />
Massaria anomia <strong>CBS</strong> 123109 GU301792 GU296130 GU349062<br />
Massaria anomia <strong>CBS</strong> 591.78 GU301839 GU296169 GU371769<br />
Massaria ariae M52 HQ599382 HQ599456 HQ599322<br />
Massaria aucupariae M49 HQ599384 HQ599455 HQ599324<br />
Massaria campestris M28 HQ599385 HQ599449 HQ599459 HQ599325<br />
Massaria conspurcata M14 HQ599393 HQ599441 HQ599333<br />
Massaria gigantispora M26 HQ599397 HQ599447 HQ599337<br />
Massaria inquinans M19 HQ599402 HQ599444 HQ599460 HQ599342<br />
Massaria lantanae M18 HQ599406 HQ599443 HQ599346<br />
Massaria macra M3 HQ599408 HQ599450 HQ599348<br />
Massaria mediterranea M45 HQ599417 HQ599452 HQ599357<br />
Massaria platanoidea M7 HQ599420 HQ599457 HQ599462 HQ599359<br />
Massaria pyri M21 HQ599424 HQ599445 HQ599363<br />
Massaria vindobonensis M27 HQ599429 HQ599448 HQ599464 HQ599368<br />
Massaria vomitoria M13 HQ599437 HQ599440 HQ599466 HQ599375<br />
Massarina cisti <strong>CBS</strong> 266.62 FJ795447 FJ795490 FJ795464<br />
Massarina eburnea <strong>CBS</strong> 473.64 GU301840 GU296170 GU371732 GU349040<br />
Massarina igniaria <strong>CBS</strong> 845.96 GU301841 GU296171 GU371793<br />
Massarina ricifera JK 5535 F GU479793 GU479759
Fungal Diversity<br />
Table 3 (continued)<br />
Species Culture/voucher 1 LSU SSU RPB2 TEF1<br />
Massariosphaeria phaeospora <strong>CBS</strong> 611.86 GU301843 GU296173 GU371794<br />
Mauritiana rhizophorae BCC 28866 GU371824 GU371832 GU371796 GU371817<br />
Mauritiana rhizophorae BCC 28867 GU371825 GU371833 GU371797 GU371818<br />
Melanomma pulvis-pyrius <strong>CBS</strong> 124080 GU456323 GU456302 GU456350 GU456265<br />
Melanomma pulvis-pyrius <strong>CBS</strong> 371.75 GU301845 GU371798 GU349019<br />
Melanomma pulvis-pyrius SMH 3291 GU385197<br />
Melanomma rhododendri ANM 73 GU385198<br />
Misturatosphaeria aurantonotata GKM1238 GU385173 GU327761<br />
Misturatosphaeria aurantonotata GKM1280 GU385174 GU327762<br />
Misturatosphaeria claviformis GKM1210 GU385212 GU327763<br />
Misturatosphaeria kenyensis GKM1195 GU385194 GU327767<br />
Misturatosphaeria kenyensis GKM L100Na GU385189 GU327766<br />
Misturatosphaeria minima GKM169N GU385165 GU327768<br />
Misturatosphaeria tennesseensis ANM911 GU385207 GU327769<br />
Misturatosphaeria uniseptata SMH4330 GU385167 GU327770<br />
Monascostroma innumerosum <strong>CBS</strong> 345.50 GU301850 GU296179 GU349033<br />
Monotosporella tuberculata <strong>CBS</strong> 256.84 GU301851 GU349006<br />
Montagnula anthostomoides <strong>CBS</strong> 615.86 GU205223 GU205246<br />
Montagnula opulenta <strong>CBS</strong> 168.34 DQ678086 AF164370 DQ677984<br />
Morosphaeria ramunculicola BCC 18405 GQ925854 GQ925839<br />
Morosphaeria ramunculicola JK 5304B GU479794 GU479760 GU479831<br />
Morosphaeria velataspora BCC 17059 GQ925852 GQ925841<br />
Morosphaeria velataspora BCC 17058 GQ925851 GQ925840<br />
Massariosphaeria grandispora <strong>CBS</strong> 613 86 GU301842 GU296172 GU371725 GU349036<br />
Massariosphaeria typhicola <strong>CBS</strong> 123126 GU301844 GU296174 GU371795<br />
Neophaeosphaeria filamentosa <strong>CBS</strong> 102202 GQ387577 GQ387516 GU371773 GU349084<br />
Neotestudina rosatii <strong>CBS</strong> 690.82 DQ384069<br />
Neottiosporina paspali <strong>CBS</strong> 331.37 EU754172 EU754073 GU371779 GU349079<br />
Ophiosphaerella herpotricha <strong>CBS</strong> 240.31 DQ767656 DQ767650 DQ767645 DQ767639<br />
Ophiosphaerella herpotricha <strong>CBS</strong> 620.86 DQ678062 DQ678010 DQ677958 DQ677905<br />
Ophiosphaerella sasicola MAFF 239644 AB524599 AB524458 AB539098 AB539111<br />
Paraconiothyrium minitans <strong>CBS</strong> 122788 EU754173 EU754074 GU371776 GU349083<br />
Paraphaeosphaeria michotii <strong>CBS</strong> 591.73 GU456326 GU456305 GU456352 GU456267<br />
Paraphaeosphaeria michotii <strong>CBS</strong> 652.86 GU456325 GU456304 GU456351 GU456266<br />
Phaeosphaeria ammophilae <strong>CBS</strong> 114595 GU301859 GU296185 GU371724 GU349035<br />
Phaeosphaeria avenaria <strong>CBS</strong> 602.86 AY544684 AY544725 DQ677941 DQ677885<br />
Phaeosphaeria avenaria DAOM 226215 AY544684 AY544725 DQ677941 DQ677885<br />
Phaeosphaeria brevispora MAFF 239276 AB524600 AB524459 AB539099 AB539112<br />
Phaeosphaeria brevispora NBRC 106240 AB524601 AB524460 AB539100 AB539113<br />
Phaeosphaeria caricis <strong>CBS</strong> 120249 GU301860 GU349005<br />
Phaeosphaeria elongata <strong>CBS</strong> 120250 GU456327 GU456306 GU456345 GU456261<br />
Phaeosphaeria eustoma <strong>CBS</strong> 573.86 DQ678063 DQ678011 DQ677959 DQ677906<br />
Phaeosphaeria luctuosa <strong>CBS</strong> 308.79 GU301861 GU349004<br />
Phaeosphaeria nigrans <strong>CBS</strong> 576.86 GU456331 GU456356 GU456271<br />
Phaeosphaeria nodorum <strong>CBS</strong> 259.49 GU456332 GU456285<br />
Phaeosphaeria oryzae <strong>CBS</strong> 110110 GQ387591 GQ387530<br />
Phaeosphaeriopsis musae <strong>CBS</strong> 120026 GU301862 GU296186 GU349037<br />
Phoma apiicola <strong>CBS</strong> 285.72 GU238040 GU238211<br />
Phoma betae <strong>CBS</strong> 109410 EU754178 EU754079 GU371774 GU349075
Fungal Diversity<br />
Table 3 (continued)<br />
Species Culture/voucher 1 LSU SSU RPB2 TEF1<br />
Phoma complanata <strong>CBS</strong> 268.92 EU754180 EU754081 GU371778 GU349078<br />
Phoma cucurbitacearum <strong>CBS</strong> 133.96 GU301863 GU371767<br />
Phoma exigua <strong>CBS</strong> 431.74 EU754183 EU754084 GU371780 GU349080<br />
Phoma glomerata <strong>CBS</strong> 528.66 EU754184 EU754085 GU371781 GU349081<br />
Phoma herbarum <strong>CBS</strong> 276.37 DQ678066 DQ678014 DQ677962 DQ677909<br />
Phoma radicina <strong>CBS</strong> 111.79 EU754191 EU754092 GU349076<br />
Phoma valerianae <strong>CBS</strong> 630.68 GU238150 GU238229<br />
Phoma vasinfecta <strong>CBS</strong> 539.63 GU238151 GU238230<br />
Phoma violicola <strong>CBS</strong> 306.68 GU238156 GU238231<br />
Phoma zeae-maydis <strong>CBS</strong> 588.69 EU754192 EU754093 GU371782 GU349082<br />
Platychora ulmi <strong>CBS</strong> 361.52 EF114702 EF114726<br />
Lophiostoma compressum GKM1048 GU385204 GU327772<br />
Lophiostoma scabridisporum BCC 22836 GQ925845 GQ925832 GU479829 GU479856<br />
Lophiostoma scabridisporum BCC 22835 GQ925844 GQ925831 GU479830 GU479857<br />
Pleomassaria siparia <strong>CBS</strong> 279.74 DQ678078 DQ678027 DQ677976 DQ677923<br />
Pleospora ambigua <strong>CBS</strong> 113979 AY787937<br />
Pleospora herbarum <strong>CBS</strong> 191.86 DQ247804 DQ247812 DQ247794 DQ471090<br />
Polyplosphaeria fusca <strong>CBS</strong> 125425 AB524607 AB524466 AB524822<br />
Polyplosphaeria fusca MAFF 239687 AB524606 AB524465<br />
Preussia funiculata <strong>CBS</strong> 659.74 GU301864 GU296187 GU371799 GU349032<br />
Preussia lignicola <strong>CBS</strong> 264.69 GU301872 GU296197 GU371765 GU349027<br />
Preussia terricola DAOM 230091 AY544686 AY544726 DQ470895 DQ471063<br />
Prosthemium betulinum <strong>CBS</strong> 127468 AB553754 AB553644<br />
Prosthemium canba JCM 16966 AB553760 AB553646<br />
Prosthemium orientale JCM 12841 AB553748 AB553641<br />
Prosthemium stellare <strong>CBS</strong> 126964 AB553781 AB553650<br />
Pseudotetraploa curviappendiculata <strong>CBS</strong> 125426 AB524610 AB524469 AB524825<br />
Pseudotetraploa curviappendiculata MAFF 239495 AB524608 AB524467<br />
Pseudotetraploa javanica MAFF 239498 AB524611 AB524470 AB524826<br />
Pseudotetraploa longissima MAFF 239497 AB524612 AB524471 AB524827<br />
Pseudotrichia guatopoensis SMH4535 GU385202 GU327774<br />
Pyrenochaeta acicola <strong>CBS</strong> 812.95 GQ387602 GQ387541<br />
Pleurophoma cava <strong>CBS</strong> 257.68 EU754199 EU754100<br />
Pyrenochaeta corn <strong>CBS</strong> 248.79 GQ387608 GQ387547<br />
Pyrenochaeta nobilis <strong>CBS</strong> 292.74 GQ387615 GQ387554<br />
Pyrenochaeta nobilis <strong>CBS</strong> 407.76 DQ678096 DQ677991 DQ677936<br />
Pyrenochaeta quercina <strong>CBS</strong> 115095 GQ387619 GQ387558<br />
Pyrenochaeta unguis-hominis <strong>CBS</strong> 378.92 GQ387621 GQ387560<br />
Pyrenochaetopsis decipiens <strong>CBS</strong> 343.85 GQ387624 GQ387563<br />
Pyrenophora phaeocomes DAOM 222769 DQ499596 DQ499595 DQ497614 DQ497607<br />
Pyrenophora tritici-repentis OSC 100066 AY544672 DQ677882<br />
Quadricrura bicornis <strong>CBS</strong> 125427 AB524613 AB524472 AB524828<br />
Quadricrura meridionalis <strong>CBS</strong> 125684 AB524614 AB524473 AB524829<br />
Quadricrura septentrionalis <strong>CBS</strong> 125428 AB524617 AB524476 AB524832<br />
Quintaria lignatilis BCC 17444 GU479797 GU479764 GU479832 GU479859<br />
Quintaria lignatilis <strong>CBS</strong> 117700 GU301865 GU296188 GU371761<br />
Quintaria submersa <strong>CBS</strong> 115553 GU301866 GU349003<br />
Repetophragma ontariense HKUCC 10830 DQ408575 DQ435077<br />
Rimora mangrovei JK 5246A GU301868 GU296193 GU371759
Fungal Diversity<br />
Table 3 (continued)<br />
Species Culture/voucher 1 LSU SSU RPB2 TEF1<br />
Rimora mangrovei JK 5437B GU479798 GU479765<br />
Roussoella hysterioides <strong>CBS</strong> 125434 AB524622 AB524481 AB539102 AB539115<br />
Roussoella hysterioides MAFF 239636 AB524621 AB524480 AB539101 AB539114<br />
Roussoella pustulans MAFF 239637 AB524623 AB524482 AB539103 AB539116<br />
Roussoellopsis tosaensis MAFF 239638 AB524625 AB539104 AB539117<br />
Saccothecium sepincola <strong>CBS</strong> 278.32 GU301870 GU296195 GU371745 GU349029<br />
Salsuginea ramicola KT 2597.1 GU479800 GU479767 GU479833 GU479861<br />
Salsuginea ramicola KT 2597.2 GU479801 GU479768 GU479834 GU479862<br />
Setomelanomma holmii <strong>CBS</strong> 110217 GU301871 GU296196 GU371800 GU349028<br />
Setosphaeria monoceras AY016368 AY016368<br />
Massaria platani <strong>CBS</strong> 221.37 DQ678065 DQ678013 DQ677961 DQ677908<br />
Sporormiella minima <strong>CBS</strong> 524.50 DQ678056 DQ678003 DQ677950 DQ677897<br />
Stagonospora macropycnidia <strong>CBS</strong> 114202 GU301873 GU296198 GU349026<br />
Tetraploa aristata <strong>CBS</strong> 996.70 AB524627 AB524486 AB524836<br />
Tetraplosphaeria nagasakiensis MAFF 239678 AB524630 AB524489 AB524837<br />
Lophiostoma macrostomoides GKM1033 GU385190 GU327776<br />
Lophiostoma macrostomoides GKM1159 GU385185 GU327778<br />
Thyridaria rubronotata <strong>CBS</strong> 419.85 GU301875 GU371728 GU349002<br />
Tingoldiago graminicola KH 68 AB521743 AB521726<br />
Trematosphaeria pertusa <strong>CBS</strong> 122368 FJ201990 FJ201991 FJ795476 GU456276<br />
Trematosphaeria pertusa <strong>CBS</strong> 122371 GU301876 GU348999 GU371801 GU349085<br />
Trematosphaeria pertusa SMH 1448 GU385213<br />
Triplosphaeria cylindrica MAFF 239679 AB524634 AB524493<br />
Triplosphaeria maxima MAFF 239682 AB524637 AB524496<br />
Triplosphaeria yezoensis <strong>CBS</strong> 125436 AB524638 AB524497 AB524844<br />
Ulospora bilgramii <strong>CBS</strong> 110020 DQ678076 DQ678025 DQ677974 DQ677921<br />
Verruculina enalia BCC 18401 GU479802 GU479770 GU479835 GU479863<br />
Verruculina enalia BCC 18402 GU479803 GU479771 GU479836 GU479864<br />
Westerdykella cylindrica <strong>CBS</strong> 454.72 AY004343 AY016355 DQ470925 DQ497610<br />
Westerdykella dispersa <strong>CBS</strong> 508.75 DQ468050 U42488<br />
Westerdykella ornata <strong>CBS</strong> 379.55 GU301880 GU296208 GU371803 GU349021<br />
Wicklowia aquatica AF289-1 GU045446<br />
Wicklowia aquatica <strong>CBS</strong> 125634 GU045445 GU266232<br />
Xenolophium applanatum <strong>CBS</strong> 123123 GU456329 GU456312 GU456354 GU456269<br />
Xenolophium applanatum <strong>CBS</strong> 123127 GU456330 GU456313 GU456355 GU456270<br />
Zopfia rhizophila <strong>CBS</strong> 207.26 DQ384104 L76622<br />
1 BCC Belgian Coordinated Collections of Microorganisms; CABI International Mycological Institute, CABI-Bioscience, Egham, Bakeham Lane,<br />
U.K.; <strong>CBS</strong> Centraalbureau voor Schimmelcultures, Utrecht, The Netherlands; DAOM Plant Research Institute, Department of Agriculture<br />
(Mycology), Ottawa, Canada; DUKE Duke University Herbarium, Durham, North Carolina, U.S.A.; HHUF Herbarium of Hirosaki University,<br />
Japan; IFRDCC Culture Collection, International Fungal Research & Development Centre, Chinese Academy of Forestry, Kunming, China;<br />
MAFF Ministry of Agriculture, Forestry and Fisheries, Japan; NBRC NITE Biological Resource Centre, Japan; OSC Oregon State University<br />
Herbarium, U.S.A.; UAMH University of Alberta Microfungus Collection and Herbarium, Edmonton, Alberta, Canada; UME Herbarium of the<br />
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.<br />
Jones; GKM G.K. Mugambi; JK J. Kohlmeyer; KT K. Tanaka; SMH S.M. Huhndorf<br />
Cucurbitariaceae/Didymosphaeriaceae, Delitschiaceae,<br />
Didymellaceae, Dothidotthiaceae, Hypsostromataceae, Lentitheciaceae,<br />
Leptosphaeriaceae, Lindgomycetaceae,<br />
Lophiostomataceae, Massariaceae, Massarinaceae, Melanommataceae,<br />
Montagnulaceae, Morosphaeriaceae,<br />
Phaeosphaeriaceae, Pleomassariaceae, Pleosporaceae,<br />
Sporormiaceae, Testudinaceae/Platystomaceae, Tetraplosphaeriaceae,<br />
Trematosphaeriaceae and Zopfiaceae (Plate 1).<br />
Of these, Lentitheciaceae, Massarinaceae, Montagnulaceae,<br />
Morosphaeriaceae and Trematosphaeriaceae form a robust
Fungal Diversity<br />
clade in the present study and in previous studies (Schoch et<br />
al. 2009; Zhang et al. 2009a, b). We thus emended the<br />
suborder, Massarineae, to accommodate them.<br />
<strong>Pleosporales</strong> suborder Massarineae Barr, Mycologia 71:<br />
948. (1979a). emend.<br />
Habitat freshwater, marine or terrestrial environment,<br />
saprobic. Ascomata solitary, scattered or gregarious, globose,<br />
subglobose, conical to lenticular, immersed, erumpent<br />
to superficial, papillate, ostiolate. Hamathecium of dense or<br />
rarely few, filliform pseudoparaphyses. Asci bitunicate,<br />
fissitunicate, cylindrical, clavate or broadly clavate, pedicellate.<br />
Ascospores hyaline, pale brown or brown, 1 to 3 or<br />
more transverse septa, rarely muriform, narrowly fusoid,<br />
fusoid, broadly fusoid, symmetrical or asymmetrical, with<br />
or without sheath.<br />
Accepted genera of <strong>Pleosporales</strong><br />
Acrocordiopsis Borse & K.D. Hyde, Mycotaxon 34: 535<br />
(1989). (<strong>Pleosporales</strong>, genera incertae sedis)<br />
Generic description<br />
Habitat marine, saprobic. Ascomata seated in blackish<br />
stroma, scattered or gregarious, superficial, conical to<br />
semiglobose, ostiolate, carbonaceous. Hamathecium of<br />
dense, long trabeculate pseudoparaphyses. Asci 8-spored,<br />
cylindrical with pedicels and conspicuous ocular chambers.<br />
Ascospores hyaline, 1-septate, obovoid to broadly fusoid.<br />
Anamorphs reported for genus: none.<br />
Literature: Alias et al. 1999; Barr 1987a; Borse and Hyde<br />
1989.<br />
Type species<br />
Acrocordiopsis patilii Borse & K.D. Hyde, Mycotaxon 34:<br />
536 (1989). (Fig. 1)<br />
Ascomata 1–2 mm high×1.8–3 mmdiam.,scatteredor<br />
gregarious, superficial, conical or semiglobose, with a<br />
flattened base not easily removed from the substrate, ostiolate,<br />
black, very brittle and carbonaceous and extremely difficult to<br />
cut (Fig. 1a and b). Peridium 250–310 μm thick, to 600 μm<br />
thick near the apex, thinner at the base, comprising three<br />
types of cells; outer cells pseudoparenchymatous, small<br />
heavily pigmented thick-walled cells of textura epidermoidea,<br />
cells 0.6–1×6–10 μm diam., cell wall 5–9 μm thick;<br />
cells near the substrate less pigmented, composed of cells of<br />
textura prismatica, cellwalls1–3(−5) μm thick; inner cells<br />
less pigmented, comprised of hyaline to pale brown thinwalled<br />
cells, merging with pseudoparaphyses (Fig. 1c, d and<br />
e). Hamathecium of dense, long trabeculate pseudoparaphyses,<br />
ca. 1 μm broad, embedded in mucilage, hyaline,<br />
Fig. 1 Acrocordiopsis patilii (from IMI 297769, holotype). a<br />
Ascomata on the host surface. b Section of an ascoma. c Section of<br />
lateral peridium. d Section of the apical peridium. e Section of the<br />
basal peridium. Note the paler cells of textura prismatica. f<br />
Cylindrical ascus. g Cylindrical ascus in pseudoparaphyses. h, i<br />
One-septate ascospores. Scale bars: a=3 mm, b=0.5 mm, c=200 μm,<br />
d, e =50 μm, f–i=20 μm<br />
anastomosing and sparsely septate. Asci 140–220×13–<br />
17 μm (x ¼ 165:3 15:6mm, n=10), 8-spored, bitunicate,<br />
fissitunicate, cylindrical, with short pedicels, 15–25(−40) μm<br />
long, with a large and conspicuous ocular chamber (Fig. 1f<br />
and g). Ascospores 17.5–25×12.5–15(−20) μm<br />
(x ¼ 21:5 13:6mm, n=10), uniseriate to partially overlapping,<br />
ovoid or ellipsoidal, hyaline, 1-septate, not constricted<br />
at the septum, smooth-walled (Fig. 1h and i).<br />
Anamorph: none reported.<br />
Material examined: INDIA, Indian Ocean, Malvan<br />
(Maharashtra), on intertidal wood of Avicennia alba Bl.,<br />
30 Oct. 1981 (IMI 297769, holotype).<br />
Notes<br />
Morphology<br />
Acrocordiopsis was formally established by Borse and<br />
Hyde (1989) as a monotypic genus represented by A. patilii<br />
based on its “conical or semiglobose superficial carbonaceous<br />
ascomata, trabeculate pseudoparaphyses, cylindrical,<br />
bitunicate, 8-spored asci, and hyaline, 1-septate, obovoid or<br />
ellipsoid ascospores”. Acrocordiopsis patilii was first<br />
collected from mangrove wood (Indian Ocean) as a marine<br />
fungus, and a second marine Acrocordiopsis species<br />
was reported subsequently from Philippines (Alias et al.<br />
1999). Acrocordiopsis is assigned to Melanommataceae<br />
(Melanommatales sensu Barr 1983) based on its ostiolate<br />
ascomata and trabeculate pseudoparaphyses (Borse and<br />
Hyde 1989). Morphologically, Acrocordiopsis is similar to<br />
Astrosphaeriella sensu stricto based on the conical ascomata<br />
and the brittle, carbonaceous peridium composed of thickwalled<br />
black cells with rows of palisade-like parallel cells at<br />
the rim area. Ascospores of Astrosphaeriella are, however,<br />
elongate-fusoid, usually brown or reddish brown and<br />
surrounded by a gelatinous sheath when young; as such<br />
they are readily distinguishable from those of Acrocordiopsis.<br />
A new family (Acrocordiaceae) was introduced by Barr<br />
(1987a) to accommodate Acrocordiopsis. This proposal,<br />
however, has been rarely followed and Jones et al. (2009)<br />
assigned Acrocordiopsis to Melanommataceae.<br />
Phylogenetic study<br />
Acrocordiopsis patilii nested within an unresolved clade<br />
within <strong>Pleosporales</strong> (Suetrong et al. 2009). Thus its familial<br />
placement is unresolved, but use of the Acrocordiaceae<br />
could be reconsidered with more data.<br />
b
Fungal Diversity
Fungal Diversity<br />
Concluding remarks<br />
Acrocordiopsis, Astrosphaeriella sensu stricto, Mamillisphaeria,<br />
Caryospora and Caryosporella are morphologically<br />
similar as all have very thick-walled carbonaceous ascomata,<br />
narrow pseudoparaphyses in a gelatinous matrix (trabeculae)<br />
and bitunicate, fissitunicate asci. Despite their similarities, the<br />
shape of asci and ascospores differs (e.g. Mamillisphaeria has<br />
sac-like asci and two types of ascospores, brown or hyaline,<br />
Astrosphaeriella has cylindro-clavate asci and narrowly fusoid<br />
ascospores, both Acrocordiopsis and Caryosporella has<br />
cylindrical asci, but ascospores of Caryosporella are reddish<br />
brown). Therefore, the current familial placement of Acrocordiopsis<br />
cannot be determined. All generic types of Astrosphaeriella<br />
sensu stricto, Mamillisphaeria and Caryospora<br />
should be recollected and isolated for phylogenetic study.<br />
Aigialus Kohlm. & S. Schatz, Trans. Br. Mycol. Soc. 85:<br />
699 (1985). (Aigialaceae)<br />
Generic description<br />
Habitat marine, saprobic. Ascomata mostly subglobose in<br />
front view, fusoid in sagittal section, rarely subglobose,<br />
scattered, immersed to erumpent, papillate, ostiolate, ostiole<br />
rounded or slit-like, periphysate. Peridium 2-layered. Hamathecium<br />
of trabeculate pseudoparaphyses. Asci 8-spored,<br />
cylindrical, pedicellate, with an ocular chamber and conspicuous<br />
apical ring. Ascospores ellipsoidal to fusoid, muriform,<br />
yellow brown to brown, with terminal appendages.<br />
Anamorphs reported for genus: none.<br />
Literature: Eriksson 2006; Jones et al. 2009; Kohlmeyer<br />
and Schatz 1985; Lumbsch and Huhndorf 2007.<br />
Type species<br />
Aigialus grandis Kohlm. & S. Schatz, Trans. Br. Mycol.<br />
Soc. 85: 699 (1985). (Fig. 2)<br />
Ascomata 1–1.25 mm high×1–1.3 mm diam. in front view,<br />
250–400 μm broad in sagittal section, vertically flattened<br />
subglobose, laterally compressed, scattered, immersed to semiimmersed,<br />
papillate, with an elongated furrow at the top of the<br />
papilla, wall black, carbonaceous, ostiolate, ostiole filled with<br />
branched or forked septate periphyses (Fig. 2a). Peridium 70–<br />
100 μm thick laterally, up to 150 μm thick at the apex, thinner<br />
at the base, comprising two cell types, outer layer composed of<br />
small heavily pigmented thick-walled pseudoparenchymatous<br />
cells, cells 1–2 μm diam., cell wall 2–5 μm thick, inner layer<br />
thin, composed of small hyaline cells (Fig. 2b). Hamathecium<br />
of dense, very long trabeculate pseudoparaphyses, 0.8–1.2 μm<br />
broad, embedded in mucilage, anastomosing and branching<br />
above the asci. Asci 450–640×22–35 μm (x ¼ 505 30mm,<br />
n=10), 8-spored, bitunicate, fissitunicate, cylindrical to<br />
cylindro-clavate, with a long furcate pedicel, 90–180 μm long,<br />
with a low truncate ocular chamber and a refractive apical<br />
apparatus (to 12 μm wide×4 μm high) (Fig. 2e and f).<br />
Ascospores 75–95×15–26 μm (x ¼ 84:3 17:5mm, n=10),<br />
obliquely uniseriate and partially overlapping, broadly fusoid<br />
to fusoid with narrowly rounded ends in front view, flat on one<br />
side from side view (14–20 μm thick), yellowish brown, apical<br />
cells usually hyaline, muriform, with 14–17(−18) transversal<br />
septa, 1–3 longitudinal septa in most cells, slightly constricted<br />
at the septa, with a gelatinous cap at each end (Fig. 2c and d).<br />
Anamorph: none reported.<br />
Material examined: BELIZE, Wee-Wee Cay, on submerged<br />
wood of roots and branches of Rhizophora mangle<br />
L., Mar. 1983, leg. J. Kohlmeyer (NY, J.K. 4332b, isotype).<br />
Notes<br />
Morphology<br />
Aigialus was formally established by Kohlmeyer and Schatz<br />
(1985) based on its immersed or semi-immersed ascomata<br />
with periphysate ostiole, trabeculate pseudoparaphyses, cylindrical<br />
and fissitunicate asci, and distinctive muriform ascospores<br />
with gelatinous sheath or caps. There are five accepted<br />
species in the genus, namely A. grandis, A. mangrovei Borse,<br />
A. parvus S. Schatz & Kohlm., A. rhizophorae Borse and A.<br />
striatispora K.D. Hyde (Jones et al. 2009). Aigialus was first<br />
assigned to the Melanommatales, but its familial status was<br />
uncertain (Kohlmeyer and Schatz 1985). Barr (1990b)<br />
included Aigialus in Massariaceae based on its conspicuous<br />
apical ring in the asci and ascospore characters, and this has<br />
subsequently been widely followed (Eriksson 2006; Hawksworth<br />
et al. 1995; Kirk et al. 2001; Lumbsch and Huhndorf 2007).<br />
Phylogenetic study<br />
The generic type of Aigialus (A. grandis) together with<br />
other three marine species, i.e. A. mangrovei, A. parvus as well<br />
as A. rhizophorae form a robust clade on the phylogenetic<br />
tree. Thus a new family, Aigialaceae, was introduced to<br />
accommodate Aigialus together with Ascocratera and Rimora<br />
(Suetrong et al. 2009).<br />
Concluding remarks<br />
The pleosporalean status of Aigialus has been phylogenetically<br />
verified, and the single branch containing Aigialus,<br />
Ascocratera and Rimora represents a familial rank of<br />
Aigialaceae (Suetrong et al. 2009).<br />
Amniculicola Yin. Zhang & K.D. Hyde, Mycol. Res. 112:<br />
1189 (2008). (Amniculicolaceae)<br />
Generic description<br />
Habitat freshwater, saprobic. Ascomata solitary, scattered,<br />
or in small groups, initially immersed, becoming erumpent,
Fungal Diversity<br />
Fig. 2 Aigialus grandis (from NY, J.K. 4332b, isotype). a Ascomata<br />
on the host surface. Note the longitudinal slit-like furrow which is the<br />
ostiole. b Section of the peridium. c, d Released ascospores. e<br />
Ascospores in ascus. Note the conspicuous apical ring. f Cylindrical<br />
ascus with a long pedicel. Scale bars: a=1 mm, b=200 μm, c–f=<br />
20 μm
Fungal Diversity<br />
to nearly superficial, globose, subglobose to conical, wall<br />
black, roughened; apex well differentiated into two tuberculate<br />
flared lips surrounding a slit-like ostiole. Peridium thin, 2-<br />
layered, outer layer composed of small heavily pigmented<br />
thick-walled cells of textura angularis, inner layer composed<br />
of hyaline thin-walled cells of textura angularis. Hamathecium<br />
of dense, long trabeculate pseudoparaphyses, embedded<br />
in mucilage, anastomosing between and above the asci. Asci<br />
8-spored, bitunicate, fissitunicate, cylindrical to narrowly<br />
fusoid, short pedicellate, with an ocular chamber and a small<br />
apical apparatus. Ascospores fusoid, hyaline, 1-septate,<br />
constricted at the septum, surrounded by an irregular hyaline<br />
gelatinous sheath.<br />
Anamorphs reported for genus: Anguillospora longissima,<br />
Spirosphaera cupreorufescens and Repetophragma<br />
ontariense (Zhang et al. 2008c, 2009c).<br />
Literature: Zhang et al. 2008c, 2009a, c.<br />
Type species<br />
Amniculicola lignicola Ying Zhang & K.D. Hyde, Mycol.<br />
Res. 112: 1189 (2008). (Fig. 3)<br />
Ascomata 350–450 μm high×300–500 μm diam.,<br />
solitary, scattered, or in small groups of 2–3, initially<br />
immersed, becoming erumpent, to nearly superficial, with<br />
basal wall remaining immersed in host tissue, globose,<br />
subglobose, broadly or narrowly conical, often laterally<br />
flattened, with a flattened base not easily removed from the<br />
substrate, wall black, roughened, often bearing remnants of<br />
wood fibers; apex well differentiated into two tuberculate<br />
flared lips surrounding a slit-like ostiole, 150–250 μm long,<br />
filled with a purplish amorphous matter, oriented in the axis<br />
of the wood fibers; underlying wood stained pale purple<br />
(Fig. 3a and b). Peridium 40–55 μm thick laterally, up to<br />
120 μm thick at the apex, thinner at the base, coriaceous,<br />
2-layered, outer layer composed of small heavily pigmented<br />
thick-walled cells of textura angularis, cells4–9 μm diam.,<br />
cell wall 2–3 μm thick, apex cells smaller and walls thicker,<br />
inner layer composed of hyaline thin-walled cells of textura<br />
angularis, 8–16 μm diam., in places with columns of textura<br />
prismatica, oriented perpendicular to the ascomatal surface,<br />
and larger, paler cells of textura prismatica towards the<br />
interior and at the base, 10–25 μm (Fig. 3c, d and e).<br />
Hamathecium of dense, long trabeculate pseudoparaphyses<br />
Fungal Diversity<br />
Fig. 3 Amniculicola lignicola (from PC 0092661, holotype). a<br />
Superficial ascomata gregarious on the host surface. b An erumpent<br />
ascoma with elongated papilla and slit-like ostiole. c Habitat section of<br />
a superficial ascoma. d, e Section of an ascoma and the partial<br />
peridium. f Cylindrical 8-spored ascus with a short pedicel. g Hyaline,<br />
1-septate broadly fusoid ascospores. Scale bars: a=1 mm, b–d=<br />
100 μm, e=50 μm, f, g=20 μm<br />
1- (rarely 2- to 3-) septate, fusoid, reddish brown, constricted<br />
at the main septum.<br />
Anamorphs reported for genus: Exosporiella (= Phanerocorynella)<br />
(Sivanesan 1983).
Fungal Diversity<br />
Fig. 4 Anomalemma epochnii (from K(M):143936, syntype). a<br />
Gregarious ascomata on the host surface. b, c Bitunicate asci. Note the<br />
wide pseudoparaphyses. d Section of the apical peridium comprising<br />
thick-walled cells of textura angularis. e–h Fusoid to broadly fusoid<br />
ascospores. Scale bars: a=0.5 mm, b–h=20 μm
Fungal Diversity<br />
Literature: Berkeley and Broome 1866; Keissler 1922;Massee<br />
1887; Saccardo 1878a; Sivanesan 1983.<br />
Type species<br />
Anomalemma epochnii (Berk. & Broome) Sivan., Trans.<br />
Br. Mycol. Soc. 81: 328 (1983). (Fig. 4)<br />
≡ Sphaeria epochnii Berk. & Broome, Ann. Mag. nat.<br />
Hist., Ser. 3 18: 128 (1866).<br />
Ascomata 340–500 μm high×170–286 μm diam., gregarious<br />
on the intertwined hyphae, superficial, papillate, wall<br />
black, coriaceous, roughened (Fig. 4a). Peridium composed of<br />
two types of cells, outer layer 17–22 μm wide, composed of<br />
heavily pigmented thick-walled cells of textura angularis,<br />
cells up to 8×13 μm diam., cell wall 1–1.5 μm thick, inner<br />
layer 30–34 μm thick, composed of hyaline thin-walled cells<br />
(Fig. 4d). Hamathecium of dense, long cellular pseudoparaphyses,<br />
2–4 μm broad, septate. Asci 75–108×9.5–12.5 μm<br />
(x ¼ 92:8 11:1mm, n=10), 8-spored, bitunicate, fissitunicate,<br />
dehiscence not observed, cylindro-clavate to clavate, with<br />
a furcate pedicel up to 6–25 μm long, with a small ocular<br />
chamber best seen in immature asci (ca. 2μm wide×1 μm<br />
high) (Fig. 4b and c). Ascospores 20–25(−30)×5–7.5 μm<br />
(x ¼ 23:1 6:3mm, n=10), obliquely uniseriate and partially<br />
overlapping to biseriate, fusoid to narrowly fusoid with narrowly<br />
rounded ends, brown, 1-septate, rarely 2- to 3-septate, deeply<br />
constricted at the median septum, smooth (Fig. 4e, f, g and h).<br />
Anamorph: Exosporiella fungorum (Fr.) P. Karst.<br />
(Sivanesan 1983).<br />
= Epochnium fungorum Fr., Syst. mycol. 3: 449 (1832).<br />
Mycelium composed of branched, septate, pale brown<br />
hyphae. Stroma none. Conidiophores macronematous or<br />
semi-macronematous, mononematous, hyaline, smooth,<br />
branched towards the apex. Conidiogenous cells monoblastic,<br />
cylindrical or doliform. Conidia cylindrical or<br />
ellipsoidal, dry, 3-4-septate, smooth, hyaline or pale brown.<br />
Material examined: UK, England, Warleigh near Bath,<br />
on fungus on bark (Epochnium sp.), Mar. 1866, leg.<br />
Warbright? (K(M):143936, syntype, ex herb. C.E.<br />
Broome).<br />
Notes<br />
Morphology<br />
Sphaeria epochnii was first described and illustrated by<br />
Berkeley and Broome (1866) from Britain and the anamorphic<br />
stage is the hyphomycetous Epochniella fungorum. Sphaeria<br />
epochnii has subsequently been transferred to Melanomma (as<br />
M. epochnii (Berk. & Broome) Sacc.; Saccardo 1878a),<br />
Byssosphaeria (as B. epochnii (Berk. & Broome) Cooke;<br />
Massee 1887) andChaetosphaeria (as C. epochnii (Berk. &<br />
Broome) Keissl.; Keissler 1922). The deposition of Sphaeria<br />
epochnii in Chaetosphaeria is obviously unacceptable, as<br />
Chaetosphaeria has unitunicate asci. Melanomma has been<br />
reported having Aposphaeria or Pseudospiropes anamorphs,<br />
which differs from Exosporiella (Sivanesan 1983). In addition,<br />
the presence of well developed prosenchymatous stroma<br />
in Sphaeria epochnii can also readily distinguish it from<br />
Melanomma (Sivanesan 1983).<br />
The gregarious ascomata and formation of prosenchymatous<br />
stroma of Anomalemma resembles those of Cucurbitaria,<br />
but the pleosporaceous dictyosporous ascospores of Cucurbitaria<br />
readily distinguish it from Anomalemma epochnii. In<br />
addition, the pseudoparenchymatous peridium, fungicolous<br />
habitat and brown 1-septate ascospores, which later becoming<br />
3-septate differ from any other pleosporalean genus. Thus a<br />
new genus, Anomalemma, was introduced to accommodate it<br />
(Sivanesan 1983). Anomalemma is presently monotypic.<br />
Phylogenetic study<br />
None.<br />
Concluding remarks<br />
Anomalemma epochnii certainly resembles Byssosphaeria<br />
in its ascomata clustering together in groups on closely<br />
intertwined hyphae and brown ascospores, and may well be<br />
included in this genus. Its fungicolous habitat, however,<br />
distinguishes it from Byssosphaeria.<br />
Appendispora K.D. Hyde, Sydowia 46: 29 (1994a).<br />
(?Didymellaceae)<br />
Generic description<br />
Habitat terrestrial, saprobic. Ascomata small, clustered,<br />
immersed, subglobose or irregularly pyriform. Peridium<br />
thin. Hamathecium of dense, long trabeculate pseudoparaphyses.<br />
Asci 8-spored, bitunicate, fissitunicate, cylindrical,<br />
apical rounded with ocular chamber and faint ring, with<br />
short pedicels. Ascospores uniseriate to partially overlapping,<br />
fusoid, brown, 1-septate, slightly constricted at the septum.<br />
Anamorphs reported for genus: none.<br />
Literature: Hyde 1994a.<br />
Type species<br />
Appendispora frondicola K.D. Hyde, Sydowia 46: 30<br />
(1994a). (Fig. 5)<br />
Ascomata 120–280 μm high×180–280 μm diam., clustered,<br />
immersed with minute ostioles visible through cracks<br />
or blackened dots on the host surface, subglobose or<br />
irregularly pyriform (Fig. 5a and b). Peridium 40 μm thick,<br />
comprising two types of cells; outer cells, small heavily<br />
pigmented thick-walled cells of textura angularis, innercells<br />
compressed, hyaline. Hamathecium of dense, very long<br />
trabeculate pseudoparaphyses, ca. 1μm broad, embedded in<br />
mucilage, hyaline, anastomosing (Fig. 5e). Asci 130–144×11–
Fungal Diversity<br />
Fig. 5 Appendispora frondicola<br />
(from BRIP 21354, holotype).<br />
a Immersed ascomata on<br />
host surface. b Valsoid configuration<br />
of the ascomata. c Cylindrical<br />
ascus. d Squash showing<br />
asci and numerous pseudoparaphyses.<br />
e Thin strands of<br />
anastomosing pseudoparaphyses.<br />
f, g Ascospores with one or two<br />
appendages. Scale bars:<br />
a, b=0.5 mm, c–g=30 μm<br />
13 μm, 8-spored, bitunicate, fissitunicate, cylindrical, with an<br />
ocular chamber and faint ring, with short pedicels (Fig. 5c and<br />
d). Ascospores 21–30×7–9 μm, uniseriate to partially overlapping,<br />
fusoid, brown, 1-septate, slightly constricted at the<br />
septum, with an irregular ridged ornamentation and 3–5<br />
narrow appendages at each end (Fig. 5f and g).<br />
Anamorph: none reported.<br />
Material examined: BRUNEL, Jalan, Muara, Simpang<br />
835, on dead rachis of Oncosperma horridum on forest floor,<br />
Nov. 1992, K.D. Hyde 1652 (BRIP 21354, holotype).<br />
Notes<br />
Morphology<br />
Appendispora was described as a saprobe of palm, and is<br />
characterized by small, immersed ascomata, bitunicate,<br />
fissitunicate asci, trabeculate pseudoparaphyses, brown, 1-<br />
septate, appendaged ascospores with irregular wall striations<br />
(Hyde 1994a).Basedonitstrabeculatepseudoparaphyses<br />
embedded within gel matrix and its brown<br />
ascospores, Appendispora was assigned to Didymosphaeriaceae<br />
(Barr 1987b; Hyde1994a).<br />
Phylogenetic study<br />
None.<br />
Concluding remarks<br />
The saprobic habitat and association with monocots,<br />
cylindrical asci, trabeculate pseudoparaphyses as<br />
well as its brown, 1-septate ascospores make it<br />
difficult to determine a better phylogenetic position than<br />
Didymellaceae.
Fungal Diversity<br />
Ascorhombispora L. Cai & K.D. Hyde, Cryptog. Mycol.<br />
28: 294 (2007). (<strong>Pleosporales</strong>, genera incertae sedis)<br />
Generic description<br />
Habitat freshwater, saprobic. Ascomata solitary or gregarious,<br />
superficial, globose to subglobose, dark brown to black, short<br />
papillate, ostiolate, coriaceous. Peridium relatively thin,<br />
textura angularis in longitudinal section, 2-layered. Hamathecium<br />
not observed. Asci 8-spored, obpyriform, broadly clavate<br />
to saccate, pedicellate, bitunicate, apex rounded, persistent.<br />
Ascospores overlapping 2-3-seriate, broadly fusoid to rhomboid,<br />
thick-walled, surrounded by mucilaginous sheath, 3-<br />
euseptate, not constricted at septa, median septum wide,<br />
forming a darker band, central cells large, trapezoid, dark<br />
brown to black, verruculose, polar end cells small and paler.<br />
Anamorphs reported for genus: none.<br />
Literature: Cai and Hyde 2007.<br />
Type species<br />
Ascorhombispora aquatica L. Cai & K.D. Hyde, Cryptog.<br />
Mycol. 28: 295 (2007). (Fig. 6)<br />
Ascomata 140–170 μm high×150–185 μm diam., solitary<br />
or gregarious, superficial, globose to subglobose, dark brown<br />
to black, short papillate, ostiolate, ostioles rounded, small,<br />
coriaceous. Peridium relatively thin, 10–18 μm wide, textura<br />
angularis in longitudinal section, composed of two layers of<br />
angular cells, outer later dark brown to black, relatively thickwalled,<br />
inner layer hyaline, relatively thin-walled (Fig. 6a and<br />
b). Hamathecium not observed. Asci 100–198×72–102 μm<br />
(x ¼ 186 88mm, n=15), 8-spored, obpyriform, broadly<br />
clavate to saccate, pedicellate, bitunicate, apex rounded,<br />
deliquescent (Fig. 6c, d and e). Ascospores 30.5–45×16–<br />
26.5 μm (x ¼ 38:5 21mm, n=25), overlapping 2-3-seriate,<br />
broadly fusoid to rhomboid, thick-walled, surrounded by<br />
mucilaginous sheath, 3-euseptate, not constricted at septa,<br />
median septum wide, forming a darker band, central cells<br />
large, trapezoid, 11–18 μm long, dark brown to black,<br />
verruculose, polar end cells small, hemispherical, 3.5–4 μm<br />
long, subhyaline to pale brown, smooth (Fig. 6f).<br />
Anamorph: none reported.<br />
Material examined: CHINA, Yunnan, Jinghong, on<br />
submerged bamboo in a small forest stream, 26 Jan. 2003,<br />
leg. det. L. Cai, CAI-1H31 (HKU(M) 10859, holotype).<br />
Notes<br />
Morphology<br />
Ascorhombispora was introduced as a monotypic<br />
genus from freshwater by Cai and Hyde (2007), and is<br />
characterized by superficial, coriaceous, non-stromatic<br />
ascomata, large, saccate asci; lack of interascal filaments<br />
and trapezoid (rhombic), 3-septate, dark brown to black<br />
ascospores with smaller end cells which are subhyaline to<br />
pale brown. Ascorhombispora is most comparable with<br />
Caryospora and Zopfia. But the globose to subglobose<br />
ascomata and thin peridium, saccate asci lacking interascal<br />
pseudoparaphyses, and the 3-septate, rhomboid ascospores<br />
with the paler end cells of Ascorhombispora differs from<br />
those of Caryospora (Cai and Hyde 2007).<br />
Phylogenetic study<br />
Phylogenetic analysis based on either SSU or LSU<br />
rDNA sequences indicated that Ascorhombispora aquatica<br />
belongs to <strong>Pleosporales</strong>, but its familial placement was left<br />
undetermined (Cai and Hyde 2007).<br />
Concluding remarks<br />
The sac-shaped asci and absence of pseudoparaphyses<br />
are uncommon in <strong>Pleosporales</strong>, especially among those<br />
from freshwater.<br />
Asteromassaria Höhn., Sber. Akad. Wiss. Wien, Math.-<br />
naturw. Kl., Abt. I 126: 368 (1917). (?Morosphaeriaceae)<br />
Generic description<br />
Habitat terrestrial, saprobic. Ascomata medium-sized,<br />
clustered, at first immersed and then breaking through<br />
the host surface and becoming superficial, globose,<br />
subglobose, coriaceous. Peridium 2-layered, thicker near<br />
the base. Hamathecium of dense, septate, cellular pseudoparaphyses<br />
which branch and anastomosing frequently<br />
between and above asci. Asci (4-)8-spored, bitunicate,<br />
cylindro-clavate to clavate, with a short truncated pedicel<br />
and a small ocular chamber. Ascospores obliquely uniseriate<br />
and partially overlapping to biseriate, fusoid to<br />
fusoid-ellipsoidal, pale brown when mature, 1-septate,<br />
some becoming 3-septate when old, constricted at the<br />
median septum.<br />
Anamorphs reported for genus: Scolicosporium (Sivanesan<br />
1984).<br />
Literature: Barr 1982a; b; 1993a; Boise 1985; Shoemaker<br />
and LeClair 1975; Sivanesan 1987; Tanaka et al. 2005.<br />
Type species<br />
Asteromassaria macrospora (Desm.) Höhn., F. von, Sber.<br />
Akad. Wiss. Wien, Math.-naturw. Kl., Abt. I 126: 368<br />
(1917). (Fig. 7)<br />
≡ Sphaeria macrospora Desm., Ann. Sci. Nat. Bot. 10:<br />
351 (1849).<br />
Ascomata 400–600 μm high×450–650 μm diam., 4–20<br />
clustered together, at first immersed and then breaking<br />
through the host surface and becoming superficial, globose,
Fungal Diversity<br />
Fig. 6 Ascorhombispora aquatica (from HKU(M) 10859, holotype).<br />
a Section of an ascoma. b Section of a partial peridium. c Immature<br />
ascus. d–f Mature asci with ascospores. Note the deliquescent ascal<br />
wall in f. Note the wide, dark band in the medium septum of<br />
ascospores in d and e and the mucilaginous sheath and paler end cells<br />
in e and f. Scale bars: a=20 μm, b–f=15 μm (figures referred to Cai<br />
and Hyde 2007)<br />
subglobose, not easily removed from the substrate, wall black,<br />
coriaceous, roughened, apex usually widely porate, with or<br />
without papilla (Fig. 7a). Peridium 70–90 μm wide, thicker<br />
near the base where it is up to 180 μm wide, comprising two<br />
cell types, outer cells composed of heavily pigmented<br />
small cells, cells 3–5 μm diam., inner layer composed of<br />
less pigmented cells of textura angularis, 10–20 μm<br />
diam. (Fig. 7b and c). Hamathecium of dense, septate, 2–<br />
3 μm broad, pseudoparaphyses which branch and anastomosing<br />
frequently between and above asci (Fig. 7d). Asci<br />
(180-)200–280×28–43 μm (x ¼ 230 35mm, n=10), 8-<br />
Fig. 7 Asteromassaria macrospora (from L, 1004). a Ascomata<br />
clustered in a group breaking through the host surface. b Section of an<br />
ascoma. c Section of a partial peridium. Note the cells of textura<br />
angularis. d Pseudoparaphyses. Note the branches. e Upper part of the<br />
ascus illustrating the ocular chamber. f Ascus with a short pedicel. g–j<br />
Ascospores. Note the mucilaginous sheath in G and minutely verruculose<br />
ornamentation in J. Scale bars: a=0.5 mm, b, c=100 μm, d–j=10 μm<br />
b
Fungal Diversity
Fungal Diversity<br />
spored (sometimes 4-spored), bitunicate, fissitunicate dehiscence<br />
not observed, cylindro-clavate to clavate, with a short<br />
truncated pedicel up to 30 μm, with a small ocular chamber<br />
(ca. 3μm wide×3 μm high) (Fig. 7e and f). Ascospores 50–<br />
58×(14-)18–21 μm (x ¼ 55:3 18:2mm, n=10), obliquely<br />
uniseriate and partially overlappingtobiseriate,fusoidto<br />
fusoid-ellipsoidal, with narrowly rounded ends, lightly<br />
brown when mature, 1-septate, some becoming 3-septate<br />
when old, constricted at the median septum, the upper<br />
cell often broader and longer than the lower one,<br />
minutely verrucose (Fig. 7g, h, i and j).<br />
Anamorph: Scolicosporium macrosporium (Berk.) B.<br />
Sutton.<br />
Acervuli immersed in bark, brown, discrete, up to<br />
250 μm diam., opening by irregular rupture of the<br />
overlaying tissues. Peridium of thin-walled angular cells.<br />
Conidiophores cylindrical, 1-2-septate, up to 30 μm long<br />
and 3–5 μm wide. Conidiogenous cells holoblastic, 1-2-<br />
annellate, cylindrical, hyaline. Conidia 100–190×12–<br />
15 μm, fusoid, pale brown with paler or hyaline ends, 7–<br />
17 transverse septate, smooth-walled, with a tapered apex<br />
and truncate base (adapted from Sivanesan 1984).<br />
Material examined: CZECH REPUBLIC, Mährisch-<br />
Welвkirchen (Hranice), Wsetin (Vsetin), Berg Čap., on<br />
Fagus sylvatica L., Aug. 1938, F. Petrak (L, 1004).<br />
Notes<br />
Morphology<br />
In this study we were unable to obtain the holotype,<br />
so we used a collection of Petrak’s. The main morphological<br />
characters of Asteromassaria are the medium- to<br />
large-sized, globose to depressed ascomata opening with a<br />
pore, clavate to oblong asci, narrowly cellular pseudoparaphyses,<br />
pale to dark brown, bipolar symmetric, mostly<br />
fusoid, distoseptate or euseptate ascospores (Barr 1993a).<br />
The bipolar symmetric ascospores of Asteromassaria can<br />
readily be distinguished from other genera of this family<br />
(Barr 1993a; Tanaka et al. 2005). Currently, it comprises<br />
12 species (Tanaka et al. 2005; http://www.mycobank.org,<br />
28-02-2009).<br />
Phylogenetic study<br />
Asteromassaria pulchra (Harkn.) Shoemaker & P.M.<br />
LeClair is basal to Morosphaeriaceae in the phylogenetic<br />
tree based on four genes, but its placement is<br />
influenced by taxon sampling that was different in<br />
several analyses.<br />
Concluding remarks<br />
Asteromassaria can be distinguished from other comparable<br />
genera, i.e. Pleomassaria and Splanchnonema by 1-<br />
septate and pale brown ascospores, thick-walled textura<br />
angularis peridium and Scolicosporium anamorphic stage<br />
(see under Pleomassaria).<br />
Astrosphaeriella Syd. & P. Syd., Annls mycol. 11: 260<br />
(1913). (?Melanommataceae)<br />
Generic description<br />
Habitat terrestrial, saprobic. Ascomata densely scattered or<br />
in small groups, erumpent through the outer layers of the<br />
host tissues to nearly superficial, reflexed pieces of the<br />
ruptured host tissue usually persisting around the base of<br />
the ascomata, often star-like, conical to semiglobose, with a<br />
central papilla. Peridium upper wall usually comprising a<br />
thick dark brittle pseudoparenchymatous layer, base usually<br />
flattened and thin-walled. Hamathecium of dense, filliform,<br />
trabeculate pseudoparaphyses, embedded in mucilage. Asci<br />
8-spored, bitunicate, fissitunicate, cylindro-clavate to narrowly<br />
fusoid. Ascospores narrowly fusoid with acute ends,<br />
hyaline, pale brown or brown, 1-3-septate.<br />
Anamorphs reported for genus: Pleurophomopsis (Hyde<br />
et al. 2011).<br />
Literature: von Arx and Müller 1975; Barr 1990a; Chen<br />
and Hsieh 2004; Hawksworth 1981; Hawksworth and<br />
Boise 1985; Hyde and Fröhlich 1998; Hyde et al. 2000;<br />
Kirk et al. 2001; Sydow and Sydow 1913; Tanaka and<br />
Harada 2005a; b; Tanaka et al. 2009.<br />
Type species<br />
Astrosphaeriella stellata Syd. & P. Syd., Annls mycol. 11:<br />
260 (1913). (Fig. 8)<br />
Ascomata 360–570 μm high×860–1150 μm diam.,<br />
densely scattered or in small groups, erumpent through the<br />
outer layers of the host tissues to nearly superficial, reflexed<br />
pieces of the ruptured host tissue usually persisting around<br />
the base of the ascomata, forming star-like flanges around the<br />
ascomata from the surface view; ascomata broadly conical,<br />
with a flattened base not easily removed from the substrate,<br />
wall black; apex with a central papilla which is black and<br />
shiny at maturity, scarcely projecting (Fig. 8a). Peridium 40–<br />
70 μm thick, carbonaceous and crisp, 1-layered, composed of<br />
very small dark brown thick-walled pseudoparenchymatous<br />
cells, cells 2–5 μm diam., cell wall 2–6 μm thick, in places at<br />
the base composed of hyaline cells of textura prismatica,<br />
cells 5×8 μm diam. (Fig. 8b). Hamathecium of dense, very<br />
long trabeculate pseudoparaphyses,
Fungal Diversity<br />
Fig. 8 Astrosphaeriella fusispora<br />
(BISH 145726). a Ascomata<br />
forming a small group on<br />
host surface. Note the remains<br />
of the host forming flanges<br />
around the ascomata. b Section<br />
of the partial peridium. Note the<br />
black peridium and wedge of<br />
palisade cells between the lateral<br />
and basal walls. c Asci in trabeculate<br />
pseudoparaphyses. d–f<br />
Narrowly fusoid ascospores.<br />
Scale bars: a=1 mm, b=<br />
100 μm, c=50 μm, d=15 μm,<br />
f=10 μm<br />
ends, hyaline, turning pale brown when mature, 1(−3)-<br />
septate, constricted at the median septum (Fig. 8d,e and f).<br />
Anamorph: none reported.<br />
Material examined: USA, Hawaii, Kapano Gulch, in<br />
bamboo culms, 5 Jun. 1947, leg. Kopf & Rogers, det. Miller<br />
(BISH 145726, as Astrosphaeriella fusispora Syd. & P. Syd.).<br />
Notes<br />
Morphology<br />
Astrosphaeriella has been treated as a synonym of Microthelia<br />
(von Arx and Müller 1975), but the large conical<br />
ascomata, numerous trabeculate pseudoparaphyses and 1-<br />
septate and elongated ascospores of Astrosphaeriella all<br />
disagree with those of Microthelia (Hawksworth 1981). It<br />
was assigned to Platystomaceae by Barr (1990a) in<strong>Pleosporales</strong><br />
or Melanommataceae by Kirk et al. (2001).<br />
Following a systematic study of Astrosphaeriella, only four<br />
species were accepted, i.e. A. aosimensis I. Hino & Katum.,<br />
A. stellata, A. trochus (Penz. & Sacc.) D. Hawksw. and A.<br />
venezuelensis M.E. Barr & D. Hawksw. (Hawksworth 1981),<br />
and it was defined as a tropical genus, occurring exclusively<br />
on palms or bamboo. Astrosphaeriella stellata was selected<br />
as the type of Astrosphaeriella,andA. fusispora was regarded
Fungal Diversity<br />
as a synonym of A. stellata (Hawksworth 1981). More taxa<br />
were subsequently added (Barr 1990a; Hawksworth and<br />
Boise 1985; Hyde and Fröhlich 1998), and the generic<br />
concept extended to include three elements: 1. typical semiimmersed<br />
to superficial ascomata with flattened base,<br />
cylindro-clavate asci with fusoid ascospores and trabeculate<br />
pseudoparaphyses, i.e. Astrosphaeriella sensu stricto (e.g.<br />
A. fusispora and A. vesuvius (Berk. & Broome) D. Hawksw.<br />
& Boise); 2. Trematosphaeria-like with rounded ascomata (e.<br />
g. A. africana D. Hawksw.); and 3. Massarina-like species<br />
with immersed ascomata (e.g. A. bakeriana (Sacc.) K.D.<br />
Hyde & J. Fröhl.) (Chen and Hsieh 2004; Tanaka and Harada<br />
2005a; b). Currently, a broad generic concept of Astrosphaeriella<br />
is accepted, and 47 taxa are included in Astrosphaeriella.<br />
Phylogenetic study<br />
Phylogenetic analysis based on LSU and SSU nurDNA<br />
sequence data indicates that Astrosphaeriella is polyphyletic,<br />
and located in the basal region of the <strong>Pleosporales</strong> between<br />
Testudinaceae and Zopfiaceae/Delitschiaceae (Tanaka et al.<br />
2009), or basal to Aigialaceae (Schoch et al. 2009). The<br />
genus is, however, clearly not related to Trematosphaeria as<br />
previously understood (Boise 1985).<br />
Concluding remarks<br />
Astrosphaeriella is currently polyphyletic and new<br />
collections of the different elements listed above are needed<br />
in order to understand the placement of various species. We<br />
suggest that some immersed bambusicolous species may<br />
belong in Tetraplospheariaceae.<br />
Asymmetricospora J. Fröhl. & K.D. Hyde, Sydowia 50:<br />
183 (1998). (?Melanommataceae)<br />
Generic description<br />
Habitat terrestrial, saprobic. Ascomata solitary or in small<br />
groups, immersed, black, lenticular in section, uni- or often<br />
multi-locular, with a central ostiole without tissue differentiation.<br />
Upper peridium carbonaceous, thicker at sides and apex.<br />
Lower peridium composed of irregular-shaped, hyaline cells.<br />
Hamathecium of trabeculate pseudoparaphyses, branching and<br />
anastomosing between and above asci, embedded in mucilage.<br />
Asci 8-spored, bitunicate, fissitunicate unknown, clavate, short<br />
pedicellate. Ascospores 1-septate, hyaline, constricted at the<br />
septum, with a broad, spreading mucilaginous sheath.<br />
Anamorphs reported for genus: none.<br />
Literature: Fröhlich and Hyde 1998.<br />
Type species<br />
Asymmetricospora calamicola J. Fröhl. & K.D. Hyde,<br />
Sydowia 50: 184 (1998). (Fig. 9)<br />
Ascomata 675–950 μm high×875–1500 μm diam.,<br />
solitary or in small groups of 2–10, immersed and forming<br />
slightly protruding domes on the substrate surface, with nearwhite<br />
rim around the central ostiole; in vertical view lenticular,<br />
multi- or rarely unilocular, individual locules 175–270 μm<br />
high×320–400 μm diam., with a flattened base, ostiole a<br />
central opening without tissue differentiation (Fig. 9a). Upper<br />
peridium 32–70 μm wide, carbonaceous, composed of a few<br />
layers of black walled cells of textura angularis. Lower<br />
peridium thinner, composed of hyaline cells of textura<br />
globulosa or textura prismatica (Fig. 9b). Hamathecium of<br />
long trabeculate pseudoparaphyses, 1.2–1.6(−2) μm wide,<br />
branching and anastomosing between and above asci,<br />
embedded in mucilage. Asci 137.5–207.5×26–35 μm<br />
(x ¼ 172:8 31:5mm, n=20), 8-spored, bitunicate, fissitunicate<br />
dehiscence not observed, clavate, with short pedicel (to<br />
25 μm), with ocular chambers (ca. 3μm wide×4 μm high)<br />
(Fig. 9c and d). Ascospores 35–55×10.5–15 μm<br />
(x ¼ 44:7 12:4mm, n=50), biseriate, navicular to obovoid,<br />
hyaline, becoming pale brown when senescent, straight or<br />
usually curved, smooth, asymmetric, 1-septate, the upper cell<br />
larger with a rounded end, basal cell with a tapering end,<br />
constricted at the septum, with spreading mucilaginous sheath<br />
(Fig. 9e, f and g) (data from Fröhlich and Hyde 1998).<br />
Anamorph: none reported.<br />
Material examined: AUSTRALIA, North Queensland,<br />
Palmerston, Palmerston National Park, on dead rattan of<br />
Calamus caryotoides A.Cunn. ex Mart., Mar. 1994, J.<br />
Fröhlich (HKU(M) 7794, holotype).<br />
Notes<br />
Morphology<br />
Asymmetricospora was introduced as a monotypic genus<br />
represented by A. calamicola based on its “absence of a<br />
subiculum, the absence of short dark setae around the papilla<br />
and its asymmetric ascospores” (Fröhlich and Hyde 1998).<br />
Because of the immersed ascomata, ostiole and peridium<br />
morphology, fissitunicate asci and trabeculate pseudoparaphyses,<br />
Asymmetricospora was assigned to Melanommataceae<br />
(sensu Barr 1990a; Fröhlich and Hyde 1998).<br />
Morphologically Asymmetricospora can be distinguished<br />
from its most comparable genus, Astrosphaeriella, by its<br />
ostiole, which is a simple opening without tissue differentiation,<br />
asymmetric ascospores, and the usually multiloculate<br />
fruiting body (Fröhlich and Hyde 1998).<br />
Phylogenetic study<br />
None.<br />
Concluding remarks<br />
The placement of Asymmetricospora under Melanommataceae<br />
remains to be confirmed.
Fungal Diversity<br />
Fig. 9 Asymmetricospora calamicola (from HKU(M) 7794, holotype).<br />
a Ascomata immersed in the substrate. b Section of the<br />
peridium. c Mature and immature asci in pseudoparaphyses (in cotton<br />
Barria Z.Q. Yuan, Mycotaxon 51: 313 (1994).<br />
(Phaeosphaeriaceae)<br />
Generic description<br />
Habitat terrestrial, parasitic. Ascomata small- to medium-sized,<br />
solitary or scattered, immersed, globose, subglobose, ostiolate,<br />
coriaceous. Apex with or without papilla and with a pore-like<br />
ostiole. Peridium 2-layered. Hamathecium of dense, long<br />
cellular pseudoparaphyses, septate, embedded in mucilage.<br />
blue). d Clavate ascus with a small ocular chamber. e–g Ascospores<br />
with sheath. Scale bars: a, b=0.5 mm, c=50 μm, d–g=20 μm<br />
Asci bitunicate, fissitunicate, cylindrical to clavate, with a short,<br />
furcate pedicel. Ascospores ellipsoid, hyaline at first, turning<br />
brown at maturity, 1-septate, strongly constricted at the septum.<br />
Anamorphs reported for genus: none.<br />
Literature: Yuan 1994.<br />
Type species<br />
Barria piceae Z.Q. Yuan, Mycotaxon 51: 314 (1994).<br />
(Fig. 10)
Fungal Diversity<br />
Fig. 10 Barria piceae (from<br />
NY 92003, isotype). a Ascoma<br />
on the host surface. Note the<br />
wide opening ostiole. b Section<br />
of the partial peridium with two<br />
types of cells. c, d Asci with<br />
ocular chambers and short pedicels.<br />
e, f Ellipsoid ascospores<br />
which are turning brown with<br />
thin sheath around them. Scale<br />
bars: a=0.5 mm, b=50 μm, c,<br />
d=20 μm, e, f=10 μm<br />
Ascomata 240–370 μm high×200–320 μm diam.,<br />
solitary, scattered, immersed, globose, subglobose, coriaceous,<br />
apex with or without papilla and with a pore-like<br />
ostiole (Fig. 10a). Peridium 20–35 μm thick, comprising<br />
two cell types, the outer cells comprising 3–4 layers of<br />
brown pseudoparenchymatous cells, cells 4–5 μm diam.,<br />
cell wall 2–3 μm thick, inner cells comprising 3–4 layers of<br />
pale brown compressed cells, cells 2×16 μm diam., cell<br />
wall 0.5–1.5 μm thick (Fig. 10b). Hamathecium of dense,<br />
long cellular pseudoparaphyses, 2–3 μm broad, septate.<br />
Asci 135–200(−220)×14–20 μm (x ¼ 156 16:6mm, n=<br />
10), 8-spored, bitunicate, fissitunicate, cylindrical to clavate,<br />
with a short, furcate pedicel, up to 22 μm long, with a<br />
large ocular chamber (ca. 4μm wide×3 μm high) (Fig. 10c<br />
and d). Ascospores 19–21.5×10–12 μm (x ¼ 20:4 11mm,<br />
n=10), uniseriate to partially overlapping, ellipsoid, hyaline<br />
or greenish with numerous small guttules at first and olive<br />
green to smoky brown at maturity, 1-septate, strongly<br />
constricted at the septum, foveolate, surrounded with sheath<br />
(Fig. 10e and f).<br />
Anamorph: none reported.<br />
Material examined: CHINA, Xinjiang Province, Uygur,<br />
Urumqi, Tianshan Mountain, on needles of Picea schrenkiana,<br />
1 Jul. 1992, Z.Q. Yuan (NY 92003, isotype).
Fungal Diversity<br />
Notes<br />
Morphology<br />
Barria was established by Yuan (1994) as a monotypic<br />
genus represented by B. piceae according to its “two-celled,<br />
pigmented ascospores, pseudoparenchymatous peridium and<br />
narrowly cellular pseudoparaphyses” thus differing in its<br />
combination of characters from all of the morphologically<br />
related dothideomycetous genera, such as Didymosphaeria,<br />
Didymopleella or Stegasphaeria. The taxon was considered<br />
to belong in Phaeosphaeriaceae. Ascomata and<br />
colour or shape of ascospores, however, readily distinguish<br />
it from other 1-septate Phaeosphaeriaceae genera,<br />
i.e. Didymella, Lautitia and Metameris (Yuan 1994).<br />
Barria piceae causes blight of spruce needles.<br />
Phylogenetic study<br />
None.<br />
Concluding remarks<br />
The status of Barria with its unusual verrucose ascospores<br />
and thick gel coating is uncertain. In many ways it<br />
resembles Belizeana, with its cylindrical asci, 1-septate,<br />
ellipsoid ascospores with sheath and verruculose surface<br />
(Kohlmeyer and Volkmann-Kohlmeyer 1987). However,<br />
the latter is a marine genus while Barria causes leaf blight<br />
of terrestrial Picea (Yuan 1994). The placement in<br />
Phaeosphaeriaceae seems logical based on the parasitic<br />
life style, thin and simple peridium, wide cellular pseudoparaphyses<br />
and brown ascospores. However, molecular<br />
data are needed to confirm this.<br />
Belizeana Kohlm. & Volkm.-Kohlm., Bot. Mar. 30: 195<br />
(1987). (<strong>Pleosporales</strong>, genera incertae sedis)<br />
Generic description<br />
Habitat marine, saprobic. Ascomata solitary, scattered, or<br />
in small groups, medium-sized, immersed to semiimmersed,<br />
subglobose to broadly ampulliform, black,<br />
ostiolate, carbonaceous. Peridium thin, comprising several<br />
layers of brown thin-walled cells of textura angularis.<br />
Hamathecium of dense, filliform pseudoparaphyses, rarely<br />
branched. Asci 8-spored, bitunicate, fissitunicate, broadly<br />
cylindrical to clavate, with a short pedicel and an ocular<br />
chamber. Ascospores uniseriate, broadly ellipsoidal, hyaline,<br />
turn pale brown when senescent, 1-septate, constricted<br />
at the septum, thick-walled, 2-layered, mature<br />
spores with tuberculate ornamentation between the two<br />
layers.<br />
Anamorphs reported for genus: Phoma-like (Kohlmeyer<br />
and Volkmann-Kohlmeyer 1987).<br />
Literature: Kohlmeyer and Volkmann-Kohlmeyer 1987.<br />
Type species<br />
Belizeana tuberculata Kohlm. & Volkm.-Kohlm., Bot.<br />
Mar. 30: 196 (1987). (Fig. 11)<br />
Ascomata 170–300 μm high×160–290 μm diam., solitary,<br />
scattered, or in small groups of 2–3, immersed to semiimmersed,<br />
subglobose to broadly ampulliform, carbonaceous,<br />
black, pale brown on the sides, ostiolate, epapillate or shortly<br />
papillate, ostiolar canal filled with a tissue of hyaline cells<br />
(Fig. 11a). Peridium 25–35 μm wide, comprising several<br />
layers thin-walled cells of textura angularis, which are<br />
hyaline inwardly, near the base composed of a hyaline<br />
hyphal mass producing asci, up to 20 μm thick (Fig. 11b, c<br />
and e). Hamathecium of dense, ca. 2 μm broad, filliform<br />
pseudoparaphyses, rarely branched, embedded in mucilage<br />
(Fig. 11g). Asci 145–170×20–30 μm (x ¼ 163 25mm, n=<br />
10), 8-spored, bitunicate, fissitunicate, broadly cylindrical to<br />
clavate with a short pedicel, thick-walled, with a small ocular<br />
chamber (Fig. 11d, f and h). Ascospores 21–26 × 13–18 μm<br />
(x ¼ 22 15mm, n = 10), uniseriate, broadly ellipsoidal,<br />
hyaline, turn pale brown when senescent, 1-septate, constricted<br />
at the septum, thick-walled, 2-layered, mature spores<br />
with tuberculate ornamentation between the two layers<br />
(Fig. 11i and j).<br />
Anamorph: Phoma-like (Kohlmeyer and Volkmann-<br />
Kohlmeyer 1987).<br />
Material examined: BELIZE, Twin Cays, on Laguncularia<br />
sp., 7 Apr. 1983, leg. & det. J. Kohlmeyer (Herb. J.<br />
Kohlmeyer No. 4398, holotype); AUSTRALIA, Towra<br />
Point, New South Wales, trunk of eroded tree with oysters<br />
and shipworms, intertidal zone, Botany Bay, 23 Aug. 1981<br />
(Herb. J. Kohlmeyer No. 4209, paratype).<br />
Notes<br />
Morphology<br />
Belizeana was formally established to accommodate B.<br />
tuberculata, an obligate marine fungus, which is characterized<br />
by verrucose ascospores (Kohlmeyer and Volkmann-<br />
Kohlmeyer 1987). Belizeana tuberculata canbeassignedto<br />
Pleosporaceae (<strong>Pleosporales</strong>) according to Luttrell’s (1973)<br />
treatment and keys of von Arx and Müller (1975), but cannot<br />
resolve a proper family based on Barr (1979a, 1983). The<br />
unique morphology together with obligate marine habitat<br />
makes B. tuberculata readily distinguishable from all other<br />
taxa of Pleosporaceae.<br />
Phylogenetic study<br />
None.<br />
Concluding remarks<br />
The ascospores of Belizeana tuberculata are most comparable<br />
with those of Acrocordiopsis patilii, but the superficial
Fungal Diversity<br />
Fig. 11 Belizeana tuberculata<br />
(from Herb. J. Kohlmeyer No.<br />
4398, holotype). a Immersed to<br />
semi-immersed ascomata. b, e<br />
Vertical section of an ascoma. c<br />
Section of a partial peridium. d<br />
Squash mounts with a large<br />
number of asci. f Broadly<br />
cylindrical ascus with a large<br />
ocular chamber. g Filliform<br />
pseudoparaphyses. h Apical part<br />
of an ascus. Note the large<br />
ocular chamber. i, j One-septate<br />
ascospores. Scale bars:<br />
a=0.3 mm, b=100 μm,<br />
c=20 μm, d, e=50 μm,<br />
f–j=10 μm<br />
conical ascomata of A. patilii are distinct from B. tuberculata.<br />
Thus, the familial placement of Belizeana is still undetermined.<br />
Biatriospora K.D. Hyde & Borse, Mycotaxon 26: 263<br />
(1986). (<strong>Pleosporales</strong>, genera incertae sedis)
Fungal Diversity<br />
Generic description<br />
Habitat marine, saprobic. Ascomata large, solitary or gregarious,<br />
immersed, subglobose to pyriform, ostiolate, papillate,<br />
periphysate, black, branching, carbonaceous. Hamathecium<br />
of dense, long trabeculate pseudoparaphyses, embedded in<br />
mucilage. Asci 8-spored, bitunicate, fissitunicate, cylindrical,<br />
with apical apparatus. Ascospores uniseriate to partially<br />
overlapping, fusoid, hyaline when young, becoming brown<br />
to dark brown at maturity, multi-septate towards each end,<br />
with a hyaline, globose refractive chamber or appendage at<br />
each end, not constricted at the septum.<br />
Anamorphs reported for genus: none.<br />
Literature: Hyde and Borse 1986; Suetrong et al. 2009.<br />
Type species<br />
Biatriospora marina K.D. Hyde & Borse, Mycotaxon 26:<br />
264 (1986). (Fig. 12)<br />
Ascomata 650–860 μm high×350–510 μm diam., solitary<br />
or gregarious, immersed, subglobose to pyriform, ostiolate,<br />
papillate, periphysate, black, carbonaceous (Fig. 12.2a). Hamathecium<br />
of dense, long trabeculate pseudoparaphyses, 1–<br />
1.5 μm broad, branching, embedded in mucilage. Asci 175–<br />
Fig. 12 1 Biatriospora marina<br />
(from IMI 297768, holotype). a,<br />
b Cylindrical asci. Note the<br />
mucilage pseudoparaphyses in<br />
(a) and the conspicuous ocular<br />
chamber in (b). c, d Ascospores<br />
with hyaline end chambers<br />
(arrowed). Scale bars: a, b=<br />
50 μm, c, d=20 μm. 2 Line<br />
drawings of Biatriospora marina<br />
(based on holotype). a Section<br />
through ascocarp showing<br />
asci and pseudoparaphyses. b<br />
Asci and pseudoparaphyses. c<br />
Ascospores. Scale bars: a=<br />
200 μm, b=40 μm, c=30 μm<br />
(figure with permission from<br />
Hyde and Borse 1986)
Fungal Diversity<br />
Fig. 12<br />
2 (continued)<br />
400×22–40 μm, 8-spored, bitunicate, fissitunicate, cylindrical,<br />
with long pedicels and apical apparatus (Fig. 12.1a, b, 2b).<br />
Ascospores 55–82×16–25 μm, uniseriate to partially overlapping,<br />
fusoid, hyaline when young, becoming brown to<br />
dark brown at maturity, 2-4-septate towards each end, and<br />
with a hyaline, globose refractive chamber or appendage<br />
at each end, 6–8×4–6 μm diam., not constricted at the<br />
septum (Fig. 12.1c, d, 2c).
Fungal Diversity<br />
Anamorph: none reported.<br />
Material examined: SEYCHELLES, 2 Jan. 1984 (Herb.<br />
IMI 297768 holotype).<br />
Notes<br />
Morphology<br />
Biatriospora was introduced to accommodate a<br />
marine fungus B. marina, which is characterized by<br />
horizontal ascomata and ascospores with polar, globose<br />
refractive chambers and polar septa (Hyde and Borse<br />
1986). Polar refractive chambers can also occur in other<br />
marine fungi, such as Lulworthia and Aigialus. The<br />
chambers have been proposed as important for spore<br />
attachment to substrates in a liquid environment (Hyde<br />
and Borse 1986).<br />
Phylogenetic study<br />
Multigene phylogenetic analysis indicated that Biatriospora<br />
marina forms a separate branch, sister to other families of<br />
<strong>Pleosporales</strong> (Suetrong et al. 2009), and maybe related to<br />
species in Roussoella (Plate 1).<br />
Concluding remarks<br />
The familial status of Biatriospora can not be determined.<br />
Bicrouania Kohlm. & Volkm.-Kohlm., Mycol. Res. 94:<br />
685 (1990). (?Melanommataceae)<br />
Generic description<br />
Habitat marine, saprobic. Ascomata immersed gregarious,<br />
erumpent to superficial, globose to subglobose, black,<br />
periphysate, coriaceous, epapillate or papillate, ostiolate.<br />
Peridium thin, 2-layered. Hamathecium of dense, long<br />
trabeculate pseudoparaphyses, branching and anastomosing<br />
between and above the asci. Asci 8-spored, bitunicate,<br />
fissitunicate, cylindrical, with a thick, furcate pedicel<br />
lacking ocular chamber. Ascospores obliquely uniseriate<br />
and partially overlapping, ellipsoidal with broadly rounded<br />
ends, reddish brown, 1-septate, thick-walled, constricted at the<br />
septum.<br />
Anamorphs reported for genus: none.<br />
Literature: Jones et al. 2009; Kohlmeyer and Volkmann-<br />
Kohlmeyer 1990.<br />
Type species<br />
Bicrouania maritima (P. Crouan & H. Crouan) Kohlm. &<br />
Volkm.-Kohlm., Mycol. Res. 94: 685 (1990). (Fig. 13)<br />
≡ Sphaeria maritima P. Crouan & H. Crouan, Florule du<br />
Finistére, Paris: 27 (1867) non Sphaeria maritima Cooke &<br />
Plowright, Grevillia 5: 120 (1877).<br />
Ascomata 320–440 μm high×370–460 μm diam.,<br />
gregarious, immersed, mostly erumpent to superficial,<br />
globose to subglobose, black, coriaceous, with a rough<br />
surface, papillate or epapillate, ostiolate, periphysate<br />
(Fig. 13a). Peridium 40–50 μm thick laterally, up to<br />
75 μm thick at the apex, thinner at the base, 2-layered,<br />
outer layer composed of small heavily pigmented<br />
pseudoparenchymatous cells, inner layer very thin,<br />
composed of hyaline thin-walled small cells, merging into<br />
pseudoparaphyses (Fig. 13a and b). Hamathecium of<br />
dense, very long trabeculate pseudoparaphyses, 0.8–<br />
1.2 μm broad, branching and anastomosing between and<br />
above the asci. Asci 170–225×17.5–22.5 μm<br />
(x ¼ 199:6 20mm, n=10), 8-spored, bitunicate, fissitunicate,<br />
cylindrical, with a thick, furcate pedicel which is up<br />
to 70 μm long, lacking ocular chamber (Fig. 13c, d and e).<br />
Ascospores 22–26×12–15 μm (x ¼ 24:5 13:3mm, n=<br />
10), obliquely uniseriate and partially overlapping, ellipsoidal<br />
with broadly rounded ends, reddish brown, 1-<br />
septate, slightly constricted at the septum, thick-walled,<br />
with a thick darkened band around the septum, smooth<br />
(Fig. 13c, d and e).<br />
Anamorph: none reported.<br />
Material examined: FRANCE, Finistère, on Halimone<br />
portulacoides (IMI 330806, isotype, as Sphaeria<br />
maritima).<br />
Notes<br />
Morphology<br />
When Kohlmeyer and Volkmann-Kohlmeyer (1990) studied<br />
the four marine Didymosphaeria species, the monotypic<br />
Bicrouania was established to accommodate B. maritima (as<br />
Didymosphaeria maritima (P. Crouan & H. Crouan) Sacc.),<br />
which could be distinguished from Didymosphaeria by its<br />
superficial ascomata lacking a clypeus, thick-walled asci and<br />
its association with algae (Kohlmeyer and Volkmann-<br />
Kohlmeyer 1990). Jones et al. (2009) agreed that it cannot<br />
be placed in Didymosphaeria based on its superficial<br />
ascomata, but that it does have many similarities with<br />
Didymosphaeria. Molecular data are required to determine<br />
its relationship with Didymosphaeria and to resolve its<br />
higher level placement.<br />
Phylogenetic study<br />
None.<br />
Concluding remarks<br />
Besides the morphological differences, its marine and<br />
substrate habitats also differ from Didymosphaeria.<br />
Bimuria D. Hawksw., Chea & Sheridan, N. Z. J. Bot. 17:<br />
268 (1979). (Montagnulaceae)
Fungal Diversity<br />
Fig. 13 Bicrouania maritima<br />
(from IMI 330806, isotype). a<br />
Section of an ascoma. b Section<br />
of papilla. Note the periphyses.<br />
c–e Eight-spored asci. Note the<br />
furcated pedicel. Scale bars: a,<br />
b=100 μm, c–e=20 μm<br />
Generic description<br />
Habitat terrestrial, saprobic. Ascomata solitary, superficial,<br />
globose, dark brown, epapillate, ostiolate. Peridium thin,<br />
pseudoparenchymatous. Hamathecium of few, cellular<br />
pseudoparaphyses, embedded in mucilage, rarely anastomosing<br />
and branching. Asci bitunicate, fissitunicate, broadly<br />
clavate with short pedicels, 2-3-spored. Ascospores muriform,<br />
broadly ellipsoid, dark brown with subhyaline end<br />
cells, verrucose.<br />
Anamorphs reported for genus: none.<br />
Literature: Barr 1987b; Hawksworth et al. 1979;<br />
Lumbsch and Huhndorf 2007.<br />
Type species<br />
Bimuria novae-zelandiae Hawksworth, Chea & Sheridan,<br />
N. Z. J. Bot. 17: 268 (1979). (Fig. 14)<br />
Ascomata (185-)200×310(-330) μm diam., solitary, scattered,<br />
semi-immersed or superficial, globose, hyaline when
Fungal Diversity<br />
Fig. 14 Bimuria novae-zelandiae (from <strong>CBS</strong> 107.79, isotype). a–c<br />
Asci with a short pedicel and small ocular chamber. d Immature ascus<br />
(in cotton blue). e Partial ascospore. Note the convex verrucae on the<br />
ascospore surface. f Released ascospores. Note the lighter end cells,<br />
germ pore and the longiseptum (arrowed). g Fissitunicate ascus<br />
dehiscent. Scale bars: a–g=20 μm<br />
young, turning dark brown to black when mature, ostiolate,<br />
the ostiole more or less sessile or raised into a very short neck.<br />
Peridium 5–8(-12) μm thick, comprising 2–3 layers of<br />
radically compressed pseudoparenchymatous cells, cells 10–<br />
15 μm diam. in surface view, cell wall 2–3 μm thick.<br />
Hamathecium consisting of few, 2.5–4 μm broad cellular<br />
pseudoparaphyses, embedded in mucilage, rarely anastomosing<br />
and branching, septate, 7–13 μm long between<br />
two septa. Asci (65-)80–95×20–32.5 μm (x ¼ 75:6<br />
29:4mm, n=10), (1-)2(-3)-spored, bitunicate, fissitunicate,<br />
broadly clavate, with a short and small knob-like pedicel<br />
which is up to 13 μm long, ocular chamber best seen in<br />
immature asci (Fig. 14a, b, c, d and g). Ascospores<br />
accumulating in a subglobose black shiny mass adhering<br />
together outside the ostiole, 55–68×25–28 μm (x ¼ 59<br />
26mm, n=10), broadly ellipsoid but becoming narrowed<br />
towards the poles, muriform with (5-)7 transverse septa, cells<br />
with (0-)l(-2) longitudinal septa in each cell, no constriction
Fungal Diversity<br />
at the septa, dark brown, the apical cells paler with no<br />
longitudinal septa, verruculose (Fig. 14e and f).<br />
Anamorph: none reported.<br />
Material examined: NEW ZEALAND, North Island,<br />
Wairarapa District, Nutty Farm, isolated from soil, 3 Mar.<br />
1978, Chea Chark Yen & J.E. Sheridan (<strong>CBS</strong> 107.79,<br />
isotype).<br />
Notes<br />
Morphology<br />
Bimuria novae-zelandiae was first isolated from soil of a<br />
barley field in New Zealand (Hawksworth et al. 1979). Based<br />
on B. novae-zelandiae, the genus is characterized by a very<br />
thin peridium, mostly 2-spored and fissitunicate asci as well as<br />
the muriform, dark brown, verrucose ascospores (Hawksworth<br />
et al. 1979). Because of its unique morphological characters,<br />
the familial placement of this genus has been debatable and it<br />
has been placed in Pleosporaceae (Hawksworth et al. 1979),<br />
in Phaeosphaeriaceae (Barr 1987b)andinMelanommataceae<br />
(Lumbsch and Huhndorf 2007).<br />
Morphologically, Bimuria is most comparable with<br />
some superficially similar or allied genera, in particular<br />
Montagnula (Hawksworth et al. 1979). However, the thick<br />
carbonaceous peridium distinguishes Montagnula from<br />
that of Bimuria (Hawksworth et al. 1979). In addition, the<br />
ascospores of Montagnula are discharged forcibly through<br />
the ostiole instead of forming a mass outside of the ostiole<br />
as in Bimuria (Hawksworth et al. 1979). Ascomauritiana<br />
lignicola V.M. Ranghoo & K.D. Hyde has somewhat<br />
similar ascospores in 4-spored asci, but this taxon has<br />
unitunicate asci (Ranghoo and Hyde 1999). The morphological<br />
characters of Bimuria, such as ascospore release<br />
and large, thick-walled ascospores may be an adaptation to<br />
its soil-borne habitat (Hawksworth et al. 1979).<br />
Phylogenetic study<br />
Bimuria novae-zelandiae was found to be closely related<br />
to Phaeodothis winteri (Niessl) Aptroot (syn. Didymosphaerella<br />
opulenta (De Not.) Checa & M.E. Barr) and<br />
Montagnula opulenta (De Not.) Aptroot in analysis of<br />
combined sequences, i.e. SSU rDNA, LSU rDNA, RPB2<br />
and TEF1 sequences (Schoch et al. 2006, 2009). These two<br />
species had been included by Barr (2001) in her new family<br />
Montagnulaceae.<br />
Concluding remarks<br />
We agree with Barr (2001) and include the genus in<br />
Montagnulaceae based on both morphological and phylogenetic<br />
characters.<br />
Bricookea M.E. Barr, Mycotaxon 15: 346 (1982).<br />
(?Phaeosphaeriaceae)<br />
Generic description<br />
Habitat terrestrial, saprobic (or parasitic?). Ascomata small- to<br />
medium-sized, solitary, scattered, or in small groups, immersed,<br />
erumpent to superficial, depressed globose, papillate,<br />
ostiolate. Peridium thin. Hamathecium filliform, cellular<br />
pseudoparaphyses, embedded in mucilage, anastomosing,<br />
septate. Asci bitunicate, fissitunicate, cylindrical, cylindroclavate<br />
or slightly obclavate, with a short knob-like pedicel,<br />
with an ocular chamber. Ascospores hyaline, ellipsoid to<br />
narrowly obovoid, 3-septate, constricted at each septum.<br />
Anamorphs reported for genus: none.<br />
Literature:Barr1982a;Berlese1896;Holm1957; Shoemaker<br />
and Babcock 1989a.<br />
Type species<br />
Bricookea sepalorum (Vleugel) M.E. Barr, Mycotaxon 15:<br />
346 (1982). (Fig. 15).<br />
≡ Metasphaeria sepalorum Vleugel, Svensk bot. Tidskr.<br />
2: 369 (1908).<br />
Ascomata 120–250 μm high×170–440 μm diam., solitary,<br />
scattered, or in small groups, or forming locules in massive<br />
stromatic tissues, initially immersed, becoming erumpent, to<br />
nearly superficial, depressed globose, black, membraneous,<br />
roughened; apex rounded, sometimes very short and almost<br />
inconspicuous, with a somewhat slit-like or Y-shaped ostiole<br />
(Fig. 15a). Peridium 16–30 μm wide, comprising two types of<br />
cells, outer cells heavily pigmented thick-walled textura<br />
angularis, cells 4.5–8 μm diam., cell wall 1–1.5 μm thick,<br />
inner cells of subhyaline thin-walled textura angularis, cells<br />
larger than outer cells (Fig. 15b). Hamathecium of long<br />
cellular pseudoparaphyses, 1.5–2 μm broad, embedded in<br />
mucilage, anastomosing, septate. Asci 63–83×9.5–11 μm<br />
(x ¼ 73:8 10:8mm, n=10), 8-spored, bitunicate, fissitunicate,<br />
oblong, cylindro-clavate or slightly obclavate, with a short<br />
knob-like pedicel which is 5–13 μm long, with an ocular<br />
chamber (Fig. 15c, d and e). Ascospores (14-)15.5–19×5–<br />
7 μm (x ¼ 16:9 5:9mm, n=10), obliquely uniseriate and<br />
partially overlapping to biseriate, ellipsoid to narrowly<br />
obovoid, hyaline, 3-septate, constricted at each septum, the<br />
cells above central septum often broader than the lower ones,<br />
smooth (Fig. 15f, g, h, i and j).<br />
Anamorph: none reported.<br />
Material examined: SWEDEN, on Juncus filliformis,<br />
Stockholm, J. Vleugel. Jul. 1907 (S, type as Metasphaeria<br />
sepalorum Vleugel).<br />
Notes<br />
Morphology<br />
Bricookea was formally established by Barr (1982a) asa<br />
monotypic genus represented by B. sepalorum basedonits
Fungal Diversity<br />
Fig. 15 Bricookea sepalorum (from S, type). a Ascomata on host<br />
surface (arrowed). b Section of partial peridium. Note thick-walled<br />
out layer and thin-walled inner layer. c–e Cylindrical to slightly<br />
obclavate asci with short knob-like pedicels. f–j Hyaline, 3-septate<br />
smooth-walled ascospores. Scale bars: a=0.5 mm, b=50 μm, c–j=<br />
10 μm
Fungal Diversity<br />
“globose to depressed ascomata, slit-like ostiole with labial<br />
cells, bitunicate asci, cellular pseudoparaphyses, and hyaline<br />
septate ascospores”. Bricookea was morphologically assigned<br />
to Phaeosphaeriaceae. Holm(1957) checked the authentic<br />
collections from North America and type material from<br />
Europe, and observed that the ascospores of collections from<br />
North America were significantly larger than those from the<br />
type material from Sweden. Thus, Shoemaker and Babcock<br />
(1989a) considered that the collections from North America<br />
represented a new species, which they introduced as B. barrae<br />
Shoemaker & C.E. Babc. Although the short slit-like<br />
ostiole has previously been reported (Shoemaker and<br />
Babcock 1989a), it is inconspicuous in the type specimen<br />
from Sweden. Currently, only two species are accommodated<br />
in this genus.<br />
Phylogenetic study<br />
None.<br />
Concluding remarks<br />
The knob-shaped pedicel, slit-like ostiole, hyaline<br />
ascospores as well as the herbaceous substrate all disagree<br />
with any current pleosporalean family. Thus, we temporarily<br />
retain this genus under Phaeosphaeriaceae until DNA<br />
sequence comparisons can be carried out.<br />
Byssolophis Clem., in Clements & Shear, Gen. fung., Edn 2<br />
(Minneapolis): 286 (1931). (<strong>Pleosporales</strong>, genera incertae<br />
sedis)<br />
Generic description<br />
Habitat terrestrial, saprobic. Ascomata medium-sized, gregarious,<br />
semi-immersed to erumpent, coriaceous, ovoid, with a<br />
conspicuous elongate slit-like ostiole on the top. Peridium not<br />
observed. Hamathecium of dense, long pseudoparaphyses,<br />
anastomosing and branching between and above the asci. Asci<br />
8-spored, bitunicate, fissitunicate, cylindrical or cylindroclavate,<br />
with a furcate pedicel. Ascospores fusoid, hyaline,<br />
turning faintly brown when old, 1-septate, with a short<br />
terminal appendage at each end.<br />
Anamorphs reported for genus: none.<br />
Literature: Clements and Shear 1931; Holm 1986; Müller<br />
and von Arx 1962.<br />
Type species<br />
Byssolophis byssiseda (Flageolet & Chenant.) Clem., Gen.<br />
Fung. (Minneapolis): 286 (1931). (Fig. 16)<br />
≡ Schizostoma byssisedum Flageolet & Chenant., in<br />
Chenantaise, Bull. Soc. mycol. Fr. 35: 125 (1919).<br />
Ascomata 300–450 μm high×600–750 μm long×<br />
350–420 μm broad, gregarious, semi-immersed to<br />
erumpent, coriaceous, ovoid with a flattened base and<br />
apex with a elongate slit-like ostiole, up to 700 μm long and<br />
200 μm wide (Fig. 16a). Peridium not observed. Hamathecium<br />
of dense, long pseudoparaphyses, up to 1.5–2.5 μm<br />
broad, anastomosing and branching between and above the<br />
asci (Fig. 16b). Asci 80–105×(5-)7.5–10 μm<br />
(x ¼ 91 8mm; n=10), 8-spored, bitunicate, fissitunicate,<br />
cylindrical or cylindro-clavate, with a furcate pedicel and a<br />
small ocular chamber (J-) (Fig. 16d). Ascospores 18–20<br />
(−28)×4.5–6(−7.5) μm (x ¼ 20:8 5:7mm, n=10), uniseriate<br />
to biseriate, fusoid, hyaline, turning faintly brown when<br />
old, 1-septate, with 1–2 distinct oil drops in each cell and<br />
usually with a short terminal appendage at each end<br />
(Fig. 16c).<br />
Anamorph: none reported.<br />
Material examined: on decaying wood (K(M):164030,<br />
isotype).<br />
Notes<br />
Morphology<br />
Byssolophis was introduced as a monotypic genus based<br />
on B. byssiseda, which is characterized by its semiimmersed,<br />
gregarious, ovoid ascomata, with a conspicuous<br />
central apical ostiolar slit (Holm 1986). Subsequently, two<br />
more species were introduced, viz. B. ampla (Berk. &<br />
Broome) L. Holm and B. sphaerioides (P. Karst.) E. Müll.<br />
(Holm 1986; Müller and von Arx 1962).<br />
Phylogenetic study<br />
The current phylogeny places Byssolophis sphaerioides<br />
in proximity of Hypsostromataceae without resolving any<br />
sister taxa (Plate 1).<br />
Concluding remarks<br />
The slit-like ostiole, cylindrical asci, hyaline and 1-<br />
septate ascospores as well as the form of pseudoparaphyses<br />
are similar to species in Lophiostoma. Thus, Byssolophis<br />
may be a synonym of Lophiostoma.<br />
Byssosphaeria Cooke, Grevillea 7: 84 (1879).<br />
(Melanommataceae)<br />
Generic description<br />
Habitat terrestrial, saprobic. Ascomata medium-sized,<br />
scattered to gregarious, superficial, globose, subglobose<br />
to turbinate, non papillate with white, orange, red or green<br />
ostiolar region, wall black. Hamathecium of dense,<br />
long trabeculate pseudoparaphyses, embedded in mucilage,<br />
anastomosing between and above the asci. Asci<br />
bitunicate, fissitunicate, clavate to nearly cylindrical,<br />
with a furcate pedicel. Ascospores fusoid with narrow
Fungal Diversity<br />
Fig. 16 Byssolophis byssiseda<br />
(from K(M):164030, isotype). a<br />
Ascomata gregarious on the host<br />
surface. b Numerous pseudoparaphyses.<br />
c Fusoid ascospores<br />
with or without terminal appendages.<br />
d Clavate ascus with a<br />
short furcate pedicel. Scale bars:<br />
a=1 mm, b–d=10 μm<br />
ends, straight or slightly curved, brown, 1-septate when<br />
young.<br />
Anamorphs reported for genus: Pyrenochaeta or Chaetophoma-like<br />
(Barr 1984; Hawksworth et al. 1995; Samuels<br />
and Müller 1978).<br />
Literature: von Arx and Müller 1975; Barr 1984; Boise<br />
1984; Bose 1961; Chen and Hsieh 2004; Cooke and<br />
Plowright 1879; Hyde et al. 2000; Luttrell 1973; Mugambi<br />
and Huhndorf 2009b; Müller and von Arx 1962; Samuels<br />
and Müller 1978.<br />
Type species<br />
Byssosphaeria keitii (Berk. & Broome) Cooke [as ‘Byssosphaeria<br />
keithii’], (1879). (Fig. 17)<br />
≡ Sphaeria keitii Berk. & Broome [as ‘Sphaeria keithii’],<br />
Ann. Mag. Nat. Hist., IV 17: 144 (1876).<br />
Ascomata 360–500(−600) μm high×420–640 μm<br />
diam., scattered or in small groups, superficial with<br />
basal subiculum anchoring on the substrate, globose,<br />
subglobose to turbinate, non-papillate with pore-like
Fungal Diversity<br />
Fig. 17 Byssosphaeria schiedermayriana (from K(M):108784,<br />
holotype). a Superficial ascomata on the host surface. b Brown, 1-<br />
septate ascospores. c Section of the lateral peridium. Note the outer<br />
textura angularis and inner textura epidermoidea cells. d, e Furcate<br />
asci with a long pedicel. f Dehiscent ascus. Scale bars: a=0.5 mm, c=<br />
50 μm, b, d–f=15 μm<br />
ostiole, ostiolar region sometimes with orange and<br />
greenish tint, wall black, roughened, coriaceous<br />
(Fig. 17a). Peridium 55–85 μm thick, peridium outside<br />
of the substrate comprising two cell types, outer layer<br />
composed of brown thick-walled cells of textura epidermoidea,<br />
cells 1–3 μm diam., inner layer composed of<br />
small hyaline cells, cells 3–5 μm diam., merging into<br />
pseudoparaphyses; peridium inside the substrate one<br />
layer, composed of large pale brown cells of textura<br />
angularis, cells6–13 μm diam. (Fig. 17c). Hamathecium<br />
of dense, long trabeculate pseudoparaphyses, 1–2 μm<br />
broad, embedded in mucilage, anastomosing between and<br />
above the asci. Asci 90–120(−148)×10–14 μm, 8-spored,<br />
bitunicate, fissitunicate, cylindro-clavate to clavate,<br />
biseriate above and uniseriate below, pedicel 15–20<br />
(−53) μm long, the immature asci usually with longer<br />
and furcate pedicel (−68 μm) (Fig. 17d,e and f).<br />
Ascospores 29–34(−38)×5.5–8(−10) μm, fusoid with<br />
narrow ends, mostly straight, sometimes slightly curved,<br />
smooth, pale brown, 1-septate, becoming 3-septate after<br />
discharge, with hyaline appendages at each acute to<br />
subacute end; in some mature spores the appendage may<br />
be absent (Fig. 17b).<br />
Anamorph: Pyrenochaeta sp. (Barr 1984; Samuels and<br />
Müller 1978).<br />
Pycnidia 70–500 μm diam. Conidiogenous cells phialidic,<br />
lining cavity, 5–8×4–6 μm to 5–10×3–6 μm. Conidia<br />
2.5–3.5(−4)×1.5–2(−3) μm, hyaline, ellipsoid or subglobose<br />
(Barr 1984).<br />
Material examined: ERIE, Dublin, Glasnevin Botanic<br />
Garden, on old rope, Jun. 1872, W. Keit (K(M):108784,<br />
holotype, asSphaeria keitii Berk. & Broome).
Fungal Diversity<br />
Notes<br />
Morphology<br />
Byssosphaeria was introduced by Cooke and Plowright<br />
(1879) based on its superficial ascomata seated on a “tomentose<br />
subiculum of interwoven threads”, which includes various<br />
species in Sphaeria and Byssisedae, and was validly typified by<br />
B. keitii (Cooke 1878). Byssosphaeria keitii was treated as a<br />
synonym of B. schiedermayeriana (Fuckel) M.E. Barr by<br />
Sivanesan (1971), and B. schiedermayeriana exclusively occurs<br />
in tropical regions or greenhouse environments in temperate<br />
regions (Barr 1984). Morphologically, B. keitii is characterized<br />
by its large ascomata with orange to reddish plain apices, and is<br />
closely related to B. rhodomphala (Berk.) Cooke (Barr 1984).<br />
For a long time, Byssosphaeria was assigned to<br />
Herpotrichia sensu lato, and Byssosphaeria schiedermayeriana<br />
was renamed as H. schiedermayeriana Fuckel<br />
(von Arx and Müller 1975; Bose 1961; Luttrell 1973;<br />
Müller and von Arx 1962; Sivanesan 1971). After studying<br />
Herpotrichia in North America, Barr (1984) accepted a<br />
relatively narrow generic concept, Herpotrichia sensu<br />
stricto, and revived Byssosphaeria; this proposal is<br />
supported by phylogenetic study (Mugambi and Huhndorf<br />
2009b). Currently Byssosphaeria comprises 32 species<br />
(http://www.mycobank.org, 08-01-2009).<br />
Phylogenetic study<br />
The monophyletic nature of Byssosphaeria is well<br />
demonstrated, as well as its familial status in Melanommataceae<br />
(Mugambi and Huhndorf 2009b).<br />
Concluding remarks<br />
Orange and greenish plain apices exist in the specimen<br />
we examined, which is different from records as “orange,<br />
bright or dull reddish plain apices” by Barr (1984). This<br />
might be because different specimens have different<br />
colours, or there may be a variation of apical colour within<br />
a single species, as both orange and green can coexist on<br />
the same ascoma (see Fig. 17a). The coloured apical rim,<br />
together with the trabeculate pseudoparaphyses as well as<br />
the presence of subiculum make Byssosphaeria readily<br />
distinguishable from other morphologically comparable<br />
genera, e.g. Herpotrichia and Keissleriella (Hyde et al. 2000).<br />
Calyptronectria Speg., Anal. Mus. nac. Hist. nat. B. Aires<br />
19: 412 (1909). (Melanommataceae)<br />
Generic description<br />
Habitat terrestrial, saprobic. Ascomata small- to mediumsized,<br />
solitary, scattered, or in small groups, immersed,<br />
lenticular to subglobose, papillate, ostiolate. Hamathecium<br />
of long, filliform pseudoparaphyses, branching and anastomosing,<br />
embedded in mucilage. Asci 4- to 8-spored,<br />
bitunicate, fissitunicate, cylindrical to cylindro-clavate, with<br />
a short, furcate pedicel. Ascospores muriform, broadly<br />
fusoid to fusoid with broadly to narrowly rounded ends,<br />
hyaline.<br />
Anamorphs reported for genus: none.<br />
Literature: Barr 1983; Rossman et al. 1999; Spegazzini<br />
1909.<br />
Type species<br />
Calyptronectria platensis Speg., Anal. Mus. nac. Hist. nat.<br />
B. Aires 19: 412 (1909). (Fig. 18)<br />
Ascomata 120–270 μm high×170–400 μm diam., solitary,<br />
scattered, immersed, lenticular to subglobose, papillate,<br />
ostiolate (Fig. 18a and b). Apex with a small and slightly<br />
protruding papilla. Peridium 18–30 μm wide, comprising two<br />
types of cells, outer layer composed of pseudoparenchymatous<br />
cells, cells 3–6 μm diam., cell wall 1–2 μm thick, inner<br />
layer comprising less pigmented cells, merging with pseudoparaphyses<br />
(Fig. 18b and c). Hamathecium of long, filliform<br />
pseudoparaphyses, 1–2 μm broad, branching and anastomosing,<br />
embedded in mucilage. Asci 98–140×12.5–20 μm<br />
(x ¼ 107 15:4mm, n=10), 8-spored, sometimes 4-spored,<br />
bitunicate, fissitunicate, cylindrical to cylindro-clavate, with<br />
a short, furcate pedicel, 12–20 μm long,withanocular<br />
chamber (to 4 μm wide×3 μm high) (Fig. 18e and f).<br />
Ascospores 17– 22.5 μ m ×(6.3-)7.5– 10 μ m<br />
(x ¼ 19:8 7:6mm, n=10), biseriate above and uniseriate<br />
below, ellipsoid to broadly fusoid with broadly to narrowly<br />
rounded ends, hyaline, usually with (3-)5 transverse septa,<br />
with or without 1–3 longitudinal septa in the central cells,<br />
constricted at the median septum, the upper cell often broader<br />
than the lower one, smooth, surrounded by an irregular<br />
hyaline gelatinous sheath up to 3 μm thick (in dry specimen)<br />
(Fig. 18d).<br />
Anamorph: none reported.<br />
Material examined: ARGENTINA, La Plata, on decaying<br />
branches of Manihot carthaginensis (Jacq.) Müll., Sept.<br />
1906, Spegazzini (LPS 1209, holotype).<br />
Notes<br />
Morphology<br />
Calyptronectria is a relatively poorly studied genus,<br />
which was formally established based on C. argentinensis<br />
Speg. and C. platensis, with C. platensis being chosen as<br />
the generic type (Spegazzini 1909). Morphologically,<br />
Calyptronectria is characterized by its immersed ascomata,
Fungal Diversity<br />
Fig. 18 Calyptronectria platensis (from LPS 1209, holotype). a<br />
Appearance of ascomata scattered in the substrate (after removing the<br />
out layer of the substrate). Note the protruding papilla. b Section of an<br />
ascoma. c Section of the partial peridium. Note the lightly pigmented<br />
trabeculate pseudoparaphyses and hyaline, muriform ascospores<br />
as well as its peridium that turns reddish brown in KOH<br />
(Rossman et al. 1999) (not shown here). Subsequently, C.<br />
indica Dhaware was introduced from India, and Barr (1983)<br />
transferred Teichospora ohiensis Ellis & Everh. to Calyptronectria<br />
as C. ohiensis (Ellis & Everh.) M.E. Barr.<br />
However, this proposal is inappropriate as the type specimen<br />
of T. ohiensis is “unitunicate” (Barr 1983; Rossman et al.<br />
1999). Subsequently, Rossman et al. (1999) transferred<br />
pseudoparenchymatous cells. d Released ascospores with mucilaginous<br />
sheath. e Eight-spored asci in hamathecium and embedded in gel<br />
matrix. f Ascus with a short pedicel. Scale bars: a=0.5 mm, b=<br />
100 μm, c=50 μm, d–f=10 μm<br />
Calyptronectria ohiensis to Thyridium (as T. ohiense (Ellis<br />
& Everh.) Rossman & Samuels).<br />
Phylogenetic study<br />
None.<br />
Concluding remarks<br />
The immersed ascomata, trabeculate pseudoparaphyses,<br />
bitunicate asci, hyaline and muriform ascospores as well as
Fungal Diversity<br />
the reaction of peridium to KOH (turns reddish brown)<br />
make it distinguishable from all other reported genera<br />
(Rossman et al. 1999). Thus Calyptronectria is a morphologically<br />
well defined genus.<br />
Carinispora K.D. Hyde, J. Linn. Soc., Bot. 110: 97 (1992).<br />
(<strong>Pleosporales</strong>, genera incertae sedis)<br />
Generic description<br />
Habitat marine, saprobic. One or two ascomata per<br />
stroma. Ascomata scattered or in small groups, developing<br />
beneath the host epidermis, erumpent, lenticular,<br />
ostiolate, lacking periphyses. Peridium pale brown,<br />
composed of thin-walled elongated cells at the sides<br />
and thick-walled cells of textura epidermoidea at the<br />
base. Hamathecium of dense, long filliform pseudoparaphyses,<br />
embedded in mucilage, anastomosing between<br />
and above the asci, rarely septate. Asci 8-spored,<br />
bitunicate, fissitunicate, clavate to cylindrical, with a<br />
short furcate pedicel, apex with an ocular chamber and<br />
apical ring. Ascospores biseriate, narrowly fusoid, yellow<br />
to pale brown, multi-septate, constricted at the septa, the<br />
two central cells being the largest, surrounded by a<br />
gelatinous sheath.<br />
Anamorphs reported for genus: none.<br />
Literature: Hyde 1992a, 1994b.<br />
Type species<br />
Carinispora nypae K.D. Hyde, J. Linn. Soc., Bot. 110: 99<br />
(1992). (Fig. 19)<br />
One or two ascomata per stroma. Ascomata up to 0.8 mm<br />
diam., scattered or in small groups, developing beneath the<br />
host epidermis, crust-like, as circular spots, wall brown, with<br />
a small central ostiole, in section 225–285 μm high×510–<br />
750 μm diam., lenticular, ostiolar canal lacking periphyses<br />
(Fig. 19a and b). Peridium 35–45 μm wide at sides, pale<br />
brown, at sides composed of a thin layer of thin-walled<br />
elongate cells, fusing with the stromatic tissue and host cells,<br />
at the base composed of thick-walled cells, forming a textura<br />
epidermoidea and fusing with host cells. A wedge of pale<br />
brown hyphae forming a textura porrecta is present at the<br />
rim (Fig. 19c). Hamathecium of dense, long filliform<br />
pseudoparaphyses 1–3 μm broad, embedded in mucilage,<br />
anastomosing between and above the asci, rarely septate.<br />
Asci 142–207×14.2–19.8 μm, 8-spored, bitunicate, fissitunicate,<br />
clavate to cylindrical, with a furcate pedicel, up to<br />
40 μm long, apex with an ocular chamber and apical ring (to<br />
2 μm wide×3 μm high, J-), developing from ascogenous<br />
tissue at the base of the ascocarp (Fig. 19d, e, f, g and h).<br />
Ascospores 42–66×7–10.6 μm, biseriate, narrowly fusoid<br />
with broadly to narrowly rounded ends, somewhat curved,<br />
yellow to pale brown, yellow in mass, 7-8-septate, constricted<br />
at the septa, the two central cells being the largest,<br />
surrounded by a gelatinous sheath; the sheath has a central<br />
“spine” and curved polar extrusions (Fig. 19i and j).<br />
Anamorph: none reported.<br />
Material examined: BRUNEI DARUSSALAM, Tungit<br />
Api Api mangrove, from decaying intertidal fronds of Nypa<br />
fruticans Wurmb., 14 Apr. 1987, K.D. Hyde (BRIP 17106,<br />
holotype).<br />
Notes<br />
Morphology<br />
Carinispora is distinguished from Phaeosphaeria by its<br />
saprobic life style and lenticular ascomata formed under the<br />
host epidermis, peridium structure and sheath surrounding the<br />
ascospores (Hyde 1992a, 1994b). Two species were reported,<br />
i.e. C. nypae and C. velatispora K.D. Hyde.<br />
Phylogenetic study<br />
Suetrong et al. (2009) could not resolve Carinispora<br />
nypae in a phylogeny based on four genes.<br />
Concluding remarks<br />
Both Carinispora nypae and C. velatispora are reported<br />
as marine fungi, which should be taken into consideration<br />
for their familial placement.<br />
Caryosporella Kohlm., Proc. Indian Acad. Sci., Pl. Sci. 94:<br />
355 (1985). (?Melanommataceae)<br />
Generic description<br />
Habitat marine, saprobic. Ascomata densely scattered or<br />
gregarious, superficial, subglobose, black, papillate, ostiolate,<br />
periphysate, carbonaceous. Peridium carbonaceous. Hamathecium<br />
of dense, long trabeculate pseudoparaphyses, anastomosing<br />
and branching above the asci. Asci 8-spored, bitunicate,<br />
fissitunicate, cylindrical. Ascospores ellipsoidal to broadly fusoid<br />
with narrowly hyaline rounded ends, deep reddish brown, thickwalled,<br />
1-septate with hyaline germ pore at each end.<br />
Anamorphs reported for genus: suspected spermatia<br />
(Kohlmeyer 1985).<br />
Literature: Eriksson 2006; Kohlmeyer 1985; Lumbsch and<br />
Huhndorf 2007.<br />
Type species<br />
Caryosporella rhizophorae Kohlm., Proc. Indian Acad.<br />
Sci., Pl. Sci. 94: 356 (1985). (Fig. 20)<br />
Ascomata 0.8–1.1 mm high×0.9–1.2 mm diam., densely<br />
scattered or gregarious, superficial with a flattened base, not<br />
easily removed from the host surface, subglobose, black, short
Fungal Diversity<br />
papillate, ostiolate, periphysate, carbonaceous (Fig. 20a<br />
and b). Peridium 120–150 μm thick at sides, up to<br />
200 μm thick at the apex, thinner at the base, 3-layered,<br />
outer layer composed of golden-yellow, very thickwalled<br />
cells of textura epidermoidea, mixed with subglobose,<br />
large cells near the surface, cells 7–15 μm<br />
diam., middle layer composed of deep brown, very thickwalled<br />
cells of textura epidermoidea, inner layer composed<br />
of hyaline, thin-walled cells of textura prismatica,<br />
up to 50×5 μm diam., merging with pseudoparaphyses<br />
(Fig. 20b, c and d). Hamathecium of dense, long<br />
trabeculate pseudoparaphyses, 1.5-2 μm wide, anastomosing<br />
and branching above the asci. Asci 225–250<br />
(−275)×14–17 μm (x ¼ 137 16:3mm, n=10), 8-spored,<br />
bitunicate, fissitunicate, cylindrical, with a long, narrowed,<br />
pedicel which is up to 75 μm long, apical<br />
characters not observed (Fig. 20e). Ascospores 25–28<br />
(−30)×9–13 μm (x ¼ 26:8 11mm, n=10), uniseriate to<br />
partially overlapping, ellipsoidal to broadly fusoid with<br />
narrow hyaline rounded ends, deep reddish brown, thickwalled,<br />
1-septate with hyaline germ pore at each end, slightly<br />
constricted at the septum, verruculose, sometimes with “net”-<br />
like ridged ornamentations (Fig. 20f and g).<br />
Anamorph: suspected spermatia (Kohlmeyer 1985).<br />
Material examined: BELIZE, Twin Cays, tip of prop<br />
root of Rhizophora mangle, 18 Mar. 1984, J. Kohlmeyer<br />
(NY. Herb. J. Kohlmeyer No. 4532a, holotype).<br />
Notes<br />
Morphology<br />
Caryosporella was formally established by Kohlmeyer<br />
(1985) based on the obligate marine fungus, C. rhizophorae,<br />
which is characterized by its superficial ascomata,<br />
3-layered peridium, filliform trabeculate pseudoparaphyses,<br />
and brown, 1-septate ascospores. Caryosporella was<br />
originally assigned to Massariaceae despite several major<br />
differences, such as the superficial ascomata, reddish<br />
brown ascospores (Kohlmeyer 1985). Subsequently,<br />
Caryosporella was assigned to Melanommataceae<br />
(Eriksson 2006; Lumbsch and Huhndorf 2007).<br />
Phylogenetic study<br />
Suetrong et al. (2009) showed that a single isolate of<br />
Caryosporella rhizophorae does not reside in <strong>Pleosporales</strong>,<br />
but is related to Lineolata rhizophorae (Kohlm. & E.<br />
Kohlm.) Kohlm. & Volkm.-Kohlm. and placed in Dothideomycetidae<br />
incertae sedis.<br />
Concluding remarks<br />
As an obligate marine fungus, the familial placement of<br />
Caryosporella rhizophorae is uncertain but it may not<br />
belong to <strong>Pleosporales</strong>.<br />
Fig. 19 Carinispora nypae (from BRIP 17106, holotype). a b<br />
Ascomata on the host surface. b Section of an ascoma. c Section of<br />
a partial peridium. d, e, g, h Asci with ocular chambers and short<br />
pedicels. f The ocular chamber and apical ring of ascus. i–j Narrowly<br />
fusoid ascospores. Scale bars: a=1 mm, b, c=100 μm, d, g, h=50 μm,<br />
e, f, i, j =10 μm<br />
Chaetomastia (Sacc.) Berl., Icon. fung. (Abellini) 1: 38<br />
(1890). (Teichosporaceae)<br />
≡ Melanomma subgen. Chaetomastia Sacc., Syll. fung.<br />
(Abellini) 2: 113 (1883).<br />
Generic description<br />
Habitat terrestrial, saprobic. Ascomata relatively small,<br />
scattered, or in small groups, superficial, globose or<br />
subglobose, black, papillate, ostiolate, coriaceous. Peridium<br />
relatively thin, 1-layered, composed of heavily<br />
pigmented cells of textura angularis. Hamathecium of<br />
dense, long cellular pseudoparaphyses, embedded in<br />
mucilage. Asci mostly 4-spored, bitunicate, fissitunicate,<br />
broadly cylindrical with a furcate pedicel, with a large<br />
ocular chamber, especially apparent in immature asci.<br />
Ascospores ellipsoid to broadly fusoid with broadly to<br />
narrowly rounded ends, brown, 3-septate, constricted at all<br />
septa.<br />
Anamorphs reported for genus: coelomycetous where<br />
known: conidia hyaline or brown, aseptate or 1-septate<br />
(Aposphaeria- orConiothyrium-like) (Barr 1989c).<br />
Literature: Barr 1987b, 1989c; 1993a; b; 2002; Berlese<br />
1890; Clements and Shear 1931; Eriksson 1999; Eriksson<br />
and Hawksworth 1987, 1998; Holm 1957; Leuchtmann<br />
1985; Saccardo 1883.<br />
Type species<br />
Chaetomastia hirtula (P. Karst.) Berl., Icon. fung. (Abellini)<br />
1: 38 (1890). (Fig. 21)<br />
≡ Sphaeria hirtula P. Karst., Fungi Fenn. Exs. N. 825 (1869).<br />
Ascomata 214–286 μm high×210–258 μm diam.,<br />
scattered or in groups, superficial, globose, wall black;<br />
apex often opening with a broad pore within slightly<br />
raised papilla, up to 30 μm diam., coriaceous (Fig. 21a).<br />
Peridium 20–26 μm thick, 1-layered, composed of<br />
heavily pigmented cells of textura angularis, cells up to<br />
5×15 μm diam., cell wall up to 3.5 μm thick (Fig. 21b).<br />
Hamathecium of dense, long cellular pseudoparaphyses,<br />
embedded in mucilage. Asci 90–130×12.5–17.5(−22.5)<br />
μm (x ¼ 111 16:3mm, n=10), mostly 4-spored, bitunicate,<br />
fissitunicate, broadly cylindrical, with a furcate<br />
pedicel, 18–48 μm long, with a large ocular chamber<br />
best seen in immature asci (to 3 μm wide×3 μm high)<br />
(Fig. 21c and d). Ascospores 20.5–27×7–10 μm
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Fungal Diversity<br />
Fig. 20 Caryosporella rhizophoriae (from NY. Herb. J. Kohlmeyer<br />
No. 4532a, holotype). a Gregarious ascomata on host surface. b<br />
Section of an ascoma. c, d Section of partial peridium at sides (c) and<br />
base (d). Note the three layers. e Asci with long peduncles in<br />
pseudoparaphyses. f, g Ascospores. Note the “net”-like ridged<br />
ornamentation of spore surface and hyaline germ pores. Scale bars:<br />
a=1 mm, b=200 μm, c–e=100 μm, f, g=10 μm<br />
(x ¼ 23:5 8:2mm, n=10), uniseriate to partially overlapping,<br />
ellipsoid to broadly fusoid with broadly to<br />
narrowly rounded ends, brown, 3-septate, verruculose,<br />
constricted at all septa, constricted at the median septum,<br />
the cell above the central septum often broader than the<br />
others (Fig. 21e and f).
Fungal Diversity<br />
Fig. 21 Chaetomastia hirtula (from H, FFE 825, kleptotype). a<br />
Superficial ascomata gregarious on the host surface. b Section of a<br />
partial peridium. Note the cells of textura angularis with relatively<br />
thick wall. c, d Cylindrical asci with long and furcate pedicels. e, f<br />
Brown, 3-septate ascospores. Scale bars: a=0.5 mm, b=50 μm, c–f=<br />
15 μm<br />
Anamorph: none reported.<br />
Material examined: FINLAND, ETELÄ-HÄME (EH/<br />
Ta), Tammela, Mustiala, På Rub. id., 8 May 1866. P.A.<br />
Karsten (H, FFE 825, kleptotype).<br />
Notes<br />
Morphology<br />
Chaetomastia was introduced by Saccardo (1883) asa<br />
subgenus of Melanomma, and five species were included,<br />
i.e. M. canescens Speg., M. cucurbitarioides Speg.,<br />
M. hirtulum (P. Karst.) Sacc., M. hispidulum Sacc. and<br />
M. pilosellum P. Karst. Berlese (1890) promoteditto<br />
genus rank. Subsequently, Chaetomastia hirtula (P.<br />
Karst.) Berl. was selected as the lectotype species of<br />
the genus (Clements and Shear 1931). Chaetomastia has<br />
been regarded as having unitunicate asci (Eriksson and<br />
Hawksworth 1986, 1998; Eriksson 1999). However its<br />
bitunicate status was confirmed by Holm (1957). Holm<br />
(1957) treated C. hirtula as Melanomma hirtulum (P.
Fungal Diversity<br />
Karst.) Sacc., and Leuchtmann (1985) transferred this<br />
species to Montagnula sensu lato based on the ascospore<br />
morphology and the hyphae surrounding the ascomata.<br />
Barr (1987b) suggested that ascoma, peridium structure<br />
and ascospore characters pointed Montagnula sensu<br />
stricto to Phaeosphaeriaceae, while the characters of<br />
ascomata and peridium structure of Chaetomastia were<br />
thought to fit the definition of Dacampiaceae (Barr<br />
1987b). In particular, the peridium and ascospore characters<br />
of C. hirtula are comparable with those of the generic<br />
type of Massariosphaeria (M. phaeospora). Thus, Barr<br />
(1989c) accepted Massariosphaeria sensu stricto and<br />
assigned the phragmosporous species of Massariosphaeria<br />
sensu lato to Chaetomastia.<br />
Barr (2002) later assigned Chaetomastia to Teichosporaceae<br />
based on its saprobic or hypersaprobic lifestyle,<br />
occurring on woody stems and peridium structure, and this<br />
is widely followed (Eriksson 2006; Lumbsch and Huhndorf<br />
2007). Currently, 11 species are accepted in this genus<br />
(http://www.indexfungorum.org/).<br />
Phylogenetic study<br />
None.<br />
Concluding remarks<br />
Familial placement of Chaetomastia is undetermined<br />
currently but has been included in the Teichosporaceae<br />
by authoritative sources (Eriksson 2006; Lumbsch and<br />
Huhndorf 2007) or the Dacampiaceae (http://www.<br />
indexfungorum.org/).<br />
Chaetoplea (Sacc.) Clem., Gen. Fung. (Minneapolis): 275<br />
(1931). (?Phaeosphaeriaceae)<br />
≡ Pyrenophora subgen. Chaetoplea Sacc., Syll. fung.<br />
(Abellini) 2: 279 (1883).<br />
Generic description<br />
Habitat terrestrial, saprobic. Ascomata small to medium,<br />
immersed, erumpent to superficial, globose to subglobose,<br />
papillate, ostiolate. Peridium not examined. Hamathecium<br />
of dense, long, narrowly cellular pseudoparaphyses. Asci 8-<br />
spored or 4-spored, bitunicate, fissitunicate, cylindroclavate,<br />
with a thick, furcate pedicel. Ascospores ellipsoid<br />
or fusoid, pale brown to brown, phragmosporous or<br />
muriform.<br />
Anamorphs reported for genus: Microdiplodia-like (Barr<br />
1990b).<br />
Literature: Barr 1981; 1987a; b; 1990b; Clements and<br />
Shear 1931; Ramaley and Barr 1995; Yuan and Barr<br />
1994.<br />
Fig. 22 Chaetoplea calvescens (from FH-81113, isotype). a, b Fourspored<br />
and 8-spored asci. c Released ascospores. Scale bars: a–c=<br />
10 μm<br />
Type species<br />
Chaetoplea calvescens (Fr.) Clem., Gen. Fung. (Minneapolis):<br />
275 (1931). (Fig. 22)<br />
≡ Sphaeria calvescens Fr. Scleromyc. Sueciae 401.<br />
Ascomata not examined. Peridium not examined.<br />
Hamathecium of dense, long, narrow cellular pseudoparaphyses,<br />
2–3 μm broad, septate, branching and anastomosing.<br />
Asci 90–110×10–12 μm, 8-spored, rarely 4-spored,<br />
bitunicate, fissitunicate, cylindro-clavate, with a thick,<br />
furcate pedicel which is up to 30 μm long (Fig. 22a<br />
and b). Ascospores 13–18×5.5–7 μm, obliquely uniseriate<br />
and partially overlapping, broadly fusoid to oblong<br />
with broadly rounded ends, pale brown, 2-3-septate,<br />
constricted at the septa, containing four refractive globules<br />
(Fig. 22c).<br />
Note: The specimen is only a slide, and no peridium or<br />
ascomata information could be obtained.<br />
Anamorph: coelomycetous, conidia yellowish, 1-<br />
septate, 9–13×4–5(−8) μm (Webster and Lucas 1959);<br />
Microdiplodia henningsii Staritz=Chaetodiplodia caudina<br />
Karst. (Sutton 1980) (referred to Barr 1990b (p50)).<br />
Material examined: SWEDEN, sub-collection: Curtis<br />
Herbarium, verified by R.A. Shoemaker, leg. E.M. Fries<br />
401 (FH-81113, isotype, microscope slide).<br />
Notes<br />
Morphology<br />
Chaetoplea was introduced based on C. calvescens,<br />
which has been regarded as similar to Pleospora or<br />
Leptosphaeria (Eriksson and Hawksworth 1987;<br />
Wehmeyer 1961; von Arx and Müller 1975). Based on<br />
the differences in ascomata, peridium structure, pseudoparaphyses<br />
as well as its anamorphic stage, Chaetoplea<br />
was maintained as a separate genus (Barr 1990b;<br />
Yuan and Barr 1994). Chaetoplea sensu lato was<br />
accepted by Barr (1990b), which included some species<br />
of Teichospora as well as the subgenus Pleospora subg.<br />
Cylindrosporeae.<br />
The following is from the label of specimen.<br />
“Sphaeria calvescens, Scler. Suecicae (Ed. 2) 401. No<br />
specimen of Scler. Suecicae 401 is now at Uppsala<br />
according to R. Santesson 1966. This Curtis Herbarium<br />
specimen in the Farlow Herbarium is isotype.<br />
Wehmeyer (1961) in his Pleospora monograph did<br />
not study any portion of the Scler. Suecicae exsiccatus<br />
b
Fungal Diversity
Fungal Diversity<br />
401, nor did Webster & Lucas in the taxonomic and<br />
life-history study (Trans. Brit. Myc. Soc. 42, 332–<br />
342. 1959) of this species.<br />
The specimen has most of the features described by<br />
Webster & Lucas including the presence of the<br />
conidial state Microdiplodia henningsii Staritz. I did<br />
not see vertical septa in the ascospores. Webster &<br />
Lucas note that vertical septa may be occasionally be<br />
lacking. The fungus is otherwise as they describe it<br />
although some perithecia collapse and appear cupulate.”—by<br />
R.A. Shoemaker.<br />
Phylogenetic study<br />
None.<br />
Concluding remarks<br />
The substrate of Chaetoplea sensu Barr (1990b) can<br />
be herbaceous stalks, decorticated wood or periderm, or<br />
old cotton cloth and string, which may indicate its<br />
heterogeneous nature. The ascospores seem very much<br />
like Phaeosphaeria which may be an earlier name; more<br />
details concerning the ascomatal, peridial and hamathecial<br />
structures are needed to make any conclusion.<br />
Cilioplea Munk, Dansk botanisk Arkiv 15: 113 (1953).<br />
(<strong>Pleosporales</strong>, genera incertae sedis)<br />
Generic description<br />
Habitat terrestrial, saprobic. Ascomata small- to medium-sized,<br />
solitary, scattered or in small groups, immersed, globose or<br />
subglobose, papilla covered with short and blackish setae,<br />
coriaceous. Peridium thin, comprising small heavily pigmented<br />
thick-walled cells of textura angularis. Hamathecium<br />
of cellular pseudoparaphyses. Asci 8-spored, bitunicate,<br />
fissitunicate, broadly clavate, with a short, furcate pedicel, and<br />
small ocular chamber. Ascospores fusoid to narrowly fusoid<br />
with narrowly rounded ends, pale brown to reddish brown,<br />
multi-transverse septa, usually with one longitudinal septum in<br />
some central cells, constricted at the primary septum.<br />
Anamorphs reported for genus: none.<br />
Literature: Barr 1990b, 1992b; Crivelli 1983; Lumbsch<br />
and Huhndorf 2007; Müller 1951; Munk 1953, 1957.<br />
Type species<br />
Cilioplea coronata (Niessl) Munk, Dansk botanisk Arkiv<br />
15: 113 (1953). (Fig. 23)<br />
≡ Pleospora coronata Niessl, Notiz. Pyr.: 16 (1876).<br />
Ascomata 170–290 μm high×200–410 μm diam., solitary,<br />
scattered, or in small groups, immersed, globose or subglobose,<br />
wall black, papilla raised, 50–80 μm high, with short<br />
and blackish setae, coriaceous (Fig. 23a). Peridium 9–15 μm<br />
thick laterally, up to 28 μm thick at the apex, thinner at the<br />
base, 1-layered, composed of small heavily pigmented thickwalled<br />
cells of textura angularis, cells up to 4×2.5 μm diam.,<br />
cell wall 2–3 μm thick, apex cells smaller and walls thicker<br />
(Fig. 23b). Hamathecium of long cellular pseudoparaphyses,<br />
2–3 μm broad. Asci (60-)80–108×10–15 μm<br />
(x ¼ 85:3 12:1mm, n=10), 8-spored, bitunicate, fissitunicate,<br />
broadly clavate, with a short, thick, furcate pedicel, 5–<br />
15 μm long, and a small ocular chamber (to 3 μm wide×<br />
2 μm high) (Fig. 23c and d). Ascospores 21–27.5×5.5–<br />
7.5 μm (x ¼ 24 6:7mm, n=10), biseriate to uniseriate at<br />
base, fusoid to narrowly fusoid with narrowly rounded ends,<br />
pale reddish brown, 5–7 transverse septa (mostly 5), usually<br />
with one longitudinal septum in some central cells, deeply<br />
constricted at the median septum, the part above the primary<br />
septum shorter and broader, smooth-walled.<br />
Anamorph: none reported.<br />
Material examined: GERMANY, Hadiberg. on Reseda<br />
lutea Hadiberg, 20 Sept. 1875, Niessl (M 175-89-290,<br />
lectotype; M 175-89-291, type).<br />
Notes<br />
Morphology<br />
Cilioplea was introduced by Müller (1951) as a<br />
subgenus of Pleospora, and this was followed by Munk<br />
(1957), who had earlier proposed it as a separate genus<br />
typified by C. coronata based on its hairy papilla, clavate<br />
asci as well as its “perfectly paraphysoid” (see Munk<br />
1953). A relatively narrow concept of Pleospora was<br />
accepted by Crivelli (1983), and four species was<br />
assigned under the separate genus Cilioplea, viz. C.<br />
coronata, C. genisticola (Fautrey & Lambotte) Crivelli,<br />
C. kansensis (Ellis & Everh.) Crivelli and C. nivalis<br />
(Niessl) Crivelli. Subsequently, another six species were<br />
added (Barr 1990b, 1992b). Currently, ten species are<br />
included under Cilioplea.<br />
Phylogenetic study<br />
None.<br />
Concluding remarks<br />
The most striking character of Cilioplea is its setose<br />
papilla, which has been shown to have no phylogenetic<br />
significance in Lentitheciaceae (Zhang et al. 2009a).<br />
Cilioplea was assigned under Lophiostomataceae (Lumbsch<br />
and Huhndorf 2007), but there is little morphological<br />
similarity with the Lophiostomataceae sensu stricto (Zhang<br />
et al. 2009a). Thus its familial placement needs further study.<br />
Crivellia Shoemaker & Inderb., in Inderbitzin, Shoemaker,<br />
O’Neill, Turgeon & Berbee, Can. J. Bot. 84: 1308 (2006).<br />
(Pleosporaceae)
Fungal Diversity<br />
Fig. 23 Cilioplea coronata (M 175-89-290, lectotype). a Immersed<br />
ascomata in small groups on the host surface (the covering host tissue<br />
was removed). b Section of a partial ascoma. Note the thin peridium. c<br />
Clavate asci within pseudoparaphyses. d Ascus with a small ocular<br />
chamber. Scale bars: a=0.5 mm, b=100 μm, c=50 μm, d=10 μm<br />
Generic description<br />
Habitat terrestrial, hemibiotrophic or parasitic. Ascomata<br />
small- to medium-sized, scattered, immersed, erumpent to<br />
nearly superficial, papillate, ostiolate. Peridium thin, composed<br />
of two cells types, outer cells of thick walled and<br />
textura angularis, inner cells thin-walled, yellow. Hamathecium<br />
of dense, long and thin pseudoparaphyses. Asci (4-)<br />
8-spored, bitunicate, fissitunicate dehiscence not observed,<br />
broadly cylindrical to cylindrical, with a short,<br />
furcate pedicel and an ocular chamber. Ascospores<br />
fusoid to broadly fusoid, pale brown, septate, sometimes<br />
with one or two vertical septa in the middle cells, constricted<br />
at the septa.<br />
Anamorphs reported for genus: Brachycladium (Inderbitzin<br />
et al. 2006).<br />
Literature: Inderbitzin et al. 2006.<br />
Type species<br />
Crivellia papaveracea (De Not.) Shoemaker & Inderb.,<br />
Can. J. Bot. 84: 1308 (2006). (Fig. 24)<br />
≡ Cucurbitaria papaveracea De Not., Sfer. Ital.: 62<br />
(1863).<br />
Ascomata 210–260 μm high×300–380 μm diam.,<br />
densely scattered, immersed, erumpent to nearly superficial,<br />
flattened globose, dark brown, papillate, ostiolate<br />
(Fig. 24a). Peridium 25–30 μm thick, thicker near the apex<br />
and thinner at the base, composed of two cell types, outer<br />
cells of thick-walled and textura angularis, cells up to 10×<br />
5 μm diam., cell wall 2–4 μm thick, inner cells thin-walled,<br />
yellow (Fig. 24b). Hamathecium of dense, long, 1–2 μm
Fungal Diversity<br />
broad, rarely septate pseudoparaphyses. Asci 85–125×10–<br />
13 μm (x ¼ 106 11mm, n=10), (4-)8-spored, bitunicate,<br />
fissitunicate dehiscence not observed, broadly cylindrical to<br />
cylindrical, with a short, furcate pedicel, with a relatively large<br />
ocular chamber (Fig. 24c and d). Ascospores (16-)19–24×5–<br />
7.5 μm (x ¼ 20:4 6:3mm, n=10), overlapping uniseriate to<br />
rarely biseriate, fusoid to broadly fusoid, pale brown, 3-<br />
septate, sometimes with one or two vertical septa in the<br />
middle cells, constricted at the septa, the upper cell often<br />
broader than the lower one, smooth-walled.<br />
Anamorph: Brachycladium penicillatum (Corda) Fr.<br />
(Inderbitzin et al. 2006).<br />
Material examined: AUSTRIA, Vienna, on decaying<br />
stems of Papaver rhoeas L., 28 Oct. 2001, W. Jaklitsch<br />
(UBC F14995, epitype).<br />
Notes<br />
Morphology<br />
Crivellia was separated from Pleospora and introduced<br />
as a new genus by Inderbitzin et al. (2006) based on their<br />
differences in ascospore morphology and anamorphic<br />
stages. Crivellia is characterized by having small- to<br />
medium-sized ascomata, and yellow, 3-septate ascospores<br />
Fig. 24 Crivellia papareracea<br />
(from UBC F14995, epitype).<br />
a Gregarious ascomata immersed<br />
within the host surface.<br />
b Section of an ascoma. c Asci<br />
within pseudoparaphyses. d<br />
Cylindrical ascus with a<br />
short pedicel. Scale bars:<br />
a=1 mm, b=100 μm, c,<br />
d=30 μm
Fungal Diversity<br />
with one or two vertical septa in central cells. Its<br />
Brachycladium anamorphic stage with phragmosporous<br />
conidia also differs from that of Stemphylium, which is<br />
the anamorphic stage of Pleospora (Inderbitzin et al. 2006).<br />
Currently, two species are included within Crivellia, i.e. C.<br />
homothallica Inderb. & Shoemaker and C. papaveracea.<br />
Phylogenetic study<br />
Crivellia papaveracea was shown to be closely related to<br />
some species of Alternaria, and its pleosporaceous status was<br />
confirmed following molecular studies (Inderbitzin et al. 2006).<br />
Concluding remarks<br />
Crivellia seems to belong to Pleosporaceae, and may be<br />
closely related to Pleospora.<br />
Decaisnella Fabre, Annls Sci. Nat., Bot., sér. 6 9:112<br />
(1878). (<strong>Pleosporales</strong>, genera incertae sedis)<br />
Generic description<br />
Habitat terrestrial, saprobic. Ascomata medium to large,<br />
immersed to erumpent, clypeate, papillate, ostiolate. Hamathecium<br />
of dense, long, cellular pseudoparaphyses, rarely<br />
septate, embedded in mucilage. Asci mostly 4- or 8-spored,<br />
rarely 2-spored, cylindrical to cylindro-clavate, with a<br />
furcate pedicel. Ascospores muriform, dark brown, oblong<br />
with broadly rounded ends.<br />
Anamorphs reported for genus: none.<br />
Literature: Barr 1986; 1990a; b; Fabre 1878; Saccardo<br />
1883.<br />
Type species<br />
Decaisnella spectabilis Fabre, Annls Sci. Nat., Bot., sér. 6<br />
9: 112 (1879). (Fig. 25)<br />
Ascomata 520–680 μm high×430–600 μm diam.,<br />
solitary, scattered, or in small groups of 2–3, immersed<br />
to erumpent, clypeate, globose or subglobose, black,<br />
roughened, with a blunt papilla up to 170 μm high,<br />
apex with a round ostiole, coriaceous (Fig. 25a).<br />
Peridium 70–90 μm thick at sides, thicker near the apex,<br />
comprising two types of cells; part immersed in host<br />
tissue, outer layer pseudoparenchymatous, 55–65 μm<br />
thick, pigmented, inner layer composed of lightly pigmented<br />
to hyaline thin-walled compressed cells, 15–<br />
23 μm thick, cells 3.5–7 μm diam., part above host<br />
tissue heavily pigmented covered by clypeus tissues<br />
(Fig. 25b). Hamathecium of dense, long, cellular pseudoparaphyses,<br />
1.5–3 μm broad, rarely septate, embedded in<br />
mucilage. Asci 150– 200×15– 25(− 33) μ m<br />
(x ¼ 181 20:6mm, n=10), (2-)4-spored, bitunicate, fissitunicate,<br />
broadly cylindrical, with a short, thick, furcate<br />
pedicel which is 20–40 μm long, no apical apparatus<br />
observed (Fig. 25e). Ascospores 37–45×12–17 μm<br />
(x ¼ 43 15mm, n=10), uniseriate and sometimes slightly<br />
overlapping, oblong with broadly rounded ends, dark<br />
brown, verrucose or smooth, 7–9 transverse septa and 1–<br />
3 longitudinal septa in some of the cells, no constriction at<br />
the septa (Fig. 25c and d).<br />
Anamorph: none reported.<br />
Material examined: GERMANY, Valsalpe in der Ramsau,<br />
Bayer, Alpen, on Rhamnus pumila Turra., Jul. 1913, Karl Arnold<br />
(NY2082, syntype as Teichospora megalocarpa Rehm).<br />
Notes<br />
Morphology<br />
Decaisnella was formally established by Fabre (1879),<br />
but was treated as a synonym of Teichospora by<br />
Saccardo (1883). This was followed by several mycologists<br />
over a long time. The main morphological differences<br />
between Decaisnella and Teichospora include the<br />
size and septation of ascospores, shape of ascomata,<br />
structure of peridium and type of pseudoparaphyses (Barr<br />
1986). Thus Barr (1986) revivedDecaisnella and assigned it<br />
to Massariaceae based on the shape of ascomata and large,<br />
distoseptate ascospores. Currently, 15 species are accepted<br />
under Decaisnella (http://www.mycobank.org/MycoTaxo.<br />
aspx). Neither the size of ascomata nor the ascospore<br />
characters have proven sufficient to place taxa at the family<br />
level in <strong>Pleosporales</strong> (Zhang et al. 2009a), and therefore<br />
familial placement of Decaisnella remains uncertain.<br />
Phylogenetic study<br />
Decaisnella formosa resided in the clade of Lophiostomataceae<br />
and in proximity to Lophiostoma macrostomoides<br />
De Not. (Plate 1).<br />
Concluding remarks<br />
The muriform ascospores, saprobic life style and 4-<br />
spored asci point Decaisnella spectabilis to Montagnulaceae,<br />
but this can only be confirmed following a molecular<br />
phylogenetic study.<br />
Delitschia Auersw., Hedwigia 5: 49 (1866). (Delitschiaceae)<br />
Generic description<br />
Habitat terrestrial, saprobic (coprophilous). Ascomata<br />
medium- to large-sized, solitary or scattered, immersed to<br />
erumpent, globose or subglobose, apex with or without<br />
papilla, ostiolate. Peridium thin, composed of compressed<br />
cells. Hamathecium of dense, long pseudoparaphyses,<br />
anastomosing and branching. Asci 8-spored, cylindrical to<br />
cylindro-clavate, with short pedicel. Ascospores uni- to
Fungal Diversity<br />
Fig. 25 Decaisnella spectabilis (NY2082, syntype). a Appearance of<br />
ascomata on the host surface. b Section of a partial peridium<br />
(immersed in the substrate). Note the pseudoparenchymatous out<br />
triseriate, pale to dark brown, ellipsoid, 1-septate, usually<br />
constricted at the septum, smooth, with a full length germ<br />
slit in each cell.<br />
Anamorphs reported for genus: none.<br />
layer. c, d Muriform ascospores. Note the minuitely verrucose<br />
ornamentation. e Ascus with a short pedicel. Scale bars: a=0.5 mm,<br />
b=100 μm, c–e=20 μm<br />
Literature: Auerswald 1866; Barr 2000; Cain 1934;<br />
Dennis 1968; Eriksson 2006; Griffiths 1901; Hyde and<br />
Steinke 1996; Kirschstein 1911; Kruys et al. 2006; Luck-<br />
Allen and Cain 1975; Lumbsch and Huhndorf 2007;
Fungal Diversity<br />
Moreau 1953; Munk 1957; Romero and Samuels 1991;<br />
Schoch et al. 2006; Winter 1887.<br />
Type species<br />
Delitschia didyma Auersw., Hedwigia 5: 49 (1866).<br />
(Fig. 26)<br />
Ascomata 400–800 μm diam., solitary or scattered,<br />
immersed, globose or subglobose,black,papillashort,70–<br />
130 μm broad, central, with a wide opening, coriaceous<br />
(Fig. 26a). Peridium ca. 15μm thick laterally, up to 35 μm<br />
thick at the apex, up to 30 μm at the base, comprising a<br />
single layer of small lightly pigmented thin-walled cells of<br />
textura angularis, cells 4–10 μm diam., cell wall
Fungal Diversity<br />
Fig. 26 Delitschia didyma<br />
(from L, 1950). a Ascomata on<br />
the substrate surface. Note the<br />
ostiolar opening. b Section of<br />
peridium. Note the small cells of<br />
textura angularis. c Released<br />
and unreleased ascospores. Note<br />
the germ slit in each cell. d, e<br />
Asci with ascospores and short<br />
pedicels with rounded ends.<br />
Scale bars: a=0.5 mm,<br />
b =30 μm, c–e=70 μm<br />
Type species<br />
Didymosphaeria futilis (Berk. & Broome) Rehm, Hedwigia<br />
18: 167 (1879). (Fig. 27)<br />
≡ Sphaeria futilis Berk. & Broome, Ann. Mag. nat.<br />
Hist., Ser. 2 9: 326 (1852).<br />
Ascomata 190–230 μm high×240–340 μm diam., scattered,<br />
or in small groups, immersed to slightly erumpent,<br />
subglobose to ovoid, membraneous, near-hyaline, under<br />
clypeus, papillate, periphysate (Fig. 27a and c). Papilla<br />
central, up to 100 μm high, black, with a pore-like ostiole<br />
(Fig. 27a and c). Peridium 30–40 μm wide upper part, 6–<br />
23 μm wide near the base, 1-layered, composed of brown<br />
pseudoparenchymatous cells of textura angularis, cell wall 2–<br />
3 μm thick (Fig. 27b). Hamathecium of dense, long<br />
trabeculate pseudoparaphyses, 0.8–1.5 μm broad, anastomosing<br />
mostly above the asci, embedded in mucilage (Fig. 27d).<br />
Asci 90–110×7.5–10 μm (x ¼ 97 9mm, n=10), 2–4-spored,<br />
rarely 8-spored, bitunicate, fissitunicate, cylindrical, with a<br />
furcate pedicel, 17.5–27.5 μm long, with a large ocular (to<br />
2.5 μm wide×4 μm high) (Fig. 27d, e and f). Ascospores 14–<br />
15.5×(5.5-) 6–7.5 μm (x ¼ 14:8 6:9mm, n=10), uniseriate,<br />
ellipsoid with obtuse ends, brown, 1-septate, distoseptate,<br />
slightly to not constricted, capitate (Fig. 27g).<br />
Anamorph: Dendrophoma sp., Fusicladiella sp. vel aff.<br />
(Sivanesan 1984).
Fungal Diversity<br />
Material examined: UK, England, Norfolk, King’s<br />
Cliffe; on dead stem (in ramis emortuis) Rosa sp., Mar.<br />
1850, M.J. Berkeley (K(M): 147683, holotype).<br />
Notes<br />
Morphology<br />
Didymosphaeria is a widely distributed genus with wide<br />
host range (Aptroot 1995). Didymosphaeria was formally<br />
established by Fuckel (1870) based on six ascomycetous<br />
species, and D. epidermidis (Fries) Fuckel (or D. peltigerae<br />
Fuckel) has been chosen as the lectotype species (see<br />
comments by Aptroot 1995). Hawksworth and David<br />
(1989: 494) proposed to conserve the genus with a lectotype<br />
specimen, Fungi Rhenani 1770. The genus had been<br />
considered as a depository to accommodate all types of<br />
didymosporous pyrenocarpous ascomycetes. Many workers<br />
have tried to redefine the genus and excluded some species.<br />
Saccardo (1882) restricted the genus to brown-spored species,<br />
and about 100 species have been excluded subsequently (Barr<br />
1989a, b, 1990a, 1992a, b, 1993b; Hawksworth1985a, b;<br />
Hawksworth and Boise 1985; Hawksworth and Diederich<br />
Fig. 27 Didymosphaeria futilis<br />
(from K(M): 147683, holotype).<br />
a Two immersed ascomata on<br />
the host surface (one of them is<br />
cut horizontally). b Section of<br />
an ascoma. Note the thin peridium.<br />
c Hand cut portion of<br />
ascoma showing habitat in<br />
wood. d Asci in pseudoparaphyses.<br />
Note the trabeculate pseudonparaphyses<br />
anastomosing<br />
above the asci. e, f Four-spored<br />
asci with long pedicels which<br />
are rounded at their bases. g<br />
Brown, 1-septate ascospores<br />
with spinulose ornamentation.<br />
Scale bars: a=0.3 mm, b, c=<br />
100 μm, d–g=10 μm
Fungal Diversity<br />
1988; Scheinpflug 1958). Over 400 epithets of Didymosphaeria<br />
were included until the monograph of Aptroot (1995).<br />
Aptroot (1995) examined more than 3000 specimens under<br />
the name Didymosphaeria. The type specimen of Didymosphaeria<br />
(Fungi Rhenani 1770) represents the widespread and<br />
common D. futilis (Aptroot 1995). In this study, we did not<br />
get the lectotype specimen, but described the type of D. futilis<br />
(Sphaeria futilis). Using a narrow concept (ignoring differences<br />
of host or country of origin), Aptroot (1995) accepted<br />
only seven species, which were closely related with the<br />
generic type of Didymosphaeria with over 100 synonyms<br />
distributed among them. Many taxa were found to belong to<br />
other groups, i.e. Aaosphaeria, Amphisphaeria, Astrosphaeriella,<br />
Dothidotthia, Flagellosphaeria, Kirschsteiniothelia, Megalotremis,<br />
Montagnula, Munkovalsaria, Mycomicrothelia,<br />
Parapyrenis or Phaeodothis. Didymosphaeria is mainly characterized<br />
by a peridium consisting of flattened or irregular cells<br />
or completely hyphae; a hamathecium consisting of narrow,<br />
trabeculate paraphysoids or paraphyses, richly anastomosing<br />
above the asci; and brown thinly distoseptate ascospores.<br />
Didymosphaeriaceae was maintained as a separated family<br />
within <strong>Pleosporales</strong> by Aptroot (1995) because of the<br />
distoseptate ascospores and trabeculate pseudoparaphyses<br />
mainly anastomosing above the asci. This proposal, however,<br />
has not received much support (Lumbsch and Huhndorf 2007).<br />
Phylogenetic study<br />
There have been few molecular investigations of<br />
Didymosphaeria when compared to the morphological<br />
studies. Didymosphaeria futilis resided in the clade of<br />
Cucurbitariaceae (or Didymosphaeriaceae) (Plate 1). The<br />
correct identification of the Didymosphaeria strain used for<br />
sequencing, however, has not been verified.<br />
Concluding remarks<br />
Didymosphaeria is a well established genus represented<br />
by D. futilis. Of particular significance are the narrow<br />
pseudoparaphyses which anastomose above the asci and<br />
brown 1-septate ascospores with indistinct distosepta. Familial<br />
placement of Didymosphaeria is unclear yet because<br />
of insufficient molecular data.<br />
Dothidotthia Höhn., Ber. Deutsch. Bot. Ges. 36: 312<br />
(1918). (Didymellaceae)<br />
Generic description<br />
Habitat terrestrial, saprobic. Ascomata medium-sized, solitary,<br />
clustered or somewhat gregarious, erumpent, subglobose,<br />
apex somewhat papillate to depressed, coriaceous. Peridium<br />
composed of a few layers of dark brown cells of textura<br />
angularis, and giving rise dark brown, thick-walled hyphae in<br />
the basal region, 2-layered. Hamathecium septate pseudoparaphyses<br />
branched in upper part above asci. Asci 8-spored,<br />
bitunicate, clavate, straight to curved. Ascospores biseriate to<br />
obliquely uniseriate, ellipsoid, pale brown, 1-septate.<br />
Anamorphs reported for genus: Dothiorella and Thyrostroma<br />
(Hyde et al. 2011; Phillips et al. 2008).<br />
Literature: Barr1989b; Phillips et al. 2008.<br />
Type species<br />
Dothidotthia symphoricarpi (Rehm) Höhn., Ber. Deutsch.<br />
Bot. Ges. 36: 312 (1918). (Fig. 28)<br />
≡ Pseudotthia symphoricarpi Rehm, Ann. Mycol. 11:<br />
169 (1913).<br />
Ascomata up to 500 μm high×550 μm diam., gregarious<br />
clustered, rarely solitary, erumpent, subglobose, apex somewhat<br />
papillate to depressed, coriaceous (Fig. 28a). Peridium 20–<br />
80 μm thick, composed of 3–6 layers of dark brown cells of<br />
textura angularis, giving rise dark brown, thick-walled hyphae<br />
in the basal region, 2-layered, outer layer wall thicker and inner<br />
layer wall thinner (Fig. 28b). Hamathecium hyaline, septate<br />
pseudoparaphyses, 2–3 μm wide, branched in upper part above<br />
asci. Asci 70–120×15–22 μm, 8-spored, bitunicate, clavate,<br />
straight to curved (Fig. 28c, d and e). Ascospores (20-)22–23<br />
(−26)×(8-)9–10(−11) μm, biseriate to obliquely uniseriate and<br />
partially overlapping, ellipsoid tapering towards subacutely<br />
rounded ends, pale brown, 1-septate, constricted at the septum,<br />
smooth (Fig. 28f) (description referred to Phillips et al. 2008).<br />
Anamorph: Thyrostroma negundinis (Phillips et al. 2008).<br />
Material examined: USA, North Dakota, on branches<br />
of Symphoricarpos occidentalis Hook. (NY, holotype);<br />
Colorado, San Juan Co, c. 0.5 mile up Engineer Mountain<br />
Trail from turnoff at mile 52.5, Hwy 550, dead twigs of<br />
Symphoricarpos rotundifolius A. Gray, 24 Jun. 2004, A.W.<br />
Ramaley 0410 (BPI 871823, epitype).<br />
Notes<br />
Morphology<br />
Dothidotthia was formally established to accommodate<br />
Pseudotthia symphoricarpi (Montagnellaceae, Dothideales)<br />
(von Höhnel 1918a). Many mycologists considered Dothidotthia<br />
closely related to a genus of Venturiaceae such as<br />
Dibotryon by Petrak (1927), or Gibbera by von Arx and<br />
Müller (1954) and Müller and von Arx (1962). Dothidotthia<br />
had been treated as a synonym of Gibbera (von Arx 1954;<br />
Müller and von Arx 1962), which was followed by<br />
Shoemaker (1963) and Eriksson and Hawksworth (1987).<br />
Based on the coelomycetous anamorphic stage and peridium<br />
structure, shape of asci, as well as morphology of<br />
pseudoparaphyses, Barr (1987b, 1989b) retrieved Dothidotthia,<br />
and considered it closely related to Botryosphaeria<br />
(Botryosphaeriaceae). Currently, 11 species are included<br />
within Dothidotthia (http://www.mycobank.org, 01–2011).
Fungal Diversity<br />
Fig. 28 Dothidotthia symphoricarpi<br />
(from NY, holotype). a<br />
Clustered ascomata on the host<br />
stubstrate. b Longitudinal section<br />
through an ascoma. c, d<br />
Asci with pale brown, 1-septate<br />
ascospores. e Immature asci. f<br />
Pale brown, 1-septate ascospores<br />
within asci. g Conidia of<br />
Thyrostroma anamorph in association<br />
with ascomata. Scale<br />
bars: a=0.5 mm, b=100 μm,<br />
c–g=10 μm. (figure with permission<br />
from Phillips et al.<br />
2008)<br />
Phylogenetic study<br />
Based on a multi-gene phylogenetic analysis, Dothidotthia<br />
formed a separate familial clade (Phillips et al. 2008).<br />
Thus Dothidotthiaceae was introduced to accommodate it<br />
(Phillips et al. 2008).<br />
Concluding remarks<br />
By comparing the morphological characters and phylogenetic<br />
dendrograms by Phillips et al. (2008) and de Gruyter et al.<br />
(2009), Dothidotthia seems closely related to Didymellaceae,<br />
but Dothidotthiaceae should still be treated as a separate family.
Fungal Diversity<br />
Dubitatio Speg., Anal. Soc. cient. argent. 12: 212<br />
(1881). (Arthopyreniaceae (or Massariaceae))<br />
Generic description<br />
Habitat terrestrial, saprobic. Ascomata medium-sized, solitary,<br />
densely scattered, or in small groups of 2–4,<br />
immersed, covered with white crystaline rim, papillate,<br />
ostiolate. Hamathecium of dense pseudoparaphyses, long,<br />
2–3 μm broad, branching and anastomosing. Asci cylindrical,<br />
pedicellate, with furcate pedicel. Ascospores 1-septate,<br />
asymmetrical, reddish to dark brown.<br />
Anamorphs reported for genus: Aplosporella-like (Rossman<br />
et al. 1999).<br />
Literature: Barr 1979b, 1987b; Müller and von Arx 1962;<br />
Rossman et al. 1999; Spegazzini 1881.<br />
Type species<br />
Dubitatio dubitationum Speg., Anal. Soc. cient. argent. 12:<br />
212 (1881). (Fig. 29)<br />
Ascomata 350–530 μm high×550–700 μm diam.,<br />
solitary, densely scattered, or in small groups of 2–4,<br />
immersed, with a protruding papilla, 110–160 μm high,<br />
160–250 μm diam., globose or subglobose, black,<br />
covered with white crystalline material which becomes<br />
hyaline and gel-like in water, ostiolate (Fig. 29a and b).<br />
Peridium 18–25 μm thick laterally (excluding the rim), up<br />
to 35 μm thick at the apex, thinner at the base, 1-layered,<br />
composed of small pale brown thin-walled cells of textura<br />
prismatica, cells 5–12×3–5 μm diam., cell wall up to<br />
1 μm thick, apex cells smaller and walls thicker<br />
(Fig. 29b). Hamathecium of dense, long pseudoparaphyses,<br />
2–3 μm broad, branching and anastomosing between<br />
and above the asci. Asci 150–190(−230)×12.5–15 μm<br />
(x ¼ 172:5 13:4mm, n =10), (6-)8-spored, rarely 4-<br />
spored, bitunicate, fissitunicate, cylindrical, with a furcate<br />
pedicel which is up to 40 μm long, ocular chamber not<br />
observed (Fig. 29c, d and e). Ascospores 19–22.5×10–<br />
12 μm (x ¼ 20:2 11:4mm, n=10), uniseriate to obliquely<br />
uniseriate and partially overlapping, broadly ellipsoid<br />
with broadly to narrowly rounded ends, reddish brown,<br />
1-septate, constricted at septum, asymmetric with a<br />
larger upper cell, thick-walled, possibly distoseptate<br />
(Fig. 29f, g and h).<br />
Anamorph: Aplosporella-like (for detailed description<br />
see Rossman et al. 1999).<br />
Conidiomata globose, ca. 300 μm diam. Conidia<br />
holoblastic, broadly fusoid, 13–15×7–10 μm, dark brown,<br />
finely spinulose (Rossman et al. 1999).<br />
Material examined: ARGENTINA, Buenos Aires,<br />
Tuyu, on Celtis tala Gill., Jan. 1881, leg. det. C. Spegazzini<br />
(NY, isotype; LPS, holotype).<br />
Notes<br />
Morphology<br />
When established Dubitatio, Spegazzini (1881) considered<br />
it as intermediate between Sphaeriaceae and Nectriaceae<br />
as has been mentioned by Rossman et al. (1999). Müller<br />
and von Arx (1962) treated Dubitatio as a synonym of<br />
Passerinula, while the differences of ascomata and ascospores<br />
could easily distinguish these two genera (Rossman et al.<br />
1999). After checking the type specimen, Dubitatio was assigned<br />
to Dothideomycetes, and considered closely related to Dothivalsaria<br />
in the Massariaceae (Barr 1979b, 1987b). Dubitatio<br />
chondrospora was assigned to Pseudomassaria (as P. chondrospora<br />
(Ces.) Jacz.) (Barr 1964; Müller and von Arx 1962).<br />
Phylogenetic study<br />
None.<br />
Concluding remarks<br />
The black ascomata with white crystalline covering and<br />
central white ostiolar region as well as the asymmetrical<br />
reddish brown ascospores are striking characters of Dubitatio<br />
dubitationum. The genus cannot be assigned to any<br />
family with certainty based on morphological characters<br />
and fresh collections are needed for sequencing.<br />
Entodesmium Reiss, Hedwigia 1: 28 (1854).<br />
(Phaeosphaeriaceae)<br />
Generic description<br />
Habitat terrestrial, saprobic (or parasitic?). Ascomata<br />
scattered or in small groups, immersed, papillate, ostiolate,<br />
periphysate. Peridium thin, comprising one cell type of<br />
pigmented pseudoparenchymatous cells. Hamathecium of<br />
dense, long pseudoparaphyses, septate, embedded in mucilage.<br />
Asci 8-spored, bitunicate, fissitunicate, cylindrical, with<br />
furcate pedicel. Ascospores ellipsoid to filliform, multiseptate,<br />
deeply constricted at the primary septum (usually<br />
near apex), breaking into partspores.<br />
Anamorphs reported for genus: none.<br />
Literature: von Arx and Müller 1975; Barr1992b; Eriksson<br />
1967a; b; Holm1957; Liewetal.2000; Shoemaker 1984a, b.<br />
Type species<br />
Entodesmium rude Reiss, Hedwigia 1: 28 (1854). (Fig. 30)<br />
Ascomata 160–250 μm high×150–300 μm diam., in groups,<br />
immersed with long and protruding cylindrical papilla,<br />
globose to subglobose, black, coriaceous (Fig. 30a). Papilla<br />
100–220 μm long, 70–120 μm broad, cylindrical, with<br />
periphysate ostiole. Peridium 25–33 μm wide, comprising<br />
pseudoparenchymatous cells, cells up to 10×7.5 μm diam., cell<br />
wall up to 2 μm thick, beak cells smaller and wall thicker
Fungal Diversity<br />
Fig. 29 Dubitatio dubitationum (from NY, isotype; LPS, holotype).<br />
a Appearance of ascomata scattered on the host surface. Note the<br />
exposed white covering around the ostioles. b, c Section of an ascoma.<br />
(Fig. 30b and c). Hamathecium of dense, long pseudoparaphyses,<br />
septate, 2–3 μm wide, embedded in mucilage. Asci<br />
100–175×8–13 μm (x ¼ 147:5 11:3mm, n =10), 8-<br />
spored, bitunicate, fissitunicate, cylindrical, with a furcate<br />
pedicel which is 18–50 μm long, and with a low ocular<br />
chamber (ca. 1μm wide×1 μm high) (Fig. 30e,f, g and h).<br />
Note the white covering (see arrow). d–f Cylindrical asci with short<br />
furcate pedicels. g, h Asymmetrical, 1-septate reddish-brown ascospores.<br />
Scale bars: a=1 mm, b=100 μm, c-e=50 μm, f–h=20 μm<br />
Ascospores 108–138×3–3.5 μm (x ¼ 123 3:2mm, n=10),<br />
filliform, brown, multi-septate, breaking into 22–28 partspores,<br />
5–7×3–3.5 μm diam. (Fig. 30d).<br />
Anamorph: none reported.<br />
Material examined: GERMANY, Königstein, on stems of<br />
Coronilla varia L., 20 May 1895, W. Krieger (H, Krieger 1070).
Fungal Diversity<br />
Fig. 30 Entodesmium rude (from H, Krieger 1070). a Ascomata in<br />
groups on the host surface. Note the erumpent papilla which is<br />
cylindrical and has an inconspicuous ostiole. b Section of part of an<br />
ascoma. Note the arrangement of asci and pseudoparaphyses. c Section<br />
of the peridium comprising cells of textura angularis. d Part-spores<br />
inside the ascus. e Relatively immature ascus with filliform ascospores<br />
and low ocular chamber. f–h Mature and immature asci with pedicels.<br />
Scale bars: a=0.5 mm, b, c=50 μm, d–h=10 μm<br />
Notes<br />
Morphology<br />
Entodesmium is characterized by having immersed<br />
ascomata dark cylindrical, periphysate papillae, numerous<br />
clavate to cylindrical asci surrounded by narrowly<br />
cellular pseudoparaphyses, and ellipsoidal to filliform<br />
multi-septate ascospores (Barr 1992b; Shoemaker<br />
1984b). Currently, five species, viz. Entodesmium eliassonii<br />
L. Holm, E. lapponicum (L. Holm) L. Holm, E.<br />
mayorii (E.Müll.)L.Holm,E. niessleanum (Rabenh. ex<br />
Niessl) L. Holm and E. rude are accepted in this genus<br />
(Holm 1957; Shoemaker 1984b). Von Arx and Müller<br />
(1975) assignedEntodesmium to the Pleosporaceae sensu<br />
lato, and Shoemaker (1976) assigned E. rude (as<br />
Ophiobolus rudis) to Ophiobolus sensu lato based on<br />
the fragmenting filliform ascospores. According to the<br />
short, blackish beak and periphysate ostiole, Barr (1992b)
Fungal Diversity<br />
assigned Entodesmium to Lophiostomataceae. Thehosts<br />
of Entodesmium are restricted to stems of legumes (Barr<br />
1992b; Shoemaker 1984b).<br />
Phylogenetic study<br />
Limited phylogenetic studies indicate that Entodesmium<br />
rude may have affinities to Phaeosphaeriaceae (Liew et al.<br />
2000; Plate 1).<br />
Concluding remarks<br />
Species of Entodesmium share several morphological<br />
characters, such as immersed, papillate ascomata, periphysate<br />
ostioles, pale yellow to light yellowish brown, multiseptate<br />
(≥ 3), narrowly fusoid to filliform ascospores, and<br />
are specific to legumes. All of the above similarities<br />
indicate a close relationship among members of Entodesmium.<br />
We do not agree with Barr (1992b) who assigned<br />
Entodesmium to Lophiostomataceae because the ascomata<br />
are immersed, the papilla are not laterally compressed<br />
and the peridium comprises a single type of cells of<br />
textura angularis. These characters plus multi-septate,<br />
lightly pigmented ascospores, which break up into<br />
partspores and host specificity to legumes support inclusion<br />
in Phaeosphaeriaceae. Entodesmium multiseptatum<br />
(G. Winter) L. Holm and E. niessleanum were originally<br />
described as Leptosphaeria species (Shoemaker 1984b)<br />
indicating their similarity to Phaeosphaeria with which<br />
Leptosphaeria is commonly confused (Shoemaker 1984a;<br />
Shoemaker and Babcock 1989b). Phylogenetic study has<br />
also shown that Entodesmium rude is related to members<br />
of Phaeosphaeriaceae (Liew et al. 2000). Thus we assign<br />
Entodesmium to Phaeosphaeriaceae as a separate genus<br />
until further phylogenetic analysis is carried out on<br />
verified specimens.<br />
Eudarluca Speg., Revta Mus. La Plata 15: 22 (1908).<br />
(?Phaeosphaeriaceae)<br />
Generic description<br />
Habitat terrestrial, parasitic. Ascomata small, solitary,<br />
scattered, immersed to erumpent, subglobose, ostiolate,<br />
papillate. Peridium thin, composed of a few layers cells of<br />
textura prismatica. Hamathecium of dense, cellular pseudoparaphyses,<br />
septate. Asci 8-spored, bitunicate, fissitunicate,<br />
cylindrical to fusoid, with a furcate pedicel.<br />
Ascospores broadly fusoid to fusoid, hyaline to pale yellow,<br />
rarely 1- or 3- septate, mostly 2-septate, constricted at the<br />
primary septum.<br />
Anamorphs reported for genus: Sphaerellopsis (Sivanesan<br />
1984).<br />
Literature: Bayon et al. 2006; Eriksson 1966; Katumoto<br />
1986; Ramakrishnan 1951; Spegazzini 1908.<br />
Type species<br />
Eudarluca australis Speg., Revta Mus. La Plata 15: 22<br />
(1908). (Fig. 31)<br />
Ascomata 160–190 μm high×180–290 μm diam., solitary,<br />
scattered, or in small groups, semi-immersed to erumpent,<br />
subglobose to broadly ellipsoid, wall black, ostiolate, apex<br />
with a short papilla, 40–70 μm broad (Fig. 31a and b).<br />
Peridium
Fungal Diversity<br />
Fig. 31 Eudarluca australis (from LPS 5.415, type). a Ascomata on<br />
the host surface. b Section of an ascoma. c Section of a partial<br />
peridium. Note the thin peridium with cells of textura angularis. d–g<br />
Asci with short pedicels. h Ascospores. Note the 2-septate hyaline<br />
ascospore. Scale bars: a, b=100 μm, c=50 μm, d–h=10 μm<br />
Falciformispora K.D. Hyde, Mycol. Res. 96: 26 (1992).<br />
(Trematosphaeriaceae)<br />
Generic description<br />
Habitat freshwater, saprobic. Ascomata small, scattered to<br />
gregarious, erumpent to nearly superficial, depressed<br />
globose to ovoid, black, ostiolate, epapillate, coriaceous.<br />
Peridium thin, comprising two cells types, outer layer<br />
composed of thick-walled cells of textura angularis, inner<br />
layer composed of hyaline compressed cells. Hamathecium<br />
long and cellular pseudoparaphyses, septate, embedded in<br />
mucilage. Asci 8-spored, bitunicate, fissitunicate, broadly<br />
clavate to fusoid, with a short, thick pedicel. Ascospores
Fungal Diversity<br />
fusoid to somewhat clavate, hyaline, usually slightly<br />
curved, multi-septate.<br />
Anamorphs reported for genus: none.<br />
Literature: Hyde 1992b; Raja and Shearer 2008.<br />
Type species<br />
Falciformispora lignatilis K.D. Hyde, Mycol. Res. 96: 27<br />
(1992). (Fig. 32)<br />
Ascomata 180–270 μm high×250–340 μm diam., scattered<br />
to gregarious, erumpent and eventually superficial, depressed<br />
globose to ovoid, black, ostiolate, epapillate, coriaceous<br />
(Fig. 32a). Peridium up to 35 μm wide, comprising two cell<br />
types, outer layer composed of thick-walled cells of textura<br />
angularis, upto8μm diam., cell wall up to 5 μm thick, inner<br />
layer composed of hyaline compressed cells, cells 12×3 μm<br />
diam., cell wall 1–1.5 μm thick (Fig. 32a). Hamathecium long<br />
and cellular pseudoparaphyses, 2–3 μm broad, septate,<br />
embedded in mucilage. Asci 115–130×23–31 μm, 8-spored,<br />
bitunicate, fissitunicate, broadly clavate to fusoid, with a short,<br />
thick pedicel, 8–15 μm long,withanocularchamber(to5μm<br />
wide×3 μm high) (Fig. 32b and c). Ascospores 42–50×8–<br />
10 μm, 2–3 seriate, fusoid to somewhat clavate, hyaline,<br />
usually slightly curved, 6–8-septate, mostly 7-septate, slightly<br />
constricted at all septa, smooth-walled, surrounded by a thin<br />
mucilaginous sheath which is longer at the base (up to 20–<br />
30 μm) (Fig.32d, e and f).<br />
Anamorph: none reported.<br />
Material examined: MEXICO, Nova Hispania, mangrove<br />
near Boca de Pascuales, saprobic on immersed<br />
intertidal mangrove wood, Mar. 1988, K.D. Hyde (BRIP<br />
16972, holotype).<br />
Notes<br />
Morphology<br />
Falciformispora was formally established by Hyde (1992b)<br />
as a monotypic genus and was assigned to Pleosporaceae by<br />
comparing with Setosphaeria, but Setosphaeria has the<br />
anamorphic stage of Exserohilum and is exclusively parasitic<br />
on Gramineae unlike Falciformispora. The setae on the<br />
ascomata of Setosphaeria could also serve as a distinguishing<br />
character from Falciformispora. Raja and Shearer (2008)<br />
also collected this species from freshwater in Florida. They<br />
considered that the species was more closely related to<br />
Chaetomastia than Setosphaeria, but that Falciformispora<br />
differed in having hyaline ascospores.<br />
Phylogenetic study<br />
Phylogenetic analyses in Schoch et al. (2009) and Suetrong et<br />
al. (2009) placed Falciformispora lignatilis in Trematosphaeriaceae<br />
in proximity to another marine species associated<br />
with mangroves, Halomassarina thalassiae.<br />
Concluding remarks<br />
Phylogenetic work confirmed that the saprobic habitat of<br />
Falciformispora is inconsistent with most other members of<br />
Pleosporaceae. The hyaline multi-septate ascospores with a<br />
mucilaginous sheath indicate affinities to Lophiostomataceae<br />
but this is not supported in DNA sequence comparisons.<br />
Carinispora is also similar and may be related.<br />
Hadrospora Boise, Mem. N. Y. bot. Gdn 49: 310 (1989).<br />
(?Phaeosphaeriaceae)<br />
Generic description<br />
Habitat terrestrial (or freshwater?), saprobic. Ascomata<br />
small- to medium-sized, solitary, scattered, or in groups,<br />
immersed to nearly superficial, globose to subglobose,<br />
papillate. Peridium thin, comprising pseudoparenchymatous<br />
cells. Hamathecium dense, narrowly cellular, embedded<br />
in mucilage. Asci bitunicate, fissitunicate, oblong to<br />
ovoid, with a short pedicel. Ascospores ellipsoid to broadly<br />
fusoid with narrow ends, reddish brown, multi-septate,<br />
constricted at the primary septum.<br />
Anamorphs reported for genus: Zalerion (Tanaka and<br />
Harada 2003a).<br />
Literature: Boise1984, 1989; Fisher and Webster 1992;Shearer<br />
and Crane 1971; Tanaka and Harada 2003a; Webster1993.<br />
Type species<br />
Hadrospora fallax (Mouton) Boise, Mem. N. Y. bot. Gdn<br />
49: 310 (1989). (Fig. 33)<br />
≡ Trematosphaeria fallax Mouton, Bull. Soc. R. Bot.<br />
Belg. 25: 155, (1886).<br />
Ascomata 130–240 μm high×200–330 μm diam., solitary,<br />
scattered or in groups, initially immersed, becoming erumpent to<br />
nearly superficial, with basal wall remaining immersed in host<br />
tissue, not easily removed from the substrate, globose or<br />
subglobose, roughened, papillate, coriaceous (Fig. 33a). Peridium<br />
30–45 μm wide, comprising cells of pseudoparenchymatous,upto12.5×9μm<br />
diam. (Fig. 33b and c). Hamathecium of<br />
dense, narrowly cellular pseudoparaphyses, 1–2 μm broad,<br />
embedded in mucilage. Asci 150–200×40–60 μm<br />
(x ¼ 171:5 48mm, n=10), 8-spored, bitunicate, fissitunicate,<br />
oblong to ovoid, with a short pedicel, 10–24 μm long,<br />
with a ocular chamber (to 5 μm wide×6 μm high) (Fig. 33d).<br />
Ascospores 55–80×16–22 μm (x ¼ 67:1 18:1mm, n=10),<br />
biseriate to 4-seriate, ellipsoid to broadly fusoid with narrow<br />
ends, reddish brown with paler end cells, 8-septate,<br />
constricted at the primary septum, smooth-walled (Fig. 33e,<br />
f, g, h and i).<br />
Anamorph: Zalerion sp. (Tanaka and Harada 2003a).<br />
Material examined: BELGIUM, Beaufays, on cut off,<br />
still hard wood. Oct. 1922, V. Mouton (BR, Capsa: K 7534,
Fungal Diversity<br />
Fig. 32 Falciformispora lignatilis<br />
(from BRIP 16972, holotype).<br />
a Section of a superficial<br />
ascoma. The peridium comprises<br />
two layers. b, c Squash<br />
mounts showing asci with wide<br />
pseudoparaphyses. The asci are<br />
cylindro-clavate with very short<br />
pedicels. d–f Hyaline multiseptate<br />
ascospores. Note the elongated<br />
appendage at the base<br />
(arrow head). Scale bars:<br />
a, b=100 μm, c=50 μm,<br />
d–f=10 μm<br />
holotype). (Note: The specimen is not in good condition,<br />
only a few ascomata left).<br />
Notes<br />
Morphology<br />
Boise (1989) formallyestablishedHadrospora to accommodate<br />
Trematosphaeria fallax and T. clarkia (Sivan.) Boise,<br />
and Hadrospora fallax (syn. T. fallax) was selected as the<br />
generic type. Hadrospora is a widely distributed species that<br />
has been reported from Belgium, China, Italy, Japan,<br />
Switzerland and the United States (Boise 1989; Fisher and<br />
Webster 1992; Shearer and Crane 1971; Tanaka and Harada<br />
2003a; Webster 1993). Hadrospora was temporarily<br />
assigned to Phaeosphaeriaceae based on its “small, thinwalled<br />
ascomata and narrow cellular pseudoparaphyses”<br />
(Boise 1989), which is distinguished from other genera of<br />
Phaeosphaeriaceae by its “large, stout ascospores that form<br />
within oblong-ovoid asci” (Boise 1989). Currently, Hadrospora<br />
includes two species, i.e. H. fallax and H. clarkii<br />
(Sivan.) Boise differentiated by ascospore size.<br />
Phylogenetic study<br />
None.<br />
Concluding remarks<br />
Hadrospora seems not closely related to Phaeosphaeriaceae.<br />
Halotthia Kohlm., Nova Hedwigia 6: 9 (1963). (?Zopfiaceae)<br />
Generic description<br />
Habitat marine, saprobic. Ascomata large, solitary, gregarious<br />
or confluent, broadly conical to subglobose,<br />
flattened at the base, carbonaceous, immersed to erumpent,<br />
ostiolate, epapillate. Peridium plectenchymatous.<br />
Hamathecium of dense, long, cellular pseudoparaphyses,<br />
septate, branching. Asci 8-spored, bitunicate, cylindrical,<br />
with a short pedicel. Ascospores uniseriate, ellipsoidal,<br />
subcylindrical or obtuse-fusoid, dark brown, 1-septate,<br />
constricted at the septum.<br />
Anamorphs reported for genus: none.<br />
Literature: Kohlmeyer 1963; Suetrong et al. 2009.
Fungal Diversity<br />
Fig. 33 Hadrospora fallax<br />
(from BR, Capsa: K 7534,<br />
holotype). a Ascomata forming<br />
a cluster on the host surface. b<br />
Section of an ascoma. Note the<br />
peridium structure. c Section of<br />
a partial peridium. Note the<br />
pseudoparenchymatous cells. d<br />
Asci in pseudoparaphyses. e–i<br />
Reddish brown multiseptate<br />
ascospores. Scale bars: a=<br />
0.5 mm, b=100 μm, c, d=<br />
50 μm, e–i=20 μm<br />
Type species<br />
Halotthia posidoniae (Durieu & Mont.) Kohlm., Nova<br />
Hedwigia 6: 9 (1963). (Fig. 34)<br />
≡ Sphaeria posidoniae Durieu & Mont. Exploration<br />
scientifique de l’Algérie, pp. 502–503, Taf. 25, Abb. 8a-i, 1849.<br />
Ascomata 0.8–1.1 mm high×1.5–2.1 mm diam., solitary,<br />
gregarious or confluent, broadly conical to subglobose,<br />
flattened at the base, carbonaceous, immersed to<br />
erumpent, ostiolate, epapillate (Fig. 34a). Peridium 165–<br />
275 μm thick at sides, thicker near the apex, plectenchymatous.<br />
Hamathecium of dense, long cellular pseudo-
Fungal Diversity<br />
paraphyses, 1.5–2 μm broad, septate, branching. Asci<br />
275–290×25–35 μm, 8-spored, bitunicate, cylindrical,<br />
with a short pedicel (Fig. 34b, c and d). Ascospores 37–<br />
60.5×16.5–26 μm, uniseriate, ellipsoidal, subcylindrical<br />
or obtuse-fusoid, dark brown, 1-septate, constricted at the<br />
septum (Fig. 34e, f, g and h) (adapted from Kohlmeyer<br />
and Kohlmeyer 1979).<br />
Anamorph: none reported.<br />
Material examined: ITALY, in rhizomes of Posidonia<br />
oceanica (Posidoniaceae), 1861, Caldesi (S, isotype of<br />
Sphaeria posidoniae)<br />
Notes<br />
Morphology<br />
Halotthia was introduced to accommodate the marine<br />
fungus, H. posidoniae (as Sphaeria posidoniae), which is<br />
characterized by immersed to erumpent, large, carbonaceous<br />
ascomata, thick peridium, bitunicate, 8-spored,<br />
cylindrical asci, ellipsoidal, 1-septate, and dark brown<br />
ascospores (Kohlmeyer 1963). Morphologically, Halotthia<br />
is most comparable with Bicrouania maritima, but the<br />
conical ascomata with flattened base of H. posidoniae can<br />
be readily distinguished from B. maritima.<br />
Phylogenetic study<br />
Phylogenetically, Halotthia posidoniae, Pontoporeia<br />
biturbinata and Mauritiana rhizophorae form a robust<br />
clade, which may represent a potential family (Suetrong et<br />
al. 2009).<br />
Concluding remarks<br />
Currently the familial status of Halotthia is unresolved<br />
(Suetrong et al. 2009).<br />
Helicascus Kohlm., Can. J. Bot. 47: 1471 (1969).<br />
(Morosphaeriaceae)<br />
Generic description<br />
Habitat marine, saprobic. Ascostromata lenticular, immersed,<br />
black, carbonaceous, enclosing several loculi,<br />
pseudoclypeus composed of host cells enclosed in black<br />
stromatic fungus material. Ascomata depressed ampulliform,<br />
horizontally arranged under a black pseudoclypeus,<br />
ostiolate, torsellioid ostioles, papillate. Peridium absent,<br />
partitions between loculi formed of brown, isodiametric or<br />
elongated cells of the stroma. Hamathecium of dense, long<br />
pseudoparaphyses. Asci 8-spored, bitunicate, subcylindrical<br />
to oblong clavate, with a short pedicel and conspicuous<br />
apical ring. Ascospores uniseriate, obovoid, brown, 1-<br />
septate, at each end with a germ pore, surrounded with<br />
dissolving sheath.<br />
Anamorphs reported for genus: none.<br />
Literature: Kohlmeyer1969; Kohlmeyer and Kohlmeyer<br />
1979; Suetrongetal.2009.<br />
Type species<br />
Helicascus kanaloanus Kohlm., Can. J. Bot. 47: 1471<br />
(1969). (Fig. 35)<br />
Ascostromata 0.6–0.78 mm high×1.25–2.75 mm diam.,<br />
lenticular, immersed, black, carbonaceous, enclosing 3–4(−5)<br />
loculi, pseudoclypeus composed of host cells enclosed in<br />
black stromatic fungus material (Fig. 35a). Ascomata 235–<br />
370 μm high×440–800 μm diam., depressed ampulliform,<br />
horizontally arranged under a black pseudoclypeus, ostioles<br />
70–170 μm diam., torsellioid ostiole (Adams et al. 2005),<br />
papilla slightly rising over the surface of the pseudoclypeus,<br />
subconical,canal filled with thick, bright orange to yellowish<br />
periphyses, 270–435 μm high, 255–300 μm diam. Peridium<br />
absent, partitions between loculi formed of brown, isodiametric<br />
or elongated cells of the stroma. Hamathecium of<br />
dense, very long pseudoparaphyses. Asci 250–335×25–<br />
30 μm, 8-spored, subcylindrical, finally oblong-clavate<br />
(400–480 μm long), with a short pedicel, bitunicate,<br />
thick-walled, physoclastic, apically multi-layered and<br />
annulate, ectoascus forming a third, thin permeable outer<br />
layer around the base, endoascus swelling in water and<br />
becoming coiled at maturity, finally stretching and<br />
pushing the ascus into the ostiolar canal (Fig. 35b).<br />
Ascospores 36.5–48.5×18–22.5 μm, uniseriate, obovoid,<br />
brown, 1-septate, at each end with a germ pore,<br />
surrounded with dissolving sheath, 2.7–5.4 μm thick,<br />
with funnel-shaped, apical indentations (Fig. 35c and d)<br />
(adapted from Kohlmeyer and Kohlmeyer 1979).<br />
Anamorph: none reported.<br />
Material examined: USA, Hawaii, Oahu, Kaneohe Bay,<br />
Heeia Swamp, on Rhizophora mangle, 4 Jun. 1968 (Herb. J.<br />
Kohlmeyer No. 2566, holotype; No. 2565, 2567, paratype).<br />
Notes<br />
Morphology<br />
Helicascus is another marine genus, which is characterized<br />
by its thin additional sheath around the base of the asci,<br />
the coiling and stretching mechanism of the basal part of the<br />
endoascus and its conspicuous apical apparatus which is not<br />
that common in bitunicate asci (Kohlmeyer 1969). The<br />
immersed stroma comprising several loculi sharing one<br />
common ostiole is another striking character of Helicascus.<br />
Phylogenetic study<br />
Multigene phylogenetic analysis indicated that both<br />
Helicascus kanaloanus and H. nypae K.D. Hyde nested<br />
within Morosphaeriaceae (Suetrong et al. 2009).
Fungal Diversity<br />
Fig. 34 Halotthia posidoniae (from S, isotype of Sphaeria posidoniae). a Ascomata gregarious on the host surface. b–d Mature or immature<br />
cylindrical asci. e–h Ellipsoidal, dark-brown, 1-septate ascospores. Scale bars: a=1 mm, b–d=50 μm, e–h=5 μm<br />
Concluding remarks<br />
Helicascus is a well defined marine genus.<br />
Herpotrichia Fuckel, Fungi rhenani exsic.: no. 2171<br />
(1868). (Melanommataceae)<br />
Generic description<br />
Habitat terrestrial, parasitic, hyperparasitic or saprobic.<br />
Ascomata medium-sized, immersed, erumpent to nearly<br />
superficial, scattered to gregarious, globose to subglobose
Fungal Diversity<br />
Fig. 35 Helicascus kanaloanus<br />
(from Herb. J. Kohlmeyer No.<br />
2566, holotype). a Section of<br />
ascostroma immersed in the host<br />
tissue. Note the torsellioid ostiole.<br />
b One-septate, brown,<br />
asymmetrical ascospores within<br />
the asci. c, d Released thickwalled<br />
ascospores. Note the<br />
germ pore at the lower end of<br />
the ascospores. Scale bars: a=<br />
0.5 mm, b–d=20 μm<br />
with a broad pore. Peridium composed of pseudoparenchymatous<br />
cells. Hamathecium of dense, long pseudoparaphyses,<br />
embedded in mucilage, septate, branching. Asci<br />
cylindrical to cylindro-clavate, with a furcated pedicel.<br />
Ascospores fusoid, ellipsoid or oblong with broadly to<br />
narrowly round ends, 1-septate, constricted at the septum,<br />
uni- to biseriate.<br />
Anamorphs reported for genus: Pyrenochaeta or Pyrenochaeta-like<br />
(Sivanesan 1984).<br />
Literature: von Arx and Müller 1975; Barr1984; Cannon<br />
1982; Freyer and van der Aa 1975; Mugambi and<br />
Huhndorf 2009b; Samuels 1973; Samuels and Müller<br />
1978; Sivanesan 1971, 1984.<br />
Type species<br />
Herpotrichia rubi Fuckel, Fungi rhenani exsic 2171.<br />
(1868). (Fig. 36)<br />
Ascomata 220–430 μm high×240–390(-530) μm<br />
diam., scattered to gregarious, immersed to erumpent,<br />
rarely superficial, globose to subglobose, wall black,<br />
coriaceous, apex with a small sometimes inconspicuous<br />
papilla, usually with a pore, lacking periphyses (Fig. 36a<br />
and b). Peridium 32–45 μm wideatthesides,upto60μm<br />
wide at the apex, basal wall thinner, all walls comprising<br />
cells of textura angularis, cells 2.5–4 μm diam., cell wall<br />
2–4(−7) μm thick, exterior cells more thick-walled and<br />
pigmented, inner cells thin-walled and less pigmented,<br />
comprising thin-walled cells up to 9 μm diam., apex cells<br />
smaller and walls thicker (Fig. 36b and c). Hamathecium<br />
of dense, long pseudoparaphyses, 2–3 μm broad, embedded<br />
in mucilage, septate, branching (Fig. 36e). Asci 105–<br />
150×12.5–15 μm (x ¼ 137:5 13:8mm, n=10), 8-spored,<br />
bitunicate, fissitunicate, cylindrical to cylindro-clavate,<br />
with a furcate pedicel that is 20–42.5 μm long, and<br />
ocular chamber up to 2.5 μm wide×2.5 μm high<br />
(Fig. 36d and f). Ascospores 17.5–25×(5.5-)6.3–9 μm<br />
(x ¼ 20:5 7:3mm, n=10), biseriate to partially overlapping<br />
uniseriate near the base, fusoid with narrowly<br />
rounded ends, hyaline when immature and becoming pale<br />
brown, 1-septate, deeply constricted at the septum, the<br />
upper cell often broader than the lower one, verruculose<br />
(Fig. 36g and h).<br />
Anamorph: Pyrenochaeta rhenana Sacc. (Sivanesan<br />
1984).<br />
Material examined: AUSTRIA, on Rubus idaeus L.,<br />
very rarely in the spring, in the Oestreicher meadow forest<br />
(G, F. rh. 2171, type).
Fungal Diversity<br />
Notes<br />
Morphology<br />
Herpotrichia was established by Fuckel (1868) comprising<br />
two species H. rhenana Fuckel and H. rubi Fuckel, but<br />
no generic type was assigned. Bose (1961) designatedH.<br />
rhenana as the lectotype species with H. rubi as a synonym.<br />
This proposal was followed by Müller and von Arx (1962)<br />
and Sivanesan (1971). Herpotrichia rubi was later assigned<br />
as the generic type (Holm 1979) as it was found to be validly<br />
published 2 years earlier than H. rhenana, thus having<br />
priority (Cannon 1982). However, Cannon (1982) reported<br />
that Sphaeria herpotrichoides Fuckel (1864, cited as a<br />
synonym of H. rhenana) was the earliest name. Thus he<br />
made a new combination as H. herpotrichoides (Fuckel) P.F.<br />
Cannon and cited H. rubi as the synonym. Herpotrichia rubi<br />
is maintained as the type of the genus (Holm 1979; Cannon<br />
1982), but the current name is H. herpotrichoides.<br />
Herpotrichia is a morphologically well studied genus<br />
(Barr 1984; Bose 1961; Müller and von Arx 1962;<br />
Pirozynski 1972; Samuels and Müller 1978; Sivanesan<br />
1971, 1984), and Herpotrichia sensu lato is characterized<br />
by having subglobose, pyriform to obpyriform ascomata<br />
and a peridium of textura angularis or comprising thickwalled<br />
polygonal cells with thin-walled hyaline cells<br />
towards the centre. Asci are clavate to cylindrical, 4–8-<br />
spored and ascospores are hyaline at first, becoming pale to<br />
dark brown, one to many septate, constricted or not at the<br />
septa and often surrounded by a mucilaginous sheath.<br />
Several morphologically distinct genera were synonymized<br />
under Herpotrichia using the above broad circumscription<br />
(Barr 1984; Müller and von Arx 1962; Sivanesan 1984). In<br />
particular, Barr kept Lojkania as a separate genus after<br />
studying its type material (Barr 1984, 1990a). Sivanesan<br />
(1984) was also of the opinion that Lojkania and Neopeckia<br />
were distinct genera as several of their characters differed.<br />
Byssosphaeria and Pseudotrichia have subsequently been<br />
assigned to Melanommataceae, Lojkania to Fenestellaceae<br />
and Neopeckia to Coccoideaceae (Barr 1984). Herpotrichia<br />
sensu stricto is represented by H. rubi and has erumpent to<br />
superficial ascomata or immersed in a subiculum, clavate to<br />
cylindrical, 4–8-spored, stalked asci with a conspicuous<br />
apical “nasse”, hyaline, 1-septate ascospores, usually becoming<br />
pale brown and several septate, constricted or not<br />
constricted at septa, usually surrounded by sheath (Sivanesan<br />
1984). Currently, about 90 species are included in this genus<br />
(http://www.indexfungorum.org/, 12/01/2009).<br />
Phylogenetic study<br />
Herpotrichia diffusa (Schwein.) Ellis & Everh., H.<br />
juniperi (Duby) Petr., H. herpotrichoides and H. macrotricha<br />
have been shown to have phylogenetic affinity with<br />
the generic types of Byssosphaeria schiedermayeriana,<br />
Melanomma pulvis-pyrius and Pleomassaria siparia, which<br />
had been assigned under Melanommataceae (Kruys et al.<br />
2006; Mugambi and Huhndorf 2009b; Schoch et al. 2006,<br />
2009; Zhang et al. 2009a). In this study, Pleomassaria<br />
siparia together with its closely related species of Prosthemium<br />
is kept in a separate family, viz Pleomassariaceae.<br />
Concluding remarks<br />
Even species under Herpotrichia sensu stricto (according<br />
to Sivanesan 1984) have diverse hosts (such as<br />
gymnosperms (H. coulteri (Peck) S.K. Bose and H. parasitica<br />
(R. Hartig) Rostr.) and angiosperms (H. diffusa and<br />
H. villosa Samuels & E. Müll.)) or substrates (like dead or<br />
living leaves, bark or decorticated wood) (Sivanesan 1984).<br />
Species of Herpotrichia sensu stricto are also reported from<br />
various locations such as Europe, Asia or America, and<br />
they have various life styles, e.g. parasitic, hyperparasitic or<br />
saprobic (Sivanesan 1984). Additional factors (like hosts or<br />
locations) may need to be considered in order to get a<br />
natural concept for Herpotrichia.<br />
Immotthia M.E. Barr, Mycotaxon 29: 504 (1987).<br />
(Teichosporaceae)<br />
Generic description<br />
Habitat terrestrial, hyperparasitic. Ascomata gregarious,<br />
globose, superficial, ostiolate, periphysate. Hamathecium<br />
of cellular pseudoparaphyses. Asci 8-spored, bitunicate,<br />
cylindrical, with a short pedicel. Ascospores 1-seriate,<br />
ellipsoidal, brown to reddish brown, 1-septate, constricted<br />
at the septum, smooth.<br />
Anamorphs reported for genus: none.<br />
Literature: Barr 1987a, 2002; Wang et al. 2004.<br />
Type species<br />
Immotthia hypoxylon (Ellis & Everh.) M.E. Barr, Mycotaxon<br />
29: 504 (1987). (Fig. 37)<br />
≡ Amphisphaeria hypoxylon Ellis & Everh., J. Mycol. 2:<br />
41 (1886).<br />
Ascomata gregarious, globose, superficial, ostiolate,<br />
periphysate, papillate (Fig. 37a). Hamathecium of cellular<br />
pseudoparaphyses, 2–2.5 μm broad, septate. Asci 60–82×<br />
7–9 μm, 8-spored, bitunicate, cylindrical, with a short<br />
pedicel (Fig. 37b, c and d). Ascospores 10–13×4.4–5.4 μm,<br />
1-seriate, ellipsoidal, brown to reddish brown, 1-septate,<br />
constricted at the septum, smooth (Fig. 37f, g and h)<br />
(adapted from Wang et al. 2004).<br />
Anamorph: none reported.<br />
Material examined: USA, Louisiana, Pointe a la<br />
Hache, on decaying wood, a branch of Carya oliviformis<br />
(Juglandaceae) lying on the ground in grass (parasitic on
Fungal Diversity<br />
Fig. 36 Herpotrichia rubi (from g, f. rh. 2171, type). a Numerous<br />
ascomata gregariously immersed in the host tissue. b Section of an<br />
ascoma. Note the central ostiole and peridium structure and also note<br />
the arrangement of asci and pseudoparaphyses. c Section of partial<br />
lateral peridium which comprises cells of textura angularis. d Part of a<br />
mature squashed ascus. e Relatively wide, septate pseudoparaphyses. f<br />
Immature ascus. Note the furcate pedicel. g–h One-septate ascospores.<br />
Note the verruculose ornamentation which is visible at the sides. Scale<br />
bars: a=0.5 mm, b=100 μm, c=50 μm, d=20 μm, e–h=10 μm
Fungal Diversity<br />
some effused Hypoxylon), 30 Dec. 1885, A.B. Langlois, No. 138<br />
(NY, holotype of Amphisphaeria hypoxylon Ellis & Everh.).<br />
Notes<br />
Morphology<br />
Immotthia was introduced to accommodate a species of<br />
Amphisphaeria (A. hypoxylon), which has bitunicate asci,<br />
and is characterized by superficial, ostiolate, periphysate,<br />
papillate ascomata, cellular pseudoparaphyses, bitunicate, 8-<br />
spored, cylindrical asci, ellipsoid, smooth, brown to reddish<br />
brown, 1-septate ascospores (Barr 1987a; Wangetal.2004).<br />
Phylogenetic study<br />
None.<br />
Concluding remarks<br />
It seems that those Amphisphaeria species with bitunicate<br />
asci should be assigned to <strong>Pleosporales</strong>. Morphologically,<br />
Immotthia is somewhat comparable with Herpotrichia.<br />
Isthmosporella Shearer & J.L. Crane, Mycologia 91: 141<br />
(1999). (<strong>Pleosporales</strong>, genera incertae sedis)<br />
Generic description<br />
Habitat freshwater, saprobic. Ascomata small- to mediumsized,<br />
scattered, immersed, erumpent to superficial, globose,<br />
papillate, ostiolate, periphysate, membranous. Peridium 2-<br />
layered, outer layer composed of brown, pseudoparenchymatic,<br />
fusoid-cylindric cells, inner layer composed of fusoid,<br />
subhyaline to pale brown, compressed cells. Hamathecium of<br />
rare, broad, septate, interascal pseudoparaphyses. Asci 8-<br />
spored, bitunicate, fissitunicate, oblong to clavate, with a<br />
short pedicel, ocular chamber not observed. Ascospores 3–4<br />
seriate, cylindrical to fusoid, isthmoid at centre, constricted at<br />
septa, isthmus 1-septate, surrounded by a gelatinous sheath.<br />
Anamorphs reported for genus: none.<br />
Literature: Shearer and Crane 1999.<br />
Type species<br />
Isthmosporella pulchra Shearer & J.L. Crane, Mycologia<br />
91: 142 (1999). (Fig. 38)<br />
Ascomata 240–330 μm diam., scattered on decorticated<br />
wood, immersed, erumpent to superficial, globose, black,<br />
papillate, papilla short, cylindrical, 60 μm long×55 μm<br />
wide, ostiolate, periphysate, membranous (Fig. 38a). Peridium<br />
2-layered, outer 3–4 cell layers composed of brown,<br />
pseudoparenchymatic, fusoid-cylindric cells, 2–6.5 μm<br />
long; inner layer composed of 5–7 rows of fusoid, subhyaline<br />
to pale brown compressed cells, 11–20×2–3.5 μm<br />
diam. (Fig. 38a and b). Hamathecium of rare, broad,<br />
septate, interascal pseudoparaphyses (Fig. 38f). Asci (95-)<br />
135–160(−175)×(25-)30–45(−60) μm, 8-spored, bitunicate,<br />
fissitunicate, oblong to clavate, with a short pedicel, ocular<br />
chamber not observed (Fig. 38c, d and e). Ascospores 80–<br />
105(−110)×(7-)8–10 μm, 3–4-seriate, cylindrical to fusoid,<br />
isthmoid at centre, sometimes bent at isthmus and becoming<br />
u- or v- shaped, end cells tapering, 12–17-phragmoseptate,<br />
constricted at septa, isthmus 1-septate, 2–5.5×2–<br />
4.5 μm diam., hyaline, frequently fragmenting to form<br />
partspores; filled with lipid droplets that merge to form<br />
large guttules; surrounded by a gelatinous sheath with a<br />
dense region near the isthmus, sheath greatly enlarging in<br />
water (Fig. 38g, h, i and j).<br />
Anamorph: none reported.<br />
Colonies on yeast soluble starch agar containing balsa<br />
wood sticks effuse, white. Hyphae hyaline, septate.<br />
Material examined: USA, New York, Adirondack Park.<br />
Piercefield. Tupper Lake at public boat launch from Rt. 30,<br />
UTM Zone 18, 539840 mE, 4892100mN; 44°10″59″N, 80°<br />
31′6″W, on submerged, decorticated wood, 7 Jul. 1994, J.L.<br />
Crane & C.A. Shearer A-254-1 (ILLS 53086, holotype).<br />
Notes<br />
Morphology<br />
Isthmosporella was described as a freshwater genus<br />
typified by I. pulchra, and is characterized by globose,<br />
pseudoparenchymatous ascomata, sparse, septate pseudoparaphyses,<br />
fissitunicate asci and hyaline, cylindrical to<br />
fusoid, phragmoseptate, isthmoid ascospores surrounded<br />
with a gelatinous sheath (Shearer and Crane 1999). Based<br />
on the morphological characters, i.e. small, globose<br />
ascomata, peridium with small pseudoparenchymatous cells<br />
and sparse pseudoparaphyses, Isthmosporella was assigned<br />
to the Phaeosphaeriaceae (Shearer and Crane 1999). The<br />
aquatic habitat of Isthmosporella, however, disagree with<br />
the Phaeosphaeriaceae. Isthmosporella seems less likely to<br />
belong to Pleosporineae.<br />
Phylogenetic study<br />
None.<br />
Concluding remarks<br />
Molecular phylogenetic studies should be conducted to<br />
explore its familial placement within <strong>Pleosporales</strong>.<br />
Kalmusia Niessl, Verh. nat. Ver. Brünn 10: 204 (1872).<br />
(Montagnulaceae)<br />
Generic description<br />
Habitat terrestrial, saprobic. Ascomata small- to mediumsized,<br />
solitary, scattered or in small groups, immersed to
Fungal Diversity<br />
Fig. 37 Immotthia hypoxylon (from holotype of Amphisphaeria hypoxylon). a Ascomata gregarious on host surface. b–d Bitunicate asci. e–<br />
h Released 1-septate ascospores. Scale bars: a=0.5 mm; b–h=10 μm<br />
erumpent, globose or subglobose, often laterally flattened,<br />
coriaceous, wall black, with or without papilla.<br />
Hamathecium of dense, filliform, delicate, septate pseudoparaphyses,<br />
branching and anastomosing between and<br />
above asci, embedded in mucilage. Asci bitunicate,<br />
fissitunicate unknown, clavate, with a long, furcate<br />
pedicel. Ascospores narrowly ovoid to clavate, pale<br />
brown, 3-septate, distoseptate.<br />
Anamorphs reported for genus: Cytoplea (Petrak and<br />
Sydow 1926).
Fungal Diversity<br />
Fig. 38 Isthmosporella pulchra<br />
(from ILLS 53086, holotype).<br />
a Section of an ascoma. b<br />
Section of a partial peridium. c–<br />
e Broadly clavate asci with short<br />
pedicels. f Pseudoparaphyses.<br />
g–j Ascospores. Note the 2-<br />
celled isthmus in J and mucilaginous<br />
sheath in G and H.<br />
Scale bars: a=50 μm, b–j=<br />
20 μm (figure with permission<br />
from http://fungi.life.illinois.<br />
edu/search/fwa_lit_refs)<br />
Literature: Barr 1987b, 1990a, 1992a; Lindau 1897; von<br />
Niessl 1872.<br />
Type species<br />
Kalmusia ebuli Niessl, Verh. nat. Ver. Brünn 10: 204<br />
(1872). (Fig. 39)<br />
Ascomata 290–360 μm high×300–520 μm diam., solitary,<br />
scattered, or in small groups, immersed to erumpent,<br />
globose or subglobose, coriaceous, wall black, with or<br />
without papilla, ostiolate (Fig. 39a). Papilla small, up to<br />
100 μm high, with small ostioles (Fig. 39a). Peridium 15–<br />
40 μm wide, comprising one cell type of small, pigmented,<br />
thick-walled cells of textura prismatica to textura angularis,<br />
cells ca. 5×3μm diam., cell wall 2–3 μm thick (Fig. 39b<br />
and c). Hamathecium of dense, delicate pseudoparaphyses, 1–<br />
1.5 μm broad, septate, branching and anastomosing between<br />
and above asci, embedded in mucilage. Asci 75–125×10–<br />
15 μm (x ¼ 90:5 12mm, n=10), 8-spored, bitunicate,<br />
fissitunicate unknown, clavate, with a long, narrowed, furcate
Fungal Diversity<br />
pedicel which is up to 45 μm long, and a low ocular chamber<br />
(ca. 2μm wide×1 μm high) (Fig. 39d, e and f). Ascospores<br />
15–18×5.5–6.5 μm (x ¼ 16:3 5:8mm, n=10), biseriate,<br />
narrowly ovoid to clavate, pale brown, 3-distoseptate, without<br />
constriction, smooth-walled (Fig. 39g, h and i).<br />
Anamorph: none reported.<br />
Material examined: BELGIUM, Dolembreux, on branchlets<br />
and pieces of stumps of Sarothamnus scoparius from<br />
woodland, Oct. 1922, V. Mouton (BR 101525–63, holotype).<br />
Notes<br />
Morphology<br />
Kalmusia was formally established by von Niessl (1872),<br />
and is mainly characterized as “immersed, sphaeroid ascoma<br />
with central, stout papilla, surrounded by hyphae in the<br />
substrate, stipitate asci with septate pseudoparaphyses, and<br />
brown, 3-septate, inequilateral ascospores” (Barr 1992a).<br />
The most morphologically comparable genus to Kalmusia<br />
is Thyridaria, which had been treated as a subgenus<br />
under Kalmusia (Lindau 1897), and was subsequently<br />
transferred to Platystomaceae in Melanommatales (Barr<br />
1987b, 1990a). Compared to Thyridaria, Kalmusia has<br />
sphaeroid ascomata, a peridium of small pseudoparenchymatous<br />
cells, basal asci and very thin pseudoparaphyses,<br />
thus it was assigned to Phaeosphaeriaceae of the <strong>Pleosporales</strong><br />
by Barr (1990a), and the genus is utilized to<br />
accommodate both K. ebuli and K. clivensis (Berk. &<br />
Broome) M.E. Barr, as well as closely related species, i.e.<br />
K. utahensis (Ellis & Everh.) Huhndorf & M.E. Barr and K.<br />
coniothyrium (Fuckel) Huhndorf (Barr 1992a). But this<br />
proposal is questionable, as the clavate, distoseptate ascospores,<br />
as well as the clavate asci with very long pedicels are<br />
uncommon in Phaeosphaeriaceae, and most recent phylogenetic<br />
study indicated that some species of Kalmusia reside<br />
outside of Phaeosphaeriaceae (Zhang et al. 2009a).<br />
Phylogenetic study<br />
Both Kalmusia scabrispora Teng Kaz. Tanaka, Y.<br />
Harada & M.E. Barr and K. brevispora (Nagas. & Y.<br />
Otani) Yin. Zhang, Kaz. Tanaka & C.L. Schoch reside in<br />
the clade of Montagnulaceae (Zhang et al. 2009a). Familial<br />
placement of Kalmusia can only be verified after the DNA<br />
sequences of the generic type (K. ebuli) are obtained.<br />
Concluding remarks<br />
Kalmusia is distinct amongst the <strong>Pleosporales</strong> as it has<br />
pale brown ascospores with indistinct distosepta and clavate<br />
asci with long pedicels. Although both K. scabrispora and<br />
K. brevispora reside in the clade of Montagnulaceae, they<br />
both lack the distoseptate ascospores that are possessed by<br />
the generic type (K. ebuli). Thus, the familial placement of<br />
Kalmusia is still undetermined.<br />
Karstenula Speg., Decades Mycologicae Italicae ad no. 94<br />
(in sched.) (1879). (Montagnulaceae)<br />
Generic description<br />
Habitat terrestrial, saprobic. Ascomata rarely small-,<br />
usually medium-sized, immersed usually under thin<br />
clypeus, scattered to gregarious, with flattened top and<br />
rounded pore-like ostiole, coriaceous. Peridium 2-layered,<br />
outer layer composed of reddish brown to dark brown<br />
small cells, inner layer of pale compressed cells. Hamathecium<br />
of dense, cellular pseudoparaphyses. Asci cylindrical<br />
to cylindro-clavate with short furcate pedicel.<br />
Ascospores muriform, ellipsoid to fusoid, reddish brown<br />
to dark brown.<br />
Anamorphs reported for the genus: Microdiplodia<br />
(Constantinescu 1993).<br />
Literature: Barr 1990a; Eriksson and Hawksworth 1991;<br />
Kodsueb et al. 2006a; Munk 1957; Zhang et al. 2009a.<br />
Type species<br />
Karstenula rhodostoma (Alb. & Schwein.) Speg., Decades<br />
Mycologicae Italicae no. 94. (1879). (Fig. 40)<br />
≡ Sphaeria rhodostoma Alb. & Schwein., Consp. fung.<br />
(Leipzig): 43 (1805).<br />
Ascomata 250–430 μm high×450–650 μm diam., scattered<br />
or gregarious, immersed in the subiculum which sometimes<br />
sloths off, globose or subglobose, black, flattened top often<br />
white or reddish and sometimes slightly protruding out of the<br />
substrate surface, usually with a wide opening ostiole after<br />
removing the cover, coriaceous (Fig. 40a and b). Peridium<br />
30–40 μm wide, comprising two cell types, outer region 1-<br />
layered, composed of relatively small heavily pigmented<br />
thick-walled compressed cells, cells 2–4×5–10 μm diam.,<br />
cell wall 2–4 μm thick, inner layer cells larger and wall<br />
thinner, comprising cells of textura angularis, merging with<br />
pseudoparaphyses (Fig. 40c and d). Hamathecium of dense,<br />
long cellular pseudoparaphyses 2–3.5 μm broad, septate,<br />
branching or anastomosing not observed. Asci 150–210×<br />
12.5–15 μm (x ¼ 182 13:1mm, n=10), 8-spored, bitunicate,<br />
fissitunicate, cylindrical, with a broad, furcate pedicel which is<br />
12–35 μm long, and with an ocular chamber (to 4 μm wide×<br />
3 μm high) (Fig. 40e and f). Ascospores 20–26×7.5–10 μm<br />
(x ¼ 22:4 8mm, n=10), obliquely uniseriate and partially<br />
overlapping, ellipsoid, reddish brown, with 3 transverse septa<br />
and a vertical septum in one or two central cells, constricted at<br />
the septa, verruculose (Fig. 40g, h and i).<br />
Anamorph: Microdiplodia frangulae Allesch. (Constantinescu<br />
1993).<br />
Conidiomata globose to subglobose, 330–495 μm diam.,<br />
in subiculum. Conidia 9–13×4–5 μm, reddish brown, 1-<br />
septate (information obtained from Barr 1990a).
Fungal Diversity<br />
Fig. 39 Kalmusia ebuli (from BR 101525–63, holotype). a Immersed<br />
to erumpent ascomata scattered on the host surface. b Section<br />
of a partial peridium. Note the compressed peridium cells. c Section of<br />
an ascoma. d–f Eight-spored asci with long pedicels. g Partial ascus in<br />
pseudoparaphyses. h, i Ascospores with 3 thick-walled septa. Scale<br />
bars: a=0.5 mm, b=50 μm, c=100 μm, d–g=20 μm, h, i=10 μm<br />
Material examined: Fries, Suecia (received by herbarium<br />
in 1834) (PH 01048835, type, as Sphaeria rhodostoma<br />
Alb. & Schwein.).<br />
Notes<br />
Morphology<br />
Karstenula is an ambiguous genus, which has been<br />
synonymized under Pleomassaria (Lindau 1897; Winter<br />
1885). Some of the ascomata characters are even comparable<br />
with those of Didymosphaeria, such as ascomata<br />
seated in subiculum or beneath a clypeal thickening, the<br />
development of apex vary in a large degree, even to the<br />
occasional formation of a blackened internal clypeus, and<br />
sometimes apical cells become reddish or orange-brown<br />
(Barr 1990a). Barr (1990a) redefined the concept of<br />
Karstenula (sensu lato), which encompasses some species<br />
of Thyridium. In her concept, however, Barr (1990a)<br />
treated Karstenula as having trabeculate pseudoparaphyses<br />
and this is clearly not the case. In most cases the
Fungal Diversity<br />
Fig. 40 Karstenula rhodostoma (from PH 01048835, type). a Line<br />
of ascomata on host surface (after remove the decaying cover). Note<br />
the wide ostiolar opening and light colored region around the ostiole.<br />
b Immersed ascoma under the decaying cover (see arrow). c, d<br />
Section of the peridium. The peridium comprises small thick-walled<br />
cells in the outer layer. The outside comprises defuse hyphae which is<br />
probably part of the subiculum. e Ascus with a short furcate pedicel. f<br />
Partial ascus showing arrangement of ascospores. g–i Released<br />
ascospores. Note the transverse and rarely vertical septa. Scale bars:<br />
a, b=0.5 mm, c=50 μm, d–f=20 μm, g–i=10 μm
Fungal Diversity<br />
ascospores were brown with transverse septa and sparse<br />
longitudinal septa.<br />
The ascomata of this species are similar to those found in<br />
Byssosphaeria and Herpotrichia, especially in the paler area<br />
around the ostiole and even in peridial structure and<br />
development under a subiculum. The numerous wide cellular<br />
pseudoparaphyses and cylindrical asci (in Herpotrichia) are<br />
also similar. The main difference of Karstenula from other<br />
two genera are the 3-septate ascospores with rare longitudinal<br />
septa (1-septate in Byssosphaeria and Herpotrichia).<br />
Phylogenetic study<br />
Karstenula forms a robust phylogenetic clade with<br />
Phaeodothis winteri (Niessl) Aptroot, Didymocrea sadasivanii,<br />
Bimuria novae-zelandiae, Montagnula opulenta, Curreya<br />
pityophila (J.C. Schmidt & Kunze) Arx & E. Müll. and<br />
some species of Letendraea and Paraphaeosphaeria (Kodsueb<br />
et al. 2006a; Zhang et al. 2009a). Consequently,<br />
Karstenula might be included in Montagnulaceae.<br />
Concluding remarks<br />
The description of the type of Karstenula here clearly<br />
excludes it from Melanommataceae as it has wide<br />
pseudoparaphyses. But its Montagnulaceae status can only<br />
be confirmed by more phylogenetic work including<br />
sequencing the generic type of Karstenula (K. rhodostoma).<br />
Katumotoa Kaz. Tanaka & Y. Harada, Mycoscience 46:<br />
313 (2005). (Lentitheciaceae)<br />
Generic description<br />
Habitat terrestrial or freshwater, saprobic. Ascomata smallto<br />
medium-sized, scattered or in small groups, immersed to<br />
erumpent, with a central protruding hairy papilla, subglobose.<br />
Peridium thin, comprising several layers of thinwalled<br />
compressed cells. Hamathecium of dense, cellular,<br />
filliform, embedded in mucilage, branching and anastomosing.<br />
Asci 8-spored, bitunicate, fissitunicate, clavate with<br />
short furcate pedicels. Ascospores apiosporous and hyaline<br />
when young, becoming 2-septate with reddish brown<br />
echinate central cell at maturity, with long gelatinous<br />
terminal appendages.<br />
Anamorphs reported for genus: none.<br />
Literature: Tanaka and Harada 2005b; Tanaka et al. 2009;<br />
Zhang et al. 2009a.<br />
Type species<br />
Katumotoa bambusicola Kaz. Tanaka & Y. Harada,<br />
Mycoscience 46: 313 (2005). (Fig. 41)<br />
Some information for the following description is from<br />
Tanaka and Harada (2005).<br />
Ascomata 240–330 μm high×260–420 μm diam.,<br />
scattered or in small groups, immersed, becoming erumpent,<br />
with a slightly protruding papilla covered with brown<br />
hyphae, subglobose (Fig. 41a). Peridium 13–30 μm thick,<br />
composed of a few layers of lightly pigmented, depressed<br />
cells. Hamathecium of dense, long cellular pseudoparaphyses,<br />
1.5–3 μm broad, embedded in mucilage, branching and<br />
anastomosing. Asci 110–160×17.5–24 μm (x ¼ 139 21mm,<br />
n=10), 8-spored, bitunicate, fissitunicate, cylindro-clavate<br />
withashortfurcatepedicelwhichisupto25μm long<br />
(Fig. 41b and d). Ascospores 39–50(−57)×7–10 μm<br />
(x ¼ 45:8 8:2mm, n=10), biseriate, fusoid to narrowly<br />
fusoid with acute ends, usually curved, apiosporus and<br />
hyaline when young, constricted at the primary septum, the<br />
upper cell longer and broader than the lower one, smooth,<br />
surrounded by a bipolar sheath which is up to 15 μm long,<br />
best seen in India ink, senescent ascospores yellowish<br />
brown, 2–4-septate (Fig. 41c).<br />
Anamorph: none reported.<br />
Material examined: JAPAN, Mt. Iwate, near Yakebashiri,<br />
Hirakasa, Nishine, Iwate, on culms of Oryza sativa L., 19<br />
Oct. 2003, K. Tanaka (HHUF 28663, holotype).<br />
Notes<br />
Morphology<br />
Katumotoa was formally established by Tanaka and<br />
Harada (2005b) to accommodate the monotypic species,<br />
K. bambusicola, which is characterized by immersed<br />
ascomata with a thin peridium comprising thin-walled<br />
compressed cells, cellular pseudoparaphyses, cylindroclavate<br />
and fissitunicate asci and fusoid ascospores with<br />
an elongated bipolar mucilaginous sheath. Based on its<br />
immersed ascomata, psuedoparenchymatous peridium<br />
cells and cellular pseudoparaphyses, Katumotoa was<br />
assigned to Phaeosphaeriaceae (Tanaka and Harada<br />
2005b; Tanaka et al. 2009), but this classification has<br />
been shown to be incorrect in subsequent phylogenetic<br />
studies (Tanaka et al. 2009; Zhangetal.2009a).<br />
Phylogenetic study<br />
Phylogenetic analysis based on five genes (LSU, SSU,<br />
RPB1, RPB2 and EF1) indicates that Katumotoa bambusicola<br />
resides in Lentitheciaceae, and this receives high<br />
bootstrap support (Zhang et al. 2009a). In particular, K.<br />
bambusicola forms a robust clade with Ophiosphaerella<br />
sasicola (Nagas. & Y. Otani) Shoemaker & C.E. Babc.,<br />
which has filliform ascospores (Shoemaker and Babcock<br />
1989b).<br />
Concluding remarks<br />
The hyaline, apiosporous ascospores which become<br />
2–4-celled with central reddish brown cells and large
Fungal Diversity<br />
Fig. 41 Katumotoa bambusicola (from HHUF 28663, holotype). a Ascomata scattered on the host surface. b Asci in pseudoparaphyses. c<br />
Hyaline ascospore with long terminal appendages. d Clavate ascus with a short pedicel. Scale bars: a=0.5 mm. b–d=20 μm<br />
unraveling appendages are the most striking features of<br />
this species and readily distinguish it from other<br />
pleosporalean taxa. Both Katumotoa bambusicola and<br />
Ophiosphaerella sasicola are associated with bambusicolous<br />
hosts, which might indicate that host spectrum in<br />
this case, has greater phylogenetic significance than some<br />
morphological characters (Zhang et al. 2009a).<br />
Keissleriella Höhn., Sber. Akad. Wiss. Wien, Math.-naturw.<br />
Kl., Abt. 1 128: 582 (1919). (Lentitheciaceae)<br />
Generic description<br />
Habitat terrestrial or freshwater, saprobic. Ascomata smallto<br />
medium-sized, immersed, erumpent to nearly superficial,<br />
globose, papillate, ostiolate. Papilla covered by dark setae<br />
or small blackened cells. Peridium thick, composed of cells<br />
of pseudoparenchymatous and inner layer composed of pale<br />
cells. Hamathecium of dense, long pseudoparaphyses,<br />
rarely septate, anastomosing and branching. Asci 4- or 8-<br />
spored, bitunicate, fissitunicate, cylindro-clavate, with a
Fungal Diversity<br />
furcate pedicel and a small ocular chamber. Ascospores<br />
hyaline to pale brown, ellipsoid to fusoid, 1-septate,<br />
constricted at the septum (Barr 1990a).<br />
Anamorphs reported for genus: Dendrophoma (Bose 1961).<br />
Literature: von Arx and Müller 1975; Bose 1961; Barr<br />
1990a; Dennis 1978; Eriksson 1967a; von Höhnel 1919;<br />
Luttrell 1973; Munk 1957; Zhang et al. 2009a.<br />
Type species<br />
Keissleriella aesculi (Höhn.) Höhn., Sber. Akad. Wiss.<br />
Wien, Math.-naturw. Kl., Abt. 1 128: 582 (1919). (Fig. 42)<br />
≡ Pyrenochaeta aesculi Höhn., Ber. dt. bot. Ges. 35: 249<br />
(1917).<br />
Ascomata ca. 250μm high×450 μm diam., gregarious,<br />
immersed to erumpent, globose or subglobose, with a<br />
small black papilla, ca. 75μm high and 110 μm broad,<br />
with short black external setae (Fig. 42a). Peridium ca.<br />
25–40 μm wide laterally, up to 70 μm near the apex,<br />
thinner at the base, comprising two types of cells which<br />
merge in the middle; outer cells composed of small<br />
heavily pigmented thick-walled cells, cells ca. 4 μm<br />
diam., cell wall up to 4 μm thick, and thick near the apex<br />
and thinner laterally and absent in the immersed part of<br />
the ascoma, inner cells less pigmented, comprising<br />
lightly pigmented to hyaline cells, 5–7 μm thick<br />
(Fig. 42a). Hamathecium of dense, long pseudoparaphyses,<br />
0.8–1.2 μm broad, rarely septate, anastomosing and<br />
branching, thicker near the base, ca. 2μm, constricted<br />
near the septum (Fig. 42b). Asci 80–120×6–11 μm (x ¼<br />
101 8:5mm, n=10), 4- or 8-spored, bitunicate, fissitunicate,<br />
cylindro-clavate, with a furcate pedicel which is<br />
up to 20–40 μm long, with a small ocular chamber<br />
(Fig. 42e and f). Ascospores 13–18×4–5.5 μm<br />
(x ¼ 14:5 4:8mm, n=10), obliquely uniseriate and partially<br />
overlapping, fusoid with narrowly rounded ends,<br />
hyaline, 1-septate, constricted at the septum, smooth<br />
(Fig. 42c and d).<br />
Anamorph: none reported.<br />
Material examined: AUSTRIA, Brentenmaistal in the<br />
Viennese forest, Aesculus hippocastanum L., 1916, Höhnel<br />
(FH, holotype of Otthiella aesculi). (Note: only two slides;<br />
setae cannot be seen from the slides but could be seen from<br />
the drawings on the cover).<br />
Notes<br />
Morphology<br />
Keissleriella is characterized by ascomata with setae in<br />
and over the papilla, asci are cylindrical and ascospores are<br />
hyaline, 1-septate. Based on the morphological characters,<br />
K. aesculi was regarded as conspecific with K. sambucina;<br />
as an earlier epithet, K. sambusina typifies the genus (see<br />
comments by Barr 1990a). Munk (1957) placed Trichometasphaeria<br />
and Keissleriella in Massarinaceae, and distinguished<br />
them by their substrates (Trichometasphaeria<br />
occurs on herbaceous plants and Keissleriella on woody<br />
substrates). Bose (1961) combined Trichometasphaeria<br />
under Keissleriella, which was followed by some workers<br />
(von Arx and Müller 1975; Dennis 1978; Eriksson 1967a;<br />
Luttrell 1973). Barr (1990a), however, maintained these as<br />
distinct genera based on the differences of peridium<br />
structure and pseudoparaphyses.<br />
Phylogenetic study<br />
The phylogeny of Keissleriella is poorly studied.<br />
Limited phylogenetic information indicates that K. cladophila<br />
forms a robust clade with other species of Lentitheciaceae<br />
(Zhang et al. 2009a).<br />
Concluding remarks<br />
The presence of black setae on the surface of papilla is a<br />
striking character of Keissleriella, but phylogenetic significance<br />
of setae is undetermined yet.<br />
Lentithecium K.D. Hyde, J. Fourn. & Yin. Zhang, Fungal<br />
Divers. 38: 234 (2009). (Lentitheciaceae)<br />
= Tingoldiago K. Hirayama & Kaz. Tanaka, Mycologia<br />
102: 740 (2010) syn. nov.<br />
Generic description<br />
Habitat freshwater, saprobic. Ascomata small, scattered<br />
or gregarious, immersed, slightly erumpent, depressed<br />
spherical to lenticular, ostiolate, papillate or epapillate.<br />
Peridium thin. Hamathecium of cellular pseudoparaphyses.<br />
Asci 8-ascospored, bitunicate, fissitunicate, clavate,<br />
short-stipitate. Ascospores broadly fusoid with broadly<br />
rounded ends, 1-septate, constricted, hyaline, usually<br />
with sheath.<br />
Anamorphs reported for genus: none.<br />
Literature: Shearer et al. 2009; Zhang et al. 2009a, b.<br />
Type species<br />
Lentithecium fluviatile (Aptroot & Van Ryck.) K.D. Hyde, J.<br />
Fourn. & Yin. Zhang, Fungal Divers. 38: 234 (2009). (Fig. 43)<br />
≡ Massarina fluviatilis Aptroot & Van Ryck., Nova<br />
Hedwigia 73: 162 (2001).<br />
Ascomata 230–260 μm high×280–325 μm diam., scattered<br />
or gregarious, immersed, slightly erumpent, subglobose to<br />
depressed spherical, under a small black pseudostroma<br />
originating from the apical part of the peridium, apex slightly<br />
papillate, ostiole rounded, 60–70 μm diam. (Fig. 43a and b).<br />
Peridium 15–20 μm thick at sides and at base, comprising 4–5<br />
layers of angular cells more thick-walled outwards, 50–55 μm
Fungal Diversity<br />
Fig. 42 Keissleriella sambucina (from FH, holotype of Otthiella<br />
aesculi). a Section of an ascoma. b Pseudoparaphyses which are<br />
narrow (less than 1.5 μm) and branch and anastomosing as<br />
trabeculate. c, d Hyaline ascospores with distinct constrictions at the<br />
septa. e Asci amongst narrow pseudoparaphyses. F. Ascus with a<br />
pedicel and ocular chamber. Scale bars: a=100 μm, b–f=10 μm<br />
thick at apex, of small very thick-walled cells. Hamathecium<br />
of cellular pseudoparaphyses, 2–2.5 μm broad (Fig. 43c and<br />
d). Asci 89–100×19–21 μm, 8-spored, bitunicate, fissitunicate,<br />
clavate, bumpy, short-stipitate, apex without obvious<br />
apical chamber (Fig. 43e). Ascospores 27–35×8.5–9.4 μm,, 2-<br />
3-seriate, broadly fusoid with broadly rounded ends, straight to<br />
slightly curved, 1-septate, slightly constricted, with four large<br />
guttules, hyaline, smooth-walled, a very thin mucilaginous<br />
sheath can be occasionally observed in India ink but in most<br />
cases no sheath can be observed (Fig. 43f and g).
Fungal Diversity<br />
Anamorph: none reported.<br />
Material examined: FRANCE, Haute Garonne: Avignonet,<br />
Lac de Rosel, artificial lake, on bark and wood of a<br />
submerged branch Populus sp., 23 Nov. 2006, leg. Michel<br />
Delpont, det. Jacques Fournier (IFRD 2039, holotype).<br />
Notes<br />
Morphology<br />
Lentithecium was introduced to accommodate some freshwater<br />
fungi previous assigned under Massarina, suchasM.<br />
arundinacea (Sowerby) Leuchtm. and M. fluviatilis (Zhang et<br />
al. 2009a). It is characterized by its immersed and<br />
lenticular ascomata, thin peridium which is almost equal<br />
in thickness, short pedicellate asci and fusoid or filliform,<br />
hyaline or rarely lightly pigmented, 1- to multi-septate<br />
ascospores (Zhang et al. 2009b). Lentitheciaceae was<br />
introduced to accommodate Lentithecium and some other<br />
related taxa (Zhang et al. 2009a).<br />
Phylogenetic study<br />
The clade of Lentitheciaceae comprises the generic type<br />
Lentithecium fluviatile, as well as L. arundinaceum<br />
(Sowerby) K.D. Hyde, J. Fourn. & Yin. Zhang, Stagonospora<br />
macropycnidia, Wettsteinina lacustris (Fuckel) Shoemaker<br />
& C.E. Babc., Keissleriella cladophila, and the<br />
bambusicolous species Katumotoa bambusicola and<br />
Ophiosphaerella sasicola, which receive high bootstrap<br />
support (Zhang et al. 2009a).<br />
Concluding remarks<br />
Tingoldiago graminicola K. Hirayama & Kaz. Tanaka<br />
form a robust clade with species of Lentithecium (Shearer et<br />
al. 2009). Tingoldiago has lenticular immersed to erumpent<br />
ascomata, numerous and septate pseudoparaphyses,<br />
cylindro-clavate asci and hyaline, 1-septate ascospores with<br />
sheath. All of these characters fit Lentithecium well. We<br />
treat Tingoldiago as a synonym of Lentithecium.<br />
Leptosphaeria Ces. & De Not., Comm. Soc. crittog. Ital. 1:<br />
234 (1863). (Leptosphaeriaceae)<br />
Generic description<br />
Habitat terrestrial, saprobic or parasitic. Ascomata small- to<br />
medium-sized, solitary, scattered or in small groups,<br />
erumpent to superficial, subglobose, broadly or narrowly<br />
conical, papillate, ostiolate. Peridium thick, comprising<br />
layers of cells of textura angularis. Hamathecium of dense<br />
cellular pseudoparaphyses, embedded in mucilage, anastomosing<br />
and branching. Asci 8-spored, bitunicate, fissitunicate<br />
unknown, cylindrical with a furcate pedicel and a large<br />
ocular chamber. Ascospores fusoid or narrowly fusoid,<br />
brown or reddish brown, 3-septate, constricted at each<br />
septum.<br />
Anamorphs reported for genus: Coniothyrium and<br />
Phoma (Hyde et al. 2011; Sivanesan 1984).<br />
Literature: von Arx and Müller 1975; Barr1987a, b; Cesati<br />
and de Notaris 1863; Crane and Shearer 1991; Dongetal.<br />
1998; Eriksson 1967a; Eriksson and Hawksworth 1986,<br />
1991; de Greuter et al. 1988; Hedjaroude1969; von Höhnel<br />
1907;Holm1957, 1975;Huhndorfetal.1990; Luttrell 1973;<br />
Müller 1950; Munk 1957; Saccardo 1878b, 1883, 1891,<br />
1895; Schochetal.2009; Shearer 1993; Shearer et al. 1990;<br />
Shoemaker 1984a; Sivanesan1984; Zhang et al. 2009a.<br />
Type species<br />
Leptosphaeria doliolum Ces. & De Not., Comm. Soc.<br />
crittog. Ital. 1: 234 (1863). (Fig. 44)<br />
≡ Sphaeria doliolum Pers., Icon. Desc. Fung. Min.<br />
Cognit. (Leipzig) 2: 39 (1800).<br />
Ascomata 340–450 μm high×380–500 μm diam., solitary,<br />
scattered or in small groups, superficial, subglobose, broadly<br />
or narrowly conical, with a flattened base on the host surface,<br />
black, usually with 2–4 ring-like ridges surrounding the<br />
ascomata surface, apex with a conical, usually shiny papilla<br />
(Fig. 44a). Peridium 85–110 μm wide at sides, thinner at the<br />
apex, comprising two types of cells, outer layer composed of<br />
small thick-walled cells of textura angularis, cells
Fungal Diversity<br />
lectotype species (de Greuter et al. 1988; Holm1975;<br />
Shearer et al. 1990). Leptosphaeria was originally<br />
defined based mainly on the characters of ascospores<br />
being ellipsoid or fusoid, one to many septa, hyaline to<br />
dark brown. These few common characters meant that<br />
Leptosphaeria comprised many species, and some of<br />
them should be assigned to either Euascomycetes or<br />
Loculoascomycetes (Crane and Shearer 1991). Leptosphaeria<br />
had been divided based on host and habitat<br />
(Saccardo 1878b, 1891, 1895) as well as the pseudothecium<br />
(glabrous, hairy, setose) and ascospore septation<br />
(see comments by Crane and Shearer 1991). von Höhnel<br />
(1907) used centrum structure in the classification of<br />
Leptosphaeria, and divided Leptosphaeria into three<br />
genera, viz. Leptosphaeria, Scleropleella and Nodulosphaeria.<br />
Müller (1950) subdivided Leptosphaeria into<br />
four sections based on pseudothecial and centrum<br />
structure as well as ascospore characters. This classification<br />
was modified by Munk (1957), who named these<br />
four sections as section I (Eu-Leptosphaeria), section II<br />
(Para-Leptosphaeria), section III (Scleropleella) and<br />
section IV (Nodulosphaeria). Holm (1957) used a<br />
relatively narrow concept for Leptosphaeria, which<br />
included species closely related to the generic type, L.<br />
doliolum. This viewpoint was accepted by some workers<br />
(Eriksson 1967a; Hedjaroude 1969; Shoemaker 1984a).<br />
Nevertheless, it still seems a heterogeneous group of<br />
fungi (see comments by Crane and Shearer 1991). Its position<br />
among the Loculoascomycetes is also debated. It has been<br />
placed in the Pleosporaceae (von Arx and Müller 1975;<br />
Luttrell 1973; Sivanesan 1984) orLeptosphaeriaceae (Barr<br />
1987a, b; Eriksson and Hawksworth 1991) orPhaeosphaeriaceae<br />
(Eriksson and Hawksworth 1986).<br />
Phylogenetic study<br />
Molecular phylogenetic analysis based on multigenes<br />
indicated that species of Leptosphaeria (including the<br />
generic type L. doliolum) and Neophaeosphaeria form a<br />
paraphyletic clade with moderate bootstrap support (Dong<br />
et al. 1998; Schoch et al. 2009; Zhang et al. 2009a), which<br />
is sister to other families of <strong>Pleosporales</strong> (Zhang et al.<br />
2009a). Thus the familial rank of the Leptosphaeriaceae<br />
could be temporarily verified, but further molecular<br />
phylogenetic study is needed in which more related taxa<br />
should be included.<br />
Concluding remarks<br />
Morphologically, Leptosphaeria is mostly comparable<br />
with Amarenomyces, Bricookea, Diapleella, Entodesmium,<br />
Melanomma, Nodulosphaeria, Paraphaeosphaeria, Passeriniella,<br />
Phaeosphaeria and Trematosphaeria. While it<br />
prefers non-woody parts of dicotyledonous hosts, its<br />
cylindrical ascus with short pedicel and smooth, fusoid<br />
Fig. 43 Lentithecium fluviatile (from IFRD 2039). a Erumpent b<br />
ascomata scattering on the host surface. b Habitat section of the<br />
immersed ascomata. c, d Section of an ascoma and a partical peridium.<br />
Note the peridium cells of textura angularis. e Clavate 8-spored ascus<br />
with a short pedicel. f, g Hyaline, 1-septate broadly fusoid ascospores.<br />
Scale bars: a, b=0.5 mm, c=100 μm, d=50 μm, e–g=20 μm<br />
and multi-septate ascospores make it readily distinguishable<br />
from all other genera (Shoemaker 1984a).<br />
Leptosphaerulina McAlpine, Fungus diseases of stonefruit<br />
trees in Australia and their treatment: 103 (1902).<br />
(Didymellaceae)<br />
Generic description<br />
Habitat terrestrial, parasitic or saprobic. Ascomata small,<br />
scattered, immersed, globose to subglobose, with a small,<br />
slightly protruding papilla, ostiolate. Peridium thin. Hamathecium<br />
of rare or decomposing cellular pseudoparaphyses.<br />
Asci bitunicate, obpyriform. Ascospores broadly clavate or<br />
cylindrical, hyaline, turning pale brown when old, asymmetrical,<br />
multi-septate, smooth-walled.<br />
Anamorphs reported for genus: Pithoascus and Pithomyces<br />
(Hyde et al. 2011).<br />
Literature: Barr 1972; Chlebicki 2002; Crivelli 1983;<br />
Kodsueb et al. 2006a; Zhang et al. 2009a.<br />
Type species<br />
Leptosphaerulina australis McAlpine, Fungus diseases of<br />
stone-fruit trees in Australia and their treatment: 103 (1902).<br />
(Fig. 45)<br />
Ascomata 140–170 μm diam., scattered, immersed,<br />
globose to subglobose, with a small slightly protruding<br />
papilla, ostiolate (Fig. 45a). Peridium thin, composed of<br />
one or two layers of large cells of textura angularis, pale<br />
brown (Fig. 45a). Hamathecium of rare or decomposing<br />
cellular pseudoparaphyses, up to 5 μm broad, filling<br />
the gaps between the asci. Asci 38–53×55–75 μm<br />
(x ¼ 67:5 43:3mm, n=10), 8-spored, without pedicel, bitunicate,<br />
fissitunicate dehiscence not observed, obpyriform, with a<br />
large ocular chamber and apical ring (Fig. 45b and c).<br />
Ascospores 30–40(-47)×11–14 μm (x ¼ 36:5 13mm, n=<br />
10), broadly clavate, hyaline, turning pale brown when old,<br />
asymmetrical, upper hemisphere usually with one transverse<br />
septum and with a somewhat narrowly rounded<br />
end, lower hemisphere usually with two transverse septa<br />
and with broadly rounded ends, slighted constricted at<br />
the primary septum, mostly with one vertical septum in<br />
each central cell, smooth, with thin gelatinous sheath<br />
when young, 2–3 μm thick (Fig. 45d and e).<br />
Anamorph: none reported.
Fungal Diversity
Fungal Diversity<br />
Fig. 44 Leptosphaeria doliolum (from L, lectotype). a Ascomata on the host surface. Note the shiny black surface. b Section of the partial peridium.<br />
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<br />
Material examined: USA, Kansas, Kansas State College,<br />
on Poa pratensis L. Grass plots, 2 Jul. 1953, leg. T.<br />
Rogerson, det. L.E. Wehmeyer (NY, C.T. Rogerson 3836).<br />
Notes<br />
Morphology<br />
Leptosphaerulina, introduced by McAlpine (1902), is<br />
characterized by small immersed ascomata, obpyriform asci<br />
with a large ocular chamber and apical ring as well as<br />
muriformly septate ascospores which may be hyaline or<br />
pigmented. Species of Leptosphaerulina may occur on<br />
monocotyledons or dicotyledons. Leptosphaerulina is most<br />
comparable with Pleospora, and the only difference between<br />
them is that Leptosphaerulina has smaller ascomata and<br />
hyaline ascospores that only become pigmented after<br />
discharge, whereas the ascospores of Pleospora become<br />
brown within the asci. Currently, about 60 names are<br />
accepted in this genus, and some even reported from marine<br />
environments, e.g. L. mangrovei (Inderbitzin et al. 2000).<br />
Phylogenetic study<br />
Based on multigene phylogenetic analysis, two putative<br />
strains of Leptosphaerulina australis, the generic type of<br />
Leptosphaerulina, from Switzerland (<strong>CBS</strong> 311.51) and<br />
Indonesia (<strong>CBS</strong> 317.83) resided within Didymellaceae (de<br />
Gruyter et al. 2009; Zhang et al. 2009a).<br />
Concluding remarks<br />
Because of its morphological confusion with Pleospora<br />
and the diversity of habitats within the genus, Leptosphaerulina<br />
sensu lato is likely to be polyphyletic. Fresh<br />
collections of this species are needed from Australia to<br />
epitypify this taxon and define the genus in a strict sense.<br />
The specimen described here is a collection from USA and<br />
therefore may not represent the type.<br />
Lewia M.E. Barr & E.G. Simmons, Mycotaxon 25: 289<br />
(1986). (Pleosporaceae)<br />
Generic description<br />
Habitat terrestrial, parasitic or saprobic? Ascomata<br />
small, scattered, erumpent to nearly superficial at maturity,<br />
subglobose to globose, black, smooth, papillate,<br />
ostiolate. Papilla short, blunt. Peridium thin. Hamathecium<br />
of pseudoparaphyses. Asci (4–6-)8-spored, bitunicate,<br />
fissitunicate, cylindrical to cylindro-clavate, with a<br />
short, furcate pedicel. Ascospores muriform, ellipsoid to<br />
fusoid.<br />
Anamorphs reported for genus: Alternaria (Simmons 1986).<br />
Literature: Kwasna and Kosiak 2003; Kwasnaetal.2006;<br />
Simmons 1986, 2007; Vieira and Barreto 2006.<br />
Type species<br />
Lewia scrophulariae (Desm.) M.E. Barr & E.G. Simmons,<br />
Mycotaxon 25: 294 (1986). (Fig. 46)<br />
≡ Sphaeria scrophulariae Desm., Plantes cryptogames<br />
du Nord de la France, ed. 1 fasc. 15:no. 718<br />
(1834).<br />
Ascomata ca. 150–200 μm diam., scattered, erumpent to<br />
nearly superficial at maturity, subglobose to globose, black,<br />
smooth, papillate. Papilla short, blunt. Peridium thin.<br />
Hamathecium of septate pseudoparaphyses, ca. 2–2.5 μm<br />
broad, anastomosing or branching not observed. Asci 100–<br />
140×13–17 μm, (4–6-)8-spored, bitunicate, fissitunicate,<br />
cylindrical to cylindro-clavate, with a short, furcate pedicel,<br />
ocular chamber unknown (Fig. 46a). Ascospores ellipsoid,<br />
5 (rarely 6 or 7) transversal septa and one longitudinal<br />
septum mostly through the central cells, yellowish brown to<br />
gold-brown, 20–24×8–10 μm (x ¼ 21:5 9:1mm, n=10),<br />
constricted at median septum, smooth or verruculose<br />
(Fig. 46b, e and f).<br />
Anamorph: Alternaria conjuncta (Simmons 1986).<br />
Primary conidiophore simple with a single conidiogenous<br />
locus; conidia produced in chains, the first conidia<br />
in chain is larger, 30–45×10–12 μm, 7 transverse septa,<br />
1–2 longitudinal or oblique septa in lower cells.<br />
Secondary conidiophore with 5–7 conidiogenous loci,<br />
sometimes branched; sporulation in chains, rarely<br />
branched.<br />
Material examined: (FH, slide from lectotype).<br />
Note: The specimen contains only a slide, so limited<br />
structures could be observed e.g. ascospores. The information<br />
about ascomata, peridium and whole asci is referred to<br />
Simmons (1986).<br />
Notes<br />
Morphology<br />
Lewia has “Pleospora-like” teleomorphs, while it has<br />
Alternaria anamorphs, which are characterized by the<br />
beakless conidia connected together with secondary<br />
conidiophore (Simmons 1986). Based on these characters,<br />
more species under this genus were subsequently reported,<br />
i.e. Lewia avenicola Kosiak & Kwaśna (Kwasna and<br />
Kosiak 2003); L. chlamidosporiformans B.S. Vieira & R.<br />
W. Barreto (Vieira and Barreto 2006); L. alternarina (M.<br />
D. Whitehead & J.G. Dicks.) E.G. Simmons and L.<br />
daucicaulis E.G. Simmons (Simmons 2007). Currently<br />
Lewia comprises 15 species (http://www.mycobank.org,<br />
24-02-2009).<br />
Phylogenetic study<br />
Phylogenetic analysis based either on SSU rDNA<br />
sequences or on multigenes indicated that Lewia species
Fungal Diversity<br />
Fig. 45 Leptosphaerulina australis (from NY, C.T. Rogerson 3836).<br />
a. Compressed ascoma. Note the obpyriform asci within the ascoma<br />
and the thin peridium. b, c. Eight-spored asci released from the<br />
ascomata. Note the apical apparatus (arrowed). d. Ascospores with<br />
thin sheath. e. An old pale brown ascospore. Scale bars: a-c=50 μm,<br />
d, e=10 μm<br />
(Allewia eureka (E.G. Simmons) E.G. Simmons=L.<br />
eureka) form a robust clade with other members of<br />
Pleosporaceae (Schoch et al. 2006; Schochetal.2009;<br />
Zhang et al. 2009a).<br />
Concluding remarks<br />
Its position in Pleosporaceae is confirmed.<br />
Lichenopyrenis Calat., Sanz & Aptroot, Mycol. Res. 105:<br />
634 (2001). (?Pleomassariaceae)<br />
Generic description<br />
Habitat terrestrial, parasitic on lichens. Ascomata mediumsized,<br />
globose or subglobose. Hamathecium of dense,<br />
filliform, branching, septate pseudoparaphyses. Asci bitunicate,<br />
fissitunicate, clavate, with a short sometimes furcate<br />
pedicel. Ascospores ellipsoidal with broadly rounded ends,<br />
pale orange-brown, 1-distoseptate.<br />
Anamorphs reported for genus: see below.<br />
Literature: Calatayud et al. 2001.
Fungal Diversity<br />
Fig. 46 Lewia scrophulariae<br />
(from FH, slide from lectotype).<br />
a Cylindrical ascus with a<br />
short pedicel. b Ascospores in<br />
asci. c–f Released muriform<br />
brown ascospores. Scale bars:<br />
a=20 μm, b–f=10 μm<br />
Type species<br />
Lichenopyrenis galligena Calat., Sanz & Aptroot, Mycol.<br />
Res. 105: 636 (2001). (Fig. 47)<br />
Ascomata 140–260 μm high×140–250 μm diam., gregarious,<br />
initially immersed in galls, later becoming erumpent,<br />
globose or subglobose, black, roughened (Fig. 47a and b).<br />
Peridium 18–25 μm wide, composed of 2–5 layers of heavily<br />
pigmented cells of textura angularis to compressed, cells<br />
6–11 μm diam., cell wall 1–3 μm thick (Fig. 47c, d and e).<br />
Hamathecium of dense, long filamentous pseudoparaphyses,<br />
2.5–4 μm broad, branching, septate. Asci 65–85×<br />
15–20 μm (x ¼ 74 18mm, n=10), 8-spored, bitunicate,<br />
fissitunicate, broadly clavate, with a short, thick, some-
Fungal Diversity<br />
times furcate pedicel, up to 13 μm long, ocular chamber<br />
not observed (Fig. 47f, g, h and k). Ascospores 16–20×9–<br />
11 μm (x ¼ 18 10mm, n=10), biseriate, ellipsoidal, pale<br />
orange-brown, 1-distoseptate, with prominent swelling at<br />
the septum, containing refractive globules, smooth (Fig. 47i,<br />
jandl).<br />
Anamorph: The following description is from Calatayud et<br />
al. (2001).<br />
Conidiomata pycnidial, arising in galls together with the<br />
ascomata, immersed, ca. 100–200 μm diam.; wall dark<br />
brown throughout, composed of 2–5 layers of angular to<br />
laterally compressed cells; cells relatively large, ca. 8–<br />
16 μm diam. in superficial view. Conidiophores formed by<br />
1–3 cells, frequently branched and with the uppermost cells<br />
bearing 1–4 conidiogenous cells; cells±cylindrical, hyaline<br />
except at the base, which are sometimes pale brown, 7–15×<br />
3–4 μm. Conidiogenous cells tapered towards the apex, 14–<br />
18×3–4 μm. Conidia 5–7×1.5–2 μm. Vegetative hyphae<br />
hyaline.<br />
Material examined: SPAIN, Andalucía, Province, Jaén,<br />
Andújar, lichenicolous on Leptochidium albociliatum<br />
(Desm.) M. Choisy on acid volcanic rock, 19 Apr. 2000,<br />
V. Calatayud (MA-Lichen 12715, holotype).<br />
Notes<br />
Morphology<br />
Lichenopyrenis was formally established by Calatayud et<br />
al. (2001) based on its “perithecioid ascomata with peridium<br />
comprising compressed cells, fissitunicate and J- asci, wide<br />
hamathecium filaments, and 1-septate pale orange-brown<br />
ascospores with distoseptate thickenings at maturity”, andis<br />
monotypic with L. galligena. The genus was temporarily<br />
assigned to Pleomassariaceae. Lichenopyrenis galligena is a<br />
parasite of lichens, occurring in galls in the thallus of the host<br />
(Calatayud et al. 2001).<br />
Phylogenetic study<br />
None.<br />
Concluding remarks<br />
This is one of the few species that are parasitic on<br />
lichens. The most comparable species are Parapyrenis<br />
lichenicola Aptroot & Diederich and Lacrymospora parasitica<br />
Aptroot (both in Requienellaceae, Pyrenulales) as<br />
well as some species from Dacampiaceae. The peridium<br />
structure, cellular pseudoparaphyses, distoseptate and<br />
smooth, orange-brown ascospores as well as the anamorphic<br />
stage of Lichenopyrenis can easily distinguish from all<br />
of them (Calatayud et al. 2001).<br />
Lineolata Kohlm. & Volkm.-Kohlm., Mycol. Res. 94: 687<br />
(1990). (<strong>Pleosporales</strong>, genera incertae sedis)<br />
Generic description<br />
Habitat marine, saprobic (or perthophytic?). Ascomata<br />
medium-sized, gregarious, immersed to erumpent, obpyriform,<br />
ostiolate, papillate. Peridium thin, comprising two<br />
types of cells; outer cells thick stratum pseudostromatic,<br />
inner stratum thin, composed of a few layers of hyaline<br />
cells of textura angularis. Hamathecium of dense, long<br />
trabeculate pseudoparaphyses, embedded in mucilage,<br />
anastomosing and septate. Asci 8-spored, bitunicate, cylindrical,<br />
with short pedicels, with an ocular chamber.<br />
Ascospores uniseriate to partially overlapping, ellipsoidal,<br />
dark brown, 1-septate.<br />
Anamorphs reported for genus: none.<br />
Literature: Kohlmeyer and Kohlmeyer 1966; Kohlmeyer<br />
and Volkmann-Kohlmeyer 1990.<br />
Type species<br />
Lineolata rhizophorae (Kohlm. & E. Kohlm.) Kohlm. &<br />
Volkm.-Kohlm., Mycol. Res. 94: 688 (1990). (Fig. 48)<br />
≡ Didymosphaeria rhizophorae Kohlm. & E. Kohlm.<br />
Icones Fungorum Maris (Lehre) 4 & 5: tab. 62a (1967).<br />
Ascomata 300–490 μm high×200–360 μm diam.,<br />
gregarious, immersed to erumpent, obpyriform, ostiolate,<br />
papillate, subcarbonaceous to subcoriaceous, blackish<br />
brown (Fig. 48a). Peridium 37–45 μm thick, comprising<br />
two types of cells; outer cells thick stratum pseudostromatic,<br />
composed of irregular or roundish, dark brown cells,<br />
on the outside with a more or less recognizable hyphal<br />
structure, enclosing some decaying cells of the host, inner<br />
stratum thin, composed of four or five layers of hyaline,<br />
polygonal, elongate, thin-walled cells with large lumina,<br />
merging into the pseudoparaphyses. Hamathecium of<br />
dense, long trabeculate pseudoparaphyses, 1–1.5 μm broad,<br />
embedded in mucilage, anastomosing and septate. Asci<br />
150–175×14–17.5 μm, 8-spored, bitunicate, cylindrical,<br />
with short pedicels, with an ocular chamber (Fig. 48b).<br />
Ascospores 23–32(−33)×9–12 μm, uniseriate to partially<br />
overlapping, ellipsoid, dark brown, 1-septate, not or slightly<br />
constricted at the septum, striate by delicate costae that run<br />
parallel or in a slight angle to the longitudinal axis of the<br />
ascospore (Fig. 48c, d, e and f) (adapted from Kohlmeyer<br />
and Kohlmeyer 1979).<br />
Anamorph: none reported.<br />
Material examined: US, Florida, Middle Torch Key, on<br />
Rhizophora mangle, 21 Nov. 1965, J. Kohlmeyer (Herb. J.<br />
Kohlmeyer No. 2390b, isotype); Pirate Grove Key, on R.<br />
mangle, 5 Jan. 1964 (Herb. J. Kohlmeyer No. 1721 paratype);<br />
Florida, Virginia Key, on R. mangle, 1 Jan. 1964, leg.<br />
E. Kohlmeyer (Herb. J. Kohlmeyer No. 1751 paratype);<br />
Florida, Torch Key, on R. mangle, 20 Nov. 1965, leg. J.<br />
Kohlmeyer (Herb. J. Kohlmeyer No. 2423 paratype).
Fungal Diversity<br />
Fig. 47 Lichenopyrenis galligena (from MA-Lichen 12715, holotype).<br />
a, b Ascomata forming in the host tissues. c, d Sections of<br />
ascomata. e Section of a partial peridium. f–h, k Broadly clavate asci.<br />
Note the short rounded pedicel. i, j, l Ascospores. Note the small<br />
swellings at the septa. Scale bars: a, b=0.5 mm, c, d=100 μm, e=<br />
50 μm, f–h, k=20 μm, i, j, l=10 μm
Fungal Diversity<br />
Notes<br />
Morphology<br />
Lineolata was monotypified by L. rhizophorae, whichwas<br />
originally introduced by Kohlmeyer and Kohlmeyer (1966)asa<br />
species of Didymosphaeria (as D. rhizophorae).Basedonthe<br />
morphology of ascomata and asci, Barr (1990a) assignedit<br />
under Lojkania (as L. rhizophorae). Kohlmeyer and Volkmann-<br />
Kohlmeyer (1990) restudied this species and noticed that the<br />
absence of clypeus, almost superficial ascomata, coloured<br />
peridium, a hamathecium with gelatinous matrix, asci with<br />
apical ring-like structure and the ornamented ascospores are<br />
quite different from the modified concept of Didymosphaeria.<br />
Thus they introduced Lineolata to accommodate D. rhizophorae<br />
(Kohlmeyer and Volkmann-Kohlmeyer 1990).<br />
Phylogenetic study<br />
Three isolates of Lineolata rhizophorae from varied geographic<br />
localities were analyzed by Suetrong et al. (2009)andshowntobe<br />
related to Caryosporella rhizophorae in Dothideomycetidae and<br />
excluded from Pleosporomycetidae and <strong>Pleosporales</strong>.<br />
Concluding remarks<br />
Based on initial molecular work it is likely that this species<br />
does not belong to <strong>Pleosporales</strong> in spite of its dense<br />
pseudoparaphyses and other characters shared with the order.<br />
Fig. 48 Lineolata rhizophorae<br />
(from Herb. J. Kolmeyer No.<br />
2390b, isotype of Didymosphaeria<br />
rhizophorae). a Ascomata<br />
immersed in the host<br />
substrate with protruding papilla.<br />
b Ascospores within an ascus.<br />
Note the ascospore<br />
arrangement. c–f One-septate<br />
ascospores. Note the striate ornamentation<br />
in (c). Scale bars:<br />
a=100 μm, a, b=20 μm, c–f=<br />
10 μm
Fungal Diversity<br />
Loculohypoxylon M.E. Barr, Mycotaxon 3: 326 (1976).<br />
(Teichosporaceae)<br />
Generic description<br />
Habitat terrestrial, saprobic. Ascomata relatively small,<br />
gregarious, immersed to erumpent, globose or subglobose,<br />
forming under a clypeus, papillate, ostiolate.<br />
Peridium thin, a single layer comprising hyaline thinwalled<br />
cells of textura angularis or textura prismatica.<br />
Hamathecium of septate pseudoparaphyses. Asci (2–4-)8-<br />
spored, bitunicate, cylindrical to cylindro-clavate, with a<br />
short, furcate pedicel, and wide ocular chamber. Ascospores<br />
broadly elliptic to subglobose, often apiculate at<br />
both ends, pale to dark brown, aseptate, with a germ<br />
slit.<br />
Anamorphs reported for genus: none.<br />
Literature: von Arx and Müller 1975; Barr 1976.<br />
Type species<br />
Loculohypoxylon grandineum (Berk. & Rav.) Barr, Mycotaxon<br />
3: 326 (1976). (Fig. 49)<br />
≡ Diatrype grandinea Berk. & Rav., in Berkeley,<br />
Grevillea 4: 95 (1876).<br />
Ascomata 85–130 μm high×75–145 μm diam., gregarious,<br />
immersed to widely erumpent, globose or subglobose,<br />
under a reddish brown to black clypeus, papillate,<br />
ostiolate (Fig. 49a and b). Peridium 18–30 μm thick<br />
laterally, 1-layered, composed of hyaline thin-walled cells<br />
of textura angularis to prismatica, cells up to 5×9 μm<br />
diam., cell wall 0.5–1 μm thick, apex cells smaller and<br />
walls thicker (Fig. 49c). Hamathecium comprising 2–3 μm<br />
broad, septate pseudoparaphyses. Asci 70–90×10–<br />
12.5 μm (x ¼ 76:5 10:9mm, n=10), (2–4-)8-spored,<br />
bitunicate, cylindrical to cylindro-clavate, with a short,<br />
furcate pedicel, up to 25 μm long, with a wide ocular chamber<br />
(Fig. 49f, g, and h). Ascospores 7.5–10×5–7 μm<br />
(x ¼ 8:3 5:9mm, n=10), uniseriate to partially overlapping<br />
at the upper part, broadly elliptic to subglobose, often<br />
apiculate at both ends, pale to dark brown, aseptate, with a<br />
germ slit (Fig. 49d and e).<br />
Anamorph: none reported.<br />
Material examined: USA, New Jersey, Newfield, on<br />
bark of Quercus coccinea, Sept. 1878, as Diatrype<br />
grandinea, Ellis N.A.F. 494 (NY, MASS); on Quercus<br />
sp. wood, Nov. 1893, as Anthostoma grandinea B. & Rav.,<br />
Ellis & Everhart, N.A.F. 494 (NY); Newfield, Oct. 1881,<br />
as Diatrype grandinea (NY); Newfield, Jan. 1882, on<br />
Quercus coccinea, asDiatrype grandinea B. & Rav, Ex<br />
Herb Ellis (NY); Newfield, Nov. 1893, as Anthostoma<br />
grandinea, on bark of fallen trunks of Quercus coccinea<br />
(NY).<br />
Notes<br />
Morphology<br />
Loculohypoxylon grandineum is one of the rare pleosporalean<br />
species having aseptate ascospores. When emphasis<br />
is given to ascospore morphology, Semidelitschia<br />
(monotypified by S. agasmatica Cain & Luck-Allen) is the<br />
most comparable genus. The large ascomata and ascospores,<br />
the mucilaginous sheath surrounding the ascospores<br />
as well as the coprophilous habitat of S. agasmatica differ<br />
from L. grandineum greatly. Thus Loculohypoxylon was<br />
introduced as a new genus.<br />
Phylogenetic study<br />
None.<br />
Concluding remarks<br />
Aseptate ascospores are rare in <strong>Pleosporales</strong>, and the<br />
position of this fungus needs further verification. The familial<br />
status of Loculohypoxylon in Teichosporaceae is questionable,<br />
as it is simply based on the similarity of living habitat, ascomata<br />
and asci with Immotthia and Teichospora (Barr 2002).<br />
Lophionema Sacc., Syll. fung. (Abellini) 2: 717 (1883).<br />
(<strong>Pleosporales</strong>, genera incertae sedis)<br />
Generic description<br />
Habitat terrestrial, saprobic? Ascomata solitary, scattered or<br />
in small groups, immersed to erumpent, globose to subglobose,<br />
with a flattened base, wall black, papillate, ostiolate.<br />
Peridium comprising two types of cells which merge in the<br />
middle. Hamathecium of trabeculate pseudoparaphyses,<br />
septate, rarely anastomosing and branching. Asci 8-spored,<br />
bitunicate, fissitunicate unknown, clavate to cylindro-clavate,<br />
with a short and furcate pedicel and a small inconspicuous<br />
ocular chamber. Ascospores filliform, hyaline to pale yellow,<br />
multi-septate, slightly constricted at each septum.<br />
Anamorphs reported for genus: none.<br />
Literature: Barr1992b; ChestersandBell1970; Ellis and<br />
Everhart 1892; Höhnel 1909; Solheim 1949.<br />
Type species<br />
Lophionema vermisporum (Ellis) Sacc., Syll. fung. (Abellini)<br />
2: 717 (1883). (Fig. 50)<br />
≡ Lophiostoma vermispora Ellis, Bull. Torrey bot. Club<br />
9: 19 (1882).<br />
Ascomata 320–430 μm high×280–350 μm diam.,<br />
solitary, scattered or in small groups of 2–3, immersed to<br />
erumpent, globose to subglobose, black, papillate, ostiolate.<br />
Papilla 80–120 μm high, up to 150 μm broad, cylindrical to<br />
somewhat vertically flattened neck; mostly with a short
Fungal Diversity<br />
Fig. 49 Loculohypoxylon grandineum (from N.A.F. 494). a Appearance<br />
of ascomata on the host surface. b Habitat section of ascomata. c<br />
Section of an ascoma. Note the pale brown thin-walled peridium cells.<br />
slot-like ostiole, periphysate (Fig. 50a). Peridium 30–45 μm<br />
wide at the sides and slightly thicker at the apex,<br />
d, e Uniseriate ascospores in asci. f–f Cylindro-clavate asci with<br />
ascospores. Note the ocular chamber in (g). Scale bars: a=100 μm,<br />
b=200 μm, c=50 μm, d–h=10 μm<br />
2-layered, lateral walls and wall adjacent to neck comprising<br />
two types of cells which merge in the middle; outer
Fungal Diversity<br />
cells small heavily pigmented thick-walled cells of textura<br />
angularis, cells 4–7 μm diam., cell wall 3.5–5 μm thick,<br />
inner cells less pigmented, comprising thin-walled compressed<br />
cells; apical wall cells smaller and walls thicker,<br />
basal wall thinner (ca. 15μm wide), composed of lightly<br />
pigmented thin-walled compressed cells (Fig. 50b and c).<br />
Hamathecium of trabeculate pseudoparaphyses, 1–2 μm<br />
broad, septate, anastomosing and branching rarely between<br />
and mostly above the asci. Asci 105–130(−150)×10–15 μm<br />
(x ¼ 123 12mm, n=10), 8-spored, bitunicate, fissitunicate<br />
dehiscence not observed, clavate to cylindro-clavate, with a<br />
short, narrow, furcate pedicel which is 10–25 μm long, and<br />
a small inconspicuous ocular chamber (to 1.5 μm wide×<br />
1 μm high) (Fig. 50d and e). Ascospores (80-)90–115×3–<br />
5 μm (x ¼ 95 3:5mm, n=10), filliform, gradually tapering<br />
towards the base, hyaline to light yellow, (6-)7(−8)-septate,<br />
slightly constricted at each septum, smooth (Fig. 50f).<br />
Anamorph: none reported.<br />
Material examined: USA, New Jersey, Newfield, on<br />
dead stems of Oenothera biennis, Aug. 1881, Ellis (NY<br />
643, holotype, NY 885, isotype).<br />
Notes<br />
Morphology<br />
Lophionema is a relatively poorly studied genus, which<br />
was formally established by Saccardo (1883) as a monotypic<br />
genus represented by L. vermisporum based on its “globose<br />
ascomata, compressed ostiole, cylindrical to clavate ascus,<br />
and filamentous, septate, subhyaline to lightly pigmented<br />
ascospores”. Lophionema vermisporum was consequently<br />
listed as the generic type (Clements and Shear 1931). Berlese<br />
(1890) placed the genus in Lophiostomataceae but mentioned<br />
that the genus was similar to Ophiobolus according to<br />
the variable apex, and Shoemaker (1976) transferred Lophionema<br />
vermisporum to Ophiobolus sensu lato. Chesters and<br />
Bell (1970) however, had regarded Lophionema as related to<br />
Lophiostoma despite the distinct ascospore morphology. Barr<br />
(1992b) assignedLophionema to Entodesmium based on the<br />
morphology of ascomata, papilla, peridium structure, pseudoparaphyses<br />
as well as the hyaline or slightly yellowish<br />
ascospores with a terminal appendage (not observed here).<br />
Species of Entodesmium, however, exclusively occur on<br />
legumes, but Lophionema vermisporum does not. We also<br />
note that the filliform ascospores, bitunicate asci, pseudoparaphyses<br />
and nature of the peridium may also be considered<br />
as typical of genera in the Tubeufiaceae (Barr 1980;<br />
Kodsueb et al. 2006b).<br />
Phylogenetic study<br />
None.<br />
Concluding remarks<br />
The immersed to erumpent ascomata, trabeculate pseudoparaphyses<br />
and laterally flattened papilla and periphysate<br />
ostioles indicate that this genus should be included in<br />
Lophiostomataceae. We do not accept the above proposals<br />
and, consider that Lophionema should be maintained as a<br />
separate genus with filliform ascospores in Lophiostomataceae<br />
until representative taxa can be sequenced and<br />
analyzed. Currently Lophionema comprises 10 species<br />
(http://www.mycobank.org, 08-01-2009). However, many<br />
of these are poorly studied and obscure.<br />
Lophiostoma Ces. & De Not., Comm. Soc. crittog. Ital. 1:<br />
219 (1863). (Lophiostomataceae)<br />
Generic description<br />
Habitat terrestrial, saprobic. Ascomata immersed to erumpent,<br />
usually with a distinct depressed papilla and a slotlike<br />
ostiole. Hamathecium of dense, long, septate pseudoparaphyses,<br />
embedded in mucilage, anastomosing and<br />
branching between and above the asci. Peridium unequal<br />
in thickness, thicker near the apex and thinner at base. Asci<br />
usually clavate. Ascospores 1-septate, multi-septate or even<br />
muriform, hyaline to deep brown, usually with terminal<br />
appendages.<br />
Anamorphs reported for genus: Pleuorphomopsis-like<br />
(Hyde et al. 2011).<br />
Literature: Barr 1990a; Chesters and Bell 1970; Holm and<br />
Holm 1988; Hyde and Aptroot 1998; Hyde et al. 2002;<br />
Tanaka and Harada 2003b; Yuan and Zhao 1994.<br />
Type species<br />
Lophiostoma macrostomum (Tode) Ces. & De Not.,<br />
Comm. Soc. crittog. Ital. 1: 219 (1863). (Fig. 51)<br />
≡ Sphaeria macrostoma Tode, Fung. mecklenb. sel.<br />
(Lüneburg) 2: 12 (1791).<br />
Ascomata 400–600 μm high×420–560 μm diam.,<br />
densely scattered to gregarious, semi-immersed to erumpent,<br />
globose or subglobose, with a small to large<br />
flattenedcrest-likeraisedareaabovetheascomatawhich<br />
is variable in shape, up to 300 μm high and 480 μm<br />
wide, with a slit-like ostiole along the full length of the<br />
crest (Fig. 51a and b). Peridium 30–45 μm thick at the<br />
sides, thicker at the apex and thinner at the base,<br />
composed of one cell type of small lightly pigmented<br />
thin-walled cells of textura prismatica, cellsca. 6–9×3–<br />
4 μm diam., apex composed of pseudoparenchymatous<br />
cells (Fig. 51b). Hamathecium of dense, filliform, up to<br />
3 μm near the base and less than 1.5 μm broad in the
Fungal Diversity<br />
Fig. 50 Lophionema vermisporum (from NY-643, holotype). a<br />
Appearance of ascomata on the host surface. Note the form of the<br />
neck. b Section of the peridium. c Peridium comprising two types of<br />
cells which merge in the middle; outer cells small heavily pigmented<br />
thick-walled cells of textura angularis, inner cells less pigmented, and<br />
comprising thin-walled compressed cells. d, e Cylindro-clavate, 8-<br />
spored asci. f A 7-septate filliform ascospore. Scale bars: a=0.5 mm,<br />
b=100 μm, c=50 μm, d–f=10 μm
Fungal Diversity<br />
upper place, septate pseudoparaphyses, embedded in<br />
mucilage, anastomosing and branching between and<br />
above the asci (Fig. 51f). Asci 110–145×10–15 μm<br />
(x ¼ 127:5 13mm, n=10), 8-spored, bitunicate, fissitunicate<br />
(ectotunica no constriction), cylindro-clavate, with a<br />
furcate pedicel and a small ocular chamber (to 1.5 μm wide×<br />
2 μm high) (J-) (Fig. 51c, d and e). Ascospores 27–38(−43)×<br />
5–7.5 μm (x ¼ 31:2 6:4mm, n=10), biseriate, fusoid,<br />
curved, hyaline, usually 1-septate, with 3–5 septaandfaintly<br />
brown when old, with (2-)3(−4) distinct oil drops in each cell<br />
and short terminal appendage at ends (Fig. 51h, i and j), and<br />
ornamented with warts when spores are senescent (Fig. 51g).<br />
Anamorph: none reported.<br />
Material examined: SWEDEN, Smaland, Femsjö par.,<br />
Femsjö, on Prunus, 2006, Elias Fries, det. Geir Mathiassen<br />
(UPS, lectotype, asSphaeria macrostoma Fr.). FRANCE,<br />
Ariège, Rimont, Las Muros, on dead stems of Vitis vinifera,<br />
2 Sept. 1996 (IFRD2005).<br />
Notes<br />
Morphology<br />
Lophiostoma is morphologically a well studied genus<br />
(Barr 1990a; Chesters and Bell 1970; Holm and Holm<br />
1988; Mugambi and Huhndorf 2009b; Yuan and Zhao<br />
1994), and currently it comprises about 30 species (Tanaka<br />
and Harada 2003b). The genus was characterized as<br />
having immersed to erumpent ascomata with a cylindrical<br />
or crest-like papilla and full length, slit-like ostiole; a<br />
peridium of unequal thickness, which was broader near the<br />
base (Lophiostoma-type); mostly clavate, bitunicate asci<br />
and 1- to several septate, hyaline to pigmented ascospores<br />
with terminal appendages or surrounded by a mucilaginous<br />
sheath (Holm and Holm 1988). This definition was<br />
followed by Barr (1990a), Yuan and Zhao (1994) and<br />
Hyde et al. (2002).<br />
The crest-like papilla has been regarded as a prominent<br />
morphological character of Lophiostoma macrostomum<br />
(Chesters and Bell 1970; Holm and Holm 1988). In the<br />
lectotype specimen, the raised area above the ascomata is<br />
up to 300 μm high and 480 μm long, and seen as a<br />
flattened or even Y-shaped crest (Fig. 51a). In Lophiostoma<br />
curtum (Fr.) De Not. and Lophiotrema boreale Math. the<br />
raised area above the ascomata varies considerably in<br />
height or is even lacking (Holm and Holm 1988). Thus<br />
the variable “crest-like raised area in Lophiostomataceae”<br />
was explained as an evolutionarily adaptation to the hard<br />
substrate within which the ascomata develop (Holm and<br />
Holm 1988). The ascospores of L. macrostomum usually<br />
turn reddish brown when mature, and minutely verrucose<br />
ornamentation was also found on the surface of the<br />
pigmented ascospores. Hyaline ascospores that became<br />
pigmented with age are common in Lophiostoma, such as<br />
in L. appendiculatum Fuckel, L. massarioides (Sacc.) L.<br />
Holm & K. Holm, L. semiliberum, L. subcorticale Fuckel<br />
and L. winteri (Holm and Holm 1988; Tanaka and Harada<br />
2003b). The phylogenetic significance of this character<br />
should be observed carefully in the future but at present its<br />
phylogenetic significance is unclear as this also occurs in<br />
some Lophiotrema species.<br />
Phylogenetic study<br />
Phylogenetic affinity with some Massarina species has<br />
been reported by Liew et al. (2002), and several Massarina<br />
species were transferred into Lophiostoma. Inasystematic<br />
study of Lophiostoma- and Massarina-related fungi conducted<br />
by Zhang et al. (2009b), Lophiostoma taxa clustered<br />
into two groups; one includes the type species L. macrostomum<br />
with crest-like ostioles, L. rugulosum Yin. Zhang, J.<br />
Fourn. & K.D. Hyde with a wide, umbilicate pore surrounded<br />
by 4–6 radial ridges, and L. glabro-tunicatus with small<br />
ostiolar pores; the other cluster comprises Lophiostoma-like<br />
taxa with slot-like ostioles lacking raised crests, which<br />
includes L. arundinis (Pers.) Ces. & De Not., L. caulium, L.<br />
compressum (Pers.) Ces. & De Not., L. crenatum (Pers.)<br />
Fuckel, L. fuckelii (Sacc.) Sacc., L. macrostomoides, L.<br />
semiliberum and L. viridarium Cooke, which seems to<br />
represent a natural group at the family level. This conclusion<br />
is tentative until verified sequences of L. macrostomum are<br />
included in analyses (see comments of Zhang et al. 2009a).<br />
Concluding remarks<br />
We tend to accept a narrow concept of Lophiostomataceae,<br />
which only comprises species of Lophiostoma sensu<br />
stricto (Zhang et al. 2009a).<br />
Lophiotrema Sacc., Michelia 1: 338 (1878). (<strong>Pleosporales</strong>,<br />
genera incertae sedis)<br />
Generic description<br />
Habitat terrestrial, saprobic. Ascomata small- to mediumsized,<br />
with or without short papilla. Hamathecium of dense,<br />
long, septate pseudoparaphyses, anastomosing and branching<br />
between and above asci. Asci cylindrical to cylindroclavate.<br />
Ascospores hyaline, 1–3-septate, usually with<br />
mucilaginous sheath.<br />
Anamorphs reported for genus: none.<br />
Literature: Barr 1990a; Chesters and Bell 1970; Holm and<br />
Holm 1988; Saccardo 1878a; Tanaka and Harada 2003c;<br />
Tang et al. 2003; Yuan and Zhao 1994.<br />
Type species<br />
Lophiotrema nucula (Fr.) Sacc., Michelia 1: 338 (1878).<br />
(Fig. 52)
Fungal Diversity<br />
≡ Sphaeria nucula Fr., Syst. mycol. (Lundae) 2: 466 (1823).<br />
Ascomata 200–240 μm high×200–280 μm diam., scattered,<br />
erumpent to nearly superficial, with basal wall remaining<br />
immersed in host tissue, globose to subglobose, often<br />
laterally flattened, with a flattened base not easily removed<br />
from the substrate, black, roughened; with a cylindrical or<br />
slightly compressed papilla. Papilla to 120 μm long and<br />
150 μm high, protruding, with a pore-like ostiole (Fig. 52a).<br />
Peridium 25–30 μm wide, very thin at the base, composed of<br />
heavily pigmented pseudoparenchymatous cells near the apex,<br />
cells 2–2×6 μm diam., wall 1–3(−4) μm thick, lower sides<br />
composed of pigmented cells of textura angularis, 3–5 μm<br />
diam., wall 0.8–1.5 μm thick, ostiole wall composed of<br />
heavily pigmented and thick-walled small cells (Fig. 52b and<br />
c). Hamathecium of dense, long, septate pseudoparaphyses,<br />
1–2 μm broad, anastomosing and branching between and<br />
above asci, embedded in mucilage (Fig. 52i). Asci 90–115×<br />
9–11.5 μm (x ¼ 99:5 11:5mm; n=10), 8-spored, bitunicate,<br />
fissitunicate, cylindrical, with a short, narrowed, furcate<br />
pedicel which is up to 10 μm long, with a small ocular<br />
chamber (ca. 1.5μm wide×1 μm high) (J-) (Fig. 52d, e, f<br />
and h). Ascospores 17–21(−25)×(4-)5–6.5 μm<br />
(x ¼ 19:5 5:5mm, n=10), obliquely uniseriate and partially<br />
overlapping to biseriate, broadly fusoid to fusoid, with<br />
narrowly rounded ends, hyaline and lightly pigmented on<br />
very rare occasions when senescent, 1-septate, 3-septate<br />
when old, constricted at the median septum, the upper cell<br />
often broader than the lower one (Fig. 52g).<br />
Anamorph: none reported.<br />
Material examined: on decaying wood (UPS, lectotype<br />
as Sphaeria nucula Fr.).<br />
Notes<br />
Morphology<br />
Holm and Holm (1988) provided a relatively strict<br />
definition for Lophiotrema after they examined several specimens<br />
including the type materialwhichtheylectotypified.<br />
Lophiotrema was mainly defined on the unique characters of<br />
small to medium ascomata, a “Lophiotrema-type” peridium<br />
and 1-septate ascospores. In Lophiotrema, Holm and Holm<br />
(1988) considered the ascomata to be small- to medium-sized,<br />
ca. pyriform but neck often reduced, even lacking and<br />
sometimes cylindrical. The peridium was of approximately<br />
equal thickness, 20–30 μm, composed of an outer textura<br />
angularis of uniformly pigmented cells, up to 12 μm, andan<br />
inner layer of very small hyaline cells, with somewhat<br />
thickened walls. Asci are cylindrical, spores hyaline, at first<br />
1-septate, becoming 3-septate, with distinct guttules, often<br />
with a mucilaginous sheath. Much emphasis was given to the<br />
1-septate ascospores by Holm and Holm (1988) when they<br />
described and distinguished the three Lophiotrema species: L.<br />
boreale, L. nucula, L. vagabundum (Sacc.) Sacc. and two<br />
Fig. 51 Lophiostoma macrostomum (a–h, j from UPS, leptotype; i b<br />
from IFRD 2005). a Appearance of ascomata on the host surface.<br />
Note the raised crest-like areas and full length germ slits. b Section of<br />
the peridium. c–e Cylindro-clavate asci with ascospores arranged in a<br />
2-3-seriate manner. f Hamathecium comprising branching and septate<br />
pseudoparaphyses. g–j Released or unreleased ascospores. Note the<br />
smooth young ascospores with terminal sheath, and the verrucose<br />
senescent ascospores. Scale bars: a=0.5 mm, b=200 μm, c–j=10 μm<br />
other unnamed species. This concept was widely accepted by<br />
later workers (Kirk et al. 2001; YuanandZhao1994). Tanaka<br />
and Harada (2003c) considered the peridium and asci to<br />
distinguish Lophiotrema from Lophiostoma, while Tang et al.<br />
(2003) introduced a new Lophiotrema species with elongated<br />
slit-like ostiole stating that the main difference between<br />
Lophiotrema and Lophiostoma were size of ascomata,<br />
structure of peridium, shape of asci and sheath of ascospores.<br />
This peridium concept however, is not supported by the<br />
lectotype specimen we examined here, which has a flattened<br />
thin-walled base. Thus the “Lophiotrema-like peridium” sensu<br />
Holm and Holm (1988) should not serve as a diagnostic<br />
character of Lophiotrema, while the ostiole, asci and<br />
ascospores might have some phylogenetic significance (Zhang<br />
et al. 2009b). No anamorph is yet known for Lophiotrema.<br />
Although the ascospores was reported by Holm and Holm<br />
(1988) to be verruculose this could not be observed in the<br />
lectotype examined under light microscope (1000 ×) in the<br />
present study.<br />
Phylogenetic study<br />
In the phylogenetic study of Lophiostoma, Massarina and<br />
related genera (Zhang et al. 2009b), Lophiotrema nucula<br />
formed a consistent and robust clade with three other<br />
Lophiotrema species: L. lignicola Yin. Zhang, J. Fourn. &<br />
K.D. Hyde, L. brunneosporum Yin. Zhang, J. Fourn. & K.D.<br />
Hyde and L. vagabundum, separate from other members of<br />
Lophiostoma and Massarina sensu stricto. This clade might<br />
represent Lophiotrema sensu stricto, however, the correctness<br />
of strains of L. vagabundum (<strong>CBS</strong> 628.86) and L.<br />
nucula (<strong>CBS</strong> 627.86) used in the phylogenetic study are not<br />
verified and warrant further study.<br />
Concluding remarks<br />
Holm and Holm (1988) distinguished Lophiostoma from<br />
Lophiotrema based on the smaller ascomata, 1-septate<br />
versus multi-septate ascospores, and peridial wall structure.<br />
However, we doubt that these distinguishing characters<br />
(size of ascomata, number of septa of ascospores) can be<br />
confidently used to separate these genera and we could not<br />
establish any characters that could reliably distinguish<br />
between these two genera. The molecular data, however,<br />
does separate Lophiostoma macrostomum and Lophiotrema<br />
nucula into separate clades and provides some support that<br />
these are separate genera. Although the strain of L. nucula
Fungal Diversity
Fungal Diversity<br />
Fig. 52 Lophiotrema nucula (from UPS, lectotype). a Ascomata on<br />
the host surface. b Section of a partial ascoma. c Peridium structure<br />
near the apex. d, h Cylindrical asci in the pseudoparaphyses. e, f<br />
Upper part of the asci, showing the small ocular chamber near the<br />
apex. h Mature ascospores. i Pseudoparaphyses. Scale bars: a=<br />
0.5 mm, b=100 μm, c, d=30 μm, e–i=10 μm
Fungal Diversity<br />
(<strong>CBS</strong> 627.86) was isolated by K. & L. Holm, who had<br />
examined the type specimen of L. nucula (Holm and Holm<br />
1988), the culture of Lophiostoma macrostomum used in<br />
the analysis are unverified (see comment by Zhang et al.<br />
2009b). For the purpose of this monograph we tentatively<br />
maintain Lophiotrema as distinct from Lophiostoma.<br />
Macroventuria Aa, Persoonia 6: 359 (1971).<br />
(Didymellaceae)<br />
Generic description<br />
Habitat terrestrial, saprobic. Ascomata small, solitary, scattered,<br />
or in groups, initially immersed, becoming erumpent, to nearly<br />
superficial, globose to subglobose, roughened with cylindrical<br />
setae erect from apex. Peridium thin, membranous. Hamathecium<br />
of cellular pseudoparaphyses, seems to easily disappear<br />
when mature. Asci bitunicate, somewhat obclavate to fusoid.<br />
Ascospores fusoid with broadly to narrowly rounded ends,<br />
hyaline, 1-septate, constricted at the septum.<br />
Anamorphs reported for genus: none.<br />
Literature: van der Aa 1971; von Arx and Müller 1975;<br />
Barr 1987a.<br />
Type species<br />
Macroventuria wentii Aa, Persoonia 6: 361 (1971). (Fig. 53)<br />
Ascomata 135–180 μm diam., rarely more than 200 μm<br />
diam., solitary, scattered or in groups, initially immersed,<br />
becoming erumpent, to nearly superficial, with basal wall<br />
remaining immersed in host tissue, globose to subglobose,<br />
broadly or narrowly conical, setae erect from the apical region of<br />
the ascomata, cylindrical or tapering to the rounded or pointed<br />
tip, brown, up to 90 μm long, 5–7.5 μm thick (Fig. 53a).<br />
Peridium, 25–35 μm thick, 2-layered, out layer composed of<br />
relatively thick-walled cells of textura angularis, cell wall up to<br />
3 μm thick; inner layer cells with a thinner wall and subhyaline;<br />
near apex cells smaller (Fig. 53a). Hamathecium of cellular<br />
pseudoparaphyses, 1–2 μm thick, evanescing not sure. Asci 75–<br />
93×24–30 μm, 8-spored, without pedicel, bitunicate, somewhat<br />
obclavate to fusoid (Fig. 53b). Ascospores 22–32×8–14 μm, 1–<br />
3 seriate, fusoid with broadly to narrowly rounded ends, hyaline,<br />
1-septate, constricted at the septum, smooth (Fig. 53b) (description<br />
adapted from van der Aa 1971).<br />
Anamorph: none reported.<br />
Material referred: USA, Nevada; Death Valley, plant<br />
litter, F.W. Went, 229, 1970 (<strong>CBS</strong> 526.71, holotype).<br />
Notes<br />
Morphology<br />
Macroventuria was formally established by van der Aa<br />
(1971) represented by M. anomochaeta and M. wentii based<br />
on its “near-hyaline, 1-septate ascospores, setose ascomata,<br />
and saprobic life style”. Almost all of the above characters<br />
(except the saprobic life style) point this group of fungi to<br />
Venturiaceae. Thus Macroventuria was assigned to this<br />
family as a relatively primitive genus (van der Aa 1971).<br />
Subsequently, von Arx and Müller (1975) assigned Macroventuria<br />
to Pseudosphaeriaceae (Dothideales), and this<br />
proposal was followed by Barr (1987a).<br />
Phylogenetic study<br />
Phylogenetic analysis based on combined SSU rDNA<br />
and LSU rDNA sequences indicated that both of Macroventuria<br />
anomochaeta and M. wentii form a robust clade<br />
with Leptosphaerulina argentinensis (Speg.) J.H. Graham<br />
& Luttr., L. australis, L. trifolii (Rostr.) Petr. and<br />
Platychora ulmi, which appear to share phylogenetic<br />
affinities with the Leptosphaeriaceae and Phaeosphaeriaceae,<br />
but detached from other members of Venturiaceae<br />
and Pleosporaceae (Kodsueb et al. 2006a). In addition,<br />
culture characters also support the close relationship<br />
between Macroventuria and Leptosphaerulina (Barr<br />
1987a). Analysis based on five genes, i.e. SSU, LSU,<br />
RPB1, RPB2 andTEF1, indicated Macroventuria anomochaeta<br />
resides in the well supported clade of Didymellaceae<br />
(Zhang et al. 2009a).<br />
Concluding remarks<br />
The morphological characters, such as small ascomata<br />
and hyaline, 1-septate ascospores all point at Didymellaceae,<br />
thus the familial status of Macroventuria is verified.<br />
Mamillisphaeria K.D. Hyde, S.W. Wong & E.B.G. Jones,<br />
Nova Hedwigia 62: 514 (1996b). (?Melanommataceae)<br />
Generic description<br />
Habitat freshwater, saprobic. Ascomata superficial, scattered<br />
or gregarious, conical, carbonaceous, papillate.<br />
Hamathecium of dense, filliform, trabeculate pseudoparaphyses.<br />
Asci broadly clavate to clavate, with small ocular<br />
chambers and short pedicels. Ascospores of two types, (1):<br />
2-4-seriate, ellipsoid, hyaline, slightly constricted at the<br />
main septum; with apical appendages at each end and<br />
around the ascospore; (2) 1-2-seriate, ellipsoid to fusoid,<br />
brown, with mucilaginous sheath around the ascospore<br />
(Hyde et al. 1996b).<br />
Anamorphs reported for genus: none.<br />
Literature: Hyde et al. 1996a, b.<br />
Type species<br />
Mamillisphaeria dimorphospora K.D. Hyde, S.W. Wong<br />
& E.B.G. Jones, Nova Hedwigia 62: 515 (1996b). (Fig. 54)
Fungal Diversity<br />
Fig. 53 Macroventuria wenti.<br />
a Ascomata. Note the setae. b<br />
Ascus and ascospores. Scale<br />
bars: a=50 μm, b=10 μm<br />
(figures referred to van der Aa<br />
1971)<br />
Following description is adapted from Hyde et al.<br />
1996a, b).<br />
Ascomata 455–650 μm high×980–1430 μm diam.,<br />
scattered or in small groups, superficial, conical, carbonaceous,<br />
papillate, under pseudostroma which forms a thin<br />
layer on the host surface, up to 50 μm thick between the<br />
ascomata and 125–250 μm thick on the ascomata surface<br />
(Fig. 54a and b). Peridium 10–25 μm thick, comprising<br />
several layers of compressed, densely packed, thin-walled,<br />
hyaline cells. A wedge-shaped area of vertically orientated<br />
hyaline palisade-like cells occurs at the periphery<br />
(Fig. 54b). Hamathecium of dense, trabeculate pseudoparaphyses,<br />
ca. 1μm broad, hyaline, branching and anastomosing,<br />
septate, embedded in mucilage (Fig. 54e). Two<br />
types of asci and ascospores exist in the same ascoma:<br />
TYPE 1: asci 185–320×40–100 μm (x ¼ 210 78mm, n=<br />
50), 8-spored, cylindro-clavate, bitunicate, fissitunicate,<br />
short-pedicellate, with an ocular chamber (to 13 μm<br />
wide×5 μm high) (Fig. 54c and d). Ascospores 66–84×<br />
20–38 μm (x ¼ 78 25mm, n=50), 2-4-seriate, hyaline,<br />
ellipsoidal, constricted at the central septum, with pad-like<br />
mucilaginous appendage at each end and with some<br />
mucilage associated around the spore, and TYPE 2: asci<br />
158–242×8–15 μm (x ¼ 182 11mm, n=50), 8-spored,<br />
cylindrical, bitunicate, fissitunicate, pedicellate, with an<br />
ocular chamber and faint apical ring, ascospores 29–42×<br />
6–9 μm (x ¼ 35 7mm, n =50), 1-2-seriate, brown,<br />
ellipsoidal-fusoid, surrounded by a thin mucilaginous<br />
sheath (Fig. 54f, g, h, i and j).<br />
Anamorph: none reported.<br />
Material examined: BRUNEI, on submerged wood,<br />
Aug. 1997, leg. K.D. Hyde (HKU(M) 7425).<br />
Notes<br />
Morphology<br />
Mamillisphaeria was established as a monotypic genus<br />
according to its bitunicate, fissitunicate asci, trabeculate<br />
pseudoparaphyses and dimorphic ascospores, which is<br />
typified by the widely distributed freshwater fungus, M.<br />
dimorphospora (Hyde et al. 1996a, b). The most striking<br />
character of this fungus is its dimorphic ascospores, i.e.<br />
one type is large and hyaline, and the other is comparatively<br />
smaller and brown. Only a few ascomycetes have<br />
been reported having dimorphic ascospores, such as<br />
Aquasphaeria dimorphospora and Nectria heterospora<br />
Speg. (Hyde 1995b; Spegazzini 1889). Dimorphic ascospores<br />
appear to have evolutionary benefits, for example<br />
the large ascospores with mucilaginous sheaths may<br />
facilitate nutrient storage for germination and enhanced<br />
collision and attachment to substrates. The smaller brown<br />
ascospores may help resist desiccation and damage by UV<br />
light and contribute to aerial dispersal, which might<br />
explain the worldwide distribution of M. dimorphospora<br />
(Hyde et al. 1996a, b).<br />
Phylogenetic study<br />
None.<br />
Concluding remarks<br />
Although in the key by Barr (1990a), M. dimorphospora<br />
can be referred to Massariaceae, itistemporarily<br />
assigned to Melanommataceae here based on its<br />
trabeculate pseudoparaphyses embedded in mucilage<br />
(Hyde et al. 1996a, b).
Fungal Diversity<br />
Fig. 54 Mamillisphaeria dimorphospora (from HKU(M) 7425,<br />
paratype?). a Ascomata scattered on the host surface. Note the small<br />
papilla. b Section of an ascoma. c, d Asci (TYPE 1). e Trabeculate<br />
pseudoparaphyses in a gelatinous matrix. f–j Ascospores (TYPE 1).<br />
Scale bars: a=0.5 mm, b–d=100 μm, e=10 μm, f–j=20 μm
Fungal Diversity<br />
Massarina Sacc., Syll. fung. (Abellini) 2: 153 (1883).<br />
emend. (Massarinaceae)<br />
Generic description<br />
Habitat terrestrial, saprobic. Ascomata immersed or superficial,<br />
scattered or clustered, globose, conical globose to<br />
lenticular, papillate or epapillate, ostiolate. Hamathecium of<br />
dense, cellular pseudoparaphyses. Asci clavate to cylindrical,<br />
with short pedicels. Ascospores ellipsoid to fusoid,<br />
hyaline, 1- to 3-septate, with or without mucilaginous<br />
sheath.<br />
Anamorphs reported for genus: Ceratophoma (Sivanesan<br />
1984).<br />
Literature: Aptroot 1998; Barr 1990a; Bose 1961; Eriksson<br />
and Yue 1986; Hyde 1995a; Hyde and Aptroot 1998;<br />
Liew et al. 2002; Saccardo 1883; Sivanesan 1984; Tanaka<br />
and Harada 2003d; Zhang et al. 2009a, b.<br />
Type species<br />
Massarina eburnea (Tul. & C. Tul.) Sacc., Syll. fung.<br />
(Abellini) 2: 153 (1883). (Fig. 55)<br />
≡ Massaria eburnea Tul. & C. Tul., Sel. Fung. Carp. 2:<br />
239 (1863).<br />
Ascomata to 250 μm high×500–700 μm diam., solitary or<br />
in small clusters, forming under raised dome-shaped areas,<br />
with blackened centres, with a central ostiole, immersed<br />
within the cortex of thin dead branches, ellipsoidal,<br />
rounded from above, clypeate, neck central, short and<br />
barely noticeable on host surface (Fig. 55a). Clypeus ca.<br />
250 μm diam., 60 μm thick, brown, comprising compact<br />
brown-walled cells of textura angularis to globulosa<br />
beneath host epidermal cells (Fig. 55b). Peridium ca.<br />
20 μm thick comprising 3–5 layers of hyaline compressed<br />
cells, fusing at the outside with the host (Fig. 55e).<br />
Hamathecium filamentous, cellular pseudoparaphyses, ca.<br />
2 μm broad, septate, embedded in mucilage, without<br />
anastomosing (Fig. 55d). Asci 108–170×18–22 μm<br />
(x ¼ 144:5 18:8mm, n=10), 8-spored, cylindro-clavate,<br />
pedunculate, bitunicate, fissitunicate, (1-)2-seriate, apically<br />
rounded, with an ocular chamber and faint ring (J-) (Fig. 55c<br />
and f). Ascospores 30–38×8–12 μm (x ¼ 32:4 8:6mm, n=<br />
10), fusoid to ellipsoid, 4-celled, constricted at the septa,<br />
hyaline, with acute rounded ends and surrounded by (5–<br />
8 μm diam.) mucilaginous sheath (Fig. 55g).<br />
Anamorph: Ceratophoma sp. (Sivanesan 1984).<br />
Material examined: FRANCE, on twig of Fagus sp.,<br />
(Desmazières 1764. P, holotype of Sphaeria pupula var<br />
minor), (Mycotheca universalis no. 1951 lectotype). AUS-<br />
TRIA, Silesia, Karlsbrunn, on dead twigs of Fagus sylvatica<br />
L., Aug. and Sept. 1890, Niessl., De Thümen, sub. Massarina<br />
eburnea, ETH. Saxonia, Königsbrunn, on twigs of Fagus<br />
sylvatica, Apr. 1882, W. Krieger, Rabenhorst & Winter, Fungi<br />
europaei no. 2767, ETH; FRANCE, on a dead twig of Fagus<br />
sylvatica, Deux Sèvres, Villiers en Bois, Forêt de Chizé,<br />
Rimbaud, 14 Apr. 2008, leg. det. Paul Leroy (IFRD 2006).<br />
Notes<br />
Morphology<br />
Massarina was introduced by Saccardo (1883) for<br />
species of pyrenocarpous ascomycetes that had previously<br />
been placed in Massaria, but typically had hyaline<br />
ascospores (Bose 1961). The family Massarinaceae was<br />
described by Munk (1956) to accommodate Massarina.<br />
This family was not commonly used and Massarina was<br />
later placed within the Lophiostomataceae in the <strong>Pleosporales</strong><br />
(Barr 1990a; Bose 1961; Eriksson and Yue 1986). Of<br />
the 160 epithets listed in his monograph, Aptroot<br />
accepted only 43 species (Aptroot 1998). The concept<br />
of Massarina was widely accepted as having single or<br />
aggregated, immersed to erumpent, spherical to hemispherical,<br />
pseudothecioid ascomata; cellular pseudoparaphyses;<br />
bitunicate, cylindrical to clavate or obpyriform<br />
asci; and hyaline, 1–3(−7)-septate, fusoid to long ellipsoid<br />
ascospores that mostly have a mucilaginous sheath or<br />
appendages (Aptroot 1998; Hyde and Aptroot 1998;<br />
Tanaka and Harada 2003d).<br />
In the holotype of Sphaeria pupula var. minor (P) and<br />
lectotype of Massarina eburnea (ETH), ascospores are<br />
reported as “not constricted at the septa” (Hyde 1995a).<br />
However, in one of our recent collections, ascospores that<br />
are constricted at their septa were observed (Fig. 55g),<br />
which was consistent with the description by Fallah and<br />
Shearer (2001). This might be because this character is not<br />
clear in the old (over 100 years) and dry herbarium<br />
specimens, or it may be variable between collections.<br />
Phylogenetic study<br />
Recent morphological, molecular and anamorphic results<br />
indicate, however, that Massarina is polyphyletic (Hyde<br />
1995a; Kirk et al. 2001; Liew et al. 2002). Based on the<br />
rDNA dataset, Massarina cisti and the type of Massarina<br />
(M. eburnea) forms a robust clade representing Massarina<br />
sensu stricto (Zhang et al. 2009a, b).<br />
Concluding remarks<br />
Massarina sensu stricto should be accepted, which<br />
seems to only include some terrestrial and saprobic species.<br />
Massariosphaeria (E. Müll.) Crivelli, Diss. Eidgenöss.<br />
Techn. Hochschule Zürich 7318: 141 (1983).<br />
(?Amniculicolaceae)<br />
≡ Leptosphaeria subgen. Massariosphaeria E. Müll.,<br />
Sydowia 4: 206 (1950).
Fungal Diversity<br />
Fig. 55 Massarina eburnea<br />
(from IFRD 2006). a Ascomata<br />
on the host surface. b Section<br />
of an ascoma. c Ascus with a<br />
short pedicel. d Cellular pseudoparaphyses.<br />
e Section of the<br />
peridium comprising a few<br />
layers of compressed cells. f<br />
Asci in pseudoparaphyses. g<br />
Three-septate ascospores. Scale<br />
bars: a=0.5 mm, b=100 μm,<br />
c–g=20 μm<br />
Generic description<br />
Habitat terrestrial or freshwater, saprobic. Ascomata<br />
medium-sized, scattered, or in small groups, immersed,<br />
erumpent to superficial, subglobose, black; apex with a<br />
wide and usually somewhat compressed papilla. Peridium<br />
thick or thin, usually thicker near the apex,<br />
composed of 2–3 layers of thick walled scleroparenchymatous<br />
cells. Hamathecium of dense, trabeculate<br />
pseudoparaphyses. Asci 8-spored, bitunicate, cylindrical<br />
to cylindro-clavate, with a short, thick, furcate pedicel.<br />
Ascospores fusoid to narrowly ellipsoid, brown or dark<br />
brown, multi-septate.<br />
Anamorphs reported for genus: none.<br />
Literature: Barr1989c; Huhndorf et al. 1990; Kohlmeyer et al.<br />
1996; Müller 1950; Tanaka and Harada 2004;Tanakaetal.2005.<br />
Type species<br />
Massariosphaeria phaeospora (E. Müll.) Crivelli, Ueber<br />
die Heterogene Ascomycetengattung Pleospora Rabh.;<br />
Vorschlag für eine Aufteilung (Diss. Eid genössischen Tech<br />
Hochsch Zürich 7318): 141 (1983). (Fig. 56)<br />
≡ Leptosphaeria phaeospora E. Müll., Sydowia 4: 208<br />
(1950).<br />
Ascomata 400–550 μm high×300–500 μm diam.,<br />
scattered, or in small groups, immersed, semi-immersed,<br />
subglobose, black, apex wide papilla, sometimes slightly<br />
compressed, 40–70(−100) μm broad (Fig. 56a). Peridium
Fungal Diversity<br />
10–20 μm wide at sides, comprising one cell type of<br />
2–3 layers of thick walled scleroparenchymatous cells,<br />
cell wall 2–5 μm thick, peridium thicker near<br />
the apex (Fig. 56b). Hamathecium of dense, trabeculate<br />
pseudoparaphyses, 1–2 μm broad, septate, branching<br />
and anastomosing. Asci 120–173×18–25 μm<br />
(x ¼ 133:2 20:5mm, n=10), 8-spored, bitunicate, fissitunicate<br />
dehiscence not observed, broadly cylindrical to<br />
cylindro-clavate, with a short, thick, furcate pedicel, up to<br />
15 μm long. Ascospores 32.5–42×10–13 μm<br />
(x ¼ 36 11:2mm, n=10), narrowly ellipsoid, usually<br />
slightly curved, dark brown, 7–9 septa, slightly constricted<br />
at the median septum (Fig. 56c and d).<br />
Anamorph: none reported.<br />
Material examined: SWITZERLAND, Kt. Wallis,<br />
Findelen, Artemisiae campestris L., 10 Sept. 1895, H.<br />
Wegelin (ZT, holotype).<br />
Notes<br />
Morphology<br />
Massariosphaeria was established by Müller (1950) as<br />
a section of Leptosphaeria based on its large, thick-walled<br />
ascospores with a mucilaginous sheath as well as its<br />
ascomata with a thick apex. Massariosphaeria was<br />
introduced as a separate genus by Crivelli (1983),<br />
characterized by its wide peridial apex comprising<br />
thick-walled cells, compressed to round papilla, and<br />
relatively large, thick-walled, reddish brown to brown,<br />
multi-septate to dictyosporous ascospores, usually surrounded<br />
by a sheath (Crivelli 1983; Huhndorf et al. 1990;<br />
Leuchtmann 1984). In particular, Crivelli (1983) emphasized<br />
that species of Massariosphaeria often stain the woody<br />
substrate (or culture) purple, and this was accepted by<br />
Leuchtmann (1984). Barr (1989c) had treated Massarios-<br />
Fig. 56 Massariosphaeria<br />
phaeospora (ZT, holotype). a<br />
Ascomata scattering on the host<br />
surface. Note the immersed to<br />
erumpent ascomata. b Section of<br />
a partial peridium. Note the<br />
peridium structure. c, d Released<br />
ascospores. Scale bars:<br />
a=1 mm, b–d=20 μm
Fungal Diversity<br />
phaeria as a synonym of Chaetomastia, but this viewpoint<br />
was rarely followed.<br />
Phylogenetic study<br />
The polyphyletic nature of Massariosphaeria is detected<br />
by analyzing SSU and LSU rDNA sequences (Wang et al.<br />
2007). The purple staining character has shown phylogenetic<br />
significance in Amniculicolaceae, a freshwater family from<br />
France (Zhang et al. 2009a). A single isolate of M.<br />
phaeospora was shown to be unrelated to Amniculicolaceae<br />
and clustered with a single isolate of Thyridaria rubronotata<br />
(Schoch et al. 2009; Zhang et al. 2009a).<br />
Concluding remarks<br />
Based on phylogenetic analysis, staining the substrate purple<br />
may have more phylogenetic significance than morphological<br />
characters (Zhang et al. 2009a). Thus, the generic circumscription<br />
of Massariosphaeria should be re-evaluated by further<br />
phylogenetic study with more relevant taxa included.<br />
Mauritiana Poonyth, K.D. Hyde, Aptroot & Peerally,<br />
Fungal Divers. 4: 102 (2000). (?Zopfiaceae)<br />
Generic description<br />
Habitat terrestrial, saprobic. Ascomata medium-sized, gregarious,<br />
ovoid, immersed, ostiolate, ostiole rounded. Peridium<br />
thin, thicker near the apex. Hamathecium of dense, cellular<br />
pseudoparaphyses, branching. Asci 8-spored, bitunicate, cylindrical<br />
to cylindro-clavate, with a short pedicel and a small<br />
ocular chamber. Ascospores 2-3-seriate, fusoid with rounded<br />
ends, dark brown with paler apical cells, multi-septate,<br />
distoseptate, slightly constricted at the primary septum.<br />
Anamorphs reported for genus: none.<br />
Literature: Hawksworth et al. 1995; Poonyth et al. 2000;<br />
Suetrong et al. 2009.<br />
Type species<br />
Mauritiana rhizophorae Poonyth, K.D. Hyde, Aptroot &<br />
Peerally, Fungal Divers. 4: 102 (2000). (Fig. 57)<br />
Ascomata 390–410 μm high×310–325 μm diam.,<br />
gregarious, ovoid, immersed, ostiolate, ostiole rounded<br />
(Fig. 57a). Peridium 40–60 μm thick laterally, thicker near<br />
the apex (Fig. 57a and b). Hamathecium of dense, long<br />
cellular pseudoparaphyses, 1.5–2 μm broad, branching. Asci<br />
130–180×20–25 μm (x ¼ 156 21:8mm, n=10), 8-spored,<br />
bitunicate, cylindrical to cylindro-clavate, with a short pedicel,<br />
with a small ocular chamber (Fig. 57c, d and e). Ascospores<br />
29–40×9–13 μm (x ¼ 35:4 11mm, n=10), 2-3-seriate,<br />
fusoid with rounded ends, dark brown with paler apical cells,<br />
9–13-distoseptate, slightly constricted at the primary septum,<br />
smooth (Fig. 57f,g,handi).<br />
Anamorph: none reported.<br />
Material examined: MAURITIUS, Grand Gaube,<br />
Melville mangrove, on dead decorticated Rhizophora<br />
mucronata Lam.woodstillattachedtolivingtree,Jan.<br />
1995, A.D. Poonyth (HKU(M)10219, holotype).<br />
Notes<br />
Morphology<br />
Mauritiana was introduced to accommodate the mangrove<br />
fungus, M. rhizophorae, which is characterized by<br />
immersed ostiolate, periphysate ascomata, thin peridium,<br />
bitunicate, 8-spored, cylindrical to cylindro-clavate asci,<br />
fusoid, smooth, hyaline to pale brown, multi-septate and<br />
distoseptate ascospores (Poonyth et al. 2000). But after<br />
carefully studying the type of M. rhizophorae, notypical<br />
distoseptate ascospores observed. The pigmented curved<br />
septum of the ascospore gives a “thickened” appearance.<br />
Based on its immersed ascomata, presence of cellular<br />
pseudoparaphyses, thick-walled, fissitunicate asci and<br />
brown, phragmosporous ascospores constricted at the primary<br />
septum, Mauritiana was assigned to the Pyrenulales sensu<br />
stricto (Melanommatales sensu lato, Dothideales sensu lato)<br />
(Hawksworth et al. 1995; Poonyth et al. 2000).<br />
Phylogenetic study<br />
Based on a multigene phylogenetic analysis, Mauritiana<br />
rhizophorae resided within a paraphyletic clade (Suetrong et<br />
al. 2009) sister to marine fungi Halotthia posidonia and<br />
Pontoporeia biturbinata. In this study, the dendrogram shows<br />
it to be closely related to the Sporormiaceae and Lophiostomataceae,<br />
which may indicate an uncircumscribed familial clade<br />
(Plate 1). Thus, its familial placement remains undetermined.<br />
Concluding remarks<br />
The “thickened” septa of ascospores of Mauritiana<br />
rhizophorae is quite unique in <strong>Pleosporales</strong>, which makes<br />
it easily distinguishable from other genera..<br />
Melanomma Nitschke ex Fuckel, Jb. nassau. Ver. Naturk.<br />
23–24: 159 (1870). (Melanommataceae)<br />
Generic description<br />
Habitat terrestrial, saprobic. Ascomata immersed, erumpent<br />
to nearly superficial, medium- to large-sized, globose to<br />
subglobose, coriaceous, gregarious, short papillate. Peridium<br />
pseudoparenchymatous cells outside with pale compressed<br />
cells inside. Asci cylindric to clavate with short<br />
pedicels. Hamathecium of dense, filamentous, branching,<br />
rarely anastomosing, septate pseudoparaphyses. Ascospores<br />
pale brown, reddish brown to olive-brown, ellipsoid to<br />
fusoid, 2 to multi-septate, constricted at the main septum.
Fungal Diversity<br />
Fig. 57 Mauritiana rhizophorae<br />
(from HKU(M)10219,<br />
holotype). a Vertical section of<br />
an ascoma. Note the thin layer of<br />
fungal tissue (pseudostroma?) on<br />
the host surface. b Section of a<br />
partial peridium. c Pseudoparaphyses<br />
and immature ascus. d<br />
Fissitunicate asci. e Asci showing<br />
thickening of the apical wall.<br />
f–i Ascospores with transverse<br />
septa and paler polar cells. Scale<br />
bars: a=40 μm, b, d–i=10 μm,<br />
c=20 μm<br />
Anamorphs reported for genus: Aposphaeria, Nigrolentilocus,<br />
Phoma-like and Pseudospiropes (Chesters 1938;<br />
Sivanesan 1984).<br />
Literature: Barr 1990a; Chesters 1938; Fuckel 1870;<br />
Saccardo 1878; Zhang et al. 2008a.<br />
Type species<br />
Melanomma pulvis-pyrius (Pers.) Fuckel, Jb. nassau. Ver.<br />
Naturk. 23–24: 160 (1870). (Fig. 58)<br />
≡ Sphaeria pulvis-pyrius Pers., Syn. meth. fung.<br />
(Göttingen) 1: 86 (1801).<br />
Ascomata 215–471 μm high×260–440 μm diam.,<br />
gregarious, substrate surface covered with a thin layer of<br />
brown psueodstroma, superficial, globose, subglobose,<br />
broadly or narrowly conical, often laterally flattened, black,<br />
roughened and irregular, often bearing remnants of wood<br />
fibers; apex short papillate, often somewhat puckered or<br />
sulcate (Fig. 58a). Peridium 70–90 μm wide, to 180 μm<br />
wide at the base, coriaceous, comprising two types of cells,<br />
outer cells small heavily pigmented thick-walled cells of<br />
textura angularis, apical cells smaller and walls thicker,<br />
individual cell walls to 6 μm thick, inner cells lightly<br />
pigmented to hyaline thin-walled cells of textura angularis,
Fungal Diversity<br />
5–8 μm diam., individual cell wall to 1.5–2 μm thick, in<br />
places with columns of textura prismatica, and larger, paler<br />
cells of textura prismatica towards the interior and at the<br />
base (Fig. 58b). Hamathecium of dense, filamentous, 1–2<br />
(−2.5) μm broad, branching, rarely anastomosing, septate<br />
pseudoparaphyses. Asci 98–123×6.5–7.5(−9) μm<br />
(x ¼ 109 7:5mm, n=10), 8-spored, bitunicate, fissitunicate,<br />
cylindrical to fusoid, with a short, furcate pedicel, to 25 μm<br />
long, with an ocular chamber (Fig. 58c, d, e, f and g).<br />
Ascospores 14–17.5(−19)×4.5–6.5 μm (x ¼ 15:8 5:2mm,<br />
n=10), obliquely uniseriate and partially overlapping, broadly<br />
fusoid to fusoid with broadly rounded ends, straight or slightly<br />
curved, smooth, olive-brown, 4-celled, slightly constricted at<br />
the septa, the second cell from the top slightly wider than the<br />
others, no sheath (Fig. 58h,i,j,kandl).<br />
Anamorph: Aposphaeria agminalis Sacc. or Phoma<br />
agminalis Sacc. (Sivanesan 1984).<br />
Colonies (of epitype) reaching 4 cm diam. after 20 days<br />
growth on PDA at 25°C, depressed to raised, cottony to woolly,<br />
with rhizoidal margin, grey, reverse darkened. Phoma-like<br />
anamorph has been reported by Chesters (1938) and<br />
Sivanesan (1984), but no anamorphic stage was observed<br />
in the cultures of IFRDCC 2044, <strong>CBS</strong> 109.77 and <strong>CBS</strong><br />
371.75 after culturing 3 months on PDA.<br />
Material examined:ondecayingwood(UPS,Scler.suec.<br />
n. 120, holotype,asSphaeria pulvis-pyrius Pers.); FRANCE,<br />
Ariège, Rimont, Saurine, on bark of Salix caprea, 10Apr.<br />
2008, Jacques Fournier (IFRD 2001, epitype).<br />
Notes<br />
Morphology<br />
Melanomma, the familial type of Melanommataceae, was<br />
formally established by Fuckel (1870, p 159) based on its small,<br />
carbonaceous ascomata, having: “sporen meist 2–3 mal septirt,<br />
selten ohne Scheidewand, braun oder wasscrhell.” (Chesters<br />
1938; Fuckel 1870). Saccardo (1878, p. 344) emended this<br />
genus as “Spores ovate or oblong, multi-septate, coloured.”<br />
Subsequently, Saccardo (1883, p. 98) extended the description<br />
of Melanomma as “Perithecia gregarious, seldom scattered,<br />
somewhat superficial, sphaerical, papillate or blunt, carbonaceous,<br />
smooth or somewhat hairy. Asci elongate, for the most<br />
part accompanied by paraphyses, 8-spored. Spores oblong or<br />
somewhat spindle-shaped, two to many septate, olive or dark<br />
brown. Species of Sphaeria belong here for the most part.”<br />
Melanomma pulvis-pyrius was erected as the lectotype<br />
species (Barr 1990a; Chesters1938). Barr (1990a) gavea<br />
detailed circumscription for Melanomma, under which<br />
Melanomma contains about 20 species (Kirk et al. 2001).<br />
Melanomma pulvis-pyrius is characterized by its gregarious,<br />
superficial ascomata with short papillate, cylindrical asci with a<br />
short pedicel and fusoid, olive-brown, 3-septate ascospores<br />
(Chesters 1938; Zhang et al. 2008a). One of the diagnostic<br />
characters of Melanommataceae is the trabeculate pseudoparaphyses,<br />
although no typical trabeculate pseudoparaphyses could<br />
be found in the neotype (Scler. suec. n. 120, UPS) and epitype<br />
(IFRD 2001) of M. pulvis-pyrius (Zhang et al. 2008a).<br />
Phylogenetic study<br />
Phylogenetic analysis based on five genes (LSU, SSU,<br />
RPB1, RPB2 andEF1) indicates that Melanomma pulvispyrius<br />
forms a robust clade with Byssosphaeria, Herpotrichia<br />
and Pleomassaria siparia (Pleomassariaceae) and<br />
likely represents a separate family (or families comprising<br />
Melanommataceae) (Zhang et al. 2008a; Mugambi and<br />
Huhndorf 2009b). A more recent phylogenetic analysis<br />
included a group of coelomycete species with stellate<br />
conidia, isolated from Fagales trees clustered in Melanommataceae<br />
(Tanaka et al. 2010; Plate 1).<br />
Concluding remarks<br />
The Melanomma concept based on ascospore morphology<br />
appears polyphyletic.<br />
Metameris Theiss. & Syd., Annls mycol. 13: 342 (1915).<br />
(Phaeosphaeriaceae)<br />
Generic description<br />
Habitat terrestrial, saprobic or parasitic. Ascostromata erumpent<br />
through the host surface in linear rows parallel to the host fibers.<br />
Ascomata small, globose to subglobose, black, coriaceous.<br />
Peridium composed of large lightly pigmented cells of textura<br />
angularis. Hamathecium of rare, broad pseudoparaphyses,<br />
septate, constricted at the septa. Asci bitunicate, fissitunicate,<br />
broadly cylindrical to slightly obclavate, with a short, thick,<br />
knob-like pedicel. Ascospores hyaline, 1- (rarely 2-) septate.<br />
Anamorphs reported for genus: none.<br />
Literature: von Arx and Müller 1975; Barr1972; Clements<br />
and Shear 1931; Eriksson 2006; Lumbsch and Huhndorf<br />
2007; TheissenandSydow1915.<br />
Type species<br />
Metameris japonica (Syd.) Syd., Annls mycol., 13(3–4):<br />
342 (1915). (Fig. 59)<br />
≡ Monographus japonicus Syd. Annls mycol. 10: 408<br />
(1912).<br />
Ascostromata erumpent through the host surface in linear<br />
rows parallel to the host fibers, 500–750 μm long and 140–<br />
200 μm wide, with three to ten ascomata arranged in a line<br />
(Fig. 59a). Ascomata 115–160 μm diam., semi-immersed in<br />
substrate to erumpent, globose, subglobose, black, coriaceous<br />
(Fig. 59b). Cells of ascostromata heavily pigmented and<br />
thick-walled, cells of peridium composed of large lightly<br />
pigmented cells of textura angularis, cells 5–15 μm diam.,
Fungal Diversity<br />
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<br />
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<br />
cell wall
Fungal Diversity<br />
Phylogenetic study<br />
None.<br />
Concluding remarks<br />
Its small-sized ascomata, broadly cylindrical to slightly<br />
obclavate asci with a short, thick, knob-like pedicel, as well<br />
as its monocotyledonous host preference point Metameris<br />
to the Phaeosphaeriaceae. But DNA comparisons are<br />
needed for confirmation.<br />
Mixtura O.E. Erikss. & J.Z. Yue, Mycotaxon 38: 203<br />
(1990). (Phaeosphaeriaceae)<br />
Generic description<br />
Habitat terrestrial, parasitic. Ascomata small-sized, scattered<br />
or clustered on the leaf spots, immersed, erumpent,<br />
minutely papillate, ostiolate. Papilla slightly raised. Peridium<br />
thin, comprising one cell type of lightly pigmented<br />
thin-walled cells of textura angularis. Hamathecium of<br />
dense, filliform, septate, cellular pseudoparaphyses, 4–<br />
6.3 μm broad, embedded in mucilage. Asci bitunicate,<br />
ovoid, with a very short stumpy pedicel. Ascospores fusoid<br />
to narrowly fusoid with broadly to narrowly rounded ends,<br />
curved, dark brown, multi-septate, distoseptate, with a germ<br />
pore at the lower end.<br />
Anamorphs reported for genus: none.<br />
Literature: Eriksson and Yue 1990.<br />
Type species<br />
Mixtura saginata (Syd.) O.E. Erikss. & J.Z. Yue, Mycotaxon<br />
38: 203 (1990). (Fig. 60)<br />
≡ Leptosphaeria saginata Syd., Annls mycol. 37: 376 (1939).<br />
Producing elongated yellow spots with brownish margins,<br />
leaf spots up to 45×3–5 mm, opposite side visible as a<br />
brownish spots (Fig. 60a). Ascomata 170–200 μm high×210–<br />
280 μm diam., scattered on the lower side of the leaf,<br />
immersed, erumpent, breaking through the epidermis, minutely<br />
papillate. Papilla central, slightly raised, ostiolate,<br />
ostiole surrounded by a white margin (Fig. 60b). Peridium<br />
22–34 μm wide, thicker at the apex, thinner at the base,<br />
comprising one cell type of lightly pigmented thin-walled<br />
cells of textura angularis, cells up to 6×8 μm diam., cell wall<br />
0.5–1.2 μm thick, apex cells smaller and walls thicker<br />
(Fig. 60c). Hamathecium of dense, filliform, septate, cellular<br />
pseudoparaphyses, 4–6.3 μm broad, embedded in mucilage.<br />
Asci 80–128×41–53(−69) μm (x ¼ 100:9 52:8mm, n =10),<br />
8-spored, bitunicate, fissitunicate dehiscence not observed,<br />
sac-like, with a very short stumpy pedicel and a small ocular<br />
chamber (Fig. 60d). Ascospores 86–94(−106)×20.5–23.5 μm<br />
(x ¼ 92:7 21:7mm, n=10), fasciculate, fusoid to narrowly<br />
fusoid, slightly curved, dark brown, 7-septate, distoseptate,<br />
with or without constriction at the primary septum, smoothwalled,<br />
with a germ pore at the lower end (Fig. 60e and f).<br />
Anamorph: none reported.<br />
Material examined: ECUADOR, Tungurahua, Hacienda<br />
San Antonio pr. Baños, Province, on the leaves of Chusqueae<br />
serrulatae Pilger., 9 Jan. 1938, H. Sydow. (S reg. nr F8934<br />
type, F8935isolectotype, asLeptosphaeria saginata).<br />
Notes<br />
Morphology<br />
Mixtura was formally established by Eriksson and<br />
Yue (1990) as a monotypic genus represented by M.<br />
saginata basedonitsimmersedandthin-walledascomata,<br />
sparse, broad pseudoparaphyses, sac-like asci with a short<br />
pedicel and thick apex. Mixtura has a “mixture” of<br />
characters found in other pleosporalean genera. The<br />
peridium structure is comparable with Phaeosphaeria,<br />
the ascospores with Trematosphaeria and asci with<br />
Wettsteinina (Eriksson and Yue 1990).Accordingtothe<br />
structure of ascomata and hamathecium, Mixtura was<br />
provisionally assigned to Phaeosphaeriaceae (Eriksson<br />
and Yue 1990).<br />
Phylogenetic study<br />
None.<br />
Concluding remarks<br />
Morphologically, the sparse broad pseudoparaphyses and<br />
sac-like asci with a thick apical structure in Mixtura seem<br />
more comparable with the generic type of Teratosphaeria<br />
(T. fibrillose Syd. & P. Syd., Teratosphaeriaceae, Capnodiales,<br />
Dothideomycetidae) than that of Phaeosphaeria (P.<br />
oryzae). The heavily pigmented, multi-septate ascospores<br />
and the persistent pseudoparaphyses of Mixtura however,<br />
differ from those of Teratosphaeria. Thus, here we assign<br />
Mixtura under Teratosphaeriaceae as a distinct genus until<br />
phylogenetic work is carried out.<br />
Montagnula Berl., Icon. fung. (Abellini) 2: 68 (1896).<br />
(Montagnulaceae)<br />
Generic description<br />
Habitat terrestrial, saprobic. Ascomata small- to mediumsized,<br />
immersed to erumpent, gregarious or grouped,<br />
globose to subglobose, black. Hamathecium of dense,<br />
narrowly cellular, septate pseudoparaphyses. Asci bitunicate,<br />
fissitunicate, usually cylindro-clavate to clavate with a<br />
long pedicel. Ascospores oblong to narrowly oblong,<br />
straight or somewhat curved, reddish brown to dark<br />
yellowish brown, muriform or phragmosporous.<br />
Anamorphs reported for genus: Aschersonia (Hyde et al. 2011).
Fungal Diversity<br />
Fig. 60 Mixtura saginata<br />
(from S reg. nr F8934, type). a,<br />
b Leaf spots in leaves of Chusquea<br />
serrulatae. Note the erumpent<br />
ascomata surrounded by<br />
white material in (b). c Section<br />
of an ascoma. Note the peridium<br />
structure which comprises cells<br />
of textura angularis. The arrangement<br />
of the asci and pseudoparaphyses<br />
can also be seen.<br />
d Immature asci in pseudoparaphyses.<br />
Note the stumpy pedicel<br />
and thickened apex with<br />
flattened ocular chamber. e, f<br />
Mature ascospores. Note the<br />
hyaline ends and distosepta.<br />
Scale bars: a=10 mm, b, c=<br />
100 μm, d=50 μm, e–f=20 μm<br />
Literature: Aptroot 1995; Barr2001; Berlese1896; Clements<br />
and Shear 1931; Crivelli1983; Leuchtmann 1984; Ramaley<br />
and Barr 1995; Schoch et al. 2006; Wehmeyer1957, 1961;<br />
Zhang et al. 2009a.<br />
Type species<br />
Montagnula infernalis (Niessl) Berl., Icon. fung. (Abellini).<br />
2: 68 (1896). (Fig. 61)<br />
≡ Leptosphaeria infernalis Niessl, Inst. Coimbra 31: 13<br />
(1883).<br />
Ascomata 220–280 μm high×250–310 μm diam., immersed<br />
to erumpent, gregarious or clustered, globose to<br />
subglobose, sometimes triangular in dried material, short<br />
ostiole always filled with hyaline closely adhering cells, black<br />
(Fig. 61a and b). Peridium 40–55 μm thick at sides, up to<br />
80 μm thick near the apex, 3-layered, outer layer composed of<br />
heavily pigmented thick-walled small cells of textura angularis,<br />
cells3–8 μm diam., wall 1.5–3 μm thick, apex thicker<br />
with smaller cells and thicker cell wall, thinner near the base;<br />
mid layer less pigmented, cells 4–13 μm diam.; innermost<br />
layer of narrow compressed rows of cells, merging with
Fungal Diversity<br />
pseudoparaphyses (Fig. 61c). Hamathecium of dense, narrow<br />
cellular pseudoparaphyses, 2–4.5 μm broad, septate (Fig. 61f).<br />
Asci 153–170(−200)×17.5–21.5 μm (including pedicel),<br />
bitunicate, fissitunicate, cylindro-clavate to clavate, pedicel<br />
28–60(−85) μm long, 8-spored, biseriate, with an ocular<br />
chamber best seen in immature ascus (to 3 μm wide×3 μm<br />
high) (Fig. 61d and e). Ascospores 24–29×9–11 μm, oblong to<br />
narrowly oblong, straight or somewhat curved, reddish brown<br />
to dark yellowish brown, verruculose, with five transverse<br />
septa and one vertical septum in each middle cells, constricted<br />
at the primary and secondary primary septa (Fig. 61g).<br />
Anamorph: none reported.<br />
Material examined: PORTUGAL, Coimbra Lusitania,<br />
on leaves of Fourcroya longava pr., Feb., 1881, leg. Moller.<br />
(M 1183, holotype).<br />
Notes<br />
Morphology<br />
Montagnula was introduced to accommodate two Pleospora<br />
species, i.e. P. infernalis (Niessl) Wehm. and P.<br />
gigantea Mont. by Berlese (1896), based on the presence of<br />
hyphal stromatic tissues over the ascomata and asci with<br />
relatively long pedicels (Barr 2001). Montagnula infernalis<br />
was selected as the lectotype species (Clements and Shear<br />
1931). Subsequently, Wehmeyer (1957, 1961) treated Montagnula<br />
as a subgenus of Pleospora. Crivelli(1983) accepted<br />
Montagnula as a separate genus, and divided it into two<br />
subgenera, i.e. Montagnula and Rubiginospora. Montagnula<br />
was characterized by having dark brown ascospores and<br />
exclusively occurring on Agavaceae, while Rubiginospora<br />
has reddish brown ascospores and occurs on Poaceae. This<br />
proposal was not accepted by many workers (Barr 2001).<br />
Subsequently, more species with various ascospores (such as<br />
phragmosporous species by Leuchtmann (1984) and didymosporous<br />
species by Aptroot (1995) were added in this<br />
genus), which has obviously become heterogenic. Barr<br />
(2001) assigned species of Montagnula into different genera,<br />
i.e. Kalmusia and Didymosphaerella, respectively and introduced<br />
Montagnulaceae to accommodate all of these genera.<br />
Phylogenetic study<br />
Montagnula opulenta forms a robust phylogenetic clade<br />
with species of Bimuria, Curreya, Didymocrea, Letendraea,<br />
Paraphaeosphaeria, Phaeodothis and Karstenula,<br />
which might represent a familial group (Schoch et al. 2006;<br />
Zhang et al. 2009a). A more convincing conclusion can<br />
only be obtained following sequence data from more<br />
verified fungi being added to the phylogenetic tree.<br />
Concluding remarks<br />
One striking character of Montagnula infernalis is the very<br />
long ascal pedicel once it is released from the ascomata.<br />
However, this character appears to have evolved more than<br />
once and can be found in Kirschsteiniothelia elaterascus<br />
Shearer which clusters with Helicascus (Shearer et al. 2009).<br />
The same ascus character is also found in Xenolophium and<br />
Ostropella in the Platystomaceae (Mugambi and Huhndorf<br />
2009b). Montagnula opulenta is a didymosporous species,<br />
but phylogenetically closely related to those dictyosporous<br />
(Karstenula rhodostoma) and phragmosporous (Paraphaeosphaeria<br />
michotii) members of Montagnulaceae (Zhang et al.<br />
2009a). This might indicate that compared to other morphological<br />
characters, ascospore type is not a valid character at<br />
family level classification.<br />
Moristroma A.I. Romero & Samuels, Sydowia 43: 246<br />
(1991). (<strong>Pleosporales</strong>, genera incertae sedis)<br />
Generic description<br />
Habitat terrestrial, saprobic. Ascomata medium-sized, solitary,<br />
scattered, or in small groups, superficial, cushion-like, circular<br />
in outline, wall black, roughened, containing numerous<br />
locules. Peridium thin, 1-layered. Hamathecium of dense, long<br />
filliform pseudoparaphyses, 2–3 μm broad, septate, branching.<br />
Asci polysporous, with a short, laterally displaced, sometimes<br />
papillate knob-shaped pedicel, apex very thick walled,<br />
bitunicate, fissitunicate, obclavate, ocular chamber not observed.<br />
Polyspores oblong to cylindrical, hyaline, non-septate.<br />
Anamorphs reported for genus: none.<br />
Literature: Eriksson 2006; Romero and Samuels 1991.<br />
Type species<br />
Moristroma polysporum A.I. Romero & Samuels, Sydowia<br />
43: 246 (1991). (Fig. 62)<br />
Ascomata 100–210 μm high×340–600 μm diam., solitary,<br />
scattered, or in small groups of 2–3, superficial, with basal<br />
wall remaining immersed in host tissue, cushion-like,<br />
circular in outline, wall black, roughened, containing<br />
numerous locules, each locule 120–240 μm diam., ostiolate<br />
(Fig. 62a and b). Peridium 14–30 μm thick, 1-layered,<br />
composed of small heavily pigmented thick-walled cells of<br />
textura angularis, cells2–4 μm diam., cell wall 1.5–3 μm<br />
thick, peridium between the locules hyaline (Fig. 62b and c).<br />
Hamathecium of dense, long filliform pseudoparaphyses, 2–<br />
3 μm broad, septate, branching. Asci 44–60×12–14 μm<br />
(x ¼ 54:3 13mm, n=10), polysporous, with a short, papillate<br />
knob-shaped pedicel, apex very thick-walled, bitunicate,<br />
fissitunicate, obclavate, ocular chamber not observed<br />
(Fig. 62d and e). Polyspores 3–4(−5)×0.6–1.2 μm, oblong<br />
to cylindrical, hyaline, non-septate, smooth (Fig. 62f).<br />
Anamorph: none reported.<br />
Material examined: ARGENTINA, Buenos Aires,<br />
Ramallo, on Eucalyptus viminalis Labill., May 1982, Romero
Fungal Diversity<br />
Fig. 61 Montagnula infernalis<br />
(from M 1183, holotype). a<br />
Appearance of ascomata immersed<br />
in host tissue. b Section<br />
of an immersed ascoma. Note<br />
the hyaline closely adhering<br />
cells in the ostiole region. c<br />
Section of the peridium comprising<br />
a few layers of cells. d<br />
An immature ascus with a long<br />
pedicel. e, g Mature muriform<br />
ascospores in asci. f Cellular<br />
pseudoparaphyses. Scale bars:<br />
a=0.5 mm, b, c=100 μm,<br />
d–g=20 μm<br />
27/4-13 (BAFC 32036, holotype); Nov. 1982, on decorticated<br />
wood, Romero 35/4-13 (BAFC 32037, paratype).<br />
Notes<br />
Morphology<br />
Moristroma was formally established by Romero and<br />
Samuels (1991) based on its “cushion-shaped ascomata<br />
containing lots of locules with numerous asci inside, asci<br />
obclavate, polysporous, with a knob-shaped pedicel”. The<br />
bitunicate asci and numerous cellular pseudoparaphyses<br />
undoubtedly point it to <strong>Pleosporales</strong>, while the familial<br />
placement of Moristroma is uncertain, and it was<br />
temporarily assigned to Dacampiaceae by Romero and<br />
Samuels (1991), but no 3-layered peridium is found.<br />
Eriksson (2006) assignedittoTeichosporaceae.<br />
Phylogenetic study<br />
None.<br />
Concluding remarks<br />
The familial status of Moristroma cannot be determined yet.
Fungal Diversity<br />
Fig. 62 Moristroma polysporum<br />
(from BAFC 32036, holotype).<br />
a Two multiculate<br />
ascostroma on the host surface.<br />
b Section of an ascostroma.<br />
Note the multilocula. c Section<br />
of the peridium. Note the thick<br />
walled cells. d, e Broadly cylindrical<br />
to fusoid asci containing<br />
numerous part spores. f Released<br />
part spores. Scale bars:<br />
a=0.5 mm, b=200 μm, c=<br />
50 μm, d–f=10 μm<br />
Morosphaeria Suetrong, Sakay., E.B.G. Jones & C.L.<br />
Schoch, Stud. Mycol. 64: 161 (2009). (Morosphaeriaceae)<br />
Generic description<br />
Habitat marine, saprobic. Ascomata large, solitary or<br />
gregarious, immersed to erumpent, subglobose or depressed<br />
with a flatted base, ostiolate, papillate, brown to black,<br />
coriaceous. Peridium thick. Hamathecium of dense, long<br />
cellular pseudoparaphyses, septate. Asci 8-spored, bitunicate,<br />
cylindrical, with short pedicels. Ascospores uniseriate<br />
to partially overlapping, ellipsoidal, hyaline, 1-3-septate,<br />
constricted at the septa, central cells larger, apical cells if<br />
present small and elongated, surrounded with mucilaginous<br />
sheath.<br />
Anamorphs reported for genus: none.<br />
Literature: Hyde and Borse 1986; Hyde 1991a, b;<br />
Suetrong et al. 2009; Zhang et al. 2009a.<br />
Type species<br />
Morosphaeria velataspora (K.D. Hyde & Borse) Suetrong,<br />
Sakay., E.B.G. Jones & C.L. Schoch, Stud. Mycol. 64: 161<br />
(2009). (Fig. 63)<br />
≡ Massarina velataspora K.D. Hyde & Borse, Mycotaxon<br />
27: 163 (1986).<br />
Ascomata 0.7–1.2 mm diam., solitary or gregarious,<br />
immersed to erumpent, subglobose or depressed, with a<br />
flattened base not easily removed from the substrate, ostiolate,<br />
epapillate or papillate, brown to black, coriaceous (Fig. 63a).<br />
Peridium thick, the upper part of the peridium composed of<br />
brown thick-walled cells of textura angularis, cells are<br />
smaller and wall thicker near the apex, at the rim is<br />
composed of vertical, parallel, brown, elongate cells,<br />
wedge-shape in section (Fig. 63a). Hamathecium of dense,<br />
long cellular pseudoparaphyses, 1.1–1.7 μm broad, septate.<br />
Asci 220–320×23–34 μm (x ¼ 251 28:2mm, n=10), 8-
Fungal Diversity<br />
spored, bitunicate, cylindrical, with short pedicels (Fig. 63b).<br />
Ascospores 45–56×14–19 μm (x ¼ 49:5 15:9mm, n=10),<br />
uniseriate to partially overlapping, ellipsoidal, hyaline, 1-3-<br />
septate, constricted at the septa, central cells larger, apical<br />
cells if present small and elongated, surrounded with<br />
mucilaginous sheath, 5–22 μm wide (Fig. 63c, d and e).<br />
Anamorph: none reported.<br />
Material examined: Jan. 1984, Herb. IMI 297770,<br />
slides 1–10 (holotype) and dried wood (isotype).<br />
Notes<br />
Morphology<br />
Two Massarina sensu lato species described from the<br />
marine environment, viz. M. ramunculicola (Sacc.) O.E.<br />
Erikss. & J.Z. Yue and M. velataspora K.D. Hyde & Borse,<br />
form a robust clade, and a new genus Morosphaeria was<br />
established for them (Suetrong et al. 2009). Together with<br />
two Helicascus species, they belong to Morosphaeriaceae<br />
(another marine family) (Suetrong et al. 2009). Morphologically,<br />
Morosphaeria is characterized by solitary to<br />
gregarious, subglobose to lenticular, immersed to superficial<br />
ascomata which are ostiolate and papillate, numerous,<br />
filliform pseudoparaphyses, 8-spored, clavate to cylindrical,<br />
bitunicate, fissitunicate asci, and hyaline, 1-3-septate,<br />
fusoid to ellipsoidal ascospores which are surrounded with<br />
mucilaginous sheath.<br />
Phylogenetic study<br />
Species of Morosphaeria form a sister group with<br />
Helicascus and both of these genera were assigned to a<br />
new family, i.e. Morosphaeriaceae (Suetrong et al. 2009).<br />
In this study, a strain of Asteromassaria pulchra, occuring<br />
on dead twigs of Prunus spinosa, is basal to other species<br />
of Morosphaeriaceae, and gets well support. Thus here we<br />
tentatively assign Asteromassaria in Morosphaeriaceae.<br />
Fig. 63 Morosphaeria velataspora (from IMI 297770, type). a Section of an ascoma. b Cylindrical asci embedded in pseudoparaphyses. c–e<br />
Hyaline, 1-3-septate, ascospores with mucilaginous sheath. Scale bars: a=100 μm, b=50 μm, c–e=20 μm
Fungal Diversity<br />
Concluding remarks<br />
The only morphological difference between M. velataspora<br />
and M. ramunculicola are their morphology of<br />
ascomata and size of ascospores (Hyde 1991b). But M.<br />
velataspora was reported staining the woody substrate (or<br />
agar in culture) purple (Hyde and Borse 1986; Hyde<br />
1991b). Although this character could not be verified in<br />
the strain used by Suetrong et al. (2009), purple staining has<br />
been reported to have phylogenetic significance at familial<br />
rank in freshwater fungi (Zhang et al. 2009a).<br />
Murispora Yin. Zhang, C.L. Schoch, J. Fourn., Crous & K.<br />
D. Hyde, Stud. Mycol. 64: 95 (2009b). (Amniculicolaceae)<br />
Generic description<br />
Habitat freshwater, saprobic. Ascomata medium-sized,<br />
scattered to gregarious, immersed, lenticular, apex slightly<br />
protruding, opening through a small rounded pore,<br />
substrate stained purple. Peridium thin, composed of a<br />
few layers cells of textura angularis, thicker at the apex<br />
with pseudoparenchymatous cells. Hamathecium of narrowly<br />
cellular pseudoparaphyses, embedded in mucilage.<br />
Asci 8-spored, bitunicate, fissitunicate, biseriate, cylindroclavate<br />
with short pedicels. Ascospores curved- fusoid<br />
with narrowly rounded ends, golden yellow turning brown<br />
when senescent, multi-septate, constricted at the septa,<br />
with one, rarely two longitudinal septa in all cells except<br />
end cells, smooth or finely verruculose, surrounded by a<br />
wide mucilaginous sheath.<br />
Anamorphs reported for genus: Phoma (Webster 1957).<br />
Literature: Zhang et al. 2009a, b.<br />
Type species<br />
Murispora rubicunda (Niessl) Yin. Zhang, J. Fourn. & K.<br />
D. Hyde, Stud. Mycol. 64: 96 (2009a). (Fig. 64)<br />
≡ Pleospora rubicunda Niessl, Notiz. Pyr.: 31 (1876).<br />
Ascomata 170–200 μm high×380–410 μm diam.,<br />
scattered to gregarious, immersed, lenticular, apex<br />
laterally flattened, black, slightly protruding, opening<br />
through a small rounded pore, substrate stained purple<br />
(Fig. 64a). Peridium 15–18 μm thick at sides, composed<br />
of 3–4 layers cells of textura angularis, upto28–30 μm<br />
thick at the apex with very thick-walled cells, pseudoparenchymatous,<br />
nearly absent at the base (Fig. 64b).<br />
Hamathecium of narrowly cellular pseudoparaphyses,<br />
1–1.7 μm broad, embedded in mucilage. Asci 124–<br />
142×19–21 μm, 8-spored, bitunicate, fissitunicate,<br />
biseriate, cylindro-clavate with a small ocular chamber,<br />
with short pedicels (Fig. 64c). Ascospores 30–38×10–<br />
12 μm, curved-fusoid with narrowly rounded ends,<br />
golden yellow turning brown when senescent, 7–9 transversally<br />
septate, constricted at the septa, with one, rarely<br />
two longitudinal septa in all cells except end cells which<br />
are often slightly paler, all cells filled with a large<br />
refractive guttule, smooth to finely verruculose, surrounded<br />
by a wide mucilaginous sheath (Fig. 64d).<br />
Anamorph: Phoma sp. (Webster 1957).<br />
Material examined: FRANCE, Haute Garonne,<br />
Avignonet, Lac de Rosel, 16 Jan. 2007, on submerged<br />
dead herbaceous stem (Dipsacus?), leg. Michel Delpont,<br />
det. Jacques Fournier (IFRD 2017).<br />
Notes<br />
Morphology<br />
Murispora was introduced based on Pleospora rubicunda<br />
which is characterized by immersed, erumpent or<br />
nearly superficial, globose to subglobose, elongated weakly<br />
papillate ascomata which stain the woody substrate purple,<br />
trabeculate pseudoparaphyses, 8-spored, bitunicate, fissitunicate,<br />
oblong to clavate asci, fusoid, pale or reddish<br />
brown, muriform ascospores (Zhang et al. 2009a). A<br />
phylogenetic study indicated that Murispora forms a robust<br />
clade with species of Amniculicola, and Amniculicolaceae<br />
was introduced to accommodate them (Zhang et al. 2009a).<br />
Phylogenetic study<br />
Murispora rubicunda forms a robust clade with species of<br />
Amniculicola and Neophaeosphaeria (Zhang et al. 2009a).<br />
Concluding remarks<br />
As has mentioned by Eriksson (1981, P. 135), the purplestaining<br />
species of Pleospora, treated by Webster (1957),<br />
should not belong to the Pleosporaceae. Both Pleospora<br />
straminis and P. rubelloides should be closely related to<br />
Murispora.<br />
Neomassariosphaeria Yin. Zhang, J. Fourn. & K.D. Hyde,<br />
Stud. Mycol. 64: 96 (2009a). (Amniculicolaceae)<br />
Generic description<br />
Habitat freshwater, saprobic. Ascomata medium-sized,<br />
scattered or in small groups, immersed, with a slightly<br />
protruding elongated papilla, ostiolate, lenticular, stain the<br />
substrate purple. Peridium thin. Hamathecium of dense,<br />
long cellular pseudoparaphyses, septate. Asci 8-spored,<br />
bitunicate, fissitunicate, cylindro-clavate, with short furcate<br />
pedicels. Ascospores 2-3-seriate, narrowly fusoid, somewhat<br />
curved, reddish brown, multi-septate, slightly constricted<br />
at the primary septum.<br />
Anamorphs reported for genus: none.<br />
Literature: Leuchtmann 1984; Zhang et al. 2009a, b.
Fungal Diversity<br />
Fig. 64 Murispora rubicunda<br />
(from IFRD 2017). a Habitat<br />
section of the immersed ascomata.<br />
b Section of an ascoma.<br />
Note the thin peridium and cells<br />
of textura angularis. c Mature<br />
and immature asci. d Muriform<br />
ascospores. Scale bars: a, b=<br />
100 μm, c, d=20 μm<br />
Type species<br />
Neomassariosphaeria typhicola (P. Karst.) Yin. Zhang, J.<br />
Fourn. & K.D. Hyde, Stud. Mycol. 64: 96 (2009a). (Fig. 65)<br />
≡ Leptosphaeria typhicola P. Karst., Bidr. Känn. Finl.<br />
Nat. Folk 23: 100 (1873).<br />
Ascomata 150–280 μm high×200–400 μm diam.,<br />
scattered or in small groups, immersed, lenticular, with a<br />
slightly protruding elongated papilla, ostiolate, stain the<br />
substrate purple (Fig. 65a). Peridium 15–30 μm thick.<br />
Hamathecium of dense, long cellular pseudoparaphyses,<br />
1.5–2.5 μm thick, septate. Asci 110–160×13–15 μm, 8-<br />
spored, bitunicate, fissitunicate, cylindro-clavate, with short<br />
furcate pedicels (Fig. 65b, c and d). Ascospores 30–48×7–<br />
11 μm, 2-3-seriate, narrowly fusoid, somewhat curved,<br />
reddish brown, 7-septate, slightly constricted at the primary<br />
septum, verruculose (Fig. 65c and d).<br />
Anamorph: none reported.<br />
Material examined: DENMARK, Sjaeland, Frederikskilde,<br />
Suserup Skove, Tystrup Lake, 25 May 2007, on<br />
submerged culm of Phragmites, leg. & det. Jacques Fournier<br />
(IFRD 2018).<br />
Notes<br />
Morphology<br />
Neomassariosphaeria is most comparable with Murispora,<br />
and is distinguished from Murispora by its phragmosporous<br />
ascospores. Both genera were assigned to<br />
Amniculicolaceae (Zhang et al. 2009a).<br />
Phylogenetic study<br />
Both Neomassariosphaeria grandispora and N. typhicola<br />
clustered with species of Murispora and Amniculicola in<br />
Amniculicolaceae (Zhang et al. 2009a,c).<br />
Concluding remarks<br />
Similar with those purple-staining species of<br />
Pleospora assigned to Murispora, the purple-staining<br />
species of Phaeosphaeria mentioned by Crivelli<br />
(1983) and Leuchtmann (1984) might be assigned to<br />
Neomassariosphaeria.<br />
Neophaeosphaeria M.P.S. Câmara, M.E. Palm & A.W.<br />
Ramaley, Mycol. Res. 107: 519 (2003). (Leptosphaeriaceae)
Fungal Diversity<br />
Fig. 65 Neomassariosphaeria<br />
typhicola (from IFRD 2018). a<br />
Immersed ascomata gregarious<br />
in the host substrate. b–d<br />
Cylindro-clavate asci embedded<br />
in pseudoparaphyses. Note the<br />
phragmosporous ascospores.<br />
Scale bars: a=200 μm,<br />
b–d=20 μm<br />
Generic description<br />
Habitat terrestrial, parasitic or saprobic. Ascomata small,<br />
forming in leaf spots, scattered or clustered, immersed,<br />
depressed globose, under clypeus, coriaceous. Peridium thin.<br />
Hamathecium of dense, cellular pseudoparaphyses, septate,<br />
embedded in mucilage. Asci 8-spored, bitunicate, fissitunicate<br />
dehiscence not observed, broadly cylindrical to oblong,<br />
with a short furcate pedicel. Ascospores obliquely uniseriate<br />
and partially overlapping, oblong, pale brown, 1-3-septate.<br />
Anamorphs reported for genus: Coniothyrium-like<br />
(Câmara et al. 2003).<br />
Literature: Câmara et al. 2001, 2003; Checa et al. 2002;<br />
Ellis and Everhart 1892.
Fungal Diversity<br />
Type species<br />
Neophaeosphaeria filamentosa (Ellis & Everh.) M.P.S.<br />
Câmara, M.E. Palm & A.W. Ramaley, Mycol. Res. 107:<br />
519 (2003). (Fig. 66)<br />
≡ Leptosphaeria filamentosa Ellis & Everh., J. Mycol. 4:<br />
64 (1888).<br />
Ascomata 115–157 μm high×115–186 μm diam., forming<br />
in leaf spots, scattered or clustered in circular areas, immersed,<br />
depressed globose, with a small ostiolar pore slightly<br />
penetrating above the surface, under clypeus, coriaceous,<br />
papilla not conspicuous (Fig. 66a). Peridium 18–30 μm thick,<br />
composed of large pigmented thin-walled cells of textura<br />
angularis, cells up to 10 μm diam. (Fig. 66c). Hamathecium<br />
of dense, cellular pseudoparaphyses 1.5–2.5 μm broad,<br />
septate, embedded in mucilage (Fig. 66b). Asci 70–105×8–<br />
10 μm (x ¼ 85:3 9:7mm, n=10), 8-spored, bitunicate,<br />
fissitunicate dehiscence not observed, broadly cylindrical to<br />
oblong, with a short, broad, furcate pedicel, 6–13 μm long,<br />
with a small ocular chamber, best seen in immature asci, up to<br />
1.5 μm wide×1 μm high (Fig. 66d, e and f). Ascospores 12–<br />
15×4–5 μm (x ¼ 13:8 5mm, n=10), obliquely uniseriate<br />
and partially overlapping, oblong, yellowish brown, (1-2-)3-<br />
septate, constricted at the primary septum, the upper second<br />
cell often broader than others, verruculose, containing four<br />
refractive globules (Fig. 66g).<br />
Anamorph: Ellis and Everhart (1892) noted that the<br />
“spermogonial stage is a Coniothyrium (C. concentricum)<br />
with small (4 μm), globose, brown sporidia.”<br />
Material examined: USA, New Jersey, Newfield, on<br />
dead parts in living leaves of Yucca filamentosa L., Jul.<br />
1888, Ellis & Everhart (NY, holotype).<br />
Notes<br />
Morphology<br />
Neophaeosphaeria was formally established by<br />
Câmara et al. (2003) by segregating Paraphaeosphaeria<br />
species with 3-4-septate ascospores and anamorphs of<br />
ovoid to ellipsoid, non-septate, brown, verrucose to<br />
punctuate conidia forming from percurrently proliferating<br />
conidiogenous cells. Neophaeosphaeria filamentosa was<br />
selected as the generic type. Currently, four species are<br />
included under Neophaeosphaeria, i.e. N. barrii, N.<br />
conglomerate (M.E. Barr) M.P.S. Câmara, M.E. Palm &<br />
A.W. Ramaley, N. filamentosa and N. quadriseptata (M.<br />
E. Barr) M.P.S. Câmara, M.E. Palm & A.W. Ramaley<br />
(Câmara et al. 2003). At present all species in Neophaeosphaeria<br />
occur on Yucca (Agavaceae).<br />
Phylogenetic study<br />
The four Neophaeosphaeria species form a monophyletic<br />
clade based on both ITS and SSU rDNA<br />
sequences (Câmara et al. 2001; Checa et al. 2002), and<br />
they fall in the group comprising members of Phaeosphaeriaceae<br />
and Leptosphaeriaceae (Câmara et al.<br />
2003). Neophaeosphaeria filamentosa, the generic type<br />
of Neophaeosphaeria, nestedinLeptosphaeriaceae with<br />
low to moderate bootstrap values (Schoch et al. 2009;<br />
Zhang et al. 2009a).<br />
Concluding remarks<br />
The familial status of Neophaeosphaeria under Leptosphaeriaceae<br />
is confirmed, although this family remains<br />
poorly supported in phylogenetic studies.<br />
Nodulosphaeria Rabenh., Klotzschii Herb. Viv. Mycol.,<br />
Edn 2: no. 725 (in sched.) (1858). (Phaeosphaeriaceae)<br />
Generic description<br />
Habitat terrestrial, saprobic or hemibiotrophic. Ascomata<br />
small, immersed to erumpent, globose or subglobose,<br />
black, papillate, ostiolate. Papilla with numerous setae in<br />
the pore-like ostiole. Peridium thin, composed of thickor<br />
thin-walled large cells. Hamathecium of cellular<br />
pseudoparaphyses, septate and branching. Asci 8-<br />
spored, bitunicate, fissitunicate, clavate to cylindroclavate,<br />
with a very short, furcate pedicel and a small<br />
ocular chamber. Ascospores filamentous, hyaline or pale<br />
brown, multi-septate, one of the upper cells swollen.<br />
Anamorphs reported for genus: none.<br />
Literature: Barr 1992a; Holm 1957, 1961; Shoemaker<br />
1984b; Shoemaker and Babcock 1987.<br />
Type species<br />
Nodulosphaeria hirta Rabenh., Klotzschii Herb. Viv.<br />
Mycol., Edn 2: no. 725 (in sched.) (1858). (Fig. 67)<br />
Ascomata 260–330 μm high×260–330 μm diam.,<br />
scattered, or in small groups, immersed to erumpent,<br />
globose or subglobose, black, papillate, ostiolate. Papilla<br />
50–80 μm high, numerous setae occur in the pore-like<br />
ostiole (Fig. 67a and b). Peridium 15–30 μm wide at the<br />
sides, thinner at the base, coriaceous, comprising two types<br />
of cells, outer cells of 1–2 layers of heavily pigmented cells<br />
of textura angularis, cells 6–8 μm diam., cell wall 1.5–<br />
3 μm thick, inner of compressed cells, 5×13–3×8 μm<br />
diam., wall 2–3 μm thick (Fig. 67c). Hamathecium of long<br />
cellular pseudoparaphyses 2–3 μm broad, septate and<br />
branching, mucilage not observed. Asci 100–123×12.5–15<br />
(−17.5) μm (x ¼ 110:8 14:3mm, n=10), 8-spored, bitunicate,<br />
fissitunicate, clavate to cylindro-clavate, with a very<br />
short, furcate pedicel, with a small ocular chamber (to 2 μm<br />
wide×1 μm high) (Fig. 67d). Ascospores 48–63×5–6.5 μm<br />
(x ¼ 55:3 5:6mm, n=10), 4-seriate, filamentous, pale
Fungal Diversity<br />
Fig. 66 Neophaeosphaeria filamentosa (from NY, holotype). a<br />
Ascomata as a circular cluster on the host surface. b Hamathecium of<br />
wide psuedoparaphyses. c Section of peridium comprising cells of textura<br />
angularis. d–f Cylindrical asci with thickened apex. Note the short furcate<br />
pedicel. g Pale brown, 3-septate ascospores. Note the verruculose<br />
ornamentation. Scale bars: a=200 μm, b, c=20 μm, d–g=10 μm
Fungal Diversity<br />
brown, 8-septate, the 4th upper cell broader than the others,<br />
smooth-walled, without sheath (Fig. 67e and f).<br />
Anamorph: none reported.<br />
Material examined: GERMANY, Dresdae, in herbarum<br />
caulibus emortuis perrara, exeunte majo, 1858 (BR<br />
101945–95, holotype, asNodulosphaeria hirta).<br />
Notes<br />
Morphology<br />
The name Nodulosphaeria was first used by Rabenhorst<br />
(1858) but was considered as a synonym of Leptosphaeria<br />
for many years (Clements and Shear 1931). The name was<br />
reinstated by Holm (1957) and was represented by N. hirta,<br />
which was concurrently treated as a synonym of N. derasa<br />
(Berk. & Broome) L. Holm. The most outstanding morphological<br />
characters of Nodulosphaeria were considered to be<br />
apex of ascomata often covered with setae, ascospore with<br />
three or more transverse septa with a supramedian enlarged<br />
cell or elongated to a scolecospore, mostly with terminal<br />
appendages (Barr 1992a; Holm 1961; Shoemaker 1984b).<br />
The ascomata are usually immersed and the peridium<br />
comprises a few layers of brown, relatively thin-walled cells<br />
of textura angularis and textura prismatica similar to those<br />
of Phaeosphaeria. Thus, Nodulosphaeria is likely to be a<br />
member of Phaeosphaeriaceae. However,thisneedstobe<br />
confirmed by molecular analysis. The boundary between<br />
Nodulosphaeria and Ophiobolus is not clear-cut, and the<br />
circumscriptions of them usually depend on the viewpoint of<br />
different mycologists. For instance, Shoemaker (1976) has<br />
assigned some Nodulosphaeria species such as N. erythrospora,<br />
N. fruticum, N. mathieui and N. megalosporus to<br />
Ophiobolus. Subsequently, more species were added to<br />
Nodulosphaeria (Barr 1992a; Shoemaker 1984b; Shoemaker<br />
and Babcock 1987). Currently, more than 60 names are<br />
included in Nodulosphaeria (http://www.mycobank.org/, 06/<br />
2010).<br />
Phylogenetic study<br />
None.<br />
Concluding remarks<br />
All species included in Nodulosphaeria have an inflated<br />
ascospore cell as mentioned above. However, it is likely<br />
that this character would have evolved more than once as it<br />
is probably an adaption for ascospore ejection from the<br />
ascus (Shoemaker 1976). It occurs in Ophiobolus species<br />
and the ascomata of these species are quite dissimilar to<br />
Nodulosphaeria species and their exclusion from Nodulosphaeria<br />
seems warranted. When considering whether a<br />
species belongs in Nodulosphaeria, one must also consider<br />
the ascomata and peridium structure until DNA sequences<br />
are available.<br />
Ohleria Fuckel, Fungi rhenani exsic.: no. 2173 (1868).<br />
(Melanommataceae)<br />
Generic description<br />
Habitat terrestrial, saprobic. Ascomata small to medium size,<br />
solitary, scattered, or in small groups, erumpent to nearly<br />
superficial, papillate, ostiolate. Peridium thin, thicker at the<br />
apex, 1-layered. Hamathecium of dense, long trabeculate<br />
pseudoparaphyses. Asci 8-spored, bitunicate, fissitunicate,<br />
cylindrical, with a short pedicel. Ascospore brown to reddish<br />
brown, broadly to narrowly fusoid, 3-septate, easily separating<br />
into two parts at the primary septum.<br />
Anamorphs reported for genus: Monodictys (Samuels 1980).<br />
Literature: Barr 1990b; Clements and Shear 1931; Patel et<br />
al. 1997; Samuels 1980.<br />
Type species<br />
Ohleria modesta Fuckel, Fungi rhenani exsic. (1868) (Fig.68)<br />
Ascomata 214–357 μm high×285–400 μm diam.,<br />
solitary, scattered, or in small groups of 2–3, erumpent to<br />
nearly superficial, coriaceous, with basal wall remaining<br />
immersed in host tissue, broadly or narrowly conical, with a<br />
flattened base not easily removed from the substrate, black;<br />
apex with a conical protruding papilla and an often porelike<br />
ostiole (Fig. 68a). Peridium 22–53 μm thick laterally,<br />
thicker at the apex, 1-layered, composed of heavily pigmented<br />
thick-walled cells of textura angularis, cells to 7 μm diam.,<br />
cell wall 1.5–3 μm thick, apex cells smaller and walls<br />
thicker, base cells walls thinner (Fig. 68b). Hamathecium of<br />
dense, long trabeculate pseudoparaphyses 1–2 μm broad,<br />
septate, branching and anastomosing (Fig. 68c). Asci 90–<br />
130×(5.5-)7–10 μm (x ¼ 107:3 8mm, n=10), 8-spored,<br />
with a short pedicel up to 20 μm long, bitunicate,<br />
fissitunicate, cylindrical, with a small ocular chamber (to<br />
1.5 μm wide×1.5 μm high) (Fig. 68c, d and e). Ascospore<br />
15–22×4–5 μm (x ¼ 20 4:4mm, n=10), biseriate near the<br />
top and uniseriate at the base, broadly fusoid to fusoid with<br />
broadly to narrowly rounded ends, brown to reddish brown,<br />
3-septum, deeply constricted at the median septum and<br />
breaking into two conical partspores, no constriction at the<br />
secondary septum, smooth (Fig. 68d and e).<br />
Anamorph: none reported.<br />
Material examined: GERMANY, on decorticated,<br />
decaying roots of Fagus sylvatica, very rare, collected in<br />
autumn (G: F. rh. 2173, isotype).<br />
Notes<br />
Morphology<br />
Ohleria is characterized by its subglobose to conic<br />
ascomata, produced on decorticated woody substrates, as
Fungal Diversity<br />
Fig. 67 Nodulosphaeria hirta (from BR 101945–95, holotype). a<br />
Appearance of ascomata on the host surface. b Vertical section of an<br />
ascoma. Note the setae at the apex and in the ostiole. c Section of a<br />
partial peridium. Note the outer layer cells of textura angularis and<br />
inner layer compressed cells. d Squash mount showing asci in<br />
pseudoparaphyses. e, f. The light brown filiform ascospores. Scale<br />
bars: a=0.5 mm, b=100 μm, c=50 μm, d=20 μm, e, f=10 μm<br />
well as its brown and phragmosporous ascospores which<br />
break into two parts at the median septum (Samuels 1980).<br />
Some species of Ohleria are widespread. For instance, O.<br />
brasiliensis is reported from New Zealand, Brazil as well as<br />
United States (Samuels 1980). Ohleria has been considered<br />
closely related to Sporormia and Preussia based on the<br />
ascosporic characters, and several species of Ohleria, suchas<br />
O. aemulans Rehm, O. haloxyli Kravtzev, O. silicata<br />
Kravtzev and O. kravtzevii Schwarzman, have been transferred<br />
to these genera. Clements and Shear (1931) treated<br />
Ohleria as a synonym of Ohleriella, despite the fact that<br />
Ohleriella is a coprophilous fungus. When the ascomata and<br />
habitats are considered, Ohleria seems closely related to<br />
Melanomma and Trematosphaeria (Samuels 1980).<br />
Phylogenetic study<br />
None.<br />
Concluding remarks<br />
To some degree, habitats show phylogenetic significance<br />
(Zhang et al. 2009a). Thus, Ohleria seems less likely related<br />
to Sporormia and Preussia. But its relationship with<br />
Melanomma is uncertain, because of their differences in<br />
hamathecium and ascospores.
Fungal Diversity<br />
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<br />
in pseudoparaphyses. d, e Cylindrical asci with short pedicels. Scale bars: a=1 mm, b, c=50 μm, d, e=20 μm<br />
Ohleriella Earle, Bull N Y Bot Gard 2: 349 (1902).<br />
(Delitschiaceae)<br />
Generic description<br />
Habitat terrestrial, saprobic. Ascomata medium to large,<br />
immersed, erumpent to nearly superficial, scattered or in<br />
small groups, usually with a wide papilla, ostiolate,<br />
coriaceous. Peridium composed of small pigmented cells<br />
of textura angularis. Asci 8-spored or fewer, cylindroclavate,<br />
with a furcate pedicel. Hamathecium of trabeculate<br />
pseudoparaphyses. Ascospores brown to dark brown,<br />
cylindrical to nearly clavate with broadly to narrowly round<br />
ends, multi-septate, easily broken into partspores, smooth,<br />
with elongated germ slit in each cell.<br />
Anamorphs reported for genus: none.<br />
Literature: Ahmed and Cain 1972; von Arx and Müller<br />
1975; Barr 1990a; Clements and Shear 1931.<br />
Type species<br />
Ohleriella neomexicana Earle, Bull N Y Bot Gard 2: 349<br />
(1902). (Fig. 69)<br />
Ascomata 330–420 μm high×400–575 μm diam.,<br />
solitary, scattered, or in small groups, immersed to<br />
erumpent, to nearly superficial, with basal wall remaining
Fungal Diversity<br />
immersed in host tissue, coriaceous, globose or subglobose,<br />
usually a somewhat thick, short papilla, up to<br />
100 μm high, with a pore-like ostiole (Fig. 69a). Peridium<br />
27–35 μm thick laterally, up to 55 μm thick at the apex, 1-<br />
layered, composed of small pigmented cells of textura<br />
angularis, cells up to 5×8 μm diam., cell wall 1.5–2 μm<br />
thick, apex cells smaller and walls thicker (Fig. 69b).<br />
Hamathecium of dense, long trabeculate pseudoparaphyses,<br />
1–1.5 μm broad, anastomosing and branching between<br />
and above the asci. Asci 150–208×17.5–25 μm<br />
(x ¼ 182:5 22mm, n=10), 8-spored, bitunicate, fissitunicate,<br />
cylindrical to cylindro-clavate, with a narrowed, furcate,<br />
thin pedicel, 15–55 μm long, 2–3.5 μm broad, with a large<br />
truncate ocular chamber best seen in immature asci (to 4 μm<br />
wide×3 μm high) (Fig. 69e, f and g). Ascospores 55–72.5×<br />
10–12 μm (x ¼ 63 10:4mm, n=10), 3–4 seriate to uniseriate<br />
near the base, cylindrical to clavate, with broadly to<br />
narrowly rounded ends, brown, 6–7 transversesepta,easily<br />
separating into partspores, with germ slits, central partspores<br />
of the ascospores shorter than broad, rectangular in vertical<br />
section, round in transverse section, 7–8×6–10 μm diam.,<br />
apical cells usually longer than broad, 11–17.5×6–7 μm<br />
diam. (Fig. 69c and d).<br />
Anamorph: none reported.<br />
Material examined: USA, Albuquerque, Bernalillo Co.,<br />
New Mexico, dry gravelly hill, on wood, 29 Nov. 1901, T.<br />
S.A. Cockerell (NY, holotype).<br />
Notes<br />
Morphology<br />
Ohleriella was formally established by Earle (1902) based<br />
on its “medium to large ascomata with a wide papilla,<br />
relatively wide peridium, cylindro-clavate asci, brown to<br />
deep brown multi-septate ascospore, with elongated germ slit<br />
on each cell”, and was monotypified by O. neomexicana<br />
(Barr 1990a). Ohleriella subsequently has been treated as a<br />
synonym of Ohleria, Sporormiella or Preussia (Ahmed and<br />
Cain 1972; vonArxandMüller1975; Clements and Shear<br />
1931). Spororminula tenerifae, the generic type of Spororminula,<br />
was assigned to Ohleriella, thus Spororminula was<br />
treated as a synonym of Ohleriella (Barr 1990a). Two new<br />
species were introduced by Barr (1990a) from North America.<br />
Currently, three species are included in this genus, i.e. O.<br />
herculean (Ellis & Everh.) M.E. Barr, O. neomexicana and O.<br />
nudilignae M.E. Barr & Malloch (http://www.index<br />
fungorum.org; http://www.mycobank.org, 01/03/2009).<br />
The generic type, O. neomexicana, is morphologically<br />
similar to the coprophilous genus Sporormiella, but is<br />
saprobic on grass stems.<br />
Phylogenetic study<br />
None.<br />
Concluding remarks<br />
Although we maintain Ohleriella as a separate genus<br />
here, its saprobic habitat on grasses and similarity to the<br />
coprophilous Sporormiella may indicate a close evolutionary<br />
relationship, with the grass saprobic possibly<br />
being an early relative of the coprophilous Sporormiella.<br />
Alternatively, the species/genera may simply occupy<br />
different ecological niches (i.e. dead grass vs dead grass<br />
in dung). Molecular studies are needed to resolve this<br />
issue.<br />
Ophiobolus Reiss, Hedwigia 1:27 (1854).<br />
(Phaeosphaeriaceae)<br />
Generic description<br />
Habitat terrestrial, saprobic or hemibiotrophic. Ascomata<br />
medium-sized, solitary, scattered, or in groups, globose or<br />
pyriform, coriaceous, black, papillate, ostiolate, periphysate.<br />
Peridium thin, thicker near the apex, thinner at the base.<br />
Hamathecium of long cellular pseudoparaphyses, septate,<br />
anastomosing or branching not observed. Asci 8-spored,<br />
bitunicate, fissitunicate dehiscence not observed, cylindrical,<br />
with a short, furcate pedicel. Ascospores filamentous,<br />
narrower toward the lower end, pale brown, multiseptate,<br />
separating into two partspores from the middle<br />
septum, from the breaking point, the second cell of each<br />
partspore enlarged.<br />
Anamorphs reported for genus: Coniothyrium-like,<br />
Rhabdospora, Phoma-like and Scolecosporiella (Hyde et<br />
al. 2011; Shoemaker 1976; Sivanesan 1984).<br />
Literature: Holm 1948, 1957; Müller 1952; Reiss 1854;<br />
Shoemaker 1976; Sivanesan 1984.<br />
Type species<br />
Ophiobolus disseminans Reiss, Hedwigia 1:27 (1854)<br />
(Fig. 70).<br />
Ascomata 220–380 μm high×290–430 μm diam., solitary,<br />
scattered, or in groups often arranged in a row, immersed with<br />
a protruding papilla, globose, pyriform, coriaceous, black,<br />
periphysate. Papilla 40–90 μm high, with a pore-like ostiole<br />
(Fig. 70a and b). Peridium 40–55 μm wide at the sides, up to<br />
70 μm thick at the apex, thinner at the base, comprising two<br />
cell types, outer layer composed of small heavily pigmented<br />
thick-walled cells of textura angularis, cells 2–5 μm diam.,<br />
cell wall 2–3 μm thick, apex cells smaller and walls thicker,<br />
inner layer composed of lightly pigmented or hyaline thinwalled<br />
cells of textura angularis, 5–7 μm diam., wall 1.5–<br />
2 μm thick, merging with pseudoparaphyses (Fig. 70c).<br />
Hamathecium of long cellular pseudoparaphyses, 2–3 μm<br />
broad, septate, anastomosing or branching not observed<br />
(Fig. 70e). Asci 150–195×8–12.5 μm (x ¼ 169:5 10:7mm,
Fungal Diversity<br />
Fig. 69 Ohleriella neomexicana (NY, holotype). a Ascoma scattering<br />
on the host surface. b Section of a partial peridium. Note the small<br />
cells of textura angularis. c Ascospore in ascus. d Ascospore breaking<br />
into part spores. Note the sigmoid germ slit. e Dehiscent ascus. f, g<br />
Asci with short pedicels. Scale bars: a=100 μm, b=50 μm, c–g=<br />
10 μm
Fungal Diversity<br />
n=10), 8-spored, bitunicate, fissitunicate dehiscence not<br />
observed, cylindrical but narrowing towards the base, with a<br />
short, furcate pedicel which is 10–25 μm long, ocular<br />
chamber not observed (Fig. 70d and e). Ascospores 110–<br />
160×2.5–4 μm (x ¼ 135:3 3mm, n=10), filamentous,<br />
narrower toward the lower end, pale brown, 22–30-<br />
septate, separating into two partspores from the middle<br />
septum, from the breaking point the second cell of each<br />
partspore enlarged.<br />
Anamorph: none reported.<br />
Material examined: GERMANY, near Kassel, on dead<br />
stem of Cirsium arvense (L.) Scop., Spring 1853 (BPI-<br />
629021, type).<br />
Notes<br />
Morphology<br />
Ophiobolus was established by Reiss (1854) as a<br />
monotypic genus represented by O. disseminans based on<br />
its “Perithecia discreta, ostiolis prominentibus: sporae ascis<br />
inclusae, binatae, filliformes, multiseptatae”.<br />
A broad generic concept was adopted for the genus by<br />
Holm (1948) and Müller (1952). Shoemaker (1976)surveyed<br />
Canadian species of Ophiobolus using the broad concept of<br />
Holm (1948) and Müller (1952). A narrower generic concept<br />
was used by Holm (1957), which only included species with<br />
ascospores separating into two halves. Holm (1957) assigned<br />
species with enlarged ascospore cells to Nodulosphaeria,and<br />
those with long spirally coiled ascospores to Leptospora<br />
(Shoemaker 1976). This left only three species accepted<br />
under Ophiobolus (Holm 1957), although this concept has<br />
rarely been followed with new species recently being<br />
described (Raja and Shearer 2008).<br />
Walker (1980) provided a detailed description from the<br />
type material and dealt with many species of scolecospored<br />
fungi that had been placed in Ophiobolus by Saccardo<br />
(1883). Thus, currently several Ophiobolus sensu lato<br />
species are separated into Acanthophiobolus, Entodesmium,<br />
Leptosphaeria and Leptospora. Ophiobolus sensu lato<br />
contains about 300 species names (Sivanesan 1984; http://<br />
www.mycobank.org/, 04/02/2009).<br />
Phylogenetic study<br />
Ophiobolus fulgidus (Cooke & Peck) Sacc. (as Leptosphaeria<br />
fulgida (Cooke & Peck) M. E. Barr in Dong et al.<br />
1998) lacks support in the clade of Leptosphaeriaceae<br />
(Dong et al. 1998). We expect it may closely related to<br />
Phaeosphaeriaceae.<br />
Concluding remarks<br />
We agree from morphological data that Ophiobolus<br />
should comprise species that have filamentous spores that<br />
break easily into two halves at the central septum, with the<br />
second cell on either side being swollen (Walker 1980) and<br />
that the genus presently comprises three species (i.e. O.<br />
anthrisci (L. Holm) L. Holm, O. ophioboloides (Sacc.) L.<br />
Holm and O. acuminatus). All other Ophiobolus species<br />
need to be re-examined and should be placed in other<br />
genera such as Nodulosphaeria and Leptospora. The genus<br />
is in need of revision and molecular phylogenetic study.<br />
Ophiosphaerella Speg., Anal. Mus. nac. Hist. nat. B. Aires<br />
19: 401–402 (1909). (Phaeosphaeriaceae)<br />
Generic description<br />
Habitat terrestrial, saprobic or hemibiotrophic. Ascomata<br />
small- to medium-sized, solitary or scattered, immersed,<br />
globose or subglobose, papillate, ostiolate. Peridium thin.<br />
Hamathecium of dense, filliform, septate pseudoparaphyses.<br />
Asci bitunicate, fissitunicate dehiscence not observed, cylindrical<br />
often narrower near the base, with a short furcate<br />
pedicel. Ascospores filamentous, pale brown, multi-septate.<br />
Anamorphs reported for genus: Scolecosporiella (Farr et<br />
al. 1989).<br />
Literature: von Arx and Müller 1975; Schoch et al. 2006,<br />
2009; Spegazzini 1909; Walker 1980; Wetzel et al. 1999;<br />
Zhang et al. 2009a.<br />
Type species<br />
Ophiosphaerella graminicola Speg., Anal. Mus. nac. Hist.<br />
nat. B. Aires 19: 401 (1909). (Fig. 71)<br />
Ascomata 280–325 μm high×250–300 μm diam., solitary<br />
or scattered, immersed with a short papilla protruding out of<br />
the substrate, globose or subglobose, often laterally flattened,<br />
dark brown to black, papillate, papilla ca. 100μm high, 140–<br />
180 μm broad, disk-like in appearance from above, periphysate<br />
(Fig. 71a and b). Peridium 11–25 μm wide, thicker near<br />
the apex, comprising two cell types of small cells, outer wall<br />
composed 6–10 layers of lightly brown flattened cells of<br />
textura angularis, inner layer composed of paler and thinwalled<br />
cells, both layers thicker near the apex (Fig. 71b).<br />
Hamathecium of dense, long pseudoparaphyses 0.8–1.5 μm<br />
broad near the apex, septate, 2–3 μm broad between the asci.<br />
Asci 105–135×5.5–10 μm (x ¼ 118:5 7mm, n=10), 8-<br />
spored, bitunicate, cylindrical and narrower near the<br />
base, with a short, furcate pedicel, up to 30 μm long,<br />
small inconspicuous ocular chamber (to 1.5 μm wide×<br />
1 μm high) (Fig. 71c, d, e and f). Ascospores 100–125×<br />
1.8–2.2 μm (x ¼ 118 2mm, n=10), filamentous, pale<br />
brown, 12–20 septa, smooth-walled.<br />
Anamorph: none reported.<br />
Material examined: ARGENTINA, Tucumán, on leaf<br />
sheath of Leptochloa virgata (L.) P. Beauv., 14 Apr. 1906,<br />
C. Spegazzini (LPS 858, holotype).
Fungal Diversity<br />
Fig. 70 Ophiobolus<br />
disseminans (from BPI-629021,<br />
type). a Immersed ascomata<br />
scattered on the host surface.<br />
Note the erumpent papilla. b<br />
Section of an ascoma. c. Section<br />
of a partial peridium. Note the<br />
thick-walled outer layer and<br />
thin-walled inner layer (orange<br />
colour due to DIC). d Ascus<br />
with a short furcate pedicel. e<br />
Squash mount showing asci in<br />
pseudoparaphyses. Scale bars:<br />
a=0.5 mm, b=100 μm,<br />
c=50 μm, d, e=20 μm<br />
Notes<br />
Morphology<br />
Ophiosphaerella was introduced by Spegazzini (1909)<br />
who described and illustrated a single new species, O.<br />
graminicola, and thus the genus was validly published<br />
(Walker 1980, p. 70). After checking the type specimen,<br />
Petrak and Sydow (1936) transferred the generic type to<br />
Ophiobolus graminicolus (Speg.) Petrak & Syd, and<br />
assigned Ophiosphaerella as a synonym of Ophiobolus.<br />
This was followed by von Arx and Müller (1975).<br />
Ophiosphaerella differs from Phaeosphaeria by its scolecospores<br />
without swollen cells or appendages, and from<br />
Ophiobolus by its ascospores without swollen cells or<br />
separating into partspores, thus was kept as a separating<br />
genus (Eriksson 1967a; Walker 1980).<br />
Phylogenetic study<br />
Ophiosphaerella forms a monophyletic group as a sister<br />
group of Phaeosphaeria located in Phaeosphaeriaceae (Schoch<br />
et al. 2006, 2009; Wetzel et al. 1999; Zhang et al. 2009a).<br />
Concluding remarks<br />
Numerous Ophiobolus species are likely to belong in<br />
Ophiosphaerella. The two genera are distinguished as<br />
Ophiobolus sensu Shoemaker (1976) has swollen central<br />
cells or breaking into partspores or with long spirally<br />
coiled ascospores, and Ophiosphaerella (sensu Walker
Fungal Diversity<br />
1980) has scolecospores without swollen central cells or<br />
breaking into partspores.The recent introduction of Ophiobolus<br />
shoemakeri Raja & Shearer (Raja and Shearer 2008)<br />
is probably incorrect since the ascospores do not split up<br />
into partspores and there is no swelling above septum<br />
either. In particular, its freshwater habitat also distinguishes<br />
it from other species of Ophiobolus. Like<br />
Ophiobolus, Ophiosphaerella is in need of phylogenetic<br />
analysis but appears to be closely related to Phaeosphaeriaceae<br />
(Schoch et al. 2006).<br />
Ostropella (Sacc.) Höhn., Annls mycol. 16: 144 (1918).<br />
(<strong>Pleosporales</strong>, genera incertae sedis)<br />
≡ Ostropa subgen. Ostropella Sacc., Syll. fung. (Abellini)<br />
2: 805 (1883).<br />
Generic description<br />
Habitat terrestrial, saprobic. Ascomata large, erumpent<br />
to superficial, solitary or gregarious, globose to subglobose,<br />
with broad and compressed papilla and slit-like<br />
ostiole. Peridium carbonaceous. Hamathecium of dense,<br />
long trabeculate pseudoparaphyses, anastomosing and<br />
branching, rarely septate, embedded in mucilage. Asci<br />
clavate with very long and thin and furcate pedicels.<br />
Ascospores pale brown, ellipsoid to fusoid, 1-septate,<br />
constricted.<br />
Anamorphs reported for genus: none.<br />
Literature: Barr 1990a; Chesters and Bell 1970; Holm and<br />
Yue 1987; Huhndorf 1993; Müller and von Arx 1962;<br />
Müller and Dennis 1965; Saccardo 1883.<br />
Fig. 71 Ophiosphaerella<br />
graminicola (from LPS 858,<br />
holotype). a Ascomata on the<br />
host surface. Note the protruding<br />
disk-like papilla. b Section<br />
of an ascoma. c Asci in<br />
pseudoparaphyses with short<br />
pedicels. d–f Cylindrical asci<br />
with short pedicels. Scale bars:<br />
a=0.5 mm,<br />
b=100 μm, c–f =10 μm
Fungal Diversity<br />
Type species<br />
Ostropella albocincta (Berk. & M.A. Curtis) Höhn., Annls<br />
mycol. 16: 144 (1918). (Fig. 72)<br />
≡ Ostropa albocincta Berk & M.A. Curtis, in Berkeley,<br />
J. Linn. Soc., Bot. 10: 372 (1868).<br />
Ascomata 1000–1730 μm high×1050–1450 μm diam.,<br />
scattered to gregarious, erumpent to superficial, globose to<br />
subglobose, roughened, often covered with white crustose<br />
covering, with subiculum, with a broad compressed papilla<br />
and long and slit-like ostiole (Fig. 72a). Peridium 100–<br />
250 μm thick, not of uniform thickness throughout entire wall<br />
area, composed of two cell types, one is of lightly pigmented<br />
thin-walled cells of textura prismatica, cells up to 17×3 μm<br />
diam., cell wall
Fungal Diversity<br />
Ascostromata black, immersed, penetrating into the<br />
substrate with dark brown hyphae. Ascomata up to<br />
680 μm high×540 μm diam., solitary, immersed or<br />
erumpent, subglobose to pyriform, subiculate or nonsubiculate,<br />
papillate or epapillate, ostiolate, periphysate, carbonaceous<br />
(Fig. 73a). Peridium thick. Hamathecium of long<br />
trabeculate pseudoparaphyses, 1–1.5 μm broad. Asci 90–<br />
130×12–17 μm (x ¼ 116 15mm, n=10), bitunicate, fissitunicate,<br />
cylindrical, 8-spored, uniseriate, with a short<br />
furcate pedicel, without apical apparatus (Fig. 73b, c and<br />
d). Ascospores 17.5–25×10–12.5 μm (x ¼ 21 11mm, n=<br />
10), ellipsoid to broadly fusoid with broadly rounded ends,<br />
1-septate, constricted at the septum, hyaline, smooth-walled,<br />
surrounded by a gelatinous sheath that contracts to form a<br />
lateral, lentiform, viscous appendage over the septum, 7.5–<br />
12.5 μm diam., 1–3 μm thick (Fig. 73e, f, g and h).<br />
Anamorph: none reported.<br />
Material examined: USA, Florida, Charlotte Harbor in<br />
Punta Garda, 10 Jan. 1964, leg., det. J. J. Kohlmeyer (Herb.<br />
J. Kohlmeyer No. 1720).<br />
Notes<br />
Morphology<br />
Paraliomyces was introduced to accommodate the<br />
marine fungus P. lentifer, which is characterized by<br />
immersed ascomata produced within the ascostroma,<br />
trabeculate pseudoparaphyses, cylindrical, 8-spored asci,<br />
ellipsoidal, hyaline, 1-septate ascospores surrounded by a<br />
gelatinous sheath, which forms a lentiform, viscous<br />
appendage over the septum (Kohlmeyer 1959).<br />
Phylogenetic study<br />
Based on analysis of SSU sequences, Paraliomyces<br />
lentifer nested within <strong>Pleosporales</strong>, but its familial status<br />
was left undetermined (Tam et al. 2003).<br />
Concluding remarks<br />
None.<br />
Phaeosphaeria I. Miyake, Bot. Mag., Tokyo 23: 93 (1909).<br />
(Phaeosphaeriaceae)<br />
Generic description<br />
Habitat terrestrial, saprobic or hemibiotrophic. Ascomata<br />
small, solitary, scattered, or in small groups, immersed,<br />
globose, subglobose, wall black. Apex with a pore-like<br />
ostiole. Peridium thin. Hamathecium of dense, filliform,<br />
septate pseudoparaphyses. Asci 8-spored, bitunicate, fissitunicate,<br />
broadly cylindrical to narrowly fusoid, with a<br />
short pedicel. Ascospores fusoid to narrowly fusoid, pale<br />
brown to brown, 3-septate.<br />
Fig. 72 Ostropella albocincta (K(M): 143941, syntype). a Ascomata<br />
gregarious on host surface. b Section of the partial peridium. Note the<br />
peridium comprising two cell types and the whitening tissue<br />
(arrowed). c Pseudoparaphyses. d, e Asci with long pedicel. f–<br />
h Ascospores, which are strongly constricted at the central septum.<br />
Scale bars: a=1 mm, b=100 μm, d, e, h=20 μm, c, g, f=10 μm<br />
Anamorphs reported for genus: Amarenographium,<br />
Hendersonia-like, Phaeoseptoria, Scolecosporiella and<br />
Stagonospora (Hyde et al. 2011; Leuchtmann 1984;<br />
Shoemaker and Babcock 1989b).<br />
Literature: von Arx and Müller 1975; Câmara et al.<br />
2002; Eriksson 1967a, 1981; Holm 1957; Khashnobish and<br />
Shearer 1996; Leuchtmann 1984; Miyake 1909; Shoemaker<br />
and Babcock 1989b.<br />
Type species<br />
Phaeosphaeria oryzae I. Miyake, Bot. Mag., Tokyo 23:<br />
136 (1909). (Fig. 74)<br />
Ascomata 120–140 μm high×100–140 μm diam.,<br />
solitary, scattered, or in small groups, immersed, globose,<br />
subglobose, wall black, forming black spots on the leaves<br />
of hosts (Fig. 74a). Apex with a pore-like ostiole. Peridium<br />
4–8 μm wide at the sides, composed of heavily pigmented<br />
thin-walled cells of textura angularis, cells 2–2.5×3–5 μm<br />
diam., cell wall less than 1 μm thick (Fig. 74b). Hamathecium<br />
of dense, long cellular pseudoparaphyses 2–2.5 μm<br />
broad, embedded in mucilage, rarely branched, septate. Asci<br />
53–80(−90)×7–10 μm (x ¼ 65:3 8:3mm, n=10), 8-<br />
spored, bitunicate, fissitunicate, broadly cylindrical to<br />
narrowly fusoid, with a short pedicel which is ca. 8μm<br />
long, with a small ocular chamber and an inconspicuous<br />
apical apparatus (to 2 μm wide×1 μm high) (Fig. 74c, d<br />
and e). Ascospores 17– 22(− 28)×4– 5 μ m<br />
(x ¼ 20:5 4:6mm, n=10), obliquely uniseriate, partially<br />
overlapping or biseriate, narrowly fusoid with rounded ends,<br />
pale brown, 3-septate, slightly constricted at primary septum,<br />
granulate (Fig. 74f and g).<br />
Anamorph: none reported.<br />
Material examined: JAPAN, Suruya, Shizuoka, on the<br />
leaves of Oryza sativa, Sept. 1907 (S nr F9572, F9573,<br />
lectotype).<br />
Notes<br />
Morphology<br />
Phaeosphaeria was introduced by Miyake (1909), but<br />
was regarded as a synonym of Leptosphaeria for a long<br />
time. Holm (1957), however, reinstated Phaeosphaeria,<br />
assigning some Leptosphaeria sensu lato species with<br />
relatively small ascomata and which occurred on monocotyledons<br />
to Phaeosphaeria. Although this division based<br />
b
Fungal Diversity
Fungal Diversity<br />
Fig. 73 Paraliomyces lentifer (from Herb. J. Kohlmeyer No. 1720). a<br />
Section of an immersed ascoma. b Eight-spored cylindrical asci<br />
embedded in pseudoparaphyses. c, d Cylindrical asci with short<br />
on host range is considered unnatural by some workers<br />
(Dennis 1978; Sivanesan 1984), it has been widely<br />
accepted (von Arx and Müller 1975; Eriksson 1967a;<br />
Hedjaroude 1969; Shoemaker and Babcock 1989b).<br />
Eriksson (1981) further revised the generic concept of<br />
pedicels. e–h One-septate hyaline ascospores. Scale bars: a=100 μm,<br />
b–d=20 μm, e–h=10 μm<br />
Phaeosphaeria by including dictyosporous taxa as well as<br />
some perisporium taxa. Phaeosphaeria was further divided<br />
into six subgenera, i.e. Ovispora, Fusispora, Phaeosphaeria,<br />
Spathispora, Vagispora and Sicispora, based on<br />
differences in ascospore shape and the number of septa
Fungal Diversity<br />
(Shoemaker and Babcock 1989b). Phaeosphaeria species<br />
are usually associated or parasitic on annual monocots,<br />
such as Cyperaceae, Juncaceae or Poaceae but have also<br />
been recorded as saprobes and on dicotyledons (e.g. P.<br />
viridella and P. vagans).<br />
Phylogenetic study<br />
The separation of Phaeosphaeria from Leptosphaeria<br />
sensu stricto was supported by phylogenetic studies based<br />
on ITS sequences. The peridium structure, pseudoparenchymatous<br />
cells in Phaeosphaeria versus scleroplectenchymatous<br />
cells in Leptosphaeria had phylogenetic<br />
significance in the distinction between these two genera,<br />
while the subgenus division was not supported by the<br />
phylogenetic results (Câmara et al. 2002; Morales et al.<br />
1995). The familial status of both Phaeosphaeriaceae and<br />
Leptosphaeriaceae was verified by multigene phylogenetic<br />
analysis (Schoch et al. 2009; Zhang et al. 2009a).<br />
Concluding remarks<br />
Phaeosphaeria was originally thought to be a synonym of<br />
Leptosphaeria (Müller 1950; Munk1957), however, molecular<br />
analysis has shown these two genera differ with<br />
Phaeosphaeria having pseudoparenchymatous peridium, Stagonospora-like<br />
anamorph and mostly monocotyledonous<br />
hosts and Leptosphaeria having scleroplectenchymatous<br />
peridium, Phoma-like anamorph and mostly dicotyledonous<br />
hosts (Câmara et al. 2002;Schochetal.2009; Shoemaker and<br />
Babcock 1989b; Zhang et al. 2009a). It is now recognized<br />
that Phaeosphaeria is the type genus of Phaeosphaeriaceae<br />
and related genera include Entodesmium and<br />
Setomelanomma and probably Ophiosphaerella (Schoch<br />
et al. 2009; Zhangetal.2009a). Paraphaeosphaeria was<br />
introduced as an off-shoot of Phaeosphaeria and differs<br />
in ascospore shape and septation as well as anamorphic<br />
stages (Eriksson 1967a, b). Similarly, Nodulosphaeria<br />
was recently reinstated and differs from Phaeosphaeria<br />
because of setae over the apex as well as its ascospores<br />
with swelling supramedian cells and terminal appendages<br />
(Holm 1957, 1961). While the newly reinstated Phaeosphaeria<br />
was confined to monocotyledons and particularly<br />
grasses, there are now many species that have been<br />
described from dicotyledons (Farr et al. 1989). Whether<br />
these taxa form a monophyletic group needs to be<br />
investigated with fresh collections and molecular data.<br />
Phaeosphaeriopsis M.P.S. Câmara, M.E. Palm & A.W.<br />
Ramaley, Mycol. Res. 107: 519 (2003). (Phaeosphaeriaceae)<br />
Generic description<br />
Habitat terrestrial, saprobic or hemibiotrophic? Ascomata<br />
small, scattered or in small groups, immersed,<br />
globose, subglobose. Peridium thin, comprising one cell<br />
type of textura angularis. Hamathecium of dense, wide<br />
cellular pseudoparaphyses. Asci 8-spored, bitunicate,<br />
cylindrical to broadly fusoid, with a short pedicel and a<br />
small ocular chamber. Ascospores obliquely uniseriate<br />
and partially overlapping to biseriate even triseriate,<br />
cylindrical, pale brown, multi-septate, primary septum<br />
submedian, with or without constriction, verrucose or<br />
baculate.<br />
Anamorphs reported for genus: Coniothyrium-like,<br />
Phaeostagonospora (Câmara et al. 2003).<br />
Literature: Câmara et al. 2003.<br />
Type species<br />
Phaeosphaeriopsis glaucopunctata (Grev.) M.P.S. Câmara,<br />
M.E. Palm & A.W. Ramaley, Mycol. Res. 107: 519 (2003).<br />
(Fig. 75)<br />
≡ Cryptosphaeria glaucopunctata Grev. Fl. Edin.: 362<br />
(1824).<br />
Ascomata 120–150 μm high×140–200 μm diam.,<br />
scattered, or in small groups, immersed, globose, subglobose<br />
(Fig. 75a). Peridium 10–25 μm wide, comprising one<br />
type of cells, composed of thick-walled cells of textura<br />
angularis, cells 4–9 μm diam., cell wall 2–3 μm thick,<br />
almost equal in thickness. Hamathecium of dense, wide<br />
cellular pseudoparaphyses, 3–5 μm broad. Asci (50-)60–<br />
110×10–15 μm (x ¼ 82:3 12mm, n =10), 8-spored,<br />
bitunicate, fissitunicate dehiscence not observe, cylindrical<br />
to broadly fusoid, with a short pedicel, with a small<br />
ocular chamber (to 0.8 μm wide×1 μm high) (Fig. 75b).<br />
Ascospores 18–28×5–7.5 μm (x ¼ 23:5 6:2mm, n=10),<br />
obliquely uniseriate and partially overlapping to biseriate<br />
even triseriate, cylindrical, pale brown, 4(−5)-<br />
septate, without constriction or slightly constricted at<br />
the basal septum, the forth cell from the apex usually<br />
slightly inflated, the basal cell often longer, baculate<br />
(Fig. 75c, d, e and f).<br />
Anamorph: none reported.<br />
Material examined: UK, Epping, Sept. 1863 (E, M.C.<br />
Cooke 166, barcode: E00074286).<br />
Notes<br />
Morphology<br />
Phaeosphaeriopsis was introduced to accommodate<br />
some species of Paraphaeosphaeria based on both morphological<br />
characters and results of SSU rDNA sequence<br />
analyses (Câmara et al. 2003). Most of the Phaeosphaeriopsis<br />
species occur on the Agavaceae, although P.<br />
glaucopunctata occurs on Liliaceae (Ruscus). Phaeosphaeriopsis<br />
is characterized by having uni- or multioculate<br />
stromata and 4- or 5-septate ascospores. Although the
Fungal Diversity<br />
Fig. 74 Phaeosphaeria oryzae<br />
(from S nr F9572, F9573, lectotype).<br />
a Appearance of ascomata<br />
on the host surface. b<br />
Section of an ascoma. c Squash<br />
mount showing asci in pseudoparaphyses.<br />
Note that asci with<br />
short pedicels. d, e Asci with<br />
short pedicels. F, G. Light<br />
brown 3-septate ascospores.<br />
Scale bars: a=100 μm, b–g=<br />
10 μm<br />
morphological characters of Phaeosphaeriopsis species is<br />
more diverse than those of Paraphaeosphaeria sensu<br />
stricto or Neophaeosphaeria, the ITS sequences are more<br />
similar to each other than those of the other two genera<br />
(Câmara et al. 2003). Currently, Phaeosphaeriopsis comprises<br />
seven species, namely P. agavensis (A.W. Ramaley,<br />
M.E. Palm & M.E. Barr) M.P.S. Câmara, M.E. Palm & A.<br />
W. Ramaley, P. amblyospora A.W. Ramaley, P. glaucopunctata,<br />
P. musae Arzanlou & Crous, P. nolinae (A.W.<br />
Ramaley) M.P.S. Câmara, M.E. Palm & A.W. Ramaley, P.<br />
obtusispora (Speg.) M.P.S. Câmara, M.E. Palm & A.W.<br />
Ramaley and P. phacidiomorpha (Ces.) D.F. Farr & M.E.<br />
Palm (http://www.mycobank.org/, 06/2010).<br />
Phylogenetic study<br />
The generic type of Phaeosphaeriopsis, P. glaucopunctata,<br />
located in Phaeosphaeriaceae based on SSU rDNA<br />
sequences (Câmara et al. 2003). Phaeosphaeriopsis musae<br />
is also shown to belong to Phaeosphaeriaceae in recent<br />
phylogenetic studies (Schoch et al. 2009; Plate 1).<br />
Concluding remarks<br />
None.<br />
Platysporoides (Wehm.) Shoemaker & C.E. Babc., Can. J.<br />
Bot. 70: 1648 (1992). (Pleosporaceae)<br />
≡ Pleospora subgenus Platysporoides Wehmeyer, A<br />
World Monograph of the genus Pleospora and its Segregates,<br />
p. 236. 1961.<br />
Generic description<br />
Habitat terrestrial, saprobic? Ascomata small, scattered,<br />
immersed, semi-immersed to nearly superficial, globose,<br />
subglobose, black, smooth; apex with a protruding<br />
papilla and pore-like ostiole, without periphyses. Perid-
Fungal Diversity<br />
ium thin, composed of a few layers of textura<br />
angularis. Hamathecium of numerous, cellular pseudoparaphyses,<br />
anastomosing, septate. Asci bitunicate, fissitunicate,<br />
cylindrical to cylindro-clavate, with a short,<br />
furcate pedicel. Ascospores broadly ellipsoid, reddish<br />
brown, muriform.<br />
Anamorphs reported for genus: none.<br />
Literature: Shoemaker and Babcock 1992; Wehmeyer<br />
1961.<br />
Type species<br />
Platysporoides chartarum (Fuckel) Shoemaker & C.E.<br />
Babc., Can. J. Bot. 70: 1650 (1992) (Fig. 76)<br />
≡ Pleospora chartarum Fuckel, Jb. nassau. Ver. Naturk.<br />
23–24: 133–134 (1870).<br />
Ascomata 150–230 μm high×180–260 μm diam.,<br />
scattered, immersed, semi-immersed to rarely superficial,<br />
globose, subglobose, black, smooth; apex with a protruding<br />
papilla, 50–85 μm long, 60–85 μm broad, ostiolate<br />
(Fig. 76a and b). Peridium 8–22 μm wide, composed of<br />
2–4 layers of brown cells of textura angularis, cells5–<br />
9 μm diam., cell wall 1–2.5 μm thick, without periphyses.<br />
Hamathecium of dense, long cellular pseudoparaphyses,<br />
2–3 μm broad, anastomosing, septate (Fig. 76c). Asci<br />
110–140×12.5–16.5 μm (x ¼ 121:5 14:7mm, n=10),<br />
(6-)8-spored, bitunicate, fissitunicate, cylindrical to<br />
cylindro-clavate, with a short, furcate pedicel, 8–17 μm<br />
long, ocular chamber not observed (Fig. 76c, d and e).<br />
Ascospores 20–26×8–11 μm (x ¼ 23:7 9mm, n=10),<br />
obliquely uniseriate and partially overlapping, flattened,<br />
broadly ellipsoid in front view, reddish brown, 3<br />
Fig. 75 Phaeosphaeriopsis<br />
glauco-punctata (from M.C.<br />
Cooke 166). a Ascomata immersed<br />
in the substrate. b Eightspored<br />
cylindrical asci. c–f. Pale<br />
brown baculate ascospores<br />
which are released from asci.<br />
Scale bars: a=200 μm,<br />
b=20 μm, c, d–f=10 μm
Fungal Diversity<br />
transverse septa, 1 longitudinal septum in each central<br />
cell, 1 oblique septum in each end cell, constricted at all<br />
septa, granulate, with a sheath 2–3 μm wide (as reported<br />
in Shoemaker and Babcock 1992) (Fig.76f, g and h).<br />
Anamorph: none reported.<br />
Material examined: GERMANY, Budenheim, Leopold<br />
Fuckel, Nassau’s Flora, on old paper (G NASSAU: 210558<br />
(a), as Sphaeria chartarum Wallr., type).<br />
Notes<br />
Morphology<br />
Platysporoides was introduced as a subgenus of Pleospora<br />
by Wehmeyer (1961) and was typified by Pleospora<br />
chartarum. Shoemaker and Babcock (1992) raised Platysporoides<br />
to generic rank and placed it in the Pleosporaceae<br />
based on its “applanodictyospore” and “terete pored<br />
beak of the ascomata”. Currently, eleven species are<br />
included in this genus (Shoemaker and Babcock 1992).<br />
Another comparable pleosporalean family is Diademaceae,<br />
which is distinguished from Platysporoides by its ascoma<br />
opening as “an intraepidermal discoid lid” (Shoemaker and<br />
Babcock 1992).<br />
Phylogenetic study<br />
None.<br />
Concluding remarks<br />
Aigialus grandis is another pleosporalean fungus<br />
with flattened and muriform ascospores as well as<br />
papilla and ostioles, which belongs to Aigialaceae, a<br />
phylogenetically well supported marine family<br />
(Suetrong et al. 2009). Thus, it is highly likely that<br />
flattened and muriform ascospores are of little phylogenetic<br />
significance.<br />
Pleomassaria Speg., Anal. Soc. cient. argent. 9: 192<br />
(1880). (Pleomassariaceae)<br />
Generic description<br />
Habitat terrestrial, saprobic. Ascomata medium to large,<br />
solitary, scattered, or in small groups, immersed, erumpent<br />
by a minute slit or a small conical swelling in the bark,<br />
flattened, papillate, ostiolate. Hamathecium of dense,<br />
cellular pseudoparaphyses, embedded in mucilage. Asci<br />
bitunicate, fissitunicate, broadly cylindrical to broadly<br />
cylindro-clavate, with a short, thick pedicel. Ascospores<br />
muriform, brown, constricted at the septa.<br />
Anamorphs reported for genus: Prosthemium and Shearia<br />
(Barr 1982b; Sivanesan 1984).<br />
Literature: Barr 1982b, 1990b, 1993a; Clements and Shear<br />
1931; Eriksson 2006; Lumbsch and Huhndorf 2007;<br />
Shoemaker and LeClair 1975; Sivanesan 1984; Tanaka<br />
et al. 2005.<br />
Type species<br />
Pleomassaria siparia (Berk. & Broome) Sacc., Syll. fung.<br />
2: 239 (1883) (Fig. 77)<br />
≡ Sphaeria siparia Berk. & Broome, Ann. Mag. nat.<br />
Hist., Ser. 2 9: 321 (1852).<br />
Ascomata 150–410 μm high×440–740 μm diam.,<br />
solitary, scattered, or in small groups, immersed, erumpent<br />
by a minute slit or a small conical swelling in the bark,<br />
depressed globose, papillalte, ostiolate (Fig. 77a). Peridium<br />
45–60 μm wide, thicker at the apex, thinner at the<br />
base, 1-layered, composed of small pigmented thickwalled<br />
compressed cells, cells ca. 15×3μm diam., cell<br />
wall 2–3.5 μm thick, apex cells larger, base composed of<br />
small pigmented thick-walled cells of textura angularis,<br />
ca. 5μm diam. (Fig. 77b). Hamathecium of dense, cellular<br />
pseudoparaphyses, 1–2 μm broad, embedded in mucilage,<br />
anastomosing or branching not observed. Asci 180–250×<br />
28–42 μm (x¼206:3 36:8mm, n=10), 8-spored, bitunicate,<br />
fissitunicate, broadly cylindrical to broadly cylindroclavate,<br />
with a short, thick pedicel, 15–45 μm long, with<br />
inconspicuous ocular chamber (Fig. 77c and d). Ascospores<br />
45–58×12.5–17.5 μm (x ¼ 50:5 14:8mm, n=10),<br />
biseriate, narrowly oblong with broadly to narrowly<br />
rounded ends, brown, muriform with 5–8 transverse septa<br />
and 1–2 vertical septa in some cells, smooth to verrucose,<br />
constricted at the septa, surrounded by a mucilaginous<br />
sheath (Fig. 77e, f and g).<br />
Anamorph: Prosthemium betulinum Kunze (Sivanesan<br />
1984).<br />
Conidia to 120 μm diam., with 3–5 arms, each arm 3–5-<br />
septate, 40–55×13–16 μm, connected to a central cell<br />
(Fig. 77h, i and j).<br />
Material examined: UK, Wiltshire, Spye Park, on<br />
branch of Betulina with Hendersonia polycystis Berk., et<br />
Br. leg. C.E. Broome, 1850? (BR, type).<br />
Notes<br />
Morphology<br />
Pleomassaria as characterized by Barr (1982b) has<br />
medium- to large-sized, immersed ascomata, cellular<br />
pseudoparaphyses, clavate to oblong asci and large,<br />
muriform ascospores (Barr 1982b; Sivanesan 1984). The<br />
muriform and somewhat asymmetrical ascospores with a<br />
submedian primary septum distinguish Pleomassaria from<br />
Asteromassaria in the family Pleomassariaceae, while in<br />
Splanchnonema ascospores have distinct bipolar asymmetry.<br />
Barr (1982b) included five North American species in<br />
the genus, while Kirk et al. (2008) listedfourspecies.
Fungal Diversity<br />
Fig. 76 Platysporoides chartarum<br />
(from G NASSAU:<br />
210558, type). a, b Ascomata<br />
scattered among fibers. Note the<br />
central ostioles. c Asci in numerous<br />
cellular pseudoparaphyses.<br />
d, e Cylindro-clavate asci<br />
with short pedicels. f–h. Muriform<br />
ascospores. Scale bars:<br />
a, b=200 μm, c–e=20 μm,<br />
f–h=10 μm<br />
Barr (1993a) treated Pleomassaria as a synonym of<br />
Splanchnonema based on a morphological cladistic analysis,<br />
but this proposal was not followed by later workers<br />
(Eriksson 2006; Lumbsch and Huhndorf 2007; Tanaka et<br />
al. 2005).<br />
Phylogenetic study<br />
Pleomassaria siparia forms a robust phylogenetic clade<br />
with Melanomma pulvis-pyrius (generic type) (Schoch et al.<br />
2009; Zhang et al. 2009a), which might represent a<br />
phylogenetic family (or suborder?).
Fungal Diversity<br />
Concluding remarks<br />
The genera Asteromassaria, Pleomassaria and Splanchnonema<br />
of Pleomassariaceae areconsideredtobeclosely<br />
related and difficult to separate (Barr 1982b; Crivelli 1983).<br />
They all have ascomata which are immersed in bark and are<br />
visible as slightly raised pustules with small ostioles, but<br />
may eventually become erumpent (e.g. Asteromassaria<br />
macrospora). Pseudoparaphyses are cellular, asci are bitunicate,<br />
while ascospores vary from 1-septate and pale brown<br />
(e.g. Asteromassaria macrospora) tomuriform(e.g.Pleomassaria<br />
siparia) and may be symmetrical (e.g. Asteromassaria<br />
macrospora) or highly asymmetrical (e.g.<br />
Splanchnonema pustulatum). The peridium ranges from<br />
thick-walled textura angularis (e.g. Asteromassaria macrospora)<br />
to thin-walled compressed cells (e.g. Splanchnonema<br />
pustulatum) and medium textura prismatica (e.g. Pleomassaria<br />
siparia). Anamorphs also vary distinctly, Prosthemium<br />
in Pleomassaria siparia, Scolicosporium in Asteromassaria<br />
macrospora but no anamorphic stage reported for Splanchnonema<br />
pustulatum. Furthermore, Asteromassaria pulchra<br />
clusters in Morosphaeriaceae in this study, thus here we<br />
tentatively assign Asteromassaria in Morosphaeriaceae<br />
(Plate 1). There seems to be considerable confusion in this<br />
family, especially when Pleomassaria siparia forms a robust<br />
phylogenetic clade with Melanomma pulvis-pyrius (Melannomataceae).<br />
Thus in this study, Pleomassariaceae is<br />
restated as a separate family from Melannomataceae.<br />
Therefore, fresh collections of the types of these genera are<br />
needed for molecular analysis and to establish which<br />
characters are important for classification.<br />
Pleophragmia Fuckel, Jb. nassau. Ver. Naturk. 23–24: 243<br />
(1870). (Sporormiaceae)<br />
Generic description<br />
Habitat terrestrial, saprobic (coprophilous). Ascomata smallto<br />
medium-sized, gregarious, immersed to erumpent, globose<br />
to subglobose, black, coriaceous; apex with a short papilla,<br />
or sometimes forming an ostiolar pore. Peridium thin,<br />
composed of several layers of thin-walled cells of textura<br />
angularis. Hamathecium of dense, delicate pseudoparaphyses.<br />
Asci 8-spored, bitunicate, fissitunicate, clavate to<br />
cylindro-clavate, with a relatively long pedicel and an ocular<br />
chamber. Ascospores muriform, narrow oblong to cylindrical<br />
with rounded ends, dark brown, constricted at each septum.<br />
Anamorphs reported for genus: none.<br />
Literature: von Arx and Müller 1975; Cain 1934.<br />
Type species<br />
Pleophragmia leporum Fuckel, Jb. nassau. Ver. Naturk.<br />
23–24 (1870) [1869–70]. (Fig. 78)<br />
Ascomata 330–480 μm high×320–430 μm diam.,<br />
gregarious, immersed to slightly erumpent, globose to<br />
subglobose, black; apex with a short papilla, sometimes<br />
forming a ostiolar pore (Fig. 78a). Peridium 25–35 μm<br />
thick at the sides, composed of one cell type of lightly<br />
pigmented thin-walled cells of textura angularis, cells 6–<br />
10 μm diam., cell wall 1.5–2 μm thick (Fig. 78b).<br />
Hamathecium of numerous, long pseudoparaphyses, 1–<br />
2 μm broad, anastomosing not observed. Asci 160–250×<br />
22.5–27.5 μm (x ¼ 203:6 25mm, n=10), 8-spored, bitunicate,<br />
fissitunicate, clavate to cylindro-clavate, with a 20–<br />
50 μm long pedicel and an ocular chamber (to 5 μm wide×<br />
2 μm high) (Fig. 78e and f). Ascospores 42–50×8–10 μm<br />
(x ¼ 46 10mm, n=10), biseriate to uniseriate and partially<br />
overlapping, narrowly oblong to cylindrical with rounded<br />
ends, dark brown, often slightly curved, with 9 transverse<br />
septa with two crossing longitudinal septa in the centre,<br />
constricted at each septum, smooth-walled (Fig. 78c, d, g<br />
and h).<br />
Anamorph: none reported.<br />
Material examined: GERMANY, between Königstein<br />
and Glashütten, on the same dung with Delitschia minuta.<br />
s.d. (G, Fungi rhenani n2272, type).<br />
Notes<br />
Morphology<br />
Pleophragmia was formally established by Fuckel<br />
(1870) and monotypified by Pleophragmia leporum. The<br />
most comparable genus to Pleophragmia is Sporormia, as<br />
ascospores of both have no germ slits and the inner layer of<br />
wall is considerably thinner than the outer layer (Barr<br />
1990a, b). But the muriform ascospores of Pleophragmia<br />
can be readily distinguished from the phragmosporous<br />
ascospores of Sporormia. Currently, only four species are<br />
accommodated under this genus (http://www.mycobank.<br />
org, 28-02-2009).<br />
Phylogenetic study<br />
None.<br />
Concluding remarks<br />
The presence of both transverse and crossing longitudinal<br />
septa is the most striking character of Pleophragmia, although<br />
the phylogenetic significance of this character is unclear.<br />
Pleoseptum A.W. Ramaley & M.E. Barr, Mycotaxon 54: 76<br />
(1995). (Phaeosphaeriaceae)<br />
Generic description<br />
Habitat terrestrial, saprobic? Ascomata medium-sized,<br />
scattered, or in small groups, immersed, globose to conoid,
Fungal Diversity<br />
Fig. 77 1 Pleomassaria siparia<br />
(from BR, type). a Ascomata on<br />
the host surface (after removing<br />
the cortices). b Section of a<br />
partial peridium. c, d Asci with<br />
short pedicels. e–g Ascospores<br />
with thin sheath. Scale bars:<br />
a=0.5 mm, b–d=50 μm, e–g=<br />
20 μm. 2 Prosthemium betulinum<br />
(from BR, type). h–j Conidia<br />
with arms. Scale bars:<br />
h–j=20 μm<br />
black, papillate, ostiolate. Peridium 1-layered. Hamathecium<br />
of dense, long cellular pseudoparaphyses, septate,<br />
branching. Asci 8-spored, bitunicate, fissitunicate, cylindrical<br />
to cylindro-clavate, with furcate pedicel. Ascospores<br />
obliquely uniseriate and partially overlapping, muriform,<br />
ellipsoid, ovoid to fusoid, yellowish to dark brown.<br />
Anamorphs reported for genus: Camarosporium (Ramaley<br />
and Barr 1995).
Fungal Diversity<br />
Fig. 77 (continued)<br />
Literature: Ramaley and Barr 1995.<br />
Type species<br />
Pleoseptum yuccaesedum A.W. Ramaley & M.E. Barr,<br />
Mycotaxon 54: 76 (1995). (Fig. 79)<br />
Ascomata 300–500 μm diam., scattered, or in small<br />
groups of 2–3, immersed with a flattened top, globose to<br />
conoid, black, papillate, ostiolate (Fig. 79a). Papilla<br />
small, slightly protruding from the host surface. Peridium<br />
30–50 μm thick at sides, up to 100 μm thick at the apex,<br />
1-layered, composed of 5–8 layers of heavily pigmented<br />
purplish-brown cells of textura angularis, cells 5–12 μm<br />
diam., cell wall 1–2 μm thick, apex cells smaller and<br />
walls thicker (Fig. 79c). Hamathecium of dense, long<br />
cellular pseudoparaphyses 1–2 μm broad, septate,<br />
branching (Fig. 79b). Asci 125–170(−195)×15–22 μm<br />
(x ¼ 153:8 19:3mm, n=10), 8-spored, bitunicate, fissitunicate,<br />
cylindrical to cylindro-clavate, with a short,<br />
narrowed, furcate pedicel which is 10–20 μm long, with<br />
an ocular chamber best seen in immature asci (to 5 μm<br />
broad×3 μm high) (Fig. 79d and e). Ascospores 22–30×<br />
11–14 μm (x ¼ 27:1 12:6mm, n=10) obliquely uniseriate<br />
and partially overlapping, ellipsoid, ovoid to fusoid,<br />
yellowish to yellowish brown, becoming reddish brown<br />
to dark brown, muriform, with 3-(4) transverse septa,<br />
constricted at the primary septum, part above central<br />
septum wider, vertical septa exist in each cell, ornamentation<br />
of foveolae in linear rows (Fig. 79f and g).<br />
Anamorph: Camarosporium yuccaesedum Fairm.<br />
(Ramaley and Barr 1995).<br />
Conidiomata 200–450 μm diam., pycnidial, immersed,<br />
scattered, subglobose to conoid, ostiolate. Macroconidiogenous<br />
cells determinate or indeterminate, enteroblastic,<br />
hyaline, smooth. Macroconidia holoblastic, 20–36×10–<br />
15 μm diam., ellipsoid to narrowly ovoid, muriform,<br />
yellowish brown, 3–7 transverse septa, constricted at the<br />
septa. Microconidiogenous cells produced near or in the<br />
ostiole, hyaline, smooth. Microconidia 5–10×5–7 μm<br />
diam., globose to ovoid, aseptate, hyaline, smooth.<br />
Material examined: USA, Colorado, Montezuma County,<br />
hillside near entrance to Mesa Verde National Park, on dead<br />
leaves of Yucca baccata, 11 Oct. 1992, Ramaley Annette<br />
(9237A) (BPI 802381, holotype).<br />
Notes<br />
Morphology<br />
Pleoseptum is a monotypic genus established by<br />
Ramaley and Barr (1995) and represented by P. yuccaesedum<br />
basedonits“immersed ascomata, thick peridium,<br />
muriform ascospores, anamorphic stage and the linoeate<br />
ornamentation of the ascospores and conidia”. Theshape<br />
of ascomata of Pleoseptum is comparable with that of<br />
Chaetoplea, but the peridium structure easily distinguishes<br />
them. Some species of Curreya, Leptosphaeria and<br />
Heptameria are comparable with Pleoseptum, but their<br />
anamorphic stages differ.<br />
Pleoseptum yuccaesedum and its Camarosporium yuccaesedum<br />
anamorph both formed in the leaves of Yucca<br />
baccata and the ascomata and conidiomata were indistinguishable.<br />
Camarosporium is the anamorph of diverse<br />
teleomorph genera included in Botryosphaeriales and<br />
Cucurbitariaceae (Kirk et al. 2008). The genus is in need<br />
of revision (Sutton 1980) and is no doubt polyphyletic.<br />
Phylogenetic study<br />
None.<br />
Concluding remarks<br />
The placement of Pleoseptum under Phaeosphaeriaceae<br />
is still tentative.<br />
Pleospora Rabenh. ex Ces. & De Not., Comm. Soc. crittog.<br />
Ital. 1: 217 (1863). (Pleosporaceae)<br />
Generic description<br />
Habitat terrestrial, saprobic or parasitic. Ascomata smallto<br />
medium-sized, immersed, erumpent to superficial,
Fungal Diversity<br />
Fig. 78 Pleophragmia leporum (from G. Fungi rhenani n2272,<br />
type). a Appearance of ascomata on the substrate surface. Note the<br />
ostiolar pore. b Section of a partial peridium. c, h Apical part of an<br />
ascus. Note the apical apparatus in (c). d Released ascospores. e, f<br />
Clavate Asci with pedicels. g Part of a broken ascospore. Note the<br />
crossing septa. Scale bars: a=0.5 mm, B=50 μm, c–f=20 μm, g,<br />
h=10 μm
Fungal Diversity<br />
Fig. 79 Pleoseptum yuccaesedum (from BPI 802381, holotype). a<br />
Appearance of ascomata scattered on the host surface. Only the<br />
upper region is visible. b Squash mount of asci in pseudoparaphyses.<br />
c Section of an ascoma. Note the peridium comprising cells of<br />
textura angularis. d, e Asci with short furcate pedicels. f, g<br />
Muriform dark-brown ascospores. Scale bars: a=0.5 mm, b=<br />
40 μm, c=100 μm, d, e=20 μm, f, g=10 μm
Fungal Diversity<br />
papillate, ostiolate. Peridium thin. Hamathecium of dense,<br />
cellular pseudoparaphyses. Asci 8-spored, bitunicate,<br />
fissitunicate, cylindrical to clavate, with furcate pedicel<br />
and small inconspicuous ocular chamber. Ascospores<br />
muriform, brown or pale brown, with or without<br />
sheath.<br />
Anamorphs reported for genus: Stemphylium (Simmons<br />
1985).<br />
Literature: Barr 1981; Frisullo and Braun 1996; Kodsueb<br />
et al. 2006a; Luttrell1951; Wehmeyer1946, 1961, 1975;<br />
Zhang et al. 2009a.<br />
Type species<br />
Pleospora herbarum (Pers.) Rabenh., Klotzschii Herb. Viv.<br />
Mycol. 2: no. 547 (1854). (Fig. 80)<br />
≡ Sphaeria herbarum Pers., Syn. meth. fung. (Göttingen)<br />
1: 78 (1801).<br />
Ascomata 130–220 μm high×250–420 μm diam.,<br />
scattered, or in small groups of 2–3, immersed, semiimmersed<br />
to erumpent, broadly to narrowly oblong and<br />
flattened, with flattened base not easily removed from the<br />
substrate, wall black, papillate, ostiolate (Fig. 80a and b).<br />
Peridium 30–50 μm thick on sides, thinner at the base,<br />
coriaceous, 2-layered, outer layer composed of one or two<br />
layers of heavily pigmented thick-walled cells of textura<br />
angularis, cells 5–10 μm diam., cell wall 2–4 μm thick,<br />
apex cells smaller and walls thicker, inner layer composed<br />
of hyaline thin-walled cells of textura angularis, 8–12 μm<br />
diam., wall hyaline, 0.5–1.5 μm thick (Fig. 80c). Hamathecium<br />
of dense, cellular pseudoparaphyses, 2–3 μm<br />
broad, filling the gaps between the asci. Asci 100–210×<br />
27.5–30 μm (x ¼ 142:2 28:3mm, n=10), 8-spored, bitunicate,<br />
fissitunicate, broadly cylindrical to clavate, with a<br />
short, thick, furcate pedicel, 8-12(−20) μm long, with<br />
small inconspicuous ocular chamber (ca. 3 μm wide×<br />
1 μm high) (Fig. 80d and e). Ascospores 28–38×12.5–<br />
15 μm (x ¼ 33 14:5mm, n=10), ellipsoidal, straight or<br />
sometimes curved, with broadly rounded ends and upper<br />
hemispore slightly shorter and broader; spores usually<br />
divided by 3 A-transsepta, all 4 segments by longisepta<br />
andthenbyonestratumofB-transsepta(maturesporesas<br />
a rule with 7 transsepta, 3A+4B), yellowish brown,<br />
smooth; each hemispore with thick gelatinous sheath, the<br />
lower one with umbilicus (sheaths fused in mature spores)<br />
(Fig. 80f, g, h, i, j and k).<br />
Anamorph: Stemphyllium herbarum E. Simmons<br />
(Simmons 1985).<br />
Material examined: GERMANY, on stalks of Melilotusalla?<br />
at the bank of the Elbe in Konigstein, 1882 (E,<br />
Krieger 683); as Sphaeria herbarum Persoon Syn. fung. p.<br />
78 (E, 81); as Sphaeria herbarum Fries, Scleromyceti<br />
Sueciae 38 (E, lectotype).<br />
Notes<br />
Morphology<br />
Pleospora was originally assigned within Sphaeriales.<br />
Subsequently, it was assigned within Pseudosphaeriales and<br />
<strong>Pleosporales</strong> (Wehmeyer 1961). Pleospora is a large group,<br />
which is widely distributed and associated with a wide range<br />
of species of monocotyledons as well as dicotyledons<br />
(Wehmeyer 1975). All species of Pleospora have muriform<br />
ascospores (Wehmeyer 1961, 1975). Pleospora has downward<br />
growing pseudoparaphyses within the ascomata of<br />
“Pleospora-type” development (Luttrell Univ. Mo. Stud.<br />
1951), which subsequently served as a diagnostic character.<br />
However, only a limited number of species had detailed<br />
studies on this character (Wehmeyer 1961). The heterogeneous<br />
nature of Pleospora has been noted, and several<br />
subgenera have been erected, such as Scleroplea to include<br />
all “sclerotioid” species of Pleospora, Teichosporoides to<br />
accommodate species of Pleospora with immersed ascomata,<br />
Pleosphaeria for those having superficial and setose ascomata<br />
(Wehmeyer 1961). Similarly, Cucurbitaria, Fenestella and<br />
Montagnula are also separated as a section from Pleospora.<br />
Most of these subgenera are currently at genus rank.<br />
Phylogenetic study<br />
The polyphyletic nature of Pleospora is clear (Kodsueb et<br />
al. 2006a), and those that stain the woody substrate purple<br />
should be assigned to Amniculicolaceae (Zhang et al. 2009a).<br />
Concluding remarks<br />
As some Pleospora species have a wide range of host<br />
spectrum, especially on both monocotyledons and dicotyledons,<br />
it is highly possible they are cryptic species.<br />
Preussia Fuckel, Hedwigia 6: 175 (1867) [1869–70].<br />
(Sporormiaceae)<br />
Generic description<br />
Habitat terrestrial, saprobic (on decaying fibers or coprophilous).<br />
Ascomata small- to medium-sized, cleistothecial or<br />
perithecial, solitary or scattered on substrate surface,<br />
globose, membraneous, black. Peridium thin, composed<br />
of thick-walled, poly-angular cells from the surface view.<br />
Pseudoparaphyses not observed. Asci (4-) 8-spored, bitunicate,<br />
clavate to broadly clavate, with a long and thin and<br />
furcate pedicel. Ascospores 3–6 seriate to uniseriate near<br />
the base, cylindrical with rounded ends, brown, septate,<br />
easily breaking into partspores, with germ slits in each cell.<br />
Anamorphs reported for genus: Phoma (von Arx 1973;<br />
Cain 1961; Malloch and Cain 1972).<br />
Literature: Ahmed and Cain 1972; Arenaletal.2005; von<br />
Arx 1973; von Arx and van der Aa 1987; Auerswald1866;
Fungal Diversity<br />
Fig. 80 Pleospora herbarium (from E, Krieger 683). a Immersed<br />
ascomata scattering on host surface. b Ascomata in small groups. Note:<br />
the surface layer of the host is removed. c Section of an ascoma. Note<br />
the peridium cells of textura angularis. d, e. Asci with short pedicels f–j<br />
Ascospores. Scale bars: a, b=0.5 mm, c=100 μm, d, e=30 μm, f–j=<br />
20 μm<br />
Barr 1987b, 1990a;Boylan1970; Cain 1961; Eriksson 1992;<br />
Fuckel 1866; Guarroetal.1981, 1997a, b; KhanandCain<br />
1979a, b; Kruys and Wedin 2009; Lodha 1971; Lorenzo<br />
1994; Luck-Allen and Cain 1975; Maciejowska and
Fungal Diversity<br />
Williams 1963; Malloch and Cain 1972; Munk 1957;<br />
Narendra and Rao 1976; Rai and Tewari 1963; Sultana<br />
and Malik 1980.<br />
Type species<br />
Preussia funiculata (Preuss) Fuckel, Jb. nassau. Ver.<br />
Naturk. 23–24: 91 (1870) [1869–70]. (Fig. 81)<br />
≡ Perisporium funiculatum Preuss, Fung. Hoyersw.: no.<br />
145 (1851).<br />
Ascomata 240–500 μm diam., cleistothecial, solitary,<br />
scattered on substrate, superficial, globose, membraneous,<br />
black (Fig. 81a). Peridium thin, composed of thick-walled,<br />
poly-angular cells in front view (Fig. 81b). Pseudoparaphyses<br />
not observed. Asci 42–65×20–25 μm (x ¼ 55:8 21:8mm, n=<br />
10), (4-)8-spored, bitunicate, broadly clavate, with a long and<br />
thin and furcate pedicel, up to 115 μm long, ocular chamber<br />
not observed (Fig. 81c and d). Ascospores 30–40×6.3–<br />
7.5 μm (x ¼ 35:6 6:9mm, n=10), 3–6 seriate to uniseriate<br />
near the base, cylindrical with rounded ends, brown, with<br />
3 transverse septa, easily breaking into partspores, central<br />
cells round in transverse section but rectangular in<br />
vertical section, with a germ slit in each cell, 6.5–8.5×<br />
4–7.5 μm broad, apical cells 8.8–10×5–7 μm broad,<br />
sheath not observed.<br />
Anamorph: none reported.<br />
Material examined: USA, Ontario, York Co., Nashville,<br />
on old jute sack on ground, 1 Jul. 1960, leg. & det. R.F.<br />
Cain (in part Preussia typharum) (TRTC 46985).<br />
Notes<br />
Morphology<br />
Preussia was introduced by Fuckel (1866) to accommodate<br />
species having cleistothecioid ascomata, bitunicate<br />
asci, multi-septate ascospores with a germ slit in<br />
each cell and with a gelatinous sheath, and occurring in<br />
soil or plant debris. Preussia, Sporormia and Sporormiella<br />
are regarded as closely related genera, which share<br />
numerous morphological characters. Sporormia can be<br />
distinguished from Preussia by its perithecioid ascomata<br />
and cylindrical asci. The only distinguishing morphological<br />
character for Preussia from Sporormiella are the<br />
cleistothecioid ascomata in Preussia (Barr 2000; Cain<br />
1961), but this has been shown to have little phylogenetic<br />
significance (von Arx 1973; Zhang et al. 2009a).<br />
Substrate preference has been used to distinguish species<br />
of Sporormiella and Preussia, with Sporormiella being<br />
restricted to a coprophilous habitat, while Preussia grows<br />
in plant debris, wood or soil (von Arx and van der Aa<br />
1987). This proposal was rejected, as P. intermedia<br />
(Clum) Cain can be isolated from either soil or dung<br />
(Guarro et al. 1997b). In a review of Preussia, Cain<br />
(1961) accepted 12 species, and some of them are coprophilous.<br />
Subsequently, numerous additional new species have<br />
been published (Arenal et al. 2005; Barr 1987b, 1990a;<br />
Boylan 1970; Eriksson1992; Guarro et al. 1981, 1997a, b;<br />
Khan and Cain 1979a; Lodha1971; Lorenzo 1994; Luck-<br />
Allen and Cain 1975; Maciejowska and Williams 1963;<br />
Malloch and Cain 1972; Narendra and Rao 1976; Rai and<br />
Tewari 1963; Sultana and Malik 1980). Currently, 84 species<br />
are listed under Preussia (http://www.mycobank.org/<br />
mycotaxo.aspx, 10/2010) and Kirk et al. (2008) estimates<br />
there are 51 species.<br />
Phylogenetic study<br />
In phylogenetic analysis based on ITS, nLSU, mtSSU<br />
and β-tubulin gene fragments, Preussia, Sporormiella and<br />
Spororminula clustered together. Thus, Sporormiella together<br />
with Spororminula are treated as synonyms of<br />
Preussia (Kruys and Wedin 2009).<br />
Concluding remarks<br />
Preussia sensu lato (including Sporormiella and Spororminula)<br />
based on both morphology and molecular data<br />
should be accepted pending further research.<br />
Quintaria Kohlm. & Volkm.-Kohlm., Bot. Mar. 34: 34<br />
(1991). (<strong>Pleosporales</strong>, genera incertae sedis)<br />
Habitat marine, saprobic. Ascomata medium-sized, scattered<br />
or loosely gregarious, immersed, mostly subglobose,<br />
rarely globose, with a protruding papilla, ostiolate. Peridium<br />
thin, 2-layered, coriaceous, thicker near the apex.<br />
Hamathecium of dense, filamentous, trabeculate pseudoparaphyses,<br />
branching and anastomosing between and<br />
above asci. Asci 8-spored, bitunicate, fissitunicate,<br />
cylindro-clavate, with a short furcate pedicel. Ascospores<br />
biseriate, broadly fusoid to fusoid, hyaline, mostly 5-<br />
septate, rarely up to 7-septate.<br />
Anamorphs reported for genus: none.<br />
Literature: Hyde and Goh 1999; Kohlmeyer and<br />
Volkmann-Kohlmeyer 1991; Suetrong et al. 2009; Zhang<br />
et al. 2008b.<br />
Type species<br />
Quintaria lignatilis (Kohlm.) Kohlm. & Volkm.-Kohlm.,<br />
Bot. Mar. 34: 35 (1991). (Fig. 82)<br />
≡ Trematosphaeria lignatilis Kohlm., Marine Ecology,<br />
[Pubblicazioni della Stazione Zoologica Napoli I] 5(4): 365<br />
(1984).<br />
Ascomata 240–500 μm diam., scattered or loosely<br />
gregarious, immersed, globose to subglobose, coriaceous,<br />
ostiolate, ostiole is encrusted with thick-walled black cells,<br />
papilla up to 400 μm long (Fig. 82a). Peridium thin, 20–<br />
30 μm wide, thinner at the base, thicker near the apex, up to
Fungal Diversity<br />
Fig. 81 Preussia funiculata<br />
(from TRTC 46985). a Superficial<br />
cleistothecoid ascomata. b<br />
Part of peridium from front<br />
view. c Squash mounts showing<br />
a large number of asci. d A<br />
clavate ascus with a long and<br />
thin pedicel. Scale bars: a=<br />
0.5 mm, b=20 μm, c, d=50 μm<br />
300 μm, 2-layered, outer layer composed of hyphoid cells,<br />
inner layer composed of compressed cells of textura<br />
angularis (Fig. 82b). Hamathecium of dense, filamentous,<br />
trabeculate pseudoparaphyses, 0.8–1.5 μm broad, branching<br />
and anastomosing between and above asci (Fig. 82e).<br />
Asci 175–250×25–35 μm (x ¼ 220 28mm, n=10), 8-<br />
spored, bitunicate, fissitunicate, cylindro-clavate, with a<br />
short, furcate pedicel, to 20 μm long (Fig. 82c and e).<br />
Ascospores 55–73×12.5–15 μm (x ¼ 63:3 13:1mm, n=<br />
10), biseriate, broadly fusoid to fusoid, usually slightly<br />
curved, smooth, hyaline, mostly 5-septate, rarely up to 7-<br />
septate, smooth-walled, lacking a sheath.<br />
Anamorph: none reported.<br />
Material examined: BELIZE, Twin Cays, on attached<br />
dead tip of prop root of Rhizophora mangle, with shipworms,<br />
3 Apr. 1983, leg. & det. J.K. Kohlmeyer (J.<br />
Kohlmeyer No. 4365a, holotype).<br />
Notes<br />
Morphology<br />
Quintaria was introduced to accommodate the marine<br />
fungus, Trematosphaeria lignatilis, based on its immersed<br />
ascomata with rounded bases, black incrustations surrounding<br />
the sides of the ostiolar canal as well as its hyaline<br />
ascospores (Kohlmeyer and Volkmann-Kohlmeyer 1991).<br />
Subsequently, three more species were introduced to this<br />
genus, viz. Q. aquatica K.D. Hyde & Goh, Q. microsporum<br />
Yin. Zhang, K.D. Hyde & J. Fourn. and Q. submerse K.D.<br />
Hyde & Goh, which are all from freshwater (Hyde and Goh<br />
1999; Zhang et al. 2008b).<br />
Phylogenetic study<br />
Multigene phylogenetic study indicated that Quintaria<br />
lignatilis forms a separate sister clade to other families of<br />
<strong>Pleosporales</strong>, which may represent a new familial linage<br />
(Suetrong et al. 2009). This was supported by phylogenetic<br />
studies which place the freshwater Q. submersa separate from<br />
Q. lignatilis (Schoch et al. 2009;Suetrongetal.2009; Plate 1).<br />
Concluding remarks<br />
The freshwater members of Quintaria should likely be<br />
excluded from this genus, and only the generic type, Q.<br />
lignatilis retained, but this needs confirmation.<br />
Roussoëlla Sacc., in Saccardo & Paoletti, Atti Inst. Veneto<br />
Sci. lett., ed Arti, Sér. 3 6: 410 (1888). (Arthopyreniaceae<br />
(or Massariaceae))
Fungal Diversity<br />
Generic description<br />
Habitat terrestrial, saprobic. Ascomata medium-sized, clustered,<br />
immersed in host tissue, forming under darkened,<br />
slightly raised, somewhat liner or dome-shaped stroma on the<br />
host, with a flush intra-epidermal papilla; immersed under<br />
clypeus, papillate, ostiolate. Peridium thin, comprising<br />
several layers of compressed cells. Hamathecium of dense,<br />
long trabeculate pseudoparaphyses, embedded in mucilage,<br />
hyaline, anastomosing and septate. Asci 8-spored, bitunicate,<br />
cylindrical, with furcate pedicel, and a conspicuous ocular<br />
chamber. Ascospores uniseriate to partially overlapping, fusoid<br />
or ellipsoidal, brown, 1-septate, constricted at the septum.<br />
Anamorphs reported for genus: Cytoplea (Hyde et al.<br />
1996a).<br />
Literature: Hyde et al. 1996a; Hyde 1997; Ju et al. 1996;<br />
Tanaka et al. 2009.<br />
Type species<br />
Roussoëlla nitidula Sacc. & Paol., Atti Ist. Veneto Sci., Ser.<br />
6, 6:410. (1888). (Fig. 83)<br />
Ascomata 160–200 μm high×400–500 μm diam., clustered,<br />
immersed in host tissue, forming under darkened,<br />
slightly raised, somewhat liner or dome-shaped stroma on the<br />
host, with a flush intra-epidermal papilla; in vertical section<br />
subglobose with a flattened base, immersed under clypeus,<br />
subglobose with a flattened base, papillate, ostiolate (Fig. 83a).<br />
Peridium up to 20 μm thick, comprising several layers of<br />
compressed cells. Hamathecium of dense, long trabeculate<br />
pseudoparaphyses, 1–1.5 μm broad, embedded in mucilage,<br />
anastomosing and septate. Asci 123–220×7–11 μm,8-spored,<br />
bitunicate, cylindrical, with furcate pedicels, and a conspicuous<br />
ocular chamber (Fig. 83b, c and d). Ascospores 17.5–<br />
22×5.5–8 μm, uniseriate to partially overlapping, fusoid or<br />
ellipsoidal, brown, 1-septate, constricted at the septum,<br />
ornamented with longitudinal wall striations and surrounded<br />
by a wide mucilaginous sheath (Fig. 83e, f, g and h).<br />
Anamorph: Cytoplea hysterioides K.D. Hyde (Hyde et<br />
al. 1996a).<br />
Material examined: MALAYSIA, Malacca, on culms of<br />
Bambusa Bar & Grill, 1885, B. Scortechini 15 (PAD,<br />
Roussoëlla nitidula Sacc. Paol. 2484, holotype, on a loose<br />
label Roussoëlla nitidula S. & P. Est Phyllachora phaeodidym./15<br />
prob. original material from Malacca Peninsula).<br />
Notes<br />
Morphology<br />
Roussoëlla was introduced by Saccardo for the single<br />
species R. nitidula Sacc. & Paol. (Saccardo and Paoletti<br />
1888). It was redescribed by Hyde et al. (1996a) and the<br />
anamorph of Roussoëlla hysterioides (Ces.) Höhn., Cytoplea<br />
hysterioides K.D. Hyde was determined and described.<br />
Roussoëlla was then reviewed by Hyde (1997) and a<br />
modified key for Roussoëlla species was provided based on<br />
the one proposed by Ju et al. (1996). Roussoëlla is<br />
characterized as having immersed ascomata containing<br />
long cylindrical asci and brown 1-septate ornamented<br />
ascospores. In this study, we have checked the type species<br />
and it matches Hyde et al. (1996a). The asci are bitunicate,<br />
but we could not observe the fissitunicate dehiscence.<br />
Phylogenetic study<br />
Species of Roussoëlla, Roussoellopsis as well as Arthopyrenia<br />
salicis form a robust phylogenetic clade, which form<br />
a sister group with pleosporalean families, but the generic<br />
type of Roussoëlla (R. nitidula) was not included in the<br />
phylogenetic study (Tanaka et al. 2009).<br />
Concluding remarks<br />
The bambusicolous habitat of Roussoëlla is a striking<br />
character at generic rank classification but its relationship to<br />
the lichenized Arthopyrenia is unexpected and will require<br />
more analysis.<br />
Saccharicola D. Hawksw. & O.E. Erikss., in Eriksson &<br />
Hawksworth, Mycologia 95: 431 (2003). (Massarinaceae)<br />
Generic description<br />
Habitat terrestrial, parasitic. Ascomata medium-sized, solitary,<br />
scattered, immersed, globose to subglobose, carbonaceous,<br />
papillate, ostiolate. Peridium relatively thin, composed of one<br />
cell type of pale brown to hyaline pseudoparenchymatous cells.<br />
Hamathecium of trabeculate pseudoparaphyses. Asci bitunicate,<br />
8-spored, cylindro-clavate to clavate. Ascospores biseriate and<br />
sometimes laterally uniseriate, fusoid with narrowly rounded<br />
ends, septate, constricted at the septa, the upper second cell<br />
becoming pigmented when mature, smooth or verruculose.<br />
Anamorphs reported for genus: Stagonospora (Eriksson<br />
and Hawksworth 2003; Kaiser et al. 1979; Leuchtmann<br />
1984).<br />
Literature: Eriksson and Hawksworth 2003.<br />
Type species<br />
Saccharicola bicolor (D.Hawksw.,W.J.Kaiser&Ndimande)<br />
D. Hawksw. & O.E. Erikss., Mycologia 95: 431 (2003).<br />
(Fig. 84)<br />
≡ Leptosphaeria bicolor D. Hawksw., W.J. Kaiser &<br />
Ndimande, Mycologia 71: 483 (1979).<br />
Ascomata 125–175 μm high×175–220 μm diam.,<br />
solitary, scattered, immersed, globose to subglobose, wall<br />
black, carbonaceous, with a protruding papilla, with a<br />
central ostiole (Fig. 84a). Peridium 15–20 μm thick
Fig. 82 Quintaria lignitalis<br />
(from J. Kohlmeyer No. 4365a,<br />
holotype). a Ascomata immersed<br />
in substrate. b Section of<br />
an ascoma. Note the thin peridium<br />
and elongated papilla. c, e<br />
Asci embedded in pseudoparaphyses.<br />
d Five septate fusoid<br />
hyaline ascospores. Scale bars:<br />
a=0.5 mm, b=200 μm, c, e=<br />
50 μm, d =20 μm<br />
Fungal Diversity
Fungal Diversity<br />
composed of one cell type of pale brown to hyaline<br />
pseudoparenchymatous cells, becoming thicker near the apex<br />
(Fig. 84a). Hamathecium of 1–2 μm broad, filliform, hyaline,<br />
septate pseudoparaphyses, branching and anastomosing in<br />
mucilage. Asci (90-)125–150×(20-)25–30 μm, 8-spored, with<br />
a short pedicel, bitunicate, cylindro-clavate to clavate, with a<br />
small ocular chamber at the apex (Fig. 84c). Ascospores 29–<br />
42×8–11 μm, biseriate and sometimes laterally uniseriate,<br />
fusoid with narrowly rounded ends, (2-)3-septate, deeply<br />
constricted at the septa, the upper second cell subhyaline to<br />
pale brown when young and becoming dark brown to almost<br />
black at maturity, smooth or verruculose (Fig. 84d). (data from<br />
the original description by Kaiser et al. (1979) because of the<br />
bad condition of the type material).<br />
Fig. 83 Roussoëlla nitidula<br />
(from PAD Paol. 2484, holotype).<br />
a Appearance of the<br />
stroma on host surface. b Asci<br />
and pseudoparaphyses. c, d<br />
Long cylindrical furcate asci.<br />
e–h. Ascospores. Note the striate<br />
ornamentation. Scale bars:<br />
a=0.5 mm, b–d=20 μm,<br />
e–h=10 μm
Fungal Diversity<br />
Anamorph: Pycnidia typical of Stagonospora (Sphaeropsidales),<br />
“scattered,arisingsinglybothonthehostandinpure<br />
culture, in culture generally surrounded by an envelope of<br />
mycelial hyphae, numerous, immersed on the host, but nearly<br />
superficial in culture, subglobose to slightly applanate, black,<br />
150–250 μm diam., with a central slightly papillate ostiole,<br />
lacking a distinct neck; walls mainly 15–20 μm thick,<br />
composed of three to six layers of pseudoparenchymatous<br />
cells, the outermost layers dark brown and inner pale brown to<br />
hyaline cells somewhat compressed radially, very variable in<br />
size, cells of the outer layers mainly 7–12 μm long×4–6 μm<br />
wide in vertically section and 10–12 μm diam. in surface<br />
view, wall not or only slightly thicked near the ostiole.<br />
Conidiogenous cells lining the inner surface of the pycnidial<br />
cavity, holoblastic, minute and difficult to distinguish from the<br />
pseudoparenchymatous cells with which they are mixed,<br />
mammiform with a flattened apex, hyaline, smooth walled,<br />
about 4–6 μm tall and 4–6 μm wide. Conidia copiously<br />
produced, ellipsoid, with somewhat truncated ends, hyaline,<br />
smooth walled, (2-)3 septate, not or slightly constricted at the<br />
septa, often guttulate, rather thin walled, (21-)24–28(−34)<br />
μm×7–8.5(−11.5) μm” (from Kaiser et al. 1979).<br />
Material examined: KENYA, near Nairobi, on leaves of<br />
Saccharum officinarum L.; 24 Aug. 1977; leg. W.J. Kaiser<br />
(IMI 215888, holotype).<br />
Notes<br />
Morphology<br />
Saccharicola was separated from Leptosphaeria as a new<br />
genus based on its Stagonospora anamorph and its biotrophic<br />
habitat in leaves of sugar cane, and two species were<br />
included, i.e. Saccharicola bicolor and S. taiwanensis (J.M.<br />
Yen & C.C. Chi) O.E. Erikss. & D. Hawksw. (Eriksson and<br />
Hawksworth 2003). Saccharicola is characterized by its<br />
parasitic habitat on monocots, small ascomata, bitunicate<br />
asci, presence of pseudoparaphyses as well as its 3-septate<br />
ascospores (Eriksson and Hawksworth 2003).<br />
Phylogenetic study<br />
Based on the limited phylogenetic analysis of SSU<br />
sequences, Saccharicola is considered to be closely related<br />
to Massarina eburnea, the generic type of Massarina<br />
(Eriksson and Hawksworth 2003). Thus, Saccharicola was<br />
assigned to Massarinaceae, which includes Keissleriella,<br />
Massarina and Saccharicola (Eriksson and Hawksworth<br />
2003).<br />
Concluding remarks<br />
Based on the parasitic habitat on monocots and its small<br />
ascomata and Stagonospora (or Cercospora? for S. taiwanensis,<br />
see Eriksson and Hawksworth 2003; Shoemaker<br />
and Babcock 1989b) anamorph, Saccharicola seems more<br />
similar to Pleosporineae. Further molecular study is needed<br />
for confirmation.<br />
Salsuginea K.D. Hyde, Bot. Mar. 34: 315 (1991). (<strong>Pleosporales</strong>,<br />
genera incertae sedis)<br />
Generic description<br />
Habitat marine, saprobic. Ascomata large, solitary, fusoid,<br />
conical or subglobose, with or without a flattened base,<br />
immersed under a darkened clypeus, papillate, ostiolate.<br />
Peridium thin, composed of round cells (in cross section) at<br />
sides, fusing at the top with the clypeus, thin at the base.<br />
Hamathecium of dense, long trabeculate pseudoparaphyses,<br />
anastomosing, embedded in mucilage. Asci 8-spored, bitunicate,<br />
fissitunicate, clavate to cylindro-clavate, pedunculate,<br />
with a large ocular chamber and conspicuous apical ring.<br />
Ascospores uniseriate, obovoid, brown to black, with hyaline<br />
apical germ pores, 1-septate, constricted at the septum, dark<br />
brown with paler apical cells, lacking sheath, smooth.<br />
Anamorphs reported for genus: none.<br />
Literature: Hyde 1991a; Suetrong et al. 2009.<br />
Type species<br />
Salsuginea ramicola K.D. Hyde, Bot. Mar. 34: 316 (1991).<br />
(Fig. 85)<br />
Ascomata 1040–2600 μm high×455–1430 μm diam.,<br />
solitary, fusoid, conical or subglobose, with or without a<br />
flattened base, immersed under a darkened clypeus, papillate,<br />
ostiolate, ostiole rounded (Fig. 85a). Peridium up to 39 μm<br />
thick, composed of round cells (in cross section) at sides,<br />
fusing at the top with the clypeus, thin at the base (Fig. 85b).<br />
Hamathecium of dense, long trabeculate pseudoparaphyses,<br />
1–2 μm broad, anastomosing, embedded in mucilage. Asci<br />
440–512×29–34 μm, 8-spored, bitunicate, fissitunicate,<br />
clavate to cylindro-clavate, pedunculate, with a large ocular<br />
chamber and conspicuous apical ring (Fig. 85c and e).<br />
Ascospores 59–72×24–30 μm, uniseriate, obovoid, brown to<br />
black, with hyaline apical germ pores, 1-septate, constricted<br />
at the septum, dark brown with paler apical cells, lacking<br />
sheath, smooth (Fig. 85d and f).<br />
Anamorph: none reported.<br />
Material examined: THAILAND, Ranong mangrove,<br />
Aegiceras corniculatum (L.) Blanco., Oct. 1988, leg. & det.<br />
K.D. Hyde (BRIP 17102, holotype).<br />
Notes<br />
Morphology<br />
Salsuginea was introduced to accommodate the mangrove<br />
fungus, S. ramicola, which is characterized by large,<br />
immersed, ostiolate and papillate ascomata under a clypeus,
Fungal Diversity<br />
Fig. 84 Saccharicola bicolor<br />
(from IMI 215888, holotype). a<br />
Section of an ascomata immersed<br />
in the host tissue. b<br />
Section of a partial pycnidia.<br />
Note the phragmosporous conidia.<br />
c Clavate ascus with ocular<br />
chamber and short pedicel. d<br />
Ascospores. Note the pigmented<br />
central cell(s). Scale bars: a, b=<br />
50 μm, c=20 μm, d=10 μm<br />
dense, trabeculate pseudoparaphyses embedded in gel matrix,<br />
fissitunicate, 8-spored, cylindrical asci with short pedicel and<br />
conspicuous apical apparatus, 1-septate, dark brown ascospores<br />
with paler apical cells (Hyde 1991a). Salsuginea is<br />
considered closely related to Helicascus and Caryospora, and<br />
they are all proposed to Melanommataceae (Hyde 1991a).<br />
Phylogenetic study<br />
Based on a multigene phylogenetic analysis, Salsuginea<br />
ramicola nested in a paraphyletic clade within <strong>Pleosporales</strong>;<br />
its familial status is undetermined (Suetrong et al. 2009).<br />
Concluding remarks<br />
It has been shown that trabeculate pseudoparaphyses has<br />
no phylogenetic significance at familial rank, so a well<br />
resolved phylogeny based on DNA comparisons will be<br />
necessary to categorize this genus.<br />
Semidelitschia Cain & Luck-Allen, Mycologia 61: 581<br />
(1969). (Delitschiaceae)<br />
Generic description<br />
Habitat terrestrial, saprobic (coprophilous). Ascomata<br />
immersed to slightly erumpent, scattered, coriaceous,<br />
papillate, ostiolate. Hamathecium of non-typical trabeculate<br />
pseudoparaphyses, thin, septate, rarely branching.<br />
Asci cylindrical, pedicellate, each with a conspicuous<br />
large apical ring. Ascospores non-septate, dark brown to<br />
nearly black, each with an elongated germ slit.<br />
Anamorphs reported for genus: none.<br />
Literature: Barr 2000; Cain and Luck-Allen 1969.<br />
Type species<br />
Semidelitschia agasmatica Cain & Luck-Allen, Mycologia<br />
61: 581 (1969). (Fig. 86)<br />
Ascomata 550–900 μm diam., solitary, immersed to<br />
erumpent, globose to subglobose, black, semicoriaceous,<br />
smooth-walled, with a protruding papilla and a conspicuous<br />
ostiole (Fig. 86a). Peridium thin, comprising multi-angular
Fungal Diversity<br />
cells from front view. Hamathecium of non-typical trabeculate<br />
pseudoparaphyses, 1–2 μm broad, septate, rarely<br />
branching, anastomosing not observed. Asci 410–505×(38-)<br />
43–50 μm (x ¼ 470:6 46:4mm, n=10), 8-spored, bitunicate,<br />
fissitunicate dehiscence not observed, cylindrical, with<br />
a thick pedicel which is up to 90 μm long, and with a large<br />
and conspicuous dome-shaped ocular chamber surrounded by<br />
apical ring (to 18 μm wide×4 μm high) (Fig. 86b and e).<br />
Ascospores 53–65×30–38 μm (x ¼ 61:3 34:1mm, n=10),<br />
uniseriate to obliquely uniseriate and partially overlapping,<br />
broad fusoid to subglobose, hyaline when<br />
young, then becoming yellowish brown, reddish brown<br />
Fig. 85 Salsuginea ramicola<br />
(from BRIP 17102, holotype).<br />
a Habitat section of an ascoma.<br />
b Section of the partial peridium.<br />
c Clavate mature and immature<br />
asci. d Ascospores<br />
within ascus. e Apical part of<br />
immature asci. f Ascospores<br />
with an apical chamber at each<br />
end. Scale bars: a=0.5 mm,<br />
b–e=50 μm, f=10μm
Fungal Diversity<br />
and nearly black and opaque when mature, non-septate,<br />
smooth-walled, with a full length germ slit, surrounded<br />
by a broad gelatinous sheath (Fig. 86c and d).<br />
Anamorph: none reported.<br />
Material examined: CANADA, Alberta, North of<br />
Beaver Mines, on sheep dung, 28 Jul. 1962, E.R. Luck-<br />
Allen, (TRTC 41607, paratype); USA, Montana: Gallatin<br />
County, 60 min S of Bozeman, on sheep dung, 2 Sept.<br />
1957, Cain (TRTC 42032, paratype); Stillwater County<br />
Columbus, on cow dung, 3 Sept. 1957, Cain (TRTC 42031,<br />
paratype); South Dakota, Meade Co.: South of Wall, on<br />
cow dung, 3 Sept. 1962, Cain (TRTC 40697, holotype).<br />
Notes<br />
Morphology<br />
Semidelitschia was formally establishedbyCainandLuck-<br />
Allen (1969) and was assigned to Sporormiaceae. Although it<br />
is similar to Delitschia, it differs as the ascospores are 1-<br />
celled, as opposed to 2-celled. Subsequently, Semidelitschia<br />
was transferred to Delitschiaceae together with Delitschia<br />
(Barr 2000). Currently, three species are listed under this<br />
genus, i.e. S. agasmatica Cain & Luck-Allen, S. nanostellata<br />
A.E. Bell & Mahoney and S. tetraspora J.H. Mirza & S.M.<br />
Khan (Index Fungorum) although the number of species in<br />
the genus are given as only two in Kirk et al. (2008).<br />
Phylogenetic study<br />
None.<br />
Concluding remarks<br />
This is a clearly defined genus that differs from<br />
Delitschia in having 1-celled ascospores. Cultures of S.<br />
agasmatica are needed for sequencing and for establishing<br />
the placement and uniqueness of the genus.<br />
Setomelanomma M. Morelet, Bull. Soc. Sci. nat. Arch.<br />
Toulon et du Var 227:15 (1980). (Phaeosphaeriaceae)<br />
Generic description<br />
Habitat terrestrial, hemibiotrophic or biotrophic. Ascomata<br />
small, solitary, scattered, immersed, erumpent to superficial,<br />
globose to subglobose, black; with or without a small<br />
papilla, apex covered with setae and a periphysate ostiole.<br />
Peridium thin, 1-layered, composed of several layers of<br />
cells of textura angularis. Hamathecium of dense, 1–2 μm<br />
broad pseudoparaphyses, septate, anastomosing. Asci 8-<br />
spored, bitunicate, broadly cylindrical. Ascospores fusoid to<br />
broadly clavate, pale brown to brown, 3-septate.<br />
Anamorphs reported for genus: none.<br />
Literature: Leonard and Suggs 1974; Morelet 1980;<br />
Rossman et al. 2002; Schochetal.2009; Zhang et al. 2009a.<br />
Type species<br />
Setomelanomma holmii M. Morelet, Bulletin de la Société<br />
des Sciences naturelles et d’Archéologie de Toulon et du<br />
Var 36 (no. 227): 15 (1980). (Fig. 87)<br />
(Some information in the following description is from<br />
Rossman et al. (2002))<br />
Ascomata 80–250 μm diam., solitary, scattered, immersed,<br />
erumpent to superficial, globose to subglobose,<br />
black, with setae; with or without a small papilla, apex<br />
covered with setae and a periphysate ostiole. Peridium 15–<br />
25 μm thick, 1-layered, composed of several layers of cells<br />
of textura angularis, cell wall thinner and more lightly<br />
pigmented towards centrum, cell wall thicker near the apex.<br />
Hamathecium of dense, 1–2 μm broad pseudoparaphyses,<br />
thicker near the base, septate, anastomosing (Fig. 87a and<br />
d). Asci 70–100×11–14 μm, 8-spored, bitunicate, broadly<br />
cylindrical with a short, thick, furcate pedicel, with a small<br />
ocular chamber (Fig. 87a, b and c). Ascospores 16–21×5–<br />
6.5 μm, obliquely uniseriate and partially overlapping to<br />
biseriate, fusoid to broadly clavate with broadly to narrowly<br />
rounded ends, pale brown to brown, 3-septate, slightly<br />
constricted at the median septum, smooth (Fig. 87e).<br />
Anamorph: none reported.<br />
Material examined: FRANCE, Leuglay, on dying twigs<br />
of Picea pungens. 8 May 1987, leg. M. Morelet (UPS F-<br />
117969 (slide), isotype).<br />
Notes<br />
Morphology<br />
Setomelanomma was formally established by Morelet<br />
(1980) as a monotypic genus represented by S. holmii,<br />
which was collected in France. The description, however, is<br />
not detailed and lacks illustrations. Rossman et al. (2002)<br />
collected this species in North America and detailed studies<br />
were conducted including both morphology and phylogeny.<br />
The bitunicate, broadly cylindrical asci, cellular pseudoparaphyses<br />
as well as the pale brown, septate ascospores with<br />
a median primary septum point Setomelanomma to<br />
Phaeosphaeriaceae as defined by Barr (1992a) and<br />
Eriksson et al. (2002) (Rossman et al. 2002). However, its<br />
setose ascomata, brown and 3-septate ascospores together<br />
with its residence in conifers distinguish it from all other<br />
genera under Phaeosphaeriaceae (Rossman et al. 2002).<br />
Setomelanomma is mostly comparable with Kalmusia and<br />
Phaeosphaeria. Setomelanomma can be distinguished from<br />
Kalmusia by its erumpent to superficial ascomata with<br />
almost no papilla, and Phaeosphaeria differs from Setomelanomma<br />
by its host spectrum and reported anamorphic<br />
stages (Rossman et al. 2002). Currently, five species are<br />
included in Setomelanomma, namely S. holmii, S. monoceras,<br />
S. prolata K.J. Leonard & Suggs, S. rostrata (K.J. Leonard) K.
Fungal Diversity<br />
Fig. 86 Semidelitschia agasmatica<br />
(from TRTC 40697, holotype).<br />
a Immersed ascomata<br />
scattered on the surface of the<br />
substrate. b Squash of ascoma.<br />
Note the numerous released asci.<br />
c Apical ring of cylindrical<br />
asci. d One-celled ascospores.<br />
Note the germ slits (see arrow).<br />
e Cylindrical ascus. Note the<br />
tapering pedicel. Scale bars:<br />
a=0.5 mm, b–e=100 μm<br />
J. Leonard & Suggs and S. turcica (Luttr.) K.J. Leonard &<br />
Suggs (http://www.mycobank.org/, 06/2010).<br />
Phylogenetic study<br />
Setomelanomma forms a well supported phylogenetic<br />
clade with other members of Phaeosphaeriaceae (Schoch<br />
et al. 2009; Zhang et al. 2009a).<br />
Concluding remarks<br />
None.<br />
Shiraia Henn., Bot. Jb. 28: 274 (1900). (<strong>Pleosporales</strong>,<br />
genera incertae sedis)<br />
Generic description<br />
Habitat terrestrial, parasitic. Ascostroma warty-like or<br />
tuber-like. Ascomata medium to large, subglobose, gregarious<br />
on the surface layer of ascostroma, immersed,<br />
ostiolate, with a small black opening seen on the surface<br />
of the ascostroma, ostiole rounded. Hamathecium of
Fungal Diversity<br />
dense, long trabeculate pseudoparaphyses, anastomosing<br />
and branching between the asci. Asci bitunicate, fissitunicate,<br />
cylindrical to cylindro-clavate, with a short furcate<br />
pedicel, with a big and truncate ocular chamber. Ascospores<br />
obliquely uniseriate and partially overlapping,<br />
narrowly fusoid to fusoid or broadly fusoid with tapering<br />
or narrowly rounded ends, hyaline to pale brown or<br />
brown, muriform.<br />
Anamorphs reported for genus: coelomycetous with<br />
muriform conidia (see Liu 2009).<br />
Literature: Cheng et al. 2004; Hino 1961; Kishi et al.<br />
1991; Liu 2009; Morakotkarn et al. 2008.<br />
Type species<br />
Shiraia bambusicola Henn., Bot. Jb. 28: 274 (1900).<br />
(Fig. 88)<br />
Ascostroma 1–1.5 cm high×1–2.5 cm diam., subglobose,<br />
oblong to irregular, slightly pink with cracking<br />
surface. Ascomata 350–800 μm high×300–700 μm diam.,<br />
subglobose, gregarious on the surface layer of ascostroma,<br />
immersed, ostiolate, with a small black opening seen on<br />
the surface of the ascostroma, ostiole rounded, the inner<br />
tissue of ascostroma carnation red (Fig. 88a and b).<br />
Hamathecium of dense, long trabeculate pseudoparaphyses,<br />
0.8–1.5 μm broad, anastomosing and branching<br />
between the asci. Asci 300– 425×20– 35 μm<br />
(x ¼ 360:5 28mm, n=10), 6-spored, bitunicate, fissitunicate,<br />
cylindrical to cylindro-clavate, with a short furcate<br />
pedicel, up to 50 μm long, with a big and truncate ocular<br />
chamber (Fig. 88c and d). Ascospores 62.5–80×17.5–<br />
22.5 μm (x ¼ 72:3 19:3mm, n=10), obliquely uniseriate<br />
and partially overlapping, narrowly fusoid to fusoid with<br />
tapering or narrowly rounded ends, hyaline turning pale<br />
brown when mature, muriform, with 9–13 transversal<br />
septa, 1–3 longitudinal septa in central cells, slightly<br />
constricted at the septa, usually with a gelatinous cap at<br />
each end (Fig. 88e, f and g).<br />
Anamorph: coelomycetous with muriform conidia (see<br />
Liu 2009).<br />
Material examined: CHINA, Zhejiang, Hangzhou,<br />
Panan, on bamboom, 15 Jun. 2009, leg. Liu Yongxiang<br />
(IFRD 2040).<br />
Notes<br />
Morphology<br />
Shiraia is reported as a parasite on branches of several<br />
genera of bamboo distributed mainly in southern regions of<br />
China and Japan (Hino 1961; Kishi et al. 1991; Liu 2009).<br />
Shiraia is characterized by its bambusicolous habitat, large<br />
ascostroma and muriform ascospores. Asci comprise 6<br />
ascospores in this study and some previous studies (Hino<br />
1961; Liu 2009). Shiraia bambusicola is well studied<br />
because of its medical effect in anticancer treatment (Kishi<br />
et al. 1991).<br />
Phylogenetic study<br />
Based on the SSU and ITS rDNA sequences analysis, its<br />
pleosporalean status was verified, and Shiraia was suggested<br />
to be closely related to Leptosphaeriaceae and/or<br />
Phaeosphaeriaceae (Pleosporineae) (Cheng et al. 2004).<br />
Based on the molecular phylogenetic analysis, another<br />
Shiraia-like fungus was reported which produced distinctive<br />
prawn-shaped conidioma-like structures (Morakotkarn<br />
et al. 2008), and differed from conidiomata in the anamorph<br />
of S. bambusicola described by Liu (2009).<br />
Concluding remarks A relatively broad species concept of<br />
Shiraia bambusicola is currently used, which could<br />
comprise several species.<br />
Sinodidymella J.Z. Yue & O.E. Erikss., Mycotaxon 24: 295<br />
(1985). (Teichosporaceae)<br />
Generic description<br />
Habitat terrestrial, saprobic? Ascomata medium to large,<br />
scattered, or in small groups, immersed, erumpent, to<br />
superficial, globose, subglobose, coriaceous, apex flattened,<br />
with radial ridges arranged around the central<br />
region. Peridium thick, 2-layered. Hamathecium of<br />
dense, broadly trabeculate pseudoparaphyses, anastomosing<br />
and branching between the asci. Asci 8-spored, with a<br />
short, furcate pedicel, bitunicate, cylindrical. Ascospores<br />
broadly ellipsoid, hyaline, becoming pale brown when<br />
mature, 1-septate, constricted at the median septum.<br />
Anamorphs reported for genus: none.<br />
Literature: Yue and Eriksson 1985.<br />
Type species<br />
Sinodidymella verrucosa (Petr.) J.Z. Yue & O.E. Erikss.,<br />
Mycotaxon 24: 295 (1985). (Fig. 89)<br />
≡ Amphididymella verrucosa Petr., Meddn Göteb. Bot.<br />
17: 129 (1947).<br />
Ascomata 620–930 μm high×800–1250 μm diam., scattered,<br />
or in small groups, immersed, becoming erumpent, to<br />
nearly superficial, globose, subglobose, coriaceous, apex<br />
flattened, with 3–6 radial ridges arranged around the central<br />
region, with a flattened base not easily removed from the<br />
substrate, wall black, roughened (Fig. 89a and b). Peridium<br />
100–150 μm thick, thinner at the base, 2-layered, outer layer<br />
thin, up to 40 μm thick, composed of small heavily pigmented<br />
thick-walled cells of textura globulosa, cells up to 5 μm<br />
diam., cell wall 3–6 μm thick, inner layer thick, composed of
Fungal Diversity<br />
hyaline small cells of textura epidermoidea, 2–4 μm diam.,<br />
cell wall 1–3 μm thick, interspersed with interwoven<br />
mycelium in places (Fig. 89b and c). Hamathecium of dense,<br />
broadly trabeculate pseudoparaphyses 1–2 μm broad, anastomosing<br />
between and above the asci (Fig. 89d). Asci 140–190<br />
(−205)×12.5–15(−17.5) μm (x ¼ 164 14:3mm, n=10), 8-<br />
spored, bitunicate, cylindrical, with a short, furcate pedicel,<br />
20–45 μm long, and an inconspicuous ocular chamber (to<br />
2 μm wide×1 μm high) (Fig. 89d and e). Ascospores 20–25×<br />
10–12 μm (x ¼ 22:1 10:3mm, n=10), obliquely uniseriate<br />
and partially overlapping, broadly ellipsoid with rounded<br />
ends, hyaline, becoming pale brown when mature, 1-septate,<br />
constricted at the median septum, smooth (Fig. 89f).<br />
Anamorph: none reported.<br />
Material examined: CHINA, Kansu Prov., between<br />
Scharakuto and Kweite, on rotten stems of Salsola gemmascens<br />
Pall., 25 Jul. 1935, G. Fenzel 2400 (W 16366, type).<br />
Notes<br />
Morphology<br />
Sinodidymella was formally established by Yue and<br />
Eriksson (1985) as they noticed that Amphididymella<br />
verrucosa Petr. was not congeneric with the generic type,<br />
A. adeana Petr., which is a pyrenolichen. Thus a new<br />
monotypic genus, Sinodidymella was introduced to accommodate<br />
it. The most outstanding morphological character of<br />
Sinodidymella is its radial ridges, which are somewhat<br />
comparable with that of Lophiostoma rugulosum Yin.<br />
Zhang, J. Fourn. & K.D. Hyde, although their pseudoparaphyses<br />
are dissimilar. Lophiostoma rugulosum has “tightly<br />
aggregated cellular pseudoparaphyses” and “apically ending<br />
into bunches of clavate cells” (Zhang et al. 2009b).<br />
Phylogenetic study<br />
None.<br />
Concluding remarks<br />
The radial ridges have little phylogenetic significance in<br />
genus level classification (Zhang et al. 2009b), but the<br />
broadly trabeculate pseudoparaphyses of Sinodidymella<br />
may fit Melanommataceae.<br />
Splanchnonema Corda, in Sturm, Deutschl. Fl., 3 Abt.<br />
(Pilze Deutschl.) 2(9), Tome 3: 115 (1829). (?Pleomassariaceae)<br />
Generic description<br />
Habitat terrestrial, saprobic. Ascomata medium to large,<br />
solitary or scattered, immersed in cortex with a pseudostromal<br />
covering, with a small ostiole appearing on the host<br />
surface, flattened subglobose. Peridium thin. Hamathecium<br />
Fig. 87 Setomelanomma holmii (from UPS F-117969 (slide), isotype). b<br />
a, b Asci with short pedicels in pseudoparaphyses. c Partial view of<br />
ascus. d Branching and septate pseudoparaphyses. a Three-septate<br />
lightly pigmented ascospores in ascus. Scale bars: a–e=10 μm<br />
of dense, cellular pseudoparaphyses, embedded in mucilage,<br />
anastomosing and branching. Asci bitunicate, fissitunicate,<br />
clavate to broadly cylindrical, with a short,<br />
narrowed, furcate pedicel. Ascospores clavate with a<br />
rounded apex and acute base, reddish brown, constricted<br />
at the septa.<br />
Anamorphs reported for genus: Myxocyclus, Steganosporium<br />
(Barr 1982b).<br />
Literature: Barr 1982b, 1993a; Boise 1985; Corda 1829;<br />
Eriksson 1981; Ramaley and Barr 1995; Shoemaker and<br />
LeClair 1975; Sivanesan 1984; Tanaka et al. 2005.<br />
Type species<br />
Splanchnonema pustulatum Corda, in Sturm, Deutschl. Fl.,<br />
3 Abt. (Pilze Deutschl.) 2(9), Tome 3: 115 (1829). (Fig. 90)<br />
Ascomata 400–600 μm high×550–1000 μm diam.,<br />
solitary or scattered, immersed in cortex with a pseudostromal<br />
covering, with a small ostiole appearing on the host surface,<br />
flattened subglobose (Fig. 90a). Peridium 15–25 μm thick,<br />
composed of small lightly pigmented thin-walled compressed<br />
cells (Fig. 90b). Hamathecium of dense, long cellular<br />
pseudoparaphyses 2–3 μm broad, embedded in mucilage,<br />
anastomosing and branching. Asci 200–250×30–<br />
45 μm (x ¼ 219:6 38:2mm, n=10), 8-spored, bitunicate,<br />
fissitunicate, clavate to broadly cylindrical, with a short,<br />
narrowed, furcate pedicel up to 35 μm long, without<br />
conspicuous ocular chamber (Fig. 90c and d). Ascospores<br />
45–53×20–24 μm (x ¼ 48:5 22:3mm, n=10), obliquely<br />
uniseriate and partially overlapping to biseriate, clavate<br />
with a rounded apex and acute base, reddish brown, 2-<br />
septate, apical cell largest, broader than the lower cells,<br />
basal cell smallest, constricted at the septa, smoothwalled,<br />
surrounded by a regular hyaline gelatinous<br />
sheath, 3–6 μm thick (Fig. 90e and f).<br />
Anamorph: none reported.<br />
Material examined: UK, Avon, nr Bath, Batheaston, on<br />
branch of Ulmus, C.E. Broome (L, No. 910.251-352, No.<br />
910.251-371).<br />
Notes<br />
Morphology<br />
A confusing outline of the history of Splanchnonema<br />
was provided by Shoemaker and LeClair (1975), which at<br />
the time was a valid, but little used name. Eriksson (1981)<br />
and Sivanesan (1984) stated (without comment) that the<br />
lectotype of Splanchnonema is S. pupula (Fr.) O. Kuntze.<br />
However, S. pustulatum is listed as the generic type in the
Fungal Diversity
Fungal Diversity<br />
Fig. 88 Shiraia bambusium<br />
(from IFRD 2040). a Ascostroma<br />
form a nubby structures<br />
on the twigs of host. b Vertical<br />
section of an ascostroma. Note<br />
the reddish staining of the inner<br />
tissue. c, d Cylindrical asci with<br />
a short pedicel. e–g Muriform<br />
fusoid hyaline ascospores. Scale<br />
bars: a=1 cm, b=1 mm, c, d=<br />
50 μm, e–g=20 μm<br />
online databases MycoBank and Index Fungorum. We<br />
assume Eriksson (1981) gained his data from Shoemaker<br />
and LeClair (1973), who considered S. pustulatum to be a<br />
synonym of S. pupula. Since we were unable to locate<br />
material of Corda or Fries we used a later collection of C.E.<br />
Broome.<br />
Splanchnonema can be distinguished from the morphologically<br />
comparable genera, i.e. Pleomassaria or<br />
Splanchospora by its depressed ascomata, and obovoid<br />
and asymmetrical ascospores (Barr 1982b). Currently,<br />
about 40 species are included in this genus. Barr<br />
(1993a) provided a key to 27 North American species,<br />
however, the inclusion of species with a range of<br />
ascospore types and immersed to superficial ascomata<br />
suggests the genus to be polyphyletic. Tanaka et al.<br />
(2005) suspected that the genus might include species of<br />
Pleomassaria, thus this genus needs further study.<br />
Phylogenetic study<br />
Splanchnonema platani (= Massaria platani) is poorly<br />
supported to be related to Lentitheciaceae (Schoch et al.<br />
2009).<br />
Fig. 89 Sinodidymella verrucosa (from W 16366, type). a Ascomata b<br />
on the host surface. Note the radial ridges around the pseudostiolar<br />
region. b Section of an ascoma. c Section of peridium. Note the<br />
hyaline small cells and interwoven hyphae. d Cylindrical asci in<br />
pseudoparaphyses. e Eight-spored ascus with short pedicel. f Hyaline,<br />
1-septate ascospores which turn pale brown when mature. Scale bars:<br />
a=1 mm, b=100 μm, c=50 μm, d–f=20 μm
Fungal Diversity
Fungal Diversity<br />
Concluding remarks<br />
Splanchnonema pustulatum has unique ascospores formed<br />
in immersed ascomata with thin walls, indicating that Splanchnonema<br />
sensu stricto shouldbeconfinedtoafewsimilar<br />
species. The type needs recollecting, sequencing and epitypifying<br />
in order to establish the phylogenetic relationships of<br />
this genus and to study what may be important defining<br />
characters. Also see entry under Pleomassaria.<br />
Sporormia De Not., Micromyc. Ital. Novi 5: 10 (1845).<br />
(Sporormiaceae)<br />
Generic description<br />
Habitat terrestrial, saprobic (coprophilous). Ascomata small,<br />
solitary, scattered, immersed to erumpent, globose, subglobose,<br />
wall black; apex without obvious papilla, ostiolate.<br />
Peridium thin. Hamathecium of rare, broad, septate pseudoparaphyses.<br />
Asci 8-spored, bitunicate, fissitunicate dehiscence<br />
not observed, short cylindrical, with a short, narrowed,<br />
furcate pedicel. Ascospores fasciculate, broadly filliform,<br />
reddish brown, multi-septate, easily separating into partspores,<br />
without visible germ-slits or pores.<br />
Fig. 90 Splanchnonema pustulatum (from L, No. 910.251–352, No.<br />
910.251–371). a Appearnce of ascomata on the host surface beneath a<br />
slightly raised area with minute ostiolar opening. b Section of the<br />
partial peridium. Note the compressed cells. c Dehiscent ascus. d<br />
Cluster of three asci joined in hymenium and pseudoparaphyses. e, f<br />
Asymmetric ascospores. Note the conspicuous sheath. Scale bars: a=<br />
1 mm, b–d=50 μm, e, f=20 μm
Fungal Diversity<br />
Anamorphs reported for genus: none.<br />
Literature: Ahmed and Asad 1968; Ahmed and Cain 1972;<br />
Kirschstein 1944; de Notaris 1849.<br />
Type species<br />
Sporormia fimetaria De Not., Micromyc. Ital. Novi 5: 10<br />
(1845). (Fig. 91)<br />
Ascomata 100–150 μm diam., solitary, scattered,<br />
immersed to erumpent, globose, subglobose, wall<br />
black; apex without obvious papilla, ostiolate<br />
(Fig. 91a). Peridium thin (other characters unknown).<br />
Hamathecium of rare, 2–3 μm wide, septate pseudoparaphyses.<br />
Asci 70–100×13–18 μm (x ¼ 86:4 14:9mm,<br />
n=10), 8-spored, bitunicate, fissitunicate dehiscence not<br />
observed, shortly cylindrical, with a short, narrowed,<br />
furcate pedicel up to 20 μm long, no apical apparatus<br />
could be observed (Fig. 91b, c and d). Ascospores 50–<br />
58×4–5 μm (x ¼ 54:7 4:8mm, n =10), fasciculate,<br />
broadly filliform, reddish brown, with 16 cells, easily<br />
separating into partspores, central cells of the ascospores<br />
shorter than broad, rectangular in vertical section, round<br />
in transverse section, 4–5×2.5–3.5 μm, without visible<br />
germ-slits or pores, apical cells usually longer than<br />
broad, 5–6.5 μm long, also without apertures (sheath is<br />
reported (Ahmed and Cain 1972), but not observed in<br />
this study) (Fig. 91e).<br />
Anamorph: none reported.<br />
Material examined: 1832, (RO, type, as Hormospora<br />
fimetaria De Not.).<br />
Notes<br />
Morphology<br />
Sporormia was formally established by de Notaris (1849),<br />
and only one species was described, i.e. S. fimetaria, which<br />
subsequently was selected as the generic type. Sporormia<br />
sensu stricto was accepted by several workers, and only<br />
includes members with a fasciculate ascospore arrangement,<br />
parallel to the ascus, and the part cells of the ascospores<br />
lacking germ-slits (Ahmed and Asad 1968; Ahmed and Cain<br />
1972; Kirschstein 1944). Species whose ascospores are not<br />
fasciculate and have partspores with germ-slits were assigned<br />
to Sporormiopsis by Kirschstein (1944) andtoSporormiella<br />
by Ahmed and Cain (1972).<br />
Phylogenetic study<br />
The generic status of Sporormia in <strong>Pleosporales</strong> was<br />
verified based on a phylogenetic analysis of ITS-nLSU<br />
rDNA, mtSSU rDNA and ß-tubulin sequences (Kruys and<br />
Wedin 2009). Sporormia clustered together with species of<br />
Westerdykella (including Eremodothis and Pycnidiophora),<br />
but lacks clear statistical support. Thus, the relationship of<br />
Sporormia with other genera of Sporormiaceae is unclear<br />
and not resolved yet.<br />
Concluding remarks<br />
Several coprophilous taxa (e.g. Chaetopreussia and<br />
Pleophragmia as well as Sporormiella nigropurpurea) in<br />
the <strong>Pleosporales</strong> were not included in the study by Kruys<br />
and Wedin (2009). Strains of these genera need to be<br />
collected and analyzed and their relationship with Sporormia<br />
established.<br />
Trematosphaeria Fuckel, Jb. nassau. Ver. Naturk. 23–24:<br />
161 (1870). (Trematosphaeriaceae)<br />
Generic description<br />
Habitat terrestrial or freshwater, saprobic. Ascomata<br />
subglobose, unilocular, erumpent to superficial, with<br />
papillate ostiole. Peridium thin, comprising several cell<br />
types. Hamathecium of dense, delicate, filliform, septate<br />
pseudoparaphyses. Asci bitunicate, fissitunicate,<br />
cylindro-clavate, normally 8-spored. Ascospores<br />
ellipsoid-fusoid to biconic, septate, smooth to finely<br />
verruculose, brown.<br />
Anamorphs reported for genus: hyphopodia-like (Zhang<br />
et al. 2008a).<br />
Literature: von Arx and Müller 1975; Barr 1979a;<br />
Boise 1985; Clements and Shear 1931; Zhang et al.<br />
2008a.<br />
Type species<br />
Trematosphaeria pertusa (Pers.) Fuckel, Jb. nassau. Ver.<br />
Naturk. 23–24: 161 (1870). (Fig. 92)<br />
≡ Sphaeria pertusa Pers., Syn. meth. fung. (Göttingen)<br />
1: 83 (1801).<br />
Ascomata 350–550 μm high×320–480 μm diam.,<br />
solitary, scattered, or in groups, initially immersed, becoming<br />
erumpent, to semi-immersed, subglobose, black; apex<br />
with a short ostiole usually slightly conical and widely<br />
porate, to 100 μm high (Fig. 92a and b). Peridium 48–<br />
55 μm wide laterally, to 80 μm at the apex, thinner at the<br />
base, 30–40 μm thick, coriaceous, 3-layered, comprising<br />
several cell types, one is of small heavily pigmented thickwalled<br />
cells of textura angularis, cells 4–8 μm diam., cell<br />
wall 1.5–3 μm thick in places with columns of textura<br />
prismatica orientated perpendicular to the ascomatal surface,<br />
apex cells smaller and walls thicker, forming thickwalled<br />
cells of textura pseudoparenchymata, and larger,<br />
paler cells of mixture of textura epidermoidea and textura<br />
angularis at the base (Fig. 92b, c and h). Hamathecium of<br />
dense, filamentous, 1.5–2.5 μm broad, septate pseudoparaphyses,<br />
embedded in mucilage, branching and anastomos-
Fungal Diversity<br />
ing between and above the asci (Fig. 92d, e and f). Asci<br />
100–145×15–17 μm (x ¼ 118 15:5mm, n=10), 8-spored,<br />
bitunicate, fissitunicate, cylindro-clavate, with a short,<br />
thick, furcate pedicel which is 12–30 μm long, with a<br />
truncate ocular chamber (Fig. 92d, e, f, g and i). Ascospores<br />
27.5–32.5×7.5–8.5 μm (x ¼ 29:5 8mm, n=10), biseriate<br />
to uniseriate near the base, fusoid with broadly to narrowly<br />
rounded ends, dark brown, 1–3-septate, secondary septum<br />
forming late or often absent, constricted at the median<br />
septum, the upper cell often shorter and broader than the<br />
lower one, smooth to finely verruculose, containing<br />
refractive globules (Fig. 92j and k).<br />
Anamorph: Only hyphopodia-like structures (or conidia?)<br />
observed (Zhang et al. 2008a).<br />
Colonies (of epitype) reaching 5 cm diam. after 20 days<br />
growth on MEA at 25°C, raised, woolly, deep grey, with<br />
irregular to rhizoidal margin, reverse darkened. Hyphopodialike<br />
structures (or conidia?) produced after 6 months, hyaline<br />
to pale brown, lobed, 4–4.5(−5) μm long and 3–3.5 μm diam.<br />
Material examined: EUROPE, Upsala, on decaying<br />
wood, designated by Boise (1985), (L-Pers 910269–172,<br />
as Sphaeria pertusa Pers., neotype); FRANCE, Deux<br />
Sèvres, Sansais, Le Vanneau, Les Grandes Mottines,<br />
swamp, on bark of a dead stump of Fraxinus excelsior,<br />
25 Apr. 2004, J. Fournier (IFRD 2002, epitype); Haute<br />
Garonne, Avignonet, Canal du Midi, on submerged wood<br />
of Platanus in a canal, 23 Nov. 2006, Michel Delpont, det.<br />
J. Fournier (IFRD2003).<br />
Notes<br />
Morphology<br />
Trematosphaeria was formally established in ‘Rhenish<br />
fungi’ by Fuckel (1870) based on the broadly pertuse<br />
ascomata, and Fries (1823) assigned it under Ascomycetes,<br />
Pyrenomycetes, Lophiostomataceae. Subsequently, Winter<br />
(1885) placed Trematosphaeria in Amphisphaeriaceae.<br />
Berlese (1890), however, treated Trematosphaeria as a<br />
synonym of Melanomma (Melanommataceae). After establishment<br />
of Loculoascomycetes (Luttrell 1955), Trematosphaeria<br />
was assigned to Pleosporaceae (Loculoascomycetes,<br />
<strong>Pleosporales</strong>) (Holm1957), and this was followed by von<br />
Arx and Müller (1975). Trematosphaeria was assigned to<br />
Melanommataceae by Barr (1979a), and this has been<br />
widely followed (Eriksson 2006; Kirketal.2001; Lumbsch<br />
and Huhndorf 2007).<br />
Trematosphaeria pertusa, the lectotype species of Trematosphaeria<br />
(Clements and Shear 1931), is characterized by<br />
having semi-immersed to erumpent ascomata, filamentous<br />
pseudoparaphyses, cylindro-clavate asci, fusoid, 1-septate<br />
reddish brown to dark brown ascospores (Zhang et al.<br />
2008a). All of these characters are quite different from those<br />
of Melanomma, the familial type of Melanommataceae.<br />
Fig. 91 Sporormia fimetaria (from RO, type). a Appearance of<br />
ascomata on the host surface. Note the scattered distribution. b–d<br />
Broad cylindrical asci with a short and thick pedicel. e Released<br />
filiform ascospores which may break up into part spores. Scale bars:<br />
a=0.5 mm, b–d=20 μm, e=10 μm<br />
Phylogenetic study<br />
Trematosphaeria pertusa forms a robust phylogenetic<br />
clade with Falciformispora lignatilis and Halomassarina<br />
thalassiae, and they are all assigned to Trematosphaeriaceae<br />
(Suetrong et al. 2009; Zhang et al. 2009a; Plate 1).<br />
Concluding remarks<br />
Trematosphaeria pertusa is a terrestrial species which<br />
can also survive in a freshwater environment. However,<br />
both Falciformispora lignatilis and Halomassarina thalassiae<br />
are marine fungi. Their habitat difference may indicate<br />
their distant relationship, at least above genus level.<br />
Verruculina Kohlm. & Volkm.-Kohlm., Mycol. Res. 94:<br />
689 (1990). (Testudinaceae)<br />
Generic description<br />
Habitat marine, saprobic. Ascomata medium-sized, solitary<br />
under clypeate, immersed to semi-immersed, subglobose to<br />
depressed ellipsoidal, papillate, ostiolate, periphysate,<br />
black, carbonaceous. Peridium thin, comprising a few<br />
layers of cells of textura angularis. Hamathecium of long<br />
cellular pseudoparaphyses, embedded in mucilage, hyaline,<br />
septate and sparsely branching. Asci 8-spored, bitunicate,<br />
fissitunicate, cylindrical, with short pedicels, ocular chamber<br />
not observed. Ascospores biseriate, ovoid or ellipsoidal,<br />
dark brown, 1-septate, constricted at the septum, verrucose<br />
or verruculose, with or without germ pore.<br />
Anamorphs reported for genus: none.<br />
Literature: Kohlmeyer and Volkmann-Kohlmeyer 1990;<br />
Suetrong et al. 2009.<br />
Type species<br />
Verruculina enalia (Kohlm.) Kohlm. & Volkm.-Kohlm.,<br />
Mycol. Res. 94: 689 (1990). (Fig. 93)<br />
≡ Didymosphaeria enalia Kohlm., Ber. dt. bot. Ges. 79:<br />
28 (1966).<br />
Ascomata 295–480 μm high×140–520 μm diam., solitary<br />
under clypeate, immersed to semi-immersed, subglobose to<br />
depressed ellipsoidal, ostiolate, papillate, periphysate, black,<br />
carbonaceous. Peridium thin, comprising a few layers of cells<br />
of textura angularis. Hamathecium of long cellular pseudoparaphyses,<br />
1.5–2 μm broad, embedded in mucilage, hyaline,<br />
septate and sparsely branching. Asci 177–135×12.5–15.5 μm,<br />
8-spored, bitunicate, fissitunicate, cylindrical, with short<br />
furcate pedicels, ocular chamber not observed (Fig. 93a).<br />
b
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Fungal Diversity<br />
Ascospores 16.5–23×7.5–10 μm, biseriate, ovoid or ellipsoidal,<br />
dark brown, 1-septate, constricted at the septum, verrucose<br />
or verruculose, with or without germ pore (Fig. 93b).<br />
Anamorph: none reported.<br />
Material examined: SEYCHELLES, Victoria, on submerged<br />
branch of Rhizophora mangle L., Mar. 2004, K.D.<br />
Hyde (KDH 2137, slide).<br />
Notes<br />
Morphology<br />
Verruculina was introduced to accommodate an obligate<br />
marine species, i.e. Verruculina enalia (Kohlmeyer<br />
and Volkmann-Kohlmeyer 1990). Verruculina is characterized<br />
by immersed, clypeate, carbonaceous, ostiolate and<br />
papillate ascomata. The peridium is composed of cells of<br />
textura angularis. Pseudoparaphyses are trabeculate and<br />
embedded in mucilage. Asci are 8-spored, cylindrical with<br />
short pedicels and ocular chamber, and ascospores are<br />
ellipsoidal, 1-septate, dark brown, verrucose or verruculose.<br />
The partly or completely immersed clypeate ascomata<br />
of V. enalia is comparable with those of<br />
Didymosphaeria futilis, but it differs from the later by<br />
the dark peridium, gelatinous matrix around the pseudoparaphyses,<br />
stipitate asci with an ocular chamber, and the<br />
verruculose ascospores (Kohlmeyer and Volkmann-<br />
Kohlmeyer 1990).<br />
Phylogenetic study<br />
Based on multigene phylogenetic analysis, Verruculina<br />
enalia nested within Testudinaceae (Suetrong et al. 2009).<br />
Thus, its familial placement seems clarified.<br />
Concluding remarks<br />
None.<br />
Westerdykella Stolk, Trans. Br. Mycol. Soc. 38: 422<br />
(1955). (Sporormiaceae)<br />
Generic description<br />
Habitat terrestrial, saprobic (coprophilous). Ascomata<br />
small, scattered on the upper layer of the culture medium,<br />
wall black. Peridium thin, composed of one layer of cells of<br />
polygonal, dark brown, thick-walled cells. Hamathecium<br />
not observed. Asci 32-spored, bitunicate nature undetermined,<br />
fissitunicate dehiscence not observed, subglobose to<br />
ellipsoid, arranged in the centre of the ascomata, with or<br />
without a short pedicel. Ascospores globose, brown, 1-celled,<br />
without germ pore.<br />
Anamorphs reported for genus: Phoma-like (von Arx 1974).<br />
Literature: von Arx 1973, 1981; Kruys et al. 2006; Kruys<br />
and Wedin 2009; Stolk 1955a.<br />
Fig. 92 Trematosphaeria pertusa (a, d, f–i from epitype, b, c, e, j b<br />
from neotype). a Ascomata on the host surface. b Section of an ascoma.<br />
c, h Section of the peridium. c shows the peridium structure at sides,<br />
and h indicates the basal peridium structure. Note the hyaline and thinwalled<br />
cells in (h). d Asci amongst pseudoparaphyses. e Ascus with<br />
pedicle. f, g Dehiscent ascus. i Upper part of the ascus, showing the<br />
ocular chamber and the mucilage covering the apex. j, k Ascospores.<br />
Scale bars: a=0.5 mm, b, c=100 μm, d–h=20 μm, i–k=10 μm<br />
Type species<br />
Westerdykella ornata Stolk, Trans. Br. Mycol. Soc. 38: 422<br />
(1955). (Fig. 94)<br />
Ascomata 100–300 μm diam., cleistothecoid, scattered<br />
on the upper layer of the culture medium, wall black<br />
(Fig. 94a). Peridium composed of one layer of cells of<br />
polygonal in front view, dark brown, thick-walled cells,<br />
ca. 5μm diam. Hamathecium not observed. Asci 25–32×<br />
16–22 μm, 32-spored, bitunicate nature undetermined,<br />
fissitunicate dehiscence not observed, subglobose to<br />
ellipsoid, arranged in the centre of the ascomata, with<br />
a short furcate pedicel best seen in immature asci<br />
(Fig. 94b, c, d and f). Ascospores 6.2–7×6–6.8 μm,<br />
globose, brown, 1-celled, ornamented with irregular spiral<br />
bands, which occur in four to five coils, without germ<br />
pore (Fig. 94e).<br />
Anamorph: none reported.<br />
On MEA colonies spreading, but somewhat erumpent,<br />
with moderate aerial mycelium and even, lobate margins;<br />
surface dirty white with luteous to orange patches; reverse<br />
orange to sienna. On PDA similar but with sparse aerial<br />
mycelium; surface with patches of orange to luteous and<br />
dirty white; reverse luteous with cream margins. On OA<br />
flat, spreading with sparse aerial mycelium; surface with<br />
luteous and dirty white patches and transparent margins;<br />
sporulating on OA, visible as black masses of aggregated<br />
ascomata; colonies reaching 4 cm diam. on all media (based<br />
on <strong>CBS</strong> 379.55).<br />
Material examined: MOZAMBIQUE, Inhaca, leg. H.J.<br />
Swart, mangrove mud (<strong>CBS</strong> 379.55, holotype).<br />
Notes<br />
Morphology<br />
Westerdykella was introduced to accommodate a<br />
coprophilous fungus, which is characterized by cleistothecioid<br />
and membraneous ascomata (Stolk 1955a). Asci<br />
are subglobose to ellipsoid, stalked, many-spored and<br />
evanescent. Ascospores are globose to subglobose,<br />
brown, ornamented with spiral bands, without germ pores<br />
(Stolk 1955a). Westerdykella was assigned under Phaeosporeae<br />
of the Eurotiaceae (Stolk 1955a), and was<br />
assigned to Sporormiaceae by von Arx and Müller<br />
(1975). Based on the spore ornamentation, von Arx and
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Fungal Diversity<br />
Fig. 93 Verruculina enalia (from KDH 2137, slide). a Cylindrical asci with short pedicels. b One-septate verruculose ascospores. Scale bars:<br />
a=20 μm, b=10 μm<br />
van der Aa (1987) andBarr(2000) acceptedWesterdykella as<br />
a separate genus, but this is not supported by molecular<br />
phylogenetic analysis (Kruys and Wedin 2009).<br />
Phylogenetic study<br />
Phylogenetic reconstructions indicated that both Pycnidiophora<br />
and Eremodothis should be treated as synonyms<br />
of Westerdykella (Kruys and Wedin 2009).<br />
Concluding remarks<br />
Westerdykella is another example where ascospore ornamentation<br />
can be phylogenetically uninformative. Westerdykella<br />
is proved a good genus of Sporormiaceae (Kruys and Wedin<br />
2009).<br />
Wettsteinina Höhn., Sber. Akad. Wiss. Wien, Math.-naturw.<br />
Kl., Abt. I 116: 126 (1907). (?Lentitheciaceae)<br />
Generic description<br />
Habitat terrestrial or freshwater? hemibiotrophic or<br />
saprobic. Ascomata generally small, scattered, immersed<br />
with a protruding broad papilla. Peridium very thin,<br />
composed of few layers of thin-walled large polygonal<br />
cells in surface view. Hamathecium deliquescing at<br />
maturity. Asci bitunicate, fissitunicate, subglobose to<br />
obpyriform, without a pedicel, with small truncate ocular<br />
chamber. Ascospores hyaline and turning pale brown<br />
when mature, septate, upper second cell enlarged, slightly<br />
constricted at the second septum, smooth, surrounded by a<br />
hyaline gelatinous sheath.<br />
Anamorph reported for genus: Stagonospora (Farr et al.<br />
1989).<br />
Literature: Barr 1972; Müller 1950; Shoemaker and<br />
Babcock 1987, 1989b.<br />
Type species<br />
Wettsteinina gigaspora Höhn., Sber. Akad. Wiss. Wien,<br />
Math.-naturw. Kl., Abt. 1 116: 126 (1907). (Fig. 95)<br />
Ascomata 150–250 μm diam., scattered, immersed<br />
with protruding broad papillae, 50–90 μm diam.<br />
Peridium thin, composed of few layers of thin-walled<br />
large polygonal cells in surface view, 6–15 μm diam.<br />
(Fig. 95a). Hamathecium deliquescing at maturity. Asci<br />
140–200×75–120 μm, 8-spored, bitunicate, fissitunicate,<br />
subglobose to obpyriform, lacking a pedicel, with a small<br />
truncate ocular chamber (to 8 μm wide×5 μm high)<br />
(Fig. 95b and c). Ascospores 90–110×25–30 μm, 2–4-<br />
seriate, hyaline and turning pale brown when mature,<br />
broadly clavate, 4-septate, primary septum distinct and<br />
constricted forming 1/3 rd from the apex of the ascospore,<br />
complete, secondary septa less distinct and slightly<br />
constricted, incomplete, with one forming above and two<br />
forming below the primary septum, largest cell the second<br />
cell from apex, smooth, surrounded by a hyaline gelatinous<br />
sheath 5–8 μm thick (Fig. 95d and e).<br />
Anamorph: none reported.<br />
Material examined: SLOVENIA, Postojna, on Genista<br />
sagittalis leg. Stapf. det. H. Rehm. (S, holotype of<br />
Massarina gigantospora).
Fungal Diversity<br />
Fig. 94 Westerdykella ornata (from <strong>CBS</strong> 379.55, holotype). a Appearance of the ascomata on culture substrate surface. b–f Mature and<br />
immature asci as well as the released ascospores. Note the spiral bands around the ascospores. Scale bars: a=1 mm, b–f=10 μm<br />
Notes<br />
Morphology<br />
Confusion exists in the generic type of Wettsteinina.<br />
Höhnel (1907) described W. gigaspora when introducing<br />
Wettsteinina, and listed it as the first species of Wettsteinina.<br />
ClementsandShear(1931) accepted W. gigaspora as<br />
the generic type of Wettsteinina, which is followed by<br />
Shoemaker and Babcock (1987). But according to http://<br />
www.indexfungorum.org (June 2011), W. gigantospora is<br />
the generic type of Wettsteinina. Both W. gigantospora<br />
and W. gigaspora were treated as the synonyms of W.<br />
mirabilis (Niessl) Höhn. http://www.indexfungorum.org<br />
(June, 2011, Synonymy Contributor: <strong>CBS</strong> (2010)). We
Fungal Diversity<br />
tentatively described the generic type of W. gigantospora<br />
as a representing of the type of W. gigaspora here.<br />
New family names, i.e. Pseudosphaeriaceae and Wettsteininaceae<br />
(as Wettsteiniaceae) and a new order, Pseudosphaeriales<br />
had been introduced to accommodate Wettsteinina and<br />
its synonym Pseudosphaeria (Höhnel 1907; Locquin 1972).<br />
After a systematic study, Wettsteinina was included in<br />
Pleosporaceae basedonits“Pleospora-type” centrum, and<br />
Pseudosphaeriaceae and Wettsteininaceae are treated as<br />
synonyms of Pleosporaceae (Shoemaker and Babcock 1987).<br />
Phylogenetic study<br />
Wettsteinina macrotheca (Rostr.) E. Müll., W. pachyasca<br />
(Niessl) Petr. and W. dryadis (Rostr.) Petr. were reported to be<br />
closely related to Pleomassaria siparia (Melanommataceae)<br />
(Kodsueb et al. 2006a), and W. lacustris (Fuckel) Shoemaker<br />
& C.E. Babc. nested within Lentitheciaceae (Schoch et al.<br />
2009). The generic type has not been sequenced.<br />
Concluding remarks<br />
The most striking character for Wettsteinina is its<br />
asymmetrical ascospores, thick-walled obpyriform asci<br />
and lack of pseudoparaphyses at maturity. These<br />
characters are comparable with genera in the Capnodiales<br />
and Venturiales. The phylogenetic significance of<br />
these characters are not fully understood, while the<br />
hemibiotrophic or saprobic life style may indicate its<br />
polyphyletic nature (Shoemaker and Babcock 1987).<br />
Strains from the genus, in particular the generic type<br />
require DNA sequence data so that the phylogenetic<br />
placement can be investigated.<br />
Wilmia Dianese, Inácio & Dorn. -Silva, Mycologia 93:<br />
1014 (2001). (Phaeosphaeriaceae)<br />
Generic description<br />
Habitat terrestrial, hemibiotrophic or biotrophic. Ascomata<br />
small, scattered, immersed, globose to subglobose, papillate.<br />
Peridium thin, composed of a few layers of brown, thickwalled<br />
cells of textura angularis to prismatica. Hamathecium<br />
comprising filliform, septate, rarely branching, evanescent,<br />
cellular pseudoparaphyses embedded in mucilage. Asci<br />
bitunicate, fissitunicate, cylindrical to clavate, with a short,<br />
furcate pedicel and ocular chamber. Ascospores fusoid, pale<br />
brown, 1-septate.<br />
Anamorphs reported for genus: see below.<br />
Literature: Dianese et al. 2001.<br />
Type species<br />
Wilmia brasiliensis Dianese, Inácio & Dorn.-Silva, Mycologia<br />
93: 1014 (2001). (Fig. 96)<br />
Ascomata 175–240 μm high×95–145 μm diam., scattered,<br />
immersed, globose to subglobose; apex with a short<br />
papilla, 40–80 μm long, ostiolate, periphysate, periphyses<br />
up to 90 μm long (Fig. 96a and b). Peridium 6–15 μm<br />
wide, 1-layered, composed of 3–7 layers of brown, thickwalled<br />
cells of textura angularis to prismatica, cells 4–<br />
9 μm diam., cell wall 2–4 μm thick (Fig. 96a and b).<br />
Hamathecium of long cellular pseudoparaphyses 2–3 μm<br />
broad, septate, rarely branching, embedded in mucilage,<br />
evanescent. Asci 65–95×9.5–14 μm (x ¼ 78:5 11:5mm,<br />
n=10), 8-spored, bitunicate, fissitunicate, cylindrical to<br />
clavate, with a short, furcate pedicel and a small ocular<br />
chamber (Fig. 96c, d and f). Ascospores 22.5–28×5–<br />
8.5 μm (x ¼ 26:5 6:8mm, n=10), biseriate, fusoid with<br />
narrowly rounded ends, pale brown, 1-septate, constricted<br />
at the septum, the upper cell often shorter and broader than<br />
the lower one, smooth, with or without sheath (Fig. 96d and<br />
e).<br />
Anamorph: Conidiomata 170–200 μm high×85–130 μm<br />
diam., eustromatic, immersed, subglobose to irregular,<br />
ostiolate, brown. Peridium thin, 1–2 walllayers,6–8 μm<br />
thick, thicker near the apex. Ostiole 50–63 μm high×30–35<br />
broad. Conidiogenous cells ampulliform or lageniform,<br />
phialidic, aseptate. Conidia 13–20×4–7 μm, ellipsoid,<br />
oblong, ovoid, hyaline (Dianese et al. 2001).<br />
Material examined: BRAZIL, Distrito Federal, Vargem<br />
Bonita, Fazenda Agua Limpa, on leaves of Memora<br />
pedunculata (Vell.) Miers, 18 May 1995, Carlos A. Inácio<br />
(UB Col. Microl 8438 holotype).<br />
Notes<br />
Morphology<br />
Wilmia was formally established by Dianese et al. (2001)<br />
as a monotypic genus represented by W. brasiliensis, which<br />
causes leaf spots on Memora pedunculata. The peridium of<br />
W. brasiliensis comprises a few layers of brown, thickwalled<br />
textura angularis to prismatica cells, and it also has<br />
cellular pseudoparaphyses, clavate asci, 1-septate pale<br />
brown ascospores (Dianese et al. 2001).<br />
Phylogenetic study<br />
None.<br />
Concluding remarks<br />
The dicotyledonous host habit of Wilmia brasiliensis seems<br />
in agreement with Leptosphaeriaceae rather than Phaeosphaeriaceae.<br />
But a verified conclusion can only be reached by<br />
further molecular phylogenetic study.<br />
Xenolophium Syd., Bulletin of the Bernice P. Bishop<br />
Museum, Honolulu, Hawaii 19: 96 (1925). (<strong>Pleosporales</strong>,<br />
genera incertae sedis)
Fungal Diversity<br />
Fig. 95 Wettsteinina gigantospora<br />
(from S, holotype of<br />
Massarina gigantospora). a<br />
Ascomata with protruding papilla<br />
scattered on the host surface.<br />
b Obpyriform thick-walled<br />
ascus with small apical apparatus.<br />
c Fissitunicate ascus. d<br />
Released hyaline ascospores.<br />
Note the distinct primary septum<br />
and less distinct secondary septa.<br />
e Ascospore with sheath.<br />
Scale bars: a=0.5 mm, b–d=<br />
100 μm, e=50 μm<br />
Generic description<br />
Habitat terrestrial, saprobic on wood. Ascomata nearly<br />
superficial, scattered to gregarious, globose, large, with a<br />
conspicuous compressed papilla and large slit-like ostiole.<br />
Peridium carbonaceous. Hamathecium of dense, long<br />
trabeculate pseudoparaphyses, branching and anastomosing<br />
between and among asci. Asci 8-spored, clavate, with very
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long furcate pedicels. Ascospores fusoid to narrowly fusoid,<br />
light to dark brown, 1-septate, constricted at the septum.<br />
Anamorphs reported for genus: none.<br />
Literature: Chesters and Bell 1970; Huhndorf 1993;<br />
Mugambi and Huhndorf 2009b; Müller and von Arx<br />
1962; Stevens 1925.<br />
Type species<br />
Xenolophium leve Syd., Bulletin of the Bernice P. Bishop<br />
Museum, Honolulu, Hawaii 19: 97 (1925) (Fig. 97)<br />
Current name: Xenolophium applanatum (Petch)<br />
Huhndorf, Mycologia 85: 493 (1993).<br />
≡ Schizostoma applanatum Petch, Ann. Roy. Bot. Gard.<br />
(Peradeniya) 6: 231 (1916).<br />
Ascomata 1–1.5 mm diam., scattered to clustered, erumpent<br />
to superficial, globose with base immersed in host tissue, wall<br />
black, carbonaceous, roughened with ridges, papillate. Apex<br />
with a conspicuous hysteriform papilla extending on the sides,<br />
1–1.4 mm long, 0.4–0.5 mm wide, 0.2–0.3 mm high, smooth,<br />
ostiole slit-like, nearly as long as papilla length (Fig. 97a).<br />
Peridium 140–160 μm thick, pseudoparenchymatous, composed<br />
of two distinct layers: outer crust 16–45 μm thick,<br />
blackish, of heavily melanized, nearly opaque thick-walled<br />
angular cells, of uneven thickness forming irregular strands<br />
extending into the inner layer; inner layer subhyaline,<br />
composed of thick-walled prismatic to angular cells, with<br />
columns or patches of darker thick-walled cells extending<br />
inwardly from the outer layer; papilla wall 200–220 μm thick,<br />
of heavily melanized angular thick-walled cells (Fig. 97b and<br />
c). Hamathecium of dense, very long trabeculate pseudoparaphyses<br />
0.8–1.5 μm broad, embedded in mucilage, anastomosing<br />
and branching between and above the asci. Asci 104–152×<br />
9–12 μm (excluding pedicel) (x ¼ 149 10:2mm, n=10), 8-<br />
spored, bitunicate, fissitunicate dehiscence not observed,<br />
clavate, with a long, narrowed, furcate pedicel which is 50–<br />
75 μm long (Fig. 97d). Ascospores 17–26×4–5.5 μm<br />
(x ¼ 22:5 4:8mm, n=10), upper biseriate and lower uniseriate,<br />
fusoid, straight to slightly curved, equally 1-septate, constricted<br />
at the septum, the upper cell slightly wider, with one or rarely<br />
two additional septa appearing on a small number of senescent<br />
ascospores, pale brown, median septum darker, constricted,<br />
smooth, without sheath or appendages (Fig. 97e, f and g).<br />
Anamorph: none reported.<br />
Material examined: MARTINIQUE, Morne Rouge, on<br />
rotten wood, leg C. Lécuru, det Jacques Fournier, 29 Aug.<br />
2007, IFRD 2038.<br />
Notes<br />
Morphology<br />
Xenolophium was formally established by Sydow (in<br />
Stevens 1925) to accommodate two species, i.e. X. leve and<br />
X. verrucosum, ofwhichX. leve is selected as the generic type<br />
(Huhndorf 1993). Because of its morphological similarity with<br />
some genera, such as Ostropella and Schizostoma, Xenolophium<br />
has been treated as a synonym of Ostropella (Müller<br />
and von Arx 1962)orevenofLophiostoma (Chesters and Bell<br />
1970). Huhndorf (1993) clarified the circumscription of<br />
Xenolophium and treated X. leve as a synonym of Schizostoma<br />
applanata. Xenolophium mainly differs from Ostropella in<br />
lack of “organized cell composition and triangular pattern of<br />
melanization” in the peridium (Huhndorf 1993).<br />
Phylogenetic study<br />
The polyphyletic nature of Xenolophium has been<br />
demonstrated (Mugambi and Huhndorf 2009b). The generic<br />
type of Xenolophium (X. leve, current name X. applanatum)<br />
clustered together with Ostropella albocincta (generic type<br />
of Ostropella), and both locate in Platystomaceae<br />
(Mugambi and Huhndorf 2009b).<br />
Concluding remarks<br />
The large ascomata with slit-like ostioles, hamathecium of<br />
numerous and trabeculate pseudoparaphyses, clavate asci with<br />
long pedicels, and the pale brown, 1-septate ascospores of<br />
Xenolophium leve are all comparable with those of Ostropella<br />
albocincta. However, the phylogenetic results do not support<br />
them being congeneric (Mugambi and Huhndorf 2009b).<br />
Synonyms<br />
Javaria Boise, J.R., Acta Amazonica 14(Supl.): 50 (1984).<br />
(Melanommataceae)<br />
Current name: Astrosphaeriella Syd. & P. Syd., Annls<br />
mycol. 11: 260 (1913).<br />
Generic description<br />
Habitat terrestrial, saprobic. Ascomata medium-sized, scattered,<br />
erumpent to nearly superficial, reflexed pieces of the<br />
ruptured host tissue usually persisting around the surface of<br />
the ascomata; ascomata broadly conical, with a flattened<br />
base not easily removed from the substrate, wall black,<br />
papillate. Peridium carbonaceous. Hamathecium of trabeculate<br />
pseudoparaphyses. Asci 8-spored, bitunicate, fissitunicate,<br />
cylindro-clavate to narrowly fusoid, with a short,<br />
narrowed, furcate pedicel. Ascospores elongate-fusoid,<br />
hyaline, 1-septate, constricted at the septum.<br />
Anamorphs reported for genus: none.<br />
Literature: Barr 1990a; Boise 1984.<br />
Type species<br />
Javaria samuelsii Boise, J.R., Acta Amazonica 14(Supl.):<br />
50 (1984) (Fig. 98)
Fungal Diversity<br />
Fig. 96 Wilmia brasiliensis (from UB Col. Microl 8438, holotype). a<br />
Section of an ascoma. Note the setae in the ostiole. b Conidioma of<br />
the coelomycetous anamorphic stage. c, d Clavate asci with short<br />
furcate pedicels. e, f Released 1-septate pale brown ascospores. Scale<br />
bars: a, b=100 μm, c, d=20 μm, e, f=10 μm
Fungal Diversity<br />
Fig. 97 Xenolophium applanatum (from IFRD 2038). a Gregarious<br />
ascomata on the host surface. Note protruding papilla and slit-like<br />
ostiole. b Vertical section of the papilla and ostiole. c Section of the<br />
partial peridium. Note the two layers of the peridium. d Eight-spored<br />
asci in trabeculate pseudoparaphyses. Note the long pedicels. e–g Pale<br />
brown ascospores. Scale bars: a=2 mm, b=200 μm, c=50 μm, d=<br />
20 μm, e–g=10 μm
Fungal Diversity<br />
Current name: Astrosphaeriella samuelsii Boise, Acta<br />
Amazon., Supl. 14(1–2, Suppl.): 50 (1986) [1984].<br />
Ascomata 300–380 μm diam., scattered, erumpent<br />
through the outer layers of the host tissues, to nearly<br />
superficial, reflexed pieces of the ruptured host tissue<br />
usually persisting around the surface of the ascomata;<br />
ascomata broadly conical, with a flattened base not easily<br />
removed from the substrate, wall black, papillate (Fig. 98a).<br />
Peridium 50–80 μm thick, carbonaceous and crisp, 1-<br />
layered. Hamathecium of dense, long trabeculate pseudoparaphyses,<br />
0.8–1.5 μm broad, embedded in mucilage,<br />
anastomosing between and above the asci. Asci 140–185×<br />
17.5–20 μm (x ¼ 158 19:4mm, n=10), 8-spored, bitunicate,<br />
fissitunicate, cylindro-clavate to narrowly fusoid, with<br />
a short, narrowed, furcate pedicel up to 20 μm long<br />
(Fig. 98b and c). Ascospores 48–55(−60)×6–7.5(−10) μm<br />
(x ¼ 52:2 7:7mm, n=10), biseriate, elongate- fusoid,<br />
gradually tapering towards the ends, hyaline, surrounded<br />
with sheath, 2–5 μm thick, 1-septate, constricted at the<br />
septum (Fig. 98d).<br />
Anamorph: none reported.<br />
Material examined: Serra Araca, 60 m, terra firme,<br />
open forest, deep litter. Dry. 10–13 Mar. 1984, det. Jean R.<br />
Boise, G.J. Samuels (isotype).<br />
Notes<br />
Morphology<br />
Javaria was introduced by Boise (1984) based on<br />
seven Amazonian collections on decaying palm petioles; it<br />
is comparable with Astrosphaeriella in numerous characters.<br />
But Javaria differs from Astrosphaeriella by its<br />
hyaline ascospores with sheath, and its apical ring can be<br />
stained with Congo Red, as well as its small ascomata.<br />
Barr (1990a) introduced a second species J. shimekii<br />
which occurs on woody substrate. Some mycologists treat<br />
Javaria as a synonym of Astrosphaeriella (Hyde and<br />
Fröhlich 1998).<br />
Phylogenetic study<br />
None.<br />
Concluding remarks<br />
The size of ascomata and pigmentation of ascospores has<br />
little significance at generic level classification (Zhang et al.<br />
2009a). Likewise, the staining of endotunica with Congo<br />
Red has not been shown to have great significance. Thus,<br />
we accept Javaria as a synonym of Astrosphaeriella.<br />
Pycnidiophora Clum, Mycologia 47: 899 (1955).<br />
(Sporormiaceae)<br />
Current name: Westerdykella Stolk, Trans. Br. Mycol.<br />
Soc. 38(4): 422 (1955).<br />
Generic description<br />
Habitat terrestrial, saprobic (coprophilous). Ascomata<br />
small, cleistothecial, scattered on surface of agar media,<br />
semi-immersed, globose to subglobose, black. Peridium<br />
thin, composed of thin-walled, polyangular cells from front<br />
view. Hamathecium not apparent. Asci numerous, irregularly<br />
arranged, bitunicate nature undetermined, fissitunicate<br />
nature undetermined, globose, without pedicel. Ascospores<br />
gathering in the globose asci, smooth.<br />
Anamorphs reported for genus: Phoma-like.<br />
Literature: Cain1961; Clum1955; Stolk 1955b; Thompson<br />
and Backus 1966.<br />
Type species<br />
Pycnidiophora dispersa Clum, Mycologia 47: 900 (1955)<br />
[1955]. (Fig. 99)<br />
Current name: Westerdykella dispersa (Clum) Cejp &<br />
Milko.<br />
Ascomata 200–290 μm diam., cleistothecial, scattered<br />
on surface of agar media, semi-immersed, globose to<br />
subglobose, black (Fig. 99a). Peridium thin, composed of<br />
thin-walled, poly-angular cells from front view (Fig. 99b).<br />
Hamathecium not apparent. Asci numerous, 11–14 μm<br />
diam. (x ¼ 12:3mm, n=10), irregularly arranged, 32-spored<br />
when mature, bitunicate nature undetermined, fissitunicate<br />
nature undetermined, globose, without pedicel (Fig. 99b<br />
and c). Ascospores 4–5.5×2.5–3 μm (x ¼ 4:7 2:8mm, n=<br />
10), in the globose asci, olivaceous, oblong, 1-celled,<br />
smooth (Fig. 99d).<br />
Anamorph: Phoma-like coelomycetes.<br />
On MEA colonies spreading, flat with sparse aerial<br />
mycelium, covering the dish after 1 month; surface smokegrey<br />
with dirty white margins; reverse olivaceous-grey with<br />
luteous patches. On PDA spreading without aerial mycelium,<br />
colonies transparent, sporulating profusely with black, globose<br />
ascomata and pycnidia of a Phoma-like anamorph. On<br />
OA similar, lacking aerial mycelium, sporulating profusely<br />
with black, globose ascomata (based on <strong>CBS</strong> 297.56).<br />
Material examined: USA, Michigan, East Lansing,<br />
Science Greenhouse, isolated from damped off Phlox seedling,<br />
Dec. 1952, F.M. Clum (No. 27) (MSC 133.118, type).<br />
Notes<br />
Morphology<br />
Pycnidiophora was formally established by Clum (1955)<br />
based on its “imperfect stage of pycnidium”, which was<br />
subsequently confirmed as the sexual stage (Cain 1961;<br />
Thompson and Backus 1966). Clum (1955) has described<br />
and tentatively assigned P. dispersa (Clum) Cain to
Fungal Diversity<br />
Fig. 98 Javaria samuelsii (from isotype). a Ascoma on the host surface. Note reflexed pieces of the ruptured host tissue. b, c Cylindro-clavate<br />
asci within narrow pseudoparaphyses in gelatinous matrix. d Released ascospore with sheath. Scale bars: a=1 mm, b=50 μm, c, d=20 μm<br />
Aspergillaceae (= Eurotiaceae), and Stolk (1955b) has<br />
proposed to assign the morphologically comparable species<br />
P. multispora Saito & Minoura ex Cain to Eurotiaceae as<br />
well. Cain (1961), however, suspected that the 32 asco-
Fungal Diversity<br />
spores are actually the disarticulated segments of eight 4-<br />
celled ascospores, thus assigned it under Preussia (Sporormiaceae).<br />
After detailed study, Thompson and Backus<br />
(1966) confirmed that the so-called “eight 4-celled ascospores”<br />
do not exist in the development of the asci in both<br />
P. dispersa and P. multisporum. Thus, Pycnidiophora was<br />
assigned to Eurotiaceae (Eurotiales) (Thompson and<br />
Backus 1966).<br />
Phylogenetic study<br />
Phylogenetic study based on the ITS-nLSU rDNA<br />
sequences indicated that Pycnidiophora dispersa nested<br />
within clade of Westerdykella (including the generic type,<br />
W. ornata) (Kruys and Wedin 2009). Morphologically, both<br />
genera have cleistothecioid ascomata, asci with short or<br />
without pedicels and ascospores 1-celled and no germ slits.<br />
Thus, Pycnidiophora is treated as a synonym of Westerdykella<br />
(Kruys and Wedin 2009).<br />
Concluding remarks<br />
Although the pleosporalean status of Pycnidiophora is<br />
verified, morphological characters such as the cleistothecioid<br />
ascomata and irregularly arranged asci, which do not<br />
show typical bitunicate or fissitunicate characters, absence<br />
of pseudoparaphyses as well as the ascospores separating<br />
into partspores very early all challenge the traditional<br />
concept of <strong>Pleosporales</strong> (Zhang et al. 2009a). Obviously,<br />
most of these morphological characters overlap with those<br />
of the Eurotiales.<br />
Sporormiella Ellis & Everh., N. Amer. Pyren.: 136 (1892).<br />
(Sporormiaceae)<br />
Current name: Preussia Fuckel, Hedwigia 6: 175<br />
(1867) [1869–70].<br />
Generic description<br />
Habitat terrestrial, saprobic (coprophilous). Ascomata<br />
medium-sized, solitary, scattered, or in small groups,<br />
semi-immersed to nearly superficial, globose, subglobose,<br />
black, coriaceous, ostiolate, periphysate. Peridium thin,<br />
composed of small heavily pigmented cells of textura<br />
angularis, apex cells smaller and walls thicker. Hamathecium<br />
of dense, septate, cellular pseudoparaphyses, embedded<br />
in mucilage. Asci 8-spored, bitunicate, fissitunicate,<br />
cylindro-clavate, with a narrowed, furcate pedicel. Ascospores<br />
cylindrical with rounded ends, brown, 3-septate,<br />
deeply constricted at each septa, with sigmoid germ slit in<br />
each cell.<br />
Anamorphs reported for genus: none.<br />
Literature: Ahmed and Cain 1972; Ellis and Everhart<br />
1892; Khan and Cain 1979a, b; Luck-Allen and Cain 1975.<br />
Type species<br />
Sporormiella nigropurpurea Ellis & Everh., N. Amer.<br />
Pyren.: 136 (1892). (Fig. 100)<br />
Current name: Preussia nigropurpurea (Ellis & Everh.)<br />
Kruys, Syst. Biod. 7: 476.<br />
Ascomata 314–528 μm high×(250-)357–500 μm diam.,<br />
solitary, scattered, or in small groups, immersed, semiimmersed<br />
to nearly superficial, globose, subglobose, wall<br />
black, coriaceous, smooth, papillate, papilla 43–115 μm long,<br />
72–157 μm broad, ostiolate, ostiole filled with periphyses<br />
(Fig. 100a and b). Peridium 20–28 μm thick laterally, up to<br />
40 μm thick at the apex, composed of small heavily pigmented<br />
cells of textura angularis, cells 5–8 μm diam., cell wall<br />
1–3 μm thick, apex cells smaller and walls thicker<br />
(Fig. 100c). Hamathecium of dense, long, septate, cellular<br />
pseudoparaphyses, 1.5–2 μm broad, embedded in mucilage.<br />
Asci (70-)110–158×9–12.5(−15) μm (x ¼ 114:3 11:1mm, n=<br />
10), 8-spored, bitunicate, fissitunicate, cylindrical to cylindroclavate,<br />
with a narrowed, furcate pedicel, 13–38 μm long,<br />
ocular chamber apparent (Fig. 100d and e). Ascospores 15–<br />
20×4–5.5 μm (x ¼ 17:3 4:9mm, n=10), obliquely uniseriate<br />
and partially overlapping to biseriate, shortly cylindrical<br />
with rounded ends, brown, 3-septate, deeply constricted at<br />
each septum, with sigmoid germ slit in each cell, smoothwalled<br />
(Fig. 100f and g).<br />
Anamorph: none reported.<br />
Material examined: USA, New field, New Jersey:<br />
Gloucester Co., on cow dung, Mar. 1891 (NY, holotype).<br />
Notes<br />
Morphology<br />
Sporormiella was formally established by Ellis and<br />
Everhart (1892) based on the single species, Sporormiella<br />
nigropurpurea, which is characterized by its “immersed to<br />
semi-immersed, papillate ascomata, cylindrical to cylindroclavate<br />
asci with a pedicel, three to multi-septate ascospores<br />
with elongated germ slits through the whole cell” (Ahmed<br />
and Cain 1972; Khan and Cain 1979a, b). Barr (1990a) has<br />
indicated that Sporormiella might be a synonym of<br />
Ohleriella, while Sporormiella is assigned to Sporormiaceae<br />
as a separate genus (Eriksson 2006; Lumbsch and<br />
Huhndorf 2007). Currently, about 90 species are included<br />
in this genus (http://www.mycobank.org).<br />
Phylogenetic study<br />
The phylogenetic analysis based on ITS-nLSU rDNA,<br />
mtSSU rDNA and ß-tubulin sequences indicated that Sporormiella<br />
nested in Preussia,andaSporormiella–Preussia complex<br />
is formed (Kruys and Wedin 2009). Thus, Sporormiella was<br />
assigned under Preussia (Kruys and Wedin 2009).
Fungal Diversity<br />
Fig. 99 Pycnidiophora dispersa (a from <strong>CBS</strong> 297.56; b-d from MSC<br />
133.118, type). a Ascomata scattering on the surface of the substrate.<br />
b Crashed ascoma. Note the numerous released asci. c Globose asci<br />
Concluding remarks<br />
It is clear that the presence or absence of an ostiole cannot<br />
distinguish Sporormiella from Preussia according to the<br />
findings of Guarro et al. (1997a, b) and Kruys and Wedin<br />
(2009). Thus, Sporormiella should be treated as a synonym<br />
of Preussia (Kruys and Wedin 2009).<br />
Spororminula Arx & Aa, Trans. Br. Mycol. Soc. 89: 117<br />
(1987). (Sporormiaceae)<br />
Current name: Preussia Fuckel, Hedwigia 6: 175<br />
(1867) [1869–70].<br />
Generic description<br />
Habitat terrestrial, saprobic (coprophilous). Ascomata small<br />
to medium, solitary, scattered, immersed to erumpent,<br />
globose, subglobose, to ovate, black, membraneous, papillate,<br />
ostiolate. Peridium thin, membraneous, composed of<br />
and released ascospores. d One-celled ascospores. Scale bars: a=<br />
200 μm, b–d=20 μm<br />
several layers of heavily pigmented, elongate cells of<br />
textura angularis. Hamathecium of dense trabeculate,<br />
aseptate, decomposing pseudoparaphyses. Asci bitunicate,<br />
broadly cylindro-clavate with a narrow furcated pedicel.<br />
Ascospores cylindrical to cylindro-clavate, with round ends,<br />
brown, multi-septate, easily breaking into partspores.<br />
Anamorphs reported for genus: none.<br />
Literature: von Arx and van der Aa 1987.<br />
Type species<br />
Spororminula tenerifae Arx & Aa, Trans. Br. Mycol. Soc.<br />
89: 117 (1987).(Fig. 101)<br />
Current name: Preussia tenerifae (Arx & Aa) Kruys,<br />
Syst. Biod. 7: 476.<br />
Ascomata 290–400 μm diam., solitary, scattered, initially<br />
immersed, becoming erumpent when mature, globose,<br />
subglobose to ovate, black, membraneous, with a cylindrical<br />
or somewhat conical beak, 90–150(−230) μm broad and 110–
Fungal Diversity<br />
Fig. 100 Sporormiella nigropurpurea (from NY, holotype). a<br />
Section of an ascoma. b Section of the papilla. Note the dense<br />
pseudoparaphyses. c Section of a partial peridium. d, e Eight-spored<br />
cylindro-clavate asci with furcate pedicels. f, g Four-celled, brown<br />
ascospores. Note the sigmoid germ slit in each cell. Scale bars: a=<br />
200 μm, b, c=50 μm, d, e=20 μm, f, g=10 μm
Fungal Diversity<br />
190 μm high (Fig. 101a). Peridium 20–33 μm thick, 1-<br />
layered, composed of several layers of heavily pigmented,<br />
elongate cells of textura angularis, cells up to 6.3×5 μm<br />
diam., cell wall 1–1.5 μm thick (Fig. 101b and c).<br />
Hamathecium of dense, long trabeculate pseudoparaphyses<br />
1–2 μm broad, hyaline, aseptate, decomposing when mature.<br />
Asci 165–220×33–42.5 μm, 8-spored, bitunicate, broadly<br />
clavate, with a small, thin and furcate pedicel, 35–50 μm long,<br />
3–5 μm broad, ocular chamber not observed (Fig. 101d and<br />
e). Ascospores 68–93×12.5–16 μm, 3–4 seriate to uniseriate<br />
near the base, cylindrical to cylindro-clavate with rounded<br />
ends, brown, (6-)7 transverse septa, easily breaking into<br />
partspores, central cells triangular in transverse section but<br />
rectangular in vertical section, 14–16.5×8–10 μm long, apical<br />
cells 12.5–15×11.5–17.5 μm long (Fig. 101f and g).<br />
Anamorph: none reported.<br />
Material examined: SPAIN, Canary Islands, Tenerifa<br />
Las Canadas, on rabbit? droppings, Mar. 1986, J.A. von<br />
Arx (H<strong>CBS</strong> 9812, holotype).<br />
Notes<br />
Morphology<br />
Spororminula was formally established by von Arx and van<br />
der Aa (1987) according to its “ostiolate ascomata, elongated<br />
ascospore separated into part cells by transverse septa and<br />
without germ slits”, and was monotypified by S. tenerifae.<br />
Currently, only one species was included in this genus.<br />
Phylogenetic study<br />
Based on a phylogenetic analysis of ITS-nLSU rDNA, mtSSU<br />
rDNA and ß-tubulin sequences, Spororminula tenerifae<br />
nested in the clade of Preussia, thus Spororminula was<br />
treated as a synonym of Preussia (Kruys and Wedin 2009).<br />
Concluding remarks<br />
To clarify its relationship with other genera of Sporormiaceae,<br />
further phylogenetic study is needed, which should<br />
include additional related taxa.<br />
Excluded and doubtful genera<br />
Kriegeriella Höhn., Annls mycol. 16: 39 (1918). (Dothideomycetes,<br />
families incertae sedis, Microthyriaceae)<br />
Generic description<br />
Habitat terrestrial, saprobic? Ascomata small, solitary,<br />
scattered, superficial, subglobose, black, roughened, apex<br />
Fig. 101 Spororminula tenerifae (from H<strong>CBS</strong> 9812, holotype). a b<br />
Appearance of ascomata on the host surface. b, c Sections of the<br />
partial peridium. Note the elongate cells of textura angularis. d, e<br />
Asci with thin pedicels. f, g Ascospores, which may break into part<br />
spores. Scale bars: a=0.5 mm, b=100 μm, c=50 μm, d–g=20 μm<br />
no obvious opening. Peridium thin, composed of a single<br />
type of lightly pigmented thin-walled cells. Hamathecium<br />
long cellular pseudoparaphyses, septate. Asci 8-spored,<br />
bitunicate, obpyriform. Ascospores hyaline, turning brown<br />
when mature, multi-septate, constricted at each septum.<br />
Anamorphs reported for genus: none.<br />
Literature: von Arx and Müller 1975; Barr1975, 1987b;<br />
Eriksson 2006; Lumbsch and Huhndorf 2007.<br />
Type species<br />
Kriegeriella mirabilis Höhn., Annls mycol. 16: 39 (1918)<br />
(Fig. 102)<br />
Ascomata 100–120 μm high×150–220 μm diam., solitary,<br />
scattered, superficial, with basal wall flattened on the surface of<br />
the substrate, subglobose, black, roughened, apex no obvious<br />
opening (Fig. 102a and e). Peridium thin, composed of a<br />
single type of lightly pigmented thin-walled cells, cells up to<br />
12×5 μm diam. in front view, cell wall less than 1 μm thick,<br />
apex cells smaller and walls thicker (Fig. 102a and f).<br />
Hamathecium long cellular pseudoparaphyses, 1.5–2 μm<br />
wide, septate. Asci 65–85×31–36 μm (x ¼ 63:1 33mm, n=<br />
10), 8-spored, bitunicate, fissitunicate undetermined, obpyriform,<br />
no pedicel, no ocular chamber was seen (Fig. 102c and<br />
g). Ascospores 28–37.5×8–11 μm (x ¼ 32:6 10mm, n=<br />
10), 8-seriate, fusoid with broadly rounded ends, hyaline,<br />
turning brown when mature, 5–6-septate, constricted at each<br />
septum, the upper first-cell longer and broader than the lower<br />
ones with semi-round shape, smooth (Fig. 102d, h, i).<br />
Anamorph: see Fig. b.<br />
Material examined: On the leaves of Faulenden nadeln<br />
von Pinus silvestris, bei Roñigstein, Sept. 1896, W. Rueges.<br />
(S reg. nr F12638, isolectotype).<br />
Notes<br />
Morphology<br />
Kriegeriella was formally established by von Höhnel (1918b)<br />
and was represented by two species, i.e. K. mirabilis and K.<br />
transiens; it was typified by K. mirabilis and assigned to<br />
Microthyriaceae. Subsequently, Kriegeriella was assigned to<br />
the subfamily of Aulographiodeae (Microthyriaceae)<br />
(Batista et al. 1959), Asterinaceae (Hemisphaeriales) (Luttrell<br />
1973) and Pseudosphaeriaceae (Dothideales) (Barr<br />
1975).<br />
After checking the original description and the type<br />
specimens of K. mirabilis and K. transiens, no significant
Fungal Diversity
Fungal Diversity<br />
difference could be observed, and both are described from<br />
rotting needles of conifers (Barr 1975; Batista et al. 1959;<br />
Höhnel 1918b). Morphologically, Extrawettsteinina is<br />
comparable with Kriegeriella. In particularly, E. pinastri<br />
could not be distinguished from K. transiens or K.<br />
mirabilis. Thus, K. transiens including Extrawettsteinina<br />
pinastri was treated as synonyms of K. mirabilis, andwas<br />
included in the section of Kriegeriella under the genus<br />
Kriegeriella (von Arx and Müller 1975; Barr1975). The<br />
other section of Kriegeriella, Extrawettsteinina, includes<br />
two previous Extrawettsteinina species, i.e. K. minuta and<br />
K. mediterranea. Barr(1987b) introduced a family, i.e.<br />
Kriegeriellaceae (Dothideales) to accommodate Kriegeriella<br />
and Extrawettsteinina. This proposal is rarely<br />
followed, and Kriegeriella is usually assigned to Pleosporaceae<br />
(<strong>Pleosporales</strong>) (Eriksson2006; Lumbsch and Huhndorf<br />
2007).<br />
Phylogenetic study<br />
None.<br />
Concluding remarks<br />
Kriegeriella might belong to Microthyriaceae, although it<br />
would be unusual in this family in having 5-6-septate<br />
ascospores. Micropeltidaceae better accommodates the<br />
ascospores, however, the parallel arrangement of cells of<br />
the upper peridium are not typical. Asterinaceae may be<br />
most suitable as Luttrell (1973) suggested.<br />
Phaeotrichum Cain & M.E. Barr, Can. J. Bot. 34: 676<br />
(1956). (Dothideomycetes, family incertae sedis,<br />
Phaeotrichaceae)<br />
Generic description<br />
Habitat terrestrial, saprobic (coprophilous). Ascomata<br />
small, cleistothecial, solitary, or in small groups,<br />
superficial, with long straight or slightly flexed, thin,<br />
black appendages evenly scattered on the surface of the<br />
ascomata, globose, black. Peridium thin, carbonaceousmembraneous,<br />
1-layered, composed of dark brown thickwalled<br />
cells of textura angularis. Hamathecium not<br />
observed. Asci bitunicate form not clear, fissitunicate<br />
dehiscence not observed, broadly clavate, with a relatively<br />
thick pedicel. Ascospores oblong to ellipsoid,<br />
hyaline when young, turning reddish brown at maturity,<br />
1-septate, deeply constricted at the septum, each end with<br />
a subhyaline and broadly rounded germ pore, readily<br />
forming partspores at the septum at maturity.<br />
Anamorphs reported for genus: none.<br />
Literature: Cain 1956; Malloch and Cain 1972.<br />
Type species<br />
Phaeotrichum hystricinum Cain & M.E. Barr, Can. J. Bot.<br />
34: 677 (1956). (Fig. 103)<br />
(Some information for the following description is from<br />
Cain 1956)<br />
Ascomata 170–280 μm diam., cleistothecial, solitary, or<br />
in small groups, superficial, with 15–20 long straight or<br />
slightly flexed, thin, black appendages evenly scattered on<br />
the surface of the ascomata, 0.5–1 mm long, 15–25 μm<br />
wide at base, tapering to less than 5 μm at the blunt apex,<br />
with few or without septa, globose, black, smooth<br />
(Fig. 103a). Peridium thin, carbonaceous-membraneous,<br />
1-layered, composed of dark brown thick-walled cells<br />
of textura angularis, cells 8–16 μm diam., cell wall<br />
0.5–1.5 μm thick (data obtained from Cain 1956)<br />
(Fig. 103b). Hamathecium not observed. Asci 42–48×<br />
14–17 μm, 8-spored, bitunicate form not typical, lacking<br />
fissitunicate dehiscence, broadly clavate, with a relatively<br />
thick pedicel which is about 18 μm (data obtained from<br />
Cain 1956). Ascospores 14–16×4–5 μm, 4-seriate, oblong<br />
to ellipsoid, hyaline when young, turning reddish<br />
brown at maturity, 1-septate, deeply constricted at the<br />
septum, each end with a subhyaline and broadly rounded<br />
germ pore, smooth, readily separating into partspores at<br />
the septum at maturity (Fig. 103c, d, e and f).<br />
Anamorph: none reported.<br />
Material examined: CANADA, Ontario, Muskoka,<br />
Stoneleigh, on porcupine dung, 18 Aug. 1932, Cain (TRTC<br />
4361, holotype).<br />
Note: the ascomata of the specimen are fragile and no<br />
asci could be obtained.<br />
Notes<br />
Morphology<br />
Phaeotrichum was formally established by Cain (1956)<br />
to accommodate two new coprophilous fungi, i.e. P.<br />
hystricinum and P. circinatum Cain, and P. hystricinum<br />
was selected as the generic type. Phaeotrichum is<br />
mainly characterized by its coprophilous habitat, superficial<br />
cleistothecial ascocarps covered by long hairy<br />
appendages, reddish brown 1-septate ascospore with a<br />
broadly rounded germ pore at each end, readily breaking<br />
into partspores (Cain 1956). According to Cain (1956),<br />
Phaeotrichum possesses untypical bitunicate ascus, and<br />
the ascospore releasing is described as “simply break<br />
down and allow the contents to become free in the<br />
cavity of the ascocarp”. This ascospore releasing<br />
mechanism is considered as evolutionarily developed<br />
compared to those that “discharge the ascospores<br />
through an apical pore” (Cain 1956). Although lacking
Fungal Diversity<br />
Fig. 102 Kriegeriella mirabilis (from S reg. nr F12638, isolectotype).<br />
a Section of a superficial ascoma. b Anamorphic stage. c<br />
Obpyriform ascus. Note the pigmented ascospores and hyaline<br />
ascospores coexisted in a single ascus. d Ascospores. Scale bars: a=<br />
50 μm, b–d=10 μm. e Ascomata on the host surface. f, g Crashed<br />
ascoma. Note the peridium structure. h, i Hyaline asymmetric<br />
ascospores. Scale bars: e, f =100 μm, g=50 μm, h, i=10 μm<br />
a typical bitunicate ascus, Phaeotrichum is still<br />
assigned to <strong>Pleosporales</strong>, because the lack of bitunicate<br />
ascus does not “taken by itself, exclude a fungus from<br />
close relationship” (Cain 1956).
Fig. 102 (continued)<br />
Fungal Diversity
Fungal Diversity<br />
Phylogenetic study<br />
A single unverified isolate of Phaeotrichum benjaminii is<br />
placed well outside of <strong>Pleosporales</strong> in a broad phylogenetic<br />
study (Schoch et al. 2009).<br />
Concluding remarks<br />
The superficial cleistothecial ascocarps covered by long hairy<br />
appendages, the absence of hamathecium as well as the<br />
nontypical bitunicate ascus are all distinct from members of<br />
Fig. 103 Phaeotrichum hystricinum<br />
(from TRTC 4361, holotype).<br />
a Superficial ascomata on<br />
host surface. Note the long and<br />
black appendages. b Part of<br />
peridium. Note the large cells in<br />
surface view. c–f Released reddish<br />
brown ascospores with hyaline<br />
end cells. Note the strongly<br />
constricted middle septum.<br />
Scale bars: a=0.5 mm,<br />
b–f=20 μm
Fungal Diversity<br />
<strong>Pleosporales</strong>, but definite conclusions could only be obtained<br />
by further molecular phylogenetic analysis. In this study, we<br />
assign it to Dothideomycetes incertae cedis.<br />
Zeuctomorpha Sivan., P.M. Kirk & Govindu, Bitunicate<br />
Ascomycetes and their Anamorphs: 572 (1984). (Venturiaceae)<br />
Generic description<br />
Habitat terrestrial, hemibiotrophic. Ascomata small, gregarious,<br />
superficial, globose to slightly flattened, ostiolate,<br />
covered with setae. Peridium thin, composed of heavily<br />
pigmented pseudoparenchymatous cells of textura angularis.<br />
Hamathecium of rare, septate, branching and anastomosing<br />
pseudoparaphyses. Asci 8-spored, with a short thick pedicel,<br />
bitunicate, fissitunicate, broadly clavate to obclavate. Ascospores<br />
ellipsoid, dark brown, 1-septate, asymmetrical, deeply<br />
constricted at the septum.<br />
Anamorphs reported for genus: Acroconidiellina (Sivanesan<br />
1984).<br />
Literature: Sivanesan 1984.<br />
Type species<br />
Zeuctomorpha arecae Sivan., P.M. Kirk & Govindu, in<br />
Sivanesan, Bitunicate Ascomycetes and their Anamorphs:<br />
572 (1984). (Fig. 104)<br />
Ascomata 175–300 μm diam., gregarious, superficial,<br />
globose to slightly flattened, collapsed at the apex when<br />
dry, ostiolate, covered with numerous long setae<br />
(Fig. 104a). Peridium up to 25 μm wide, composed of<br />
heavily pigmented pseudoparenchymatous cells of textura<br />
angularis, to7μm diam. Hamathecium of rare, 2–5 μm<br />
broad, septate, branching and anastomosing pseudoparaphyses.<br />
Asci 83–185×29–40(−50) μm (x ¼ 134 35:3mm, n=<br />
10), 8-spored, bitunicate, fissitunicate, broadly clavate to<br />
obclavate, with a short thick pedicel, up to 40 μm long,<br />
apically rounded, with a small ocular chamber (to 4 μm<br />
wide×7 μm high) (Fig. 104b, c and d). Ascospores 35–43×<br />
12.5–18 μm (x ¼ 36:5 15:4mm, n=10), 2–4 seriate, ellipsoid,<br />
dark brown, 1-septate, deeply constricted at the septum,<br />
usually slightly asymmetric, smooth (Fig. 104e,f,g,handi).<br />
Anamorph: Acroconidiellina arecae (Sivanesan 1984).<br />
Material examined: INDIA, Shimogee, on Areca<br />
catechu L. leaf, 1 Nov. 1979, H.C. Govindu (IMI 246067,<br />
holotype).<br />
Notes<br />
Morphology<br />
Zeuctomorpha was formally established by Sivanesan<br />
(1984) based on its superficial setose ascomata, clavate<br />
asci, ellipsoid and 1-septate ascospores, and presence of<br />
pseudoparaphyses, and was monotypified by Z. arecae.<br />
Zeuctomorpha arecae is widely distributed in tropical<br />
regions of East South Asia exclusively on the leaves of<br />
Areca catechu (Sivanesan 1984).<br />
Phylogenetic study<br />
None.<br />
Concluding remarks<br />
This taxon is unusual amongst the Pleosporaceae as it<br />
has hairy superficial ascomata, few pseudoparaphyses,<br />
broadly clavate to obclavate asci and 1-septate pigmented<br />
ascospores. All of these morphological characters are<br />
most comparable with species of Acantharia, which<br />
might be closely related to Venturiaceae (Zhang et al.<br />
data unpublished).<br />
Muroia I. Hino & Katum., J. Jap. Bot. 33: 79 (1958).<br />
(Ascomycota)<br />
Generic description<br />
Habitat terrestrial, saprobic or parasitic. Ascostromata erumpent<br />
through the host surface in linear rows parallel to the<br />
host fibers. Ascomata small- to medium-sized, semiimmersed<br />
to erumpent, subglobose to rectangular, black,<br />
coriaceous, cells of ascostromata pseudoparenchymatous,<br />
cells of peridium composed of pigmented cells of textura<br />
angularis. Hamathecium of rare, pseudoparaphyses. Asci<br />
bitunicate, clavate to cylindro-clavate. Ascospores oblong to<br />
elongated oblong, hyaline, 1-celled, usually slightly curved.<br />
Anamorphs reported for genus: none.<br />
Literature: Hino and Katumoto 1958.<br />
Type species<br />
Muroia nipponica I. Hino & Katum., J. Jap. Bot. 33: 79<br />
(1958). (Fig. 105)<br />
Ascostroma 1–6 mm long, 360–470 μm broad, linear<br />
parallel to the host fibers with several linearly arranged<br />
ascomata (Fig. 105a). Ascomata 250–400 μm diam.,<br />
semi-immersed in substrate to erumpent, subglobose to<br />
rectangular with a furrow-shaped ostiole, black, coriaceous,<br />
cells of ascostromata pseudoparenchymatous.<br />
Peridium composed of pigmented cells of textura<br />
angularis. Hamathecium of rare, 3–4.5 μm broad<br />
pseudoparaphyses. Asci (120-)150–190×30–45 μm, 8-<br />
spored, bitunicate, fissitunicate dehiscence not observed,<br />
clavate to cylindro-clavate, with a short, thin, knob-like<br />
pedicel, lacking an ocular chamber (Fig. 105b). Ascospores<br />
43–50×13–18 μm (x ¼ 46:6 15:2mm, n=10), biseriate, ob-
Fungal Diversity<br />
Fig. 104 Zeuctomorpha arecae (from IMI 246067, holotype). a<br />
Gregarious ascomata on host surface. Note the numerous setae on the<br />
surface of ascomata. b Asci with ocular chamber and short peduncles.<br />
c, d Ascus with ocular chamber and knob-like pedicel. e–i One septate<br />
ascospores which are slightly asymmetrical. Scale bars: a=0.5 mm,<br />
b–i=20 μm
Fungal Diversity<br />
long to elongated oblong, hyaline, 1-celled, usually slightly<br />
curved (Fig. 105c,d and e).<br />
Anamorph: none reported.<br />
Material examined: JAPAN, Province Ugo. on moribund<br />
culm of Sasa kurilensis, 4 Aug. 1957, coll. H. Muroi,<br />
Det. I. Hino & K. Katumoto (TNS-F-230252, isotype).<br />
Notes<br />
Morphology<br />
Muroia was introduced based on M. nipponica, which is a<br />
parasite on the lower part of Sasa kurilensis (Hino and<br />
Katumoto 1958). Muroia is characterized by its 1-celled<br />
ascospores. Considering the perithecial structure and linear<br />
ostiole, it was assigned to the Lophiostomataceae, and was<br />
regarded as closely related to the amerosporous Lophiella<br />
(Hino and Katumoto 1958).<br />
Phylogenetic study<br />
None.<br />
Concluding remarks<br />
The linear ascostroma and 1-celled, hyaline ascospores<br />
make it less likely to fit the concept of Lophiostomataceae.<br />
Because of the condition of the specimen, its bitunicate<br />
nature could not be confirmed.<br />
Genera not studied<br />
Aglaospora De Not., G. bot. ital. 2: 43 (1844).<br />
Type species: Aglaospora profusa (Fr.) De Not., G. bot.<br />
ital. 2: 43 (1844).<br />
Aglaospora, which was introduced by de Notaris<br />
(1844), has 35 species epithets (http://www.mycobank.<br />
org/mycotaxo.aspx) and was considered to be a synonym<br />
of Massaria (Voglmayr and Jaklitsch 2011) orseparate<br />
(Barr 1990a). In a recent phylogenetic study, Voglmayr and<br />
Jaklitsch (2011) confirmed that Aglaospora is a synonym of<br />
Massaria and is treated as such here. The immersed<br />
ascomata with short beaks, together with ascostroma under<br />
pseudostromatic tissues, cylindrical asci with a large and<br />
refractive apical ring, trabeculate pseudoparaphyses within a<br />
gel matrix, and distoseptate ascospores, are all similar to<br />
species of Massaria. The large and conspicuous apical ring<br />
of the ascus of Aglaospora has the appearance of being<br />
unitunicate, and thus Shoemaker and Kokko (1977) treatedit<br />
as a unitunicate taxon. Currently, its bitunicate status is<br />
widely accepted.<br />
Allewia E.G. Simmons, Mycotaxon 38: 260 (1990).<br />
Type species: Allewia proteae E.G. Simmons, Mycotaxon<br />
38: 262 (1990).<br />
Allewia was introduced by Simmons (1990) toaccommodate<br />
Lewia-like species but with Embellisia anamorphs.<br />
Embellisia differs from other similar genera by a combination<br />
of characters including the percentage of dictyoconidia,<br />
shape of conidia, thickness of septa, umbilicate sites of<br />
conidiophore geniculation, proliferating chlamydospores and<br />
hyphal coils in culture (Simmons 1971). Based on multigene<br />
phylogenetic analysis, A. eureka, which is closely related to<br />
A. proteae, clustered together with species of Alternaria.<br />
Thus, Allewia should be treated as a synonym of Lewia.<br />
Anteaglonium Mugambi & Huhndorf, System. Biodivers.<br />
7: 460 (2009).<br />
Type species: Anteaglonium abbreviatum (Schwein.)<br />
Mugambi & Huhndorf, System. Biodivers. 7: 460 (2009).<br />
≡ Hysterium abbreviatum Schwein., Trans. Am. phil.<br />
Soc., New Series 4: no. 2094 (1832).<br />
Anteaglonium was introduced to accommodate a monophyletic<br />
hysterothecial clade within <strong>Pleosporales</strong>, and four<br />
species (A. abbreviatum, A. globosum Mugambi & Huhndorf,<br />
A. parvulum (W.R. Gerard) Mugambi & Huhndorf<br />
and A. latirostrum Mugambi & Huhndorf) are included<br />
(Mugambi and Huhndorf 2009a). Anteaglonium is characterized<br />
by erumpent to superficial, globose to subglobose or<br />
elongate, fusoid to oblong ascomata, which are brown to<br />
shiny black, opening by a pronounced or indistinct<br />
longitudinal slit running entire length of fruit body or apex<br />
raised and laterally compressed; asci cylindrical with short<br />
pedicel, 8-spored, uniseriate or biseriate; ascospores fusoid<br />
to oblong, septate, constricted at the primary septum,<br />
hyaline or pigmented. A phylogenetic analysis based on<br />
DNA comparisons indicated that Anteaglonium resides as a<br />
separate clade but related to Tetraplosphaeria, Lophiotrema<br />
and other species without clear resolution. Therefore, the<br />
familial placement of Anteaglonium remains unclear<br />
(Mugambi and Huhndorf 2009a).<br />
Arthopyrenia A. Massal., Ric. auton. lich. crost. (Verona):<br />
165 (1852).<br />
Type species: Arthopyrenia rhyponta (Ach.) A. Massal.,<br />
Ric. auton. lich. crost. (Verona): 166, fig. 329 (1852).<br />
≡ Verrucaria rhyponta Ach., K. Vetensk-Acad. Nya<br />
Handl.: 150 (1809).<br />
Arthopyrenia is a lichen genus with a Trentepohlia<br />
photobiont and is characterized by dimidiate perithecoid<br />
ascomata, which are scattered to irregularly confluent, and<br />
have an upper thick clypeate wall composed of periderm<br />
cells intermixed with dark hyphae. The pseudoparaphyses are<br />
branched and asci are obpyriform, obclavate to subcylindrical<br />
and 8-spored. Ascospores are oblong, ovoid, slipper-shaped,<br />
1-3-septate, hyaline and smooth-walled (Coppins 1988;
Fungal Diversity<br />
Fig. 105 Muroia nipponica (TNS-F-230252, isotype). a Linear ascostroma parallel to the host fibers. b Crashed ascus with ascospores released.<br />
c–e Released hyaline ascospores. Scale bars: a=5 mm, b–e=30 μm<br />
Upreti and Pant 1993). Multigene phylogenetic studies<br />
indicated that Arthopyrenia salicis, a typical species of<br />
Arthopyrenia, is located within <strong>Pleosporales</strong> in close<br />
proximity to bambusicolous species in the genus Roussoella,<br />
with its familial status remaining undetermined (Del Prado et<br />
al. 2006; Schochetal.2009; Zhang et al. 2009a).<br />
Ascocratera Kohlm., Can. J. Bot. 64: 3036 (1986).<br />
Type species: Ascocratera manglicola Kohlm., Can. J. Bot.<br />
64(12): 3036 (1986).<br />
Ascocratera is a monotypic obligate marine fungus and<br />
is characterized by conical, crater-like, erumpent to<br />
superficial and carbonaceous ascomata, a depressed<br />
ostiole, a thick peridium, trabeculate pseudoparaphyses,<br />
bitunicate, fissitunicate and cylindrical asci, and ellipsoidal,<br />
hyaline, 1-septate (3-septate when senescent) ascospores<br />
surrounded by a sheath (Kohlmeyer 1986).<br />
Ascocratera was reported to be one of the most common<br />
marine fungi of the upper intertidal zone of dead<br />
mangrove roots, trunks and branches (Kohlmeyer 1986).<br />
Based on a multigene phylogenetic analysis, Ascocratera<br />
nested within the clade of Aigialaceae (Schoch et al.<br />
2009; Suetrong et al. 2009).<br />
Atradidymella M.L. Davey & Currah, Am. J. Bot. 96: 1283<br />
(2009).<br />
Type species: Atradidymella muscivora M.L. Davey &<br />
Currah, Am. J. Bot. 96: 1283 (2009).<br />
Atradidymella was introduced as a pleosporalean genus<br />
parasitic on boreal bryophytes, and is characterized by
Fungal Diversity<br />
minute, unilocular, setose pseudothecia with 2–3 wall<br />
layers; brown, fusoid, 1-septate ascospores, and an anamorphic<br />
stage (Phoma muscivora M.L. Davey & Currah)<br />
(Davey and Currah 2009). Based on an ITS rDNA<br />
sequences analysis, Atradidymella nested within Didymellaceae<br />
(Davey and Currah 2009).<br />
Bertiella (Sacc.) Sacc. & P. Syd., in Saccardo, Syll. fung.<br />
(Abellini) 14: 19 (1899).<br />
≡ Bertia subgen. Bertiella Sacc., Syll. fung. (Abellini) 1:<br />
584 (1882).<br />
Type species: Bertiella macrospora (Sacc.) Sacc. &<br />
Traverso, Syll. fung. (Abellini) 19: 147 (1910).<br />
≡ Bertia macrospora Sacc., Michelia 1(no. 8): 452 (1882).<br />
Bertia subg. Bertiella was raised to generic rank by Saccardo<br />
(1899), and is typified by B. macrospora. After studying the<br />
type specimen of B. macrospora, Eriksson and Yue (1986)<br />
assigned it to Massarina (as M. macrospora (Sacc.) O.E. Erikss.<br />
& J.Z. Yue). Concurrently, Bertiella is treated as a synonym of<br />
Massarina. Hydeetal.(2002) assigned Bertia macrospora to<br />
Lophiostoma as (L. bertiellum Aptroot & K.D. Hyde).<br />
The superficial ascomata, cylindro-clavate asci and<br />
hyaline 1-septate ascospores which may become 3-septate<br />
and pale brown when senescent and, in particular, the<br />
woody habitat indicate that B. macrospora may be related<br />
to Lophiostoma sensu Holm and Holm (1988). A single<br />
isolate of Bertiella macrospora clusters with Byssosphaeria<br />
in the Melanommataceae in a recent DNA based phylogeny<br />
(Mugambi and Huhndorf 2009b). The relationship between<br />
Bertiella and Byssosphaeria needs further study.<br />
Byssothecium Fuckel, Bot. Ztg. 19: 251 (1861).<br />
Type species: Byssothecium circinans Fuckel, Bot. Ztg.<br />
19: 251 (1861).<br />
The isotype of Byssothecium circinans is in FH as<br />
exiccatae (Fungi rhenani 730c); it was described by Boise<br />
(1983) and could not be loaned. Byssothecium circinans is<br />
regarded as a saprobe or weak parasite of Medicago sativa<br />
(Semeniuk 1983), and a Pleospora-type centrum was<br />
observed (Boise 1983). A Chaetophoma-like anamorph<br />
was produced in culture, however, no culture or herbarium<br />
specimen is listed (Boise 1983). Boise (1983) regarded<br />
Byssothecium circinans as closely related to Teichospora,<br />
however, confirmation is required. An isolate of Byssothecium<br />
circinans was sequenced and a multigene phylogeny<br />
placed it in close proximity to members of Massarinaceae<br />
(Schoch et al. 2009; Zhang et al. 2009a; Plate 1).<br />
Caryospora De Not., Micr. Ital. Novi 9: 7 (1855).<br />
Type species: Caryospora putaminum (Schwein.) De Not.,<br />
Micr. Ital., Dec. 9: 7 (1855).<br />
After studying the Caryospora species in North America,<br />
Barr (1979b) indicated that species of Caryospora may<br />
closely relate to Trematosphaeria. Boise (1985) distinguished<br />
Caryospora from Trematosphaeria based on the<br />
structure of ascospores. Currently, 17 taxa, from freshwater,<br />
marine, or terrestrial habitats (Raja and Shearer 2008), are<br />
included within Caryospora and might be polyphyletic.<br />
Celtidia J.D. Janse, Ann. Jard. Bot. Buitenzorg 14: 202 (1897).<br />
Type species: Celtidia duplicispora J.D. Janse, Ann. Jard.<br />
Bot. Buitenzorg 14: 202 (1897).<br />
Celtidia is a monotypic genus, which is characterized by<br />
its echinulate ascospores (Hawksworth 1979). It is only<br />
known from an illustration accompanying the original<br />
description from root nodules of Celtis in Java. A new<br />
collection is needed for further study of this genus.<br />
Chaetopreussia Locq.-Lin., Revue Mycol., Paris 41: 185<br />
(1977).<br />
Type species: Chaetopreussia chadefaudii Locq.-Lin.,<br />
Revue Mycol., Paris 41: 187 (1977).<br />
Chaetopreussia is a monotypic genus characterized by<br />
cleistothecioid ascomata with seta, and 3-septate ascospores<br />
without germ slits. Recent molecular analysis has shown that<br />
cleistothecioid ascomata and the presence of germ slits lack<br />
significance at the generic rank (Kruys and Wedin 2009).<br />
Chaetopreussia is possibly another synonym of Preussia.<br />
Clathrospora Rabenh., Hedwigia 1(18): 116 (1857).<br />
Type species: Clathrospora elynae Rabenh., Hedwigia 1:<br />
116 (1857).<br />
The most striking character of Clathrospora is its<br />
ascomata opening with an intraepidermal discoid lid and<br />
muriform applanate ascospores with more than one row of<br />
longitudinal septa (Shoemaker and Babcock 1992). The<br />
form of opening and applanate ascospores, however, might<br />
have limited significance at generic rank and thus,<br />
Clathrospora may be closely related to Pleosporaceae.<br />
Phylogenetic analysis based on nLSU, nSSU and mtSSU<br />
indicate that C. diplospora (Ellis & Everh.) Sacc. &<br />
Traverso nests in Pleosporaceae (Kruys et al. 2006).<br />
Clathrospora elynae is saprobic on monocots (Shoemaker<br />
and Babcock 1992).<br />
Cochliobolus Drechsler, Phytopathology 24: 973 (1934).<br />
Type species: Cochliobolus heterostrophus (Drechsler)<br />
Drechsler, Phytopathology 24: 973 (1934).<br />
Cochliobolus and its asexual relatives are well studied<br />
taxa in <strong>Pleosporales</strong> because of their economic importance.<br />
Cochliobolus includes both saprobic and pathogenic species<br />
that are significant monocot pathogens worldwide, which<br />
attack corn, rice, barley, sugarcane, wheat, and oats, all<br />
major cereal crops. Cochliobolus is characterized by globose<br />
or subglobose ascomata with a well defined long ostiolar<br />
papilla or cylindrical neck, a peridium composed of
Fungal Diversity<br />
pseudoparenchymatous cells, filliform, septate and branched<br />
pseudoparaphyses, and thin-walled cylindrical or broadly<br />
clavate asci. Ascospores are distinctively hyaline or pale<br />
brown, filliform, and strongly helicoid to loosely coiled in<br />
the asci (Sivanesan 1984). The anamorphs of Cochliobolus<br />
belong to Bipolaris and Curvularia (Sivanesan 1984).<br />
Bipolaris and Curvularia can be distinguished by characters<br />
of conidial morphology, conidial germination, hilum structure,<br />
conidial septum and wall structure, conidial septum<br />
ontogeny (Sivanesan 1987). Multigene phylogenetic analysis<br />
indicated that Cochliobolus heterostrophus and C. sativus (S.<br />
Ito & Kurib.) Drechsler ex Dastur nested within the clade of<br />
Pleosporaceae (Zhang et al. 2009a; Plate 1). Thus, its<br />
familial placement is confirmed.<br />
Comoclathris Clem., Gen. fung. (Minneapolis): 37, 173<br />
(1909).<br />
Type species: Comoclathris lanata Clem. [as ‘Comochlatris’],<br />
Gen. fung. (Minneapolis) (1909).<br />
Comoclathris is temporarily placed in Diademaceae, and<br />
its pivotal characters are the circular lid-like opening and<br />
applanate reddish-brown to dark reddish-brown muriform<br />
ascospores with single longitudinal septa (versus two or<br />
more rows of longitudinal septa of Clathrospora) (Shoemaker<br />
and Babcock 1992). Barr (1990b) treated it as a synonym of<br />
Graphyllium. Comoclathris has been linked with an Alternaria-like<br />
anamorphs (Simmons 1952), which may suggest<br />
its close relationship with Pleosporaceae.<br />
Coronopapilla Kohlm. & Volkm.-Kohlm., Mycol. Res. 94:<br />
686 (1990).<br />
Type species: Coronopapilla avellina Kohlm. & Volkm.-<br />
Kohlm., Mycol. Res. 94: 687 (1990).<br />
Coronopapilla is characterized by immersed ascomata<br />
with a conical papilla, thin peridium, 8-spored and thickwalled,<br />
cylindrical and fissitunicate asci. Ascospores are<br />
ellipsoidal, 1-3-septate, brown and distoseptate. Coronopapilla<br />
avellina is an obligate marine species, and was originally<br />
assigned to Didymosphaeriaceae (Kohlmeyer and Volkmann-<br />
Kohlmeyer 1990). The marine habitat of Coronopapilla<br />
makes it readily distinguishable from Didymosphaeria futilis<br />
(the generic type of Didymosphaeria). Thus, the familial<br />
placement of Coronopapilla is yet to be determined.<br />
Cucurbitaria Gray, Nat. Arr. Brit. Pl. (London) 1: 508, 519<br />
(1821).<br />
Type species: Cucurbitaria berberidis (Pers.) Gray, Nat.<br />
Arr. Brit. Pl. (London) 1: 508, 519 (1821).<br />
≡ Sphaeria berberidis Pers., Neues Mag. Bot. 1: 83<br />
(1794).<br />
A narrow generic concept of Cucurbitaria was accepted<br />
by Welch (1926), who restricted Cucurbitaria to five<br />
closely related species, which have turbinate ascomata that<br />
develop cespitosely in a massive subiculum or over<br />
compressed stromatic tissues and have a thick and obconoid<br />
base. A broader generic concept was accepted by Mirza<br />
(1968), who also included species with globose or ovoid to<br />
pyriform ascomata that are gregarious on the substrate with<br />
only sparse subiculum and lack an obconoid region in the<br />
base of the locule. Barr (1990b) accepted an intermediate<br />
concept, and described 11 related species from North<br />
America. Currently, 450 species are accepted in Cucurbitaria<br />
(http://www.mycobank.org/mycotaxo.aspx), and the<br />
genus was assigned to Cucurbitariaceae. In this study, an<br />
isolate of C. berberidis clustered with some species of<br />
Pyrenochaeta and Didymosphaeria futilis, and they get<br />
moderate bootstrap support (Plate 1). Cucurbitariaceae<br />
may be another family within Pleosporineae.<br />
Curreya Sacc., Syll. fung. (Abellini) 2: 651 (1883).<br />
Type species: Curreya conorum (Fuckel) Sacc., Syll. fung.<br />
(Abellini) 2: 651 (1883).<br />
Curreya is a contentious genus which had been assigned to<br />
Pleospora (Barr 1981). von Arx and van der Aa (1983),<br />
however, maintained it as distinct, because of its Coniothyrium<br />
anamorph, and considered Curreya should be closely<br />
related to Didymosphaeria, Melanomma, Paraphaeosphaeria<br />
or Massarina. Because of the small sclerotial cells of its<br />
peridium, the narrower, thinner-walled asci and its Coniothyrium-like<br />
anamorph, Barr (1990b) assigned it to the<br />
Leptosphaeriaceae. Previous phylogenetic studies indicated<br />
that a strain of Curreya pityophila (J.C. Schmidt & Kunze)<br />
Petr. nested within Massarineae (Kruys et al. 2006).<br />
Decorospora Inderb., Kohlm. & Volkm.-Kohlm., Mycologia<br />
94: 657 (2002).<br />
Type species: Decorospora gaudefroyi (Pat.) Inderb.,<br />
Kohlm. & Volkm.-Kohlm., Mycologia 94: 657 (2002).<br />
≡ Pleospora gaudefroyi Pat., Tabl. analyt. Fung. France<br />
(Paris) 10: 40 (no. 602) (1886).<br />
Decorospora gaudefroyi (as Pleospora gaudefroyi) had<br />
been considered a synonym of Pleospora herbarum,<br />
despite its striking sheath of ascospores (Wehmeyer<br />
1961). Molecular phylogenetic analysis based on partial<br />
SSU and ITS rDNA sequences indicated that Decorospora<br />
gaudefroyi was a sister taxon in the Pleosporaceae<br />
represented by Alternaria alternata (Fr.) Keissl., Cochliobolus<br />
sativus, Pleospora herbarum, Pyrenophora triticirepentis<br />
(Died.) Drechsler and Setosphaeria rostrata K.J.<br />
Leonard (Inderbitzin et al. 2002). Decorospora was<br />
introduced as a monotypic genus represented by Decorospora<br />
gaudefroyi, which is characterized by black ascomata<br />
becoming superficial on the substrate at maturity,<br />
septate and branched pseudoparaphyses, fissitunicate, clavate<br />
asci, as well as yellowish brown ascospores with seven<br />
transverse septa and one to three longitudinal septa in each
Fungal Diversity<br />
segment, enclosed in a sheath with 4–5 apical extensions<br />
(Inderbitzin et al. 2002). Decorospora gaudefroyi is an<br />
obligate marine fungus, growing at or above the high water<br />
mark (Inderbitzin et al. 2002).<br />
Diadema Shoemaker & C.E. Babc., Can. J. Bot. 67: 1349<br />
(1989).<br />
Type species: Diadema tetramerum Shoemaker & C.E.<br />
Babc. [as ‘tetramera’], Can. J. Bot. 67: 1354 (1989).<br />
During their study of Leptosphaeria and Phaeosphaeria,<br />
Shoemaker and Babcock (1989c) found some alpine fungi<br />
with typical pleosporalean characters (such as perithecoid<br />
ascomata, bitunicate asci and presence of pseudoparaphyses)<br />
having relatively large, very dark brown ascospores, mostly<br />
with a peculiar disc-like opening (as reported in some<br />
species of Wettsteinina, Shoemaker and Babcock 1987).<br />
Thus, they introduced a new genus Diadema (typified by D.<br />
tetramerum) to accommodate them (Shoemaker and<br />
Babcock 1989c). Currently, Diadema is assigned to Diademaceae,<br />
and differs from other genera in the family in having<br />
ascospores which lack longitudinal septa (Shoemaker and<br />
Babcock 1992). The large, dark brown ascospores and the<br />
disc-like opening, however, may be an adaptation to<br />
environmental factors.<br />
Diademosa Shoemaker & C.E. Babc., Can. J. Bot. 70: 1641<br />
(1992).<br />
Type species: Diademosa californiana (M.E. Barr) Shoemaker<br />
& C.E. Babc. [as ‘californianum’], Can. J. Bot. 70: 1641 (1992).<br />
≡ Graphyllium californianum M.E. Barr, Mem. N. Y.<br />
bot. Gdn 62: 40 (1990).<br />
Diademosa is the only genus in Diademaceae that has<br />
terete (cylindrical, circular in cross section) ascospores<br />
(Shoemaker and Babcock 1992).<br />
Didymella Sacc., Michelia 2(no. 6): 57 (1880).<br />
Type species: Didymella exigua (Niessl) Sacc., Syll. fung.<br />
(Abellini) 1: 553 (1882).<br />
≡ Didymosphaeria exigua Niessl, Öst. bot. Z.: 165<br />
(1875).<br />
The type specimen of Didymella (D. exigua) is lost and a<br />
neotype specimen was selected by de Gruyter et al. (2009).<br />
Didymella was characterized by the immersed or erumpent,<br />
globose or flattened and ostiolate ascomata with dense, rare<br />
(or lack?) of pseudoparaphyses. Asci are cylindrical,<br />
clavate or saccate, and 8-spored. Ascospores are hyaline,<br />
1-septate (symmetrical or asymmetrical) and constricted at<br />
the septum. Didymella has been assigned under Mycosphaerellaceae,<br />
<strong>Pleosporales</strong> (Sivanesan 1984), Phaeosphaeriaceae<br />
(Barr 1979a; Silva-Hanlin and Hanlin 1999),<br />
Venturiaceae (Reddy et al. 1998) or<strong>Pleosporales</strong> genera<br />
incertae sedis (Lumbsch and Huhndorf 2007). Based on a<br />
multigene phylogenetic analysis, the Didymella clade forms<br />
a familial rank within Pleosporineae, thus the Didymellaceae<br />
was introduced (Aveskamp et al. 2010; de Gruyter et<br />
al. 2009; Zhang et al. 2009a; Plate 1). Anamorphs of<br />
Didymellaceae include Ascochyta, Ampelomyces, Boeremia,<br />
Chaetasbolisia, Dactuliochaeta, Epicoccum, Microsphaeropsis,<br />
Peyronellaea, Phoma, Piggotia, Pithoascus,<br />
Pithomyces and Stagonosporopsis (Aveskamp et al. 2010;<br />
de Gruyter et al. 2009; Hyde et al. 2011).<br />
Didymocrea Kowalski, Mycologia 57: 405 (1965).<br />
Type species: Didymocrea sadasivanii (T.K.R. Reddy)<br />
Kowalski, Mycologia 57: 405 (1965).<br />
≡ Didymosphaeria sadasivanii T.K.R. Reddy, Mycologia<br />
53: 471 (1962).<br />
Didymocrea is a monotypic genus, and was separated<br />
from Didymosphaeria based on its “unitunicate asci”,<br />
presence of pseudoparaphyses and absence of spermatia,<br />
and assigned under Hypocreales (Kowalski 1965). Following<br />
Kowalski (1965), Luttrell (1975) also studied the<br />
centrum development of Didymocrea, and concluded that it<br />
should be a true pleosporalean fungus with functionally<br />
unitunicate asci, and retained it in Didymosphaeria. After<br />
studying the type specimen of Didymocrea sadasivanii,<br />
Aptroot (1995) concluded that it should be closely related to<br />
the loculoascomycetous genus Zopfia. Rossman et al. (1999)<br />
also kept it as a unique genus in <strong>Pleosporales</strong>. Based on a<br />
multigene phylogenetic analysis, D. sadasivanii nests within<br />
Montagnulaceae (Kruys et al. 2006; Schoch et al. 2009).<br />
Dothivalsaria Petr., Sydowia 19: 283 (1966) [1965].<br />
Type species: Dothivalsaria megalospora (Auersw.) Petr.,<br />
Sydowia 19: 283 (1966) [1965].<br />
≡ Valsaria megalospora Auersw., Leipzig. Bot.<br />
Tauschver. 5. (1866).<br />
Dothivalsaria is monotypic and is represented by D.<br />
megalospora (Petrak 1965). The taxon is characterized by<br />
immersed, medium- to large-sized ascomata which usually<br />
aggregate under blackened stromatic tissues and have<br />
trabeculate pseudoparaphyses. Asci are cylindrical, while<br />
ascospores are brown, ellipsoid, and 1-septate and uniseriate<br />
in the asci (Barr 1990a). The ascostroma of D. megalospora is<br />
comparable with those of Aglaospora profusa as has been<br />
mentioned by Barr (1990a), but their relationships are unclear.<br />
Epiphegia G.H. Otth, Mitt. naturf. Ges. Bern: 104 (1870).<br />
Type species: Epiphegia alni G.H. Otth, Mitt. naturf. Ges.<br />
Bern: 104 (1870).<br />
Epiphegia was reinstated to accommodate a species<br />
which has Phragmoporthe-like ascocarps and Massarinalike<br />
asci, pseudoparaphyses and ascospores (Aptroot 1998).<br />
Ascomata are grouped within stromatic tissues, pseudoparaphyses<br />
are cellular, asci are bitunicate and ascospores are<br />
hyaline and trans-septate (Aptroot 1998).
Fungal Diversity<br />
Eremodothis Arx, Kavaka 3: 34 (1976) [1975] (IMI 90223=<br />
<strong>CBS</strong> 610.74 type).<br />
Type species: Eremodothis angulata (A.C. Das) Arx,<br />
Kavaka 3: 34 (1976) [1975].<br />
≡ Thielavia angulata A.C. Das, Trans. Br. Mycol. Soc.<br />
45: 545 (1962).<br />
The type species of Eremodothis (E. angulata) was<br />
originally isolated from rice field soil in Fulta, India and it<br />
was assigned to Sporormiaceae because of the orange<br />
pigmentation of the colony (von Arx 1976). The cleistothecoid<br />
ascomata, sphaerical asci and 1-celled ascospores of E.<br />
angulata are comparable with those of Pycnidiophora.<br />
Based on a multigene phylogenetic study, both Eremodothis<br />
and Pycnidiophora were treated as synonyms of Westerdykella<br />
(Kruys and Wedin 2009).<br />
Extrawettsteinina M.E. Barr, Contr. Univ. Mich. Herb. 9<br />
(8): 538 (1972).<br />
Type species: Extrawettsteinina minuta M.E. Barr, Contr.<br />
Univ. Mich. Herb. 9(8): 538 (1972).<br />
Extrawettsteinina was introduced to accommodate three<br />
species, i.e. E. minuta, E. pinastri M.E. Barr and E. mediterranea<br />
(E. Müll.) M.E. Barr, which are saprobic on the dead leaves<br />
of gymnosperms and angiosperms, in North America and<br />
Europe (Barr 1972). Subsequently, a fourth species was<br />
introduced, viz. E. andromedae (Auersw.) M.E. Barr (Barr<br />
1987a). Extrawettsteinina is characterized by superficial, conical<br />
ascomata, containing a few saccate bitunicate asci, ellipsoidal,<br />
obovate-clavate, septate, smooth and hyaline ascospores which<br />
turn dull brown at maturity (Barr 1972). The diagnostic<br />
character of Extrawettsteinina is its conic ascocarps which are<br />
superficial on the substrate, and radiating arrangement of wall<br />
cells, which makes it distinguishable from comparable genera<br />
such as Stomatogene and Wettsteinina.<br />
Graphyllium Clem., Botanical Survey of Nebraska 5: 6 (1901).<br />
Type species: Graphyllium chloës Clem., Botanical Survey<br />
of Nebraska 5: 6 (1901).<br />
Graphyllium was first described as a hysteriaceous<br />
fungus with elongate ascomata, but von Höhnel (1918b,<br />
1919) recognized its similarity to Clathrospora. Petrak<br />
(1952) transferred Graphyllium to Pleospora, andnoted<br />
that the elongate ascomata and closely grouped rows of<br />
small ascomata are not sufficient to recognize the genus.<br />
Barr (1987b, 1990b) supported this proposal and considered<br />
Graphyllium differs from Clathrospora by shape,<br />
septation and pigmentation of ascospores. A narrow<br />
generic concept of Graphyllium was adapted by Shoemaker<br />
and Babcock (1992), which is characterized by hysterothecia,<br />
applanate ascospores that are at least 3-septate in side view<br />
and have some longitudinal septa in front view, and it was<br />
assigned under Hysteriaceae (order <strong>Pleosporales</strong>, Shoemaker<br />
and Babcock 1992). But subsequent classification systems<br />
tend to assign it to Diademaceae (e.g. Lumbsch and<br />
Huhndorf 2007, 2010). This seems unlikely as pointed out<br />
by Zhang et al. (2011) and the genus could be placed in one<br />
of five families containing hysteriotheciod ascomata. Recollection<br />
and molecular studies are needed.<br />
Halomassarina Suetrong, Sakay., E.B.G. Jones, Kohlm.,<br />
Volkm.-Kohlm. & C.L. Schoch, Stud. Mycol. 64: 161 (2009).<br />
Type species: Halomassarina thalassiae (Kohlm. & Volkm.-<br />
Kohlm.) Suetrong, Sakay., E.B.G. Jones, Kohlm., Volkm.-<br />
Kohlm. & C.L. Schoch, Stud. Mycol. 64: 161 (2009).<br />
≡ Massarina thalassiae Kohlm. & Volkm.-Kohlm., Can.<br />
J. Bot. 65: 575 (1987).<br />
Halomassarina is another marine genus which morphologically<br />
fits Massarina sensu lato, and is typified by H.<br />
thalassiae, which is characterized by subglobose to pyriform,<br />
immersed or erumpent, ostiolate, periphysate, papillate<br />
or epapillate, coriaceous ascomata, simple, rarely anastomosing<br />
pseudoparaphyses, 8-spored, cylindrical to clavate,<br />
pedunculate, thick-walled, fissitunicate asci, and ellipsoidal,<br />
(1-)3-septate, hyaline ascospores. Based on a multigene<br />
phylogenetic analysis, Halomassarina thalassiae clustered<br />
together with Trematosphaeria pertusa and another marine<br />
fungus Falciformispora lignatilis, and they are all assigned<br />
to Trematosphaeriaceae (Suetrong et al. data unpublished).<br />
Hypsostroma Huhndorf, Mycologia 84: 750 (1992).<br />
Type species: Hypsostroma saxicola Huhndorf, Mycologia<br />
84: 750 (1992).<br />
Hypsostroma was introduced as a tropical woodinhabiting<br />
genus by Huhndorf (1992). Hypsostroma has<br />
several striking characters including the large superficial<br />
ascomata which form on a subiculum, pseudoparenchymatous<br />
peridial cells, trabeculate pseudoparaphyses, clavate<br />
asci with long pedicels and a conspicuous apical apparatus,<br />
and ascospores that separate into partspores with a germ slit<br />
in each partspore (Huhndorf 1992). Phylogenetic study<br />
indicated that Hypsostroma should be a new genus and the<br />
Hypsostromataceae was reinstated to accommodate Hypsostroma<br />
(Mugambi and Huhndorf 2009b; Plate 1).<br />
Julella Fabre, Annls Sci. Nat., Bot., sér. 6 9: 113 (1879) [1878].<br />
Type species: Julella buxi Fabre, Annls Sci. Nat., Bot., sér.<br />
6 9: 113 (1879) [1878].<br />
Julella has been assigned to Thelenellaceae, a family of<br />
Ostropomycetidae (Lumbsch and Huhndorf 2007), and<br />
Arthopyreniaceae (= Xanthopyreniaceae sensu O. Eriksson,<br />
<strong>Pleosporales</strong>) (Barr 1985). Julella is characterized by its<br />
immersed, medium-sized ascomata with pseudoparenchymatous<br />
peridial cells, cellular pseudoparaphyses, and<br />
hyaline and muriform ascospores (Barr 1985). With the<br />
exception of hyaline ascospores, these characters are typical<br />
of Montagnulaceae. The taxonomic affinity of the generic
Fungal Diversity<br />
type of Julella needs confirmation following recollection.<br />
Julella avicenniae (Borse) K.D. Hyde is a marine fungus. A<br />
DNA based phylogeny containing most currently accepted<br />
families placed two isolates of J. avicenniae as sister to the<br />
families in the Pleosporineae with good support, which might<br />
suggest a novel family within <strong>Pleosporales</strong> (Suetrong et al.<br />
2009). However, J. avicenniae is not the generic type and<br />
therefore this conclusion must be treated with caution as only<br />
J. avicenniae can be considered pleosporalean.<br />
Lautitia S. Schatz, Can. J. Bot. 62: 31 (1984).<br />
Type species: Lautitia danica (Berl.) S. Schatz, Can. J. Bot.<br />
62: 31 (1984).<br />
≡ Leptosphaeria danica Berl., Icon. fung. (Abellini) 1:<br />
87 (1892).<br />
Lautitia is monotypified by L. danica, which is characterized<br />
by subglobose, immersed, ostiolate ascomata with a<br />
pseudoclypeus, a thin peridium, broad, cellular pseudoparaphyses,<br />
and 8-spored, bitunicate, cylindrical to clavate asci.<br />
Ascospores are hyaline, 1-septate, and obovate and the fungus<br />
is parasitic on algae (Schatz 1984). Marine or maritime fungi<br />
have been reported in Phaeosphaeria, suchasP. spartinae<br />
(Ellis & Everh.) Shoemaker & C.E. Babc. and P. ammophilae<br />
(Lasch) Kohlm. & E. Kohlm. (Zhang et al. 2009a). In<br />
addition, the prosenchymatous peridium of L. danica agrees<br />
with that of Phaeosphaeriaceae (Schatz 1984).<br />
Lepidosphaeria Parg.-Leduc, C. r. hebd. Séanc. Acad. Sci.,<br />
Paris, Sér. D 270: 2786 (1970).<br />
Type species: Lepidosphaeria nicotiae Parg.-Leduc, Pubbl.<br />
Staz. Zool. Napoli, 1 270: 2786 (1970).<br />
Lepidosphaeria is a genus likely in Testudinaceae, which<br />
is distinguished from other genera of this family by its<br />
smaller ascospores, which lack furrows, and have minute<br />
granulate ornamentation (Hawksworth 1979). In DNA<br />
sequence-based phylogenies, L. nicotiae clustered with<br />
species of Ulospora and Verruculina (Schoch et al. 2009;<br />
Zhang et al. 2009a), but more recent work including<br />
species of Platystomaceae lacks support (Plate 1).<br />
Letendraea Sacc., Michelia 2: 73 (1880).<br />
Type species: Letendraea eurotioides Sacc., Michelia 2: 73<br />
(1880).<br />
Letendraea was introduced for L. eurotioides, which is<br />
characterized by superficial, globose to conical ascomata,<br />
filliform pseudoparaphyses, obclavate to cylindrical, 8-<br />
spored asci, and fusoid to oblong, 1-septate ascospores<br />
(Saccardo 1880). Because L. helminthicola (Berk. &<br />
Broome) Weese clustered with Karstenula rhodostoma,<br />
Letendraea was assigned to Melanommataceae (Kodsueb<br />
et al. 2006b). But subsequent multigene phylogenetic<br />
analysis indicated that both L. helminthicola and L. padouk<br />
Nicot & Parg.-Leduc nested within Montagnulaceae<br />
(Schoch et al. 2009; Zhang et al. 2009a; Plate 1), and its<br />
familial status seems confirmed.<br />
Lindgomyces K. Hirayama, Kaz. Tanaka & Shearer,<br />
Mycologia 102: 133 (2010).<br />
Type species: Lindgomyces ingoldianus (Shearer & K.D.<br />
Hyde) K. Hirayama, Kaz. Tanaka & Shearer, Mycologia<br />
102: 733 (2010).<br />
≡ Massarina ingoldiana Shearer & K.D. Hyde, Mycologia<br />
89: 114 (1997).<br />
Lindgomyces was introduced to accommodate a freshwater<br />
lineage, which belongs to Massarina ingoldiana sensu lato,<br />
and is characterized by scattered, subglobose to globose,<br />
erumpent, papillate, ostiolate ascomata, cellular pseudoparaphyses,<br />
and 8-spored, fissitunicate, cylindrical to clavate<br />
asci. Ascospores are fusoid to narrowly fusoid, hyaline and 1-<br />
septate but become 3–5-septate when senescent (Hirayama et<br />
al. 2010). A new family, Lindgomycetaceae, wasintroduced<br />
to accommodate Lindgomyces (Hirayama et al. 2010).<br />
Lophiella Sacc., Michelia 1: 337 (1878).<br />
Type species: Lophiella cristata (Pers.) Sacc., Michelia 1:<br />
337 (1878).<br />
≡ Sphaeria cristata Pers., Syn. meth. fung. (Göttingen)<br />
1: 54 (1801).<br />
The generic type of Lophiella, L. cristata, was treated as<br />
a synonym of Lophiostoma angustilabrum var. crenatum<br />
(Pers.) Chesters & A.E. Bell (see http://www.indexfungorum.<br />
org/names/Names.asp).<br />
Loratospora Kohlm. & Volkm.-Kohlm., Syst. Ascom. 12:<br />
10 (1993).<br />
Type species: Loratospora aestuarii Kohlm. & Volkm.-<br />
Kohlm., Syst. Ascom. 12: 10 (1993).<br />
Loratospora was introduced as a marine genus and is<br />
monotypified by L. aestuarii (Kohlmeyer and Volkmann-<br />
Kohlmeyer 1993). The generic type is characterized by<br />
ellipsoid, immersed to erumpent, carbonaceous ascomata,<br />
which are ostiolate, and with or without a papilla. Pseudoparaphyses<br />
comprise small subglobose cells forming irregular<br />
chains and finally breaking apart, and asci are 8-spored,<br />
clavate to ellipsoidal, and fissitunicate. Ascospores are<br />
hyaline, cylindrical, 3-septate and surrounded by a mucilaginous<br />
sheath (Kohlmeyer and Volkmann-Kohlmeyer 1993).<br />
The distinctive pseudoparaphyses of Loratospora aestuarii<br />
makes it readily distinguishable from other taxa. Based on a<br />
multigene phylogenetic analysis, Loratospora aestuarii nested<br />
within the clade of Phaeosphaeriaceae (Schoch et al. 2009;<br />
Suetrong et al. 2009; Plate 1), and ascospores of L. aestuarii<br />
are in agreement with those of Phaeosphaeria as has been<br />
mentioned by Kohlmeyer and Volkmann-Kohlmeyer (1993).<br />
Macrospora Fuckel, Jb. nassau. Ver. Naturk. 23–24: 139<br />
(1870) [1869–70].
Fungal Diversity<br />
Type species: Macrospora scirpicola (DC.) Fuckel, Jb.<br />
nassau. Ver. Naturk. 23–24: 139 (1870) [1869–70].<br />
≡ Sphaeria scirpicola DC., in Lamarck & de Candolle,<br />
Fl. franç., Edn 3 (Paris) 2: 300 (1805).<br />
Macrospora had been assigned to Diademaceae based<br />
on its applanate and muriform ascospores with 1-row of<br />
longitudinal septa, with a sheath, 2–3 μm wide and<br />
constricted at first septum and ascospores are paler and<br />
larger than those of Comoclathris (Shoemaker and Babcock<br />
1992). Macrospora was however, considered as a synonym<br />
of Pyrenophora by Eriksson and Hawksworth (1991) which<br />
was assigned in Pleosporaceae, and this proposal was<br />
widely followed (Eriksson 2006; Lumbsch and Huhndorf<br />
2010). Nimbya anamorphs were reported for Macrospora<br />
(Johnson et al. 2002).<br />
Massaria De Not., G. bot. ital. 1: 333 (1844).<br />
Type species: Massaria inquinans (Tode) De Not., G. bot.<br />
ital. 1: 333 (1844).<br />
≡ Sphaeria inquinans Tode, Fung. mecklenb. sel.<br />
(Lüneburg) 1: Fig. 85 (1790).<br />
Colonies on MEA erumpent, not spreading; surface<br />
irregular, folded; margins even, feathery; surface olivaceous<br />
grey, with thin, umber margin; reverse olivaceous-grey. On<br />
PDA similar; surface olivaceous grey, margin dirty white;<br />
reverse smoke-grey to olivaceous grey; colonies reaching<br />
1 cm diam. On OA similar, surface olivaceous grey in<br />
centre, margins wide, dirty white; colonies reaching 12 mm<br />
diam. on all media tested; colonies sterile (based on <strong>CBS</strong><br />
125591).<br />
Massaria was formally established by de Notaris (1844),<br />
and is typified by M. inquinans. Numerous fungi with<br />
brown septate ascospores surrounded by gelatinous sheath<br />
were included in the genus (Barr 1979b; Shoemaker and<br />
LeClair 1975). Shoemaker and LeClair (1975) accepted a<br />
narrow concept for Massaria, with only a few species<br />
characterized by large, symmetric, 4-celled ascospores<br />
surrounded by a massive gelatinous sheath. Barr (1979b,<br />
1990a) had considered Aglaospora a separate genus, but this<br />
subsequently proved congeneric with Massaria (Voglmayr<br />
and Jaklitsch 2011). Based on intensive sample collection<br />
and multi-gene phylogenetic analysis, Voglmayr and<br />
Jaklitsch (2011) acceptedMassaria as the sole genus within<br />
Massariaceae, which is characterized by a set of well<br />
defined morphological and ecological characters; Europe is<br />
regarded as the centre of diversity.<br />
Misturatosphaeria Mugambi & Huhndorf, Stud. Mycol.<br />
64: 108 (2009).<br />
Type species: Misturatosphaeria aurantonotata Mugambi &<br />
Huhndorf, Stud. Mycol. 64: 108 (2009).<br />
Misturatosphaeria was introduced to accommodate a<br />
group of fungi which are phylogenetically closely related to<br />
Amniculicolaceae, Lophiostomataceae sensu stricto and<br />
Sporormiaceae (Mugambi and Huhndorf 2009b; Zhang et<br />
al. 2009a). Species of Misturatosphaeria are characterized<br />
by erumpent to superficial ascomata which are scattered or<br />
in groups, with or without papilla; asci cylindrical or<br />
clavate, 8-spored; pseudoparaphyses numerous, septate,<br />
ascospores brown or hyaline, phragmosporous or dictyosporous,<br />
with or without sheath. The terrestrial saprobic<br />
habitat on wood, as well as its distinct morphological<br />
characters may indicate that this genus belongs to an<br />
undescribed family. A close relationship with the marine<br />
anamorphic species Floricola striata is unexpected and<br />
may suggest that some of the species in this genus could<br />
have marine affinities (Plate 1).<br />
Navicella Fabre, Annls Sci. Nat., Bot., sér. 6 9: 96 (1879)<br />
[1878].<br />
Type species: Navicella julii Fabre, Annls Sci. Nat., Bot.,<br />
sér. 6 9: 96 (1879) [1878].<br />
Navicella is characterized by medium- to large-sized,<br />
immersed to erumpent, globose ascomata, apex elongated or<br />
rarely rounded, asci clavate or cylindrical, pseudoparaphyses<br />
trabeculate, ascospores reddish to dark brown, ellipsoid to<br />
fusoid, multi-septate, the primary septum is euseptate, and<br />
others distoseptate, obliquely uniseriate or biseriate (Barr<br />
1990a). Navicella is saprobic on bark, and was considered<br />
closely related to the Lophiostomataceae (Holm and Holm<br />
1988). Based on the wide endotunica, thin apical ring and<br />
distoseptate ascospores, Barr (1990a) transferred it to the<br />
Massariaceae. The morphological characters of Navicella do<br />
not match the Massariaceae sensu stricto (Voglmayr and<br />
Jaklitsch 2011).<br />
Neotestudina Segretain & Destombes, C. r. hebd. Séanc.<br />
Acad. Sci., Paris 253: 2579 (1961).<br />
Type species: Neotestudina rosatii Segretain &Destombes,<br />
C. r. hebd. Séanc. Acad. Sci., Paris 253: 2579 (1961).<br />
Neotestudina is characterized by medium- to large-sized,<br />
superficial, gregarious, cleistothecioid and globose ascomata<br />
which split on opening. Asci are 4- or 8-spored, and cylindrical<br />
or oblong, pseudoparaphyses are sparse and trabeculate, and<br />
ascospores are dark brown, ellipsoid, 1-septate, with a small<br />
germ pore at each end, and uniseriate or crowded in the asci<br />
(Barr 1990a). Based on the cleistothecioid ascomata, Neotestudina<br />
was assigned under Zopfiaceae (von Arx and Müller<br />
1975) or Testudinaceae (Hawksworth 1979). Barr (1990a)<br />
assigned it to Didymosphaeriaceae basedonitsascospore<br />
morphology. A DNA based phylogeny showed that sequence<br />
obtained from Neotestudina rosatii resides as sister to<br />
Ulospora bilgramii (D. Hawksw., C. Booth & Morgan-<br />
Jones) D. Hawksw., Malloch & Sivan. and other species that<br />
may represent Testudinaceae or Platystomaceae (Kruys et al.<br />
2006; Plate 1).
Fungal Diversity<br />
Paraphaeosphaeria O.E. Erikss., Ark. Bot., Ser. 2 6: 405<br />
(1967).<br />
Type species: Paraphaeosphaeria michotii (Westend.) O.E.<br />
Erikss., Cryptogams of the Himalayas 6: 405 (1967).<br />
≡ Sphaeria michotii Westend., Bull. Acad. R. Sci. Belg.,<br />
Cl. Sci., sér. 2 7: 87 (1859).<br />
Paraphaeosphaeria was separated from Leptosphaeria<br />
(Eriksson 1967a), and it is also quite comparable with<br />
Phaeosphaeria. Paraphaeosphaeria can be distinguished<br />
from Phaeosphaeria by its ascospores. Ascospores of Paraphaeosphaeria<br />
michotii have two septa, and they are biseriate,<br />
straight, subcylindrical with broadly rounded ends, rather dark<br />
brown and punctate. The primary septum is laid down closer<br />
to the distal end than to the proximal, and the larger, proximal<br />
hemispore is divided by one transversal septum. There are<br />
more septa in the proximal hemispore of other species such as<br />
Par. castagnei (Durieu & Mont.) O.E. Erikss., Par. obtusispora<br />
(Speg.) O.E. Erikss. and Par. vectis (Berk. & Broome)<br />
Hedjar. Anamorphic characters can also distinguish Paraphaeosphaeria<br />
and Phaeosphaeria. Paraphaeosphaeria has<br />
Paraconiothyrium or Coniothyrium-related anamorphs, but<br />
Phaeosphaeria has Hendersonia-Phaeoseptoria anamorphs<br />
(Eriksson 1967a). Shoemaker and Babcock (1985) redescribed<br />
some Canadian and extralimital species, and excluded<br />
Par. longispora (Wegelin) Crivelli and Par. oblongata<br />
(Niessl) Crivelli from Paraphaeosphaeria based on their<br />
longitudinal septa as well as beak-like papilla and wall<br />
structures. Molecular phylogenetic results based on multigenes<br />
indicated that Paraphaeosphaeria should belong to<br />
Montagnulaceae (Zhang et al. 2009a; Plate 1).<br />
Passeriniella Berl., Icon. fung. (Abellini) 1: 51 (1890).<br />
Type species: Passeriniella dichroa (Pass.) Berl., Icon.<br />
fung. (Abellini) 1: 51 (1890).<br />
≡ Leptosphaeria dichroa Pass.<br />
Passeriniella was introduced by Berlese in 1890 based on<br />
the black, ostiolate and papillate ascomata, 8-spored asci, as<br />
well as transverse septate ascospores, with pigmented central<br />
cells and hyaline terminal cells. Two species were included, i.<br />
e. P. dichroa and P. incarcerata (Berk. & M.A. Curtis) Berl.<br />
(Berlese 1890). Subsequently, more species were introduced<br />
including some marine taxa such as P. mangrovei G.L. Maria<br />
&K.R.Sridhar,P. obiones (P. Crouan & H. Crouan) K.D.<br />
Hyde & Mouzouras and P. savoryellopsis K.D. Hyde &<br />
Mouzouras (Hyde and Mouzouras 1988; Maria and Sridhar<br />
2002). Currently, eight species are included (http://www.<br />
mycobank.org, Jan. 2011). Both P. dichroa and P. incarcerata<br />
were considered as synonyms of Leptosphaeria obiones<br />
(P. Crouan & H. Crouan) Sacc. (Kohlmeyer and Kohlmeyer<br />
1979). The familial placement of the marine species P.<br />
savoryellopsis could not be resolved in a DNA based<br />
phylogeny but it did suggest a close relationship to<br />
Acrocordiopsis patilii (Suetrong et al. 2009) in<strong>Pleosporales</strong>.<br />
Peridiothelia D. Hawksw., Bull. Br. Mus. nat. Hist., Bot.<br />
14: 120 (1985).<br />
Type species: Peridiothelia fuliguncta (Norman) D.<br />
Hawksw., Bull. Br. Mus. nat. Hist., Bot. 14: 121 (1985).<br />
≡ Microthelia fuliguncta Norman, Öfvers. kongl.<br />
Svensk. Vetensk.-Akad. Förhandl., Stockholm 41(no. 8):<br />
36 (1884).<br />
When dealing with the names under Microthelia, Peridiothelia<br />
was introduced to accommodate species having nonclypeate<br />
peridium which composed cells of textura globulosa<br />
but sometimes angularis, “dark reddish brown except<br />
below the generative locule where the wall is poorly<br />
developed or almost absent at maturity, colour not changed<br />
significantly in potassium hydroxide, centrum turning blue in<br />
iodine” (Hawksworth 1985a). Three species were included, i.<br />
e. P. grandiuscula (Anzi) D. Hawksw., P. fuliguncta and P.<br />
oleae (Körb.) D. Hawksw., and Peridiothelia was referred to<br />
Phaeosphaeriaceae (Hawksworth 1985a, b). However, its<br />
familial placement is not confirmed yet.<br />
Phaeodothis Syd. & P. Syd., Annls mycol. 2: 166 (1904).<br />
Type species: Phaeodothis tricuspidis Syd. & P. Syd.,<br />
Annls mycol. 2: 166 (1904).<br />
Phaeodothis is characterized by its 1-septate euseptate<br />
ascospores with a sparse hamathecium consisting of thin<br />
pseudoparaphyses and immersed to superficial ascomata<br />
(Aptroot 1995).Thegenushadbeenpreviouslyassignedto<br />
Didymosphaeria, butAptroot(1995) consideredittobeclosely<br />
related to Phaeosphaeriaceae. A strain named Phaeodothis<br />
winteri (a synonym of P. tricuspidis Syd. & P. Syd.) nested<br />
within the clade of Montagnulaceae (Schoch et al. 2009).<br />
Platychora Petr., Annls mycol. 23: 102 (1925).<br />
Type species: Platychora ulmi (Schleich.) Petr., Annls<br />
mycol. 23(1/2): 103 (1925).<br />
Platychora is characterized by immersed to erumpent crustlike<br />
ascostroma with globose locules scattered inside (Barr<br />
1968). Asci are oblong to saccate or nearly cylindrical and<br />
bitunicate, and ascospores are hyaline 1-septate apiosporous<br />
and turn olivaceous when old. Platychora hadbeenpreviously<br />
assigned to Venturiaceae by Barr (1968), but molecular<br />
phylogenetic analysis indicated that a strain named Platychora<br />
ulmi (the generic type of Platychora) belongs to Didymellaceae<br />
(Winton et al. 2007; Plate 1). The generic type needs<br />
recollecting and epitypifying to stabilize the generic name.<br />
Platystomum Trevis., Bull. Soc. R. Bot. Belg. 16: 16 (1877).<br />
Type species: Platystomum compressum (Pers.) Trevis.,<br />
Bull. Soc. R. Bot. Belg. 16: 16 (1877).<br />
≡ Sphaeria compressa Pers., Syn. meth. fung. (Göttingen)<br />
1: 56 (1801).<br />
Platystomum was introduced by Trevisan in 1877, and<br />
has been considered a synonym of Lophidium, as the
Fungal Diversity<br />
ascospores of Platystomum have both transverse and<br />
vertical septa (Barr 1990a, b; ChestersandBell1970).<br />
However, the boundary between Lophiostoma and Platystomum<br />
is not clear (Chesters and Bell 1970). Holm and<br />
Holm (1988) treated Platystomum as a synonym of<br />
Lophiostoma, and concurrently, the Platystomaceae<br />
should be treated as a synonym of Lophiostomataceae.<br />
Based on a phylogenetic analysis, however, the generic<br />
type of Platystomum (P. compressum) separated from<br />
other species of Lophiostoma, and nested with the clade of<br />
Platystomaceae (Mugambi and Huhndorf 2009b) which<br />
may be closely related to species in the Testiduniaceae<br />
(Plate 1).<br />
Polyplosphaeria Kaz. Tanaka & K. Hirayama, Stud.<br />
Mycol. 64: 192 (2009).<br />
Type species: Polyplosphaeria fusca Kaz. Tanaka & K.<br />
Hirayama, Stud. Mycol. 64: 193 (2009).<br />
Polyplosphaeria is characterized by globose ascomata<br />
surrounded by numerous brown hyphae and a reddish<br />
pigment on the host surface around the ascomata (Tanaka et<br />
al. 2009). Asci are cylindro-clavate with fissitunicate<br />
dehiscence and ascospores are narrowly fusoid surrounded<br />
by a sheath. The anamorph is Piricauda-like (Tanaka et al.<br />
2009). The cylindro-clavate asci, narrowly fusoid ascospores<br />
as well as its thin and numerous pseudoparaphyses are<br />
comparable with those of Massarina sensu lato, especially<br />
Lentithecium (Zhang et al. 2009a). The terrestrial and<br />
bambusicolous habitat of Polyplosphaeria and Piricauda<br />
anamorph readily distinguishes the genus from Lentithecium.<br />
Pontoporeia Kohlm., Nova Hedwigia 6: 5 (1963).<br />
Type species: Pontoporeia biturbinata (Durieu & Mont.)<br />
Kohlm., Nova Hedwigia 6: 5 (1963)<br />
≡ Sphaeria biturbinata Durieu & Mont., Flora Algéricae<br />
1: 497 (1849).<br />
Pontoporeia was introduced by Kohlmeyer in 1963,<br />
and is monotypified by P. biturbinata. Pontoporeia was<br />
treated as a synonym of Zopfia (Malloch and Cain 1972),<br />
which is followed by Hawksworth and Booth (1974).<br />
Based on its asci originating at the periphery of the<br />
subglobose locus, filaments occupying the center of the<br />
ascocarps, the irregular peridial structure, the ascospores<br />
having 2-layered walls with a germ pore at each end and<br />
its marine habitat, Pontoporeia was kept as a separate<br />
genus within Pleosporaceae (Kohlmeyer and Kohlmeyer<br />
1979). A DNA based phylogeny placed an isolate on a<br />
long branch in relationship with other marine species,<br />
Halotthia posidoniae and Mauritiana rhizophorae, but a<br />
familial placement awaits further resolution (Suetrong et<br />
al. 2009).<br />
Pseudotrichia Kirschst., Annls mycol. 37: 125 (1939).<br />
Type species: Pseudotrichia stromatophila Kirschst., Annls<br />
mycol. 37: 125 (1939).<br />
Pseudotrichia can be distinguished from Byssosphaeria,<br />
Herpotrichia and Lojkania by its lacking of subiculum,<br />
larger ascomata usually with compressed apices, the<br />
peripheral arrangement of asci and trabeculate pseudoparaphyses<br />
(Barr 1984). Phylogenetic study of strains Pseudotrichia<br />
mutabilis and some Herpotrichia species indicated<br />
that these species are closely related, and both nested within<br />
Melanommataceae (Mugambi and Huhndorf 2009b). But in<br />
this study, Pseudotrichia guatopoensis nested in the<br />
Testudinaceae (or Platystomaceae) (Plate 1). The types of<br />
both Herpotrichia and Pseudotrichia need recollecting,<br />
redescribing and epitypifying in order to stabiles the use of<br />
these generic names and clarify their familial status.<br />
Pseudoyuconia Lar.N. Vassiljeva, Nov. sist. Niz. Rast. 20:<br />
71 (1983).<br />
Type species: Pseudoyuconia thalictri (G. Winter) Lar. N.<br />
Vassiljeva [as ‘thalicti’], Nov. sist. Niz. Rast. 20: 71 (1983).<br />
≡ Leptosphaeria thalictri G. Winter, Hedwigia 10: 40<br />
(1872).<br />
Pseudoyuconia was introduced by Vassiljeva (1983), and<br />
was monotypified by P. thalictri. Currently, Pseudoyuconia<br />
is included in Pleosporaceae (Lumbsch and Huhndorf<br />
2010).<br />
Pyrenophora Fr., Summa veg. Scand., Section Post.<br />
(Stockholm): 397 (1849).<br />
Type species: Pyrenophora phaeocomes (Rebent.) Fr.,<br />
Summa veg. Scand., Section Post. (Stockholm): 397<br />
(1849).<br />
≡ Sphaeria phaeocomes Rebent., Prodr. fl. neomarch.<br />
(Berolini): 338 (1804).<br />
Pyrenophora is characterized by immersed, erumpent to<br />
nearly superficial ascomata, indefinite pseudoparaphyses,<br />
clavate to saccate asci usually with a large apical ring, and<br />
muriform terete ascospores. Morphologically, the terete<br />
ascospores of Pyrenophora can be readily distinguished<br />
from Clathrospora and Platyspora. The indefinite pseudoparaphyses<br />
and smaller ascospores of Pyrenophora can be<br />
readily distinguished from those of Pleospora (Sivanesan<br />
1984). Based on both morphology and molecular phylogeny,<br />
Pyrenophora is closely related to Pleosporaceae<br />
(Zhang et al. 2009a).<br />
Rechingeriella Petr., in Rechinger et al. Annln naturh. Mus.<br />
Wien 50: 465 (1940).<br />
Type species: Rechingeriella insignis Petr., Annln naturh.<br />
Mus. Wien, Ser. B, Bot. Zool. 50: 465 (1940).<br />
Rechingeriella is characterized by its erumpent to<br />
superficial, cleistothecioid ascomata and thin, branching<br />
pseudoparaphyses (Hawksworth and Booth 1974). Asci are
Fungal Diversity<br />
obovate, thick-walled, bitunicate and evanescent, and<br />
ascospores are globose, simple, dark brown to black (based<br />
on the type specimen of R. insignis) (Hawksworth and<br />
Booth 1974). Based on these characters, R. insignis was<br />
treated as a species of Zopfia (as Z. insignis (Petr.) D.<br />
Hawksw. & C. Booth). Rechingeriella has been assigned to<br />
Botryosphaeriaceae by von Arx and Müller (1975). Further<br />
study should be conducted on the type specimen of R.<br />
insignis in order to clarify its taxonomic status and fresh<br />
collections are needed for epitypification.<br />
Rhytidiella Zalasky, Can. J. Bot. 46: 1383 (1968).<br />
Type species: Rhytidiella moriformis Zalasky, Can. J. Bot.<br />
46: 1383 (1968).<br />
Rhytidiella was introduced based on R. moriformis, which<br />
causes perennial rough-bark of Populus balsamifera<br />
(Zalasky 1968), and produces macroconidia belonging to<br />
Phaeoseptoria. Subsequently, three more species were<br />
introduced, viz. R. baranyayi A. Funk & Zalasky, R. hebes<br />
P.R. Johnst. and R. beloniza (Stirt.) M.B. Aguirre (Aguirre-<br />
Hudson 1991; Funk and Zalasky 1975;Johnston2007), Both<br />
R. baranyayi and R. hebes seem closely related to R.<br />
moriformis on both biology and morphology (Funk and<br />
Zalasky 1975;Johnston2007), but R. beloniza is saprobic on<br />
Cordyline australis bark (Aguirre-Hudson 1991). Rhytidiella<br />
was temporarily assigned to Cucurbitariaceae (Barr 1987b).<br />
Richonia Boud., Revue mycol., Toulouse 7: 224 (1885).<br />
Type species: Richonia variospora Boud., Revue mycol.,<br />
Toulouse 7: 265 (1885).<br />
Richonia is characterized by its 1-septate, relatively large<br />
ascospores which are broadly rounded at both ends, and<br />
have a thick ornamented undulating sheath giving an<br />
irregularly ridged appearance to mature spores (Hawksworth<br />
1979). Richonia variospora has been isolated from several<br />
localities in France, but it is rare (Hawksworth 1979).<br />
Richonia was assigned under Zopfiaceae (von Arx and<br />
Müller 1975; Hawksworth1979), and there are presently no<br />
better suggestions for its familial placement. The taxon needs<br />
recollecting and epitypifying.<br />
Rimora Kohlm., Volkm.-Kohlm., Suetrong, Sakay. & E.B.<br />
G. Jones, Stud. Mycol. 64: 166 (2009).<br />
Type species: Rimora mangrovei (Kohlm. & Vittal)<br />
Kohlm., Volkm.-Kohlm., Suetrong, Sakay. & E.B.G. Jones,<br />
Stud. Mycol. 64: 166 (2009).<br />
≡ Lophiostoma mangrovei Kohlm. & Vittal [as ‘mangrovis’],<br />
Mycologia 78: 487 (1986).<br />
Rimora was introduced based on a marine fungus R.<br />
mangrovei (syn. Lophiostoma mangrovei), and is characterized<br />
by its erumpent ascomata with elongated flat tops,<br />
cellular pseudoparaphyses and cylindrical asci (Suetrong et<br />
al. 2009). Ascospores are fusoid, hyaline, 3-septate and<br />
surrounded with an evanescent sheath (Kohlmeyer and<br />
Vittal 1986; Suetrong et al. 2009). Rimora forms a robust<br />
clade with other marine fungi, such as species of Aigialus<br />
and Ascocratera, and a new family, Aigialaceae was<br />
introduced to accommodate them (Suetrong et al. 2009).<br />
Roussoellopsis I. Hino & Katum., J. Jap. Bot. 40: 86 (1965).<br />
Type species: Roussoellopsis japonica (I. Hino & Katum.)<br />
I. Hino & Katum., J. Jap. Bot. 40: 86 (1965).<br />
≡ Didymosphaeria japonica I. Hino & Katum., Bulletin<br />
of the Faculty of Agriculture, Yamaguchi University 5: 229<br />
(1954).<br />
Roussoellopsis was introduced by Hino and Katumoto<br />
(1965) based on three bambusicolous fungal species, i.e. R.<br />
japonica, R. macrospora (I. Hino & Katum.) I. Hino &<br />
Katum. and R. tosaensis (I. Hino & Katum.) I. Hino &<br />
Katum. These three species have immersed and gregarious<br />
ascomata, clavate to cylindro-clavate asci, numerous and<br />
filliform pseudoparaphyses, and 1-septate, asymmetrical<br />
ascospores (Hino and Katumoto 1965). All these characters<br />
point Roussoellopsis to <strong>Pleosporales</strong>, but its familial<br />
placement cannot be determined.<br />
Saccothecium Fr., Fl. Scan.: 349 (1836).<br />
Type species: Saccothecium sepincola (Fr.) Fr. [as ‘saepincola’],<br />
Summa veg. Scand., Section Post. (Stockholm): 398<br />
(1849).<br />
≡ Sphaeria sepincola Fr. [as ‘saepincola’], Observ.<br />
mycol. (Havniae) 1: 181 (1815).<br />
Saccothecium is characterized by its subglobose, immersed<br />
to erumpent ascomata, absence of pseudoparaphyses<br />
and hyaline, muriform to phragmosporous ascospores. It has<br />
been assigned to the Dothioraceae (Barr 1987b; Müllerand<br />
von Arx 1950). Molecular phylogenetic analysis indicated<br />
that a strain named S. sepincola nested within Didymellaceae<br />
(Schoch et al. 2009; Plate 1). The generic type needs<br />
recollecting, redescribing and epitypifying.<br />
Setosphaeria K.J. Leonard & Suggs, Mycologia 66: 294<br />
(1974).<br />
Type species: Setosphaeria turcica (Luttr.) K.J. Leonard &<br />
Suggs, Mycologia 66: 295 (1974).<br />
≡ Trichometasphaeria turcica Luttr., Phytopathology 48:<br />
282 (1958).<br />
Setosphaeria was segregated from Keissleriella on the<br />
basis of lacking a clypeus, lysigenous development of the<br />
ostiole, occurrence of setae on the perithecial wall, the<br />
absence of periphyses in the ostiole, and the hyphomycetous<br />
conidial states, and four species were included, i.e. S.<br />
prolata, S. holmii, S. pedicellata (R.R. Nelson) K.J.<br />
Leonard & Suggs and S. turcica (Leonard and Suggs<br />
1974). Currently, nine species are included in Setosphaeria<br />
(http://www.mycobank.org, Jan/2011). Setosphaeria
Fungal Diversity<br />
monoceras Alcorn nested within Pleosporaceae based on<br />
multigene phylogenetic analysis (Schoch et al. 2009;<br />
Plate 1).<br />
Syncarpella Theiss. & Syd., Annls mycol. 13: 631 (1915).<br />
Type species: Syncarpella tumefaciens (Ellis & Harkn.)<br />
Theiss. & Syd., Annls mycol. 13(5/6): 633 (1915).<br />
≡ Sphaeria tumefaciens Ellis & Harkn., J. Mycol. 2: 41<br />
(1886).<br />
Syncarpella was introduced by Theissen and Sydow<br />
(1915) as a genus of Montagnellaceae within Dothideales.<br />
A detailed description of S. tumefaciens can be seen in Barr<br />
and Boise (1989). Syncarpella was considered closely<br />
related to Leptosphaeria, and was treated as a synonym<br />
(Clements and Shear 1931). Syncarpella is characterized by<br />
its abundant globose, ovoid to turbinate ascomata with<br />
minute papillae which are seated on a common basal stroma<br />
and which are erumpent through fissures in the host tissues<br />
(Barr and Boise 1989). The peridium is thicker at the base,<br />
pseudoparaphyses are cellular, and asci are bitunicate,<br />
clavate to oblong with a furcate pedicel. Ascospores are<br />
pale brown to brown, oblong to narrowly obovoid, ends<br />
obtuse, transversely septate, smooth-walled. All these<br />
characters fit Cucurbitariaceae, where Barr and Boise<br />
(1989) transferred Syncarpella.<br />
Teichospora Fuckel, Jb. nassau. Ver. Naturk. 23–24: 160<br />
(1870) [1869–70].<br />
Type species: Teichospora trabicola Fuckel, Jb. nassau.<br />
Ver. Naturk. 23–24: 161 (1870) [1869–70].<br />
Teichospora was introduced by Fuckel (1870), and was<br />
typified by T. trabicola, with four more species included,<br />
i.e. T. brevirostris Fuckel, T. dura Fuckel, T. morthieri<br />
Fuckel and T. obducens (Schumach.) Fuckel. Only T.<br />
brevirostris and T. trabicola were kept in Teichospora<br />
(Barr 1987b). After studying the type specimens, Barr<br />
(1987b) indicated that Teichospora was different from<br />
Strickeria with Teichospora belonging to <strong>Pleosporales</strong>,<br />
and Strickeria closely related to Melanomma (Melanommatales).<br />
Currently, more than 250 names are included<br />
within Teichospora (http://www.mycobank.org, Jan/2011),<br />
but almost no molecular phylogenetic study has been<br />
conductedonthisgenus.<br />
Testudina Bizz., Atti Inst. Veneto Sci. lett., ed Arti, Sér. 6<br />
3: 303 (1885).<br />
Type species: Testudina terrestris Bizz., Fungi venet. nov.<br />
vel. Crit. 3: 303 (1885).<br />
Testudina terrestris is characterized by its reticulately<br />
ridged ascospores, which readily distinguish it from other<br />
genera of Zopfiaceae (Hawksworth 1979). The species is<br />
usually associated with other fungi, or on the wood of Abies?<br />
and Pinus or on the fallen leaves of Taxus in Europe<br />
(Hawksworth and Booth 1974; Hawksworth1979).<br />
Tetraplosphaeria Kaz. Tanaka & K. Hirayama, Stud.<br />
Mycol. 64: 177 (2009).<br />
Type species: Tetraplosphaeria sasicola Kaz. Tanaka & K.<br />
Hirayama, Stud. Mycol. 64: 180 (2009).<br />
Tetraplosphaeria was introduced by Tanaka et al. (2009)<br />
to accommodate bambusicolous fungi with immersed to<br />
erumpent, globose to subglobose and smaller (mostly<<br />
300 μm) ascomata. The peridium is thin, and is composed<br />
of thin-walled cells of textura angularis. The pseudoparaphyses<br />
are cellular, and asci are fissitunicate, 8-spored,<br />
cylindrical to clavate with short pedicels. Ascospores are<br />
narrowly fusoid, hyaline and surrounded with a sheath.<br />
Species of Tetraplosphaeria have Tetraploa sensu stricto<br />
anamorphic stage, which is quite unique in Tetraplosphaeriaceae<br />
(Tanaka et al. 2009).<br />
Tingoldiago K. Hirayama & Kaz. Tanaka, Mycologia 102:<br />
740 (2010).<br />
Type species: Tingoldiago graminicola K. Hirayama &<br />
Kaz. Tanaka, Mycologia 102(3): 740 (2010).<br />
Tingoldiago is a genus of freshwater ascomycetes characterized<br />
by flattened, globose, immersed to erumpent ascomata,<br />
and numerous cellular pseudoparaphyses (Hirayama et al.<br />
2010). Asci are fissitunicate and cylindrical, and ascospores<br />
are 1-septate, which usually turn 3-septate and pale brown<br />
when old, usually with a sheath (Hirayama et al. 2010).<br />
Based on both morphology and multigene phylogenetic<br />
analysis, Tingoldiago should be treated as a synonym of<br />
Lentithecium (Shearer et al. 2009a; Zhang et al. 2009a).<br />
Tremateia Kohlm., Volkm.-Kohlm. & O.E. Erikss., Bot.<br />
Mar. 38: 165 (1995).<br />
Type species: Tremateia halophila Kohlm., Volkm.-Kohlm.<br />
& O.E. Erikss., Bot. Mar. 38: 166 (1995).<br />
Tremateia was introduced as a facultative marine genus<br />
which is characterized by depressed globose, immersed<br />
ascomata, numerous and cellular pseudoparaphyses, fissitunicate<br />
and clavate asci, ellipsoid muriform ascospores,<br />
and a Phoma-like anamorph (Kohlmeyer et al. 1995).<br />
These characters point Tremateia to Pleosporaceae<br />
(Kohlmeyer et al. 1995). DNA sequence based phylogenies<br />
placed T. halophila as sister to Bimuria novae-zelandiae<br />
in Montagnulaceae (Schoch et al. 2009; Suetrong et al.<br />
2009).<br />
Triplosphaeria Kaz. Tanaka & K. Hirayama, Stud. Mycol.<br />
64: 186 (2009).<br />
Type species: Triplosphaeria maxima Kaz. Tanaka & K.<br />
Hirayama, Stud. Mycol. 64: 188 (2009).<br />
Triplosphaeria was introduced as a bambusicolous genus<br />
characterized by immersed ascomata, numerous cellular
Fungal Diversity<br />
pseudoparaphyses, bitunicate, cylindrical to clavate asci with<br />
a short pedicel, fusoid, hyaline, 1-septate ascospores surrounded<br />
with a sheath, and with a Tetraploa-like anamorph<br />
(Tanaka et al. 2009). Together with Tetraplosphaeria,<br />
Pseudotetraploa, Quadricrura and Polyplosphaeria, Triplosphaeria<br />
was assigned to the Tetraplosphaeriaceae (Tanaka<br />
et al. 2009).<br />
Ulospora D. Hawksw., Malloch & Sivan., in Hawksworth,<br />
Can. J. Bot. 57: 96 (1979).<br />
Type species: Ulospora bilgramii (D. Hawksw., C. Booth<br />
& Morgan-Jones) D. Hawksw., Malloch & Sivan., Can. J.<br />
Bot. 57: 96 (1979).<br />
Ulospora was introduced as a monotypic genus to<br />
accommodate taxa of Testudinaceae whose ascospore has<br />
3–6 fissures (Hawksworth 1979). Genera of Testudinaceae<br />
are distinguished based on the morphology of ascospores,<br />
although the validity of this classification needs to be<br />
confirmed by molecular study. DNA sequence based<br />
phylogenies placed sequences from an unverified culture<br />
of U. bilgramii in a clade together with Verruculina enalia,<br />
and Lepidosphaeria nicotiae and it may have a close<br />
relationship to species in Platystomaceae (Mugambi and<br />
Huhndorf 2009b; Schoch et al. 2009; Plate 1).<br />
Zopfia Rabenh., Fungi europ. exsicc.: no. 1734 (1874).<br />
Type species: Zopfia rhizophila Rabenh., Fungi europ.<br />
exsicc.: no. 1734 (1874).<br />
Zopfia was introduced by Rabenhorst (1874) as a<br />
monotypic genus (typified by Z. rhizophila), and it was<br />
assigned to the Perisporiaceae by Saccardo (1882) and<br />
Winter (1884). Arnaud (1913) described the Zopfiaceae to<br />
accommodate Zopfia, and considered that it should be<br />
excluded from the Perisporiaceae. A relatively broad<br />
generic concept was accepted by Hawksworth and Booth<br />
(1974), in which they take the ascospore size and<br />
ornamentation variation as criteria under generic rank<br />
classification, and they treat Celtidia, Lepidosphaeria,<br />
Marchaliella, Neotestudina, Pontoporeia, Pseudophaeotrichum,<br />
Rechingeriella, Richonia and Testudina as synonyms<br />
of Zopfia. A narrow generic concept was adopted by<br />
Hawksworth (1979), and Zopfia is characterized by 1-<br />
septate ascospores, which are apiculate at both ends,<br />
smooth-walled by light microscope, with minute irregular<br />
pitting by SEM, and larger than other species of Zopfia<br />
sensu Hawksworth and Booth (1974). Three species were<br />
accepted, viz. Z. albiziae Farr, Z. biturbinata (Dur. &<br />
Mont.) Malloch & Cain and Z. rhizophila, and they all<br />
occur on roots of plants (Hawksworth 1979). DNA<br />
sequences from an unverified culture of Zopfia rhizophila<br />
placed it in close proximity to species in Delitschiaceae<br />
without strong statistical support (Kruys et al. 2006; Schoch<br />
et al. 2009; Plate 1).<br />
Zopfiofoveola D. Hawksw., Can. J. Bot. 57: 98 (1979).<br />
Type species: Zopfiofoveola punctata (D. Hawksw. & C.<br />
Booth) D. Hawksw., Can. J. Bot. 57: 98 (1979).<br />
≡ Zopfia punctata D. Hawksw. & C. Booth, Mycol. Pap.<br />
153: 23 (1974).<br />
Zopfiofoveola was hesitantly separated from Zopfia as a<br />
monotypic new genus based on its evenly distributed<br />
ornamentation with pale minute pits readily visible under<br />
the light microscope, and the more elongate shape and less<br />
pronounced apical papilla than those of Zopfia (Hawksworth<br />
1979). The type specimen of this species however, cannot be<br />
redescribed, because “the type species is only known from a<br />
microscopic preparation obtained from earthworm excrements<br />
in Sweden” as has been mentioned by Hawksworth (1979).<br />
General discussion<br />
Molecular phylogenetic studies based on four to five genes<br />
indicate that 20 families should be included in <strong>Pleosporales</strong><br />
(Schoch et al. 2009; Shearer et al. 2009; Suetrong et al.<br />
2009; Tanaka et al. 2009; Zhang et al. 2009a). Together<br />
with five unverified families (marked with “?”), 26 families<br />
are currently assigned under <strong>Pleosporales</strong> (Table 4). The<br />
Phaeotrichaceae lacks pseudoparaphyses, has cleistothecial<br />
ascomata with long setae, and conspicuous ascospores with<br />
germ pores at each end. These characters do not agree with<br />
the current concept of <strong>Pleosporales</strong> (Zhang et al. 2009a),<br />
and therefore Phaeotrichaceae is excluded from <strong>Pleosporales</strong><br />
(Table 4).<br />
Families in <strong>Pleosporales</strong><br />
Based on LSU and SSU rDNA, RPB1, RPB2 and TEF1<br />
sequence analysis, Pleosporineae is emended, and in this<br />
study, seven families are tentatively included, i.e. Cucurbitariaceae,<br />
Didymellaceae, Didymosphaeriaceae, Dothidotthiaceae,<br />
Leptosphaeriaceae, Phaeosphaeriaceae and<br />
Pleosporaceae (Zhang et al. 2009a; Plate1). In this study,<br />
Massarineae was emended to accommodate another five<br />
families, viz. Lentitheciaceae, Massarinaceae, Montagnulaceae,<br />
Morosphaeriaceae, Trematosphaeriaceae. The subordinal<br />
affinity of other families remained undetermined.<br />
Most of the families accepted within <strong>Pleosporales</strong> received<br />
high bootstrap support (Plate 1). The characters used to<br />
define a family, however, do not appear to have clear cut<br />
boundaries, as the ascomatal and hamathecial characters also<br />
seem to be poorly defined in some families. For example,<br />
both trabeculate and cellular pseudoparaphyses coexist in the<br />
Amniculicolaceae. Pycnidiophora, a genus of Sporormiaceae,<br />
has cleistothecial ascomata with spherical asci irregularly<br />
arranged in it. Brown phragmosporous ascospores are<br />
reported in Amniculicolaceae, Leptosphaeriaceae, Lophios-
Fungal Diversity<br />
tomataceae, Melanommataceae, Montagnulaceae, Phaeosphaeriaceae<br />
and Pleosporaceae. Similarly muriform ascospores<br />
occur in Aigialaceae, Amniculicolaceae,<br />
Didymellaceae, Lophiostomataceae, Montagnulaceae, Pleosporaceae<br />
and Sporormiaceae. Anamorphsof<strong>Pleosporales</strong><br />
are also variable to a large degree at the family level. Both<br />
hyphomycetous and coelomycetous anamorphs co-exist in<br />
Didymellaceae, Melanommataceae or Pleosporaceae.<br />
Phoma and Phoma-like anamorphs exist in Didymellaceae,<br />
Leptosphaeriaceae, Phaeosphaeriaceae, Pleosporaceae and<br />
Melanommataceae (de Gruyter et al. 2009; Zhang et al.<br />
2009a). It is clear that some characters, e.g. cleistothecial or<br />
perithecial ascomata, shape, colour and septation of ascospores,<br />
shape or arrangement (regular or irregular) of asci, or<br />
even presence or absence of pseudoparaphyses have evolved<br />
on numerous occasions which make the use of morphological<br />
characters in segregating families complicated. It is<br />
therefore unclear with our present state of knowledge which<br />
characters are taxonomically important at the family level or<br />
whether a suit of characters are necessary to define a family.<br />
DNA sequence comparisons are essential in delineating<br />
these taxa in combination with other characters. It is hoped<br />
that additional characters, i.e. biochemical, genomic and<br />
subcellular will be used to further distinguish these groups<br />
into natural taxa. Below we discuss each of the families,<br />
their genera and their considered important characteristics.<br />
Aigialaceae Suetrong, Sakay., E.B.G. Jones, Kohlm.,<br />
Volkm.-Kohlm. & C.L. Schoch 2010<br />
The Aigialaceae was introduced by Suetrong et al.<br />
(2009) based on its carbonaceous ascomata without papilla,<br />
cylindrical asci with apical apparatus, trabeculate pseudoparaphyses<br />
and ascospores with a sheath. The type genus<br />
(Aigialus) of the Aigialaceae was previously incorporated<br />
within the Massariaceae (Lumbsch and Huhndorf 2007).<br />
Currently, three genera are assigned under Aigialaceae, viz.<br />
Ascocratera, Aigialus and Rimora (Suetrong et al. 2009).<br />
The genera included in Aigialaceae have a wide range of<br />
morphological variation, with very few shared features as<br />
mentioned above, but all are found in mangrove habitats<br />
(Suetrong et al. 2009). The ascospores, however, vary<br />
widely from having 1 to 3 transverse septa and being hyaline<br />
to muriformly septate and brown (Suetrong et al. 2009). It is<br />
still unclear which characters unify the family and therefore<br />
placement of unsequenced genera is difficult. Further<br />
molecular work is needed to better understand this family.<br />
Amniculicolaceae Yin. Zhang, C.L. Schoch, J. Fourn.,<br />
Crous & K.D. Hyde 2009<br />
Members of Amniculicolaceae form a well supported<br />
clade, and all are freshwater fungi which usually stain the<br />
woody substrate purple (Zhang et al. 2009a, c). Genera of<br />
Amniculicolaceae have ascomata with compressed papilla<br />
Table 4 Families currently accepted in <strong>Pleosporales</strong> (syn. Melanommatales)<br />
with included genera<br />
<strong>Pleosporales</strong> subordo. Pleosporineae<br />
?Cucurbitariaceae<br />
Cucurbitaria Gray<br />
Curreya Sacc.<br />
?Rhytidiella Zalasky<br />
Syncarpella Theiss. & Syd.<br />
Didymellaceae<br />
?Appendispora K.D. Hyde<br />
Didymella Sacc. ex D. Sacc.<br />
Didymosphaerella Cooke<br />
Leptosphaerulina McAlpine<br />
Macroventuria Aa<br />
?Platychora Petr.<br />
Didymosphaeriaceae<br />
Didymosphaeria Fuckel<br />
Phaeodothis Syd. & P. Syd.<br />
Dothidotthiaceae<br />
Dothidotthia Höhn.<br />
Leptosphaeriaceae<br />
Leptosphaeria Ces. & De Not.<br />
Neophaeosphaeria Câmara, M.E. Palm & A.W. Ramaley<br />
Phaeosphaeriaceae<br />
Barria Z.Q. Yuan<br />
?Bricookea M.E. Barr<br />
?Chaetoplea (Sacc.) Clem.<br />
?Eudarluca Speg.<br />
Entodesmium Reiss<br />
?Hadrospora Boise<br />
Lautitia S. Schatz<br />
Loratospora Kohlm. & Volkm.-Kohlm.<br />
Metameris Theiss. & Syd.<br />
Mixtura O.E. Erikss. & J.Z. Yue<br />
Nodulosphaeria Rabenh.<br />
Ophiobolus Reiss<br />
Ophiosphaerella Speg.<br />
Phaeosphaeria I. Miyake<br />
Phaeosphaeriopsis Câmara, M.E. Palm & A.W.<br />
Ramaley<br />
Pleoseptum A.W. Ramaley & M.E. Barr<br />
Setomelanomma M. Morelet<br />
Wilmia Dianese, Inácio & Dornelo-Silva<br />
Pleosporaceae<br />
Cochliobolus Drechsler<br />
Crivellia Shoemaker & Inderbitzin<br />
Decorospora Inderbitzin, Kohlm. & Volkm.-Kohlm.<br />
Extrawettsteinina M.E. Barr<br />
Lewia M.E. Barr & E.G. Simmons<br />
Macrospora Fuckel<br />
Platysporoides (Wehm.) Shoemaker & C.E. Babc.<br />
Pleospora Rabenh. ex Ces. & De Not.<br />
Pseudoyuconia Lar. N. Vasiljeva<br />
Pyrenophora Fr.<br />
Setosphaeria K.J. Leonard & Suggs<br />
<strong>Pleosporales</strong> subordo. Massarineae<br />
Lentitheciaceae<br />
Lentithecium K.D. Hyde, J. Fourn. & Yin. Zhang
Fungal Diversity<br />
Table 4 (continued)<br />
Katumotoa Kaz. Tanaka & Y. Harada<br />
Keissleriella Höhn.<br />
?Wettsteinina Höhn.<br />
Massarinaceae<br />
Byssothecium Fuckel<br />
Massarina Sacc.<br />
Saccharicola D. Hawksw. & O.E. Erikss.<br />
Montagnulaceae<br />
Bimuria D. Hawksw., Chea & Sheridan<br />
?Didymocrea Kowalsky<br />
Kalmusia Niessl<br />
Karstenula Speg.<br />
Letendraea Sacc.<br />
Montagnula Berl.<br />
Paraphaeosphaeria O.E. Erikss.<br />
Tremateia Kohlm., Volkm.-Kohlm. & O.E. Erikss.<br />
Morosphaeriaceae<br />
?Asteromassaria Höhn<br />
Helicascus Kohlm.<br />
Morosphaeria Suetrong, Sakay., E.B.G. Jones & C.L. Schoch<br />
Trematosphaeriaceae<br />
Falciformispora K.D. Hyde<br />
Halomassarina Suetrong, Sakay., E.B.G. Jones, Kohlm., Volkm.-Kohlm. & C.L.<br />
Schoch<br />
Trematosphaeria Fuckel<br />
Other families<br />
Aigialaceae<br />
Aigialus Kohlm. & S. Schatz<br />
Ascocratera Kohlm.<br />
Rimora Kohlm., Volkm.-Kohlm., Suetrong, Sakay. & E.B.G. Jones<br />
Amniculicolaceae<br />
Amniculicola Y. Zhang & K.D. Hyde<br />
Murispora Yin. Zhang, C.L. Schoch, J. Fourn., Crous & K.D. Hyde<br />
?Massariosphaeria (E. Müll.) Crivelli<br />
Neomassariosphaeria Yin. Zhang, J. Fourn. & K.D. Hyde<br />
?Arthopyreniaceae (Massariaceae)<br />
Arthopyrenia A. Massal.<br />
Dothivalsaria Petr.<br />
?Dubitatio Speg.<br />
Massaria De Not.<br />
Navicella Fabre<br />
Roussoëlla Sacc.<br />
?Roussoellopsis I. Hino & Katum.<br />
Delitschiaceae<br />
Delitschia Auersw.<br />
Ohleriella Earle<br />
Semidelitschia Cain & Luck-Allen<br />
?Diademaceae<br />
Clathrospora Rabenh.<br />
Comoclathris Clem.<br />
Diadema Shoemaker & C.E. Babc.<br />
Diademosa Shoemaker & C.E. Babc.<br />
Graphyllium Clem.<br />
Hypsostromataceae<br />
Hypsostroma Huhndorf<br />
Lindgomycetaceae<br />
Lindgomyces K. Hirayama, Kaz. Tanaka & Shearer 2010<br />
Lophiostomataceae<br />
Table 4 (continued)<br />
Lophiostoma Ces. & De Not.<br />
Melanommataceae<br />
?Astrosphaeriella Syd. & P. Syd. (Syn. Javaria)<br />
?Anomalemma Sivan.<br />
?Asymmetricospora J. Fröhl. & K.D. Hyde<br />
Bertiella (Sacc.) Sacc. & P. Syd.<br />
?Bicrouania Kohlm. & Volkm.-Kohlm.<br />
Byssosphaeria Cooke<br />
Calyptronectria Speg.<br />
?Caryosporella Kohlm.<br />
Herpotrichia Fuckel<br />
?Mamillisphaeria K.D. Hyde, S.W. Wong & E.B.G. Jones<br />
Melanomma Nitschke ex Fuckel<br />
Ohleria Fuckel<br />
Pseudotrichia Kirschst.<br />
Pleomassariaceae<br />
?Lichenopyrenis Calatayud, Sanz & Aptroot<br />
?Splanchnonema Corda<br />
?Peridiothelia D. Hawksw.<br />
Pleomassaria Speg.<br />
Sporormiaceae<br />
Chaetopreussia Locq.-Lin.<br />
Eremodothis Arx<br />
Pleophragmia Fuckel<br />
Preussia Fuckel<br />
Sporormia De Not.<br />
Westerdykella Stolk<br />
?Teichosporaceae<br />
Chaetomastia (Sacc.) Berl<br />
Immotthia M.E. Barr<br />
Loculohypoxylon M.E. Barr<br />
Sinodidymella J.Z. Yue & O.E. Erikss.<br />
Teichospora Fuckel<br />
Tetraplosphaeriaceae<br />
Polyplosphaeria Kaz. Tanaka & K. Hirayama<br />
Tetraplosphaeria Kaz. Tanaka & K. Hirayama<br />
Triplosphaeria Kaz. Tanaka & K. Hirayama<br />
?Zopfiaceae (syn Testudinaceae)<br />
Caryospora De Not.<br />
Celtidia J.M. Janse<br />
?Coronopapilla Kohlm. & Volkm.-Kohlm.<br />
Halotthia Kohlm.<br />
Lepidosphaeria Parg.-Leduc<br />
?Mauritiana Poonyth, K.D. Hyde, Aptroot & Peerally<br />
Pontoporeia Kohlm.<br />
?Rechingeriella Petr.<br />
Richonia Boud.<br />
Testudina Bizz.<br />
Verruculina Kohlm. & Volkm.-Kohlm.<br />
Ulospora D. Hawksw., Malloch & Sivan.<br />
Zopfia Rabenh.<br />
Zopfiofoveola D. Hawksw.<br />
<strong>Pleosporales</strong> genera incertae sedis<br />
Acrocordiopsis Borse & K.D. Hyde<br />
Aglaospora De Not.<br />
Anteaglonium Mugambi & Huhndorf<br />
Ascorhombispora L. Cai & K.D. Hyde<br />
Atradidymella Davey & Currah
Fungal Diversity<br />
Table 4 (continued)<br />
Belizeana Kohlm. & Volkm.-Kohlm.<br />
Biatriospora K.D. Hyde & Borse<br />
Byssolophis Clem.<br />
Carinispora K.D. Hyde<br />
Cilioplea Munk<br />
Decaisnella Fabre<br />
Epiphegia Nitschke ex G.H. Otth<br />
Isthmosporella Shearer & Crane<br />
Julella Fabre<br />
Lineolata Kohlm. & Volkm.-Kohlm.<br />
Lophiella Sacc.<br />
Lophionema Sacc.<br />
Lophiotrema Sacc.<br />
Moristroma A.I. Romero & Samuels<br />
Neotestudina Segretain & Destombes<br />
Ostropella (Sacc.) Höhn.<br />
Paraliomyces Kohlm.<br />
Passeriniella Berl.<br />
Quintaria Kohlm. & Volkm.-Kohlm.<br />
Saccothecium Fr.<br />
Salsuginea K.D. Hyde<br />
Shiraia P. Henn.<br />
Xenolophium Syd.<br />
Family excluded<br />
Phaeotrichaceae<br />
Echinoascotheca Matsush.<br />
Phaeotrichum Cain & M.E. Barr<br />
Trichodelitschia Munk<br />
Genera excluded<br />
Kriegeriella Höhn.<br />
Muroia I. Hino & Katum.<br />
Zeuctomorpha Sivan., P.M. Kirk & Govindu<br />
and cylindrical to cylindro-clavate asci. Neomassariosphaeria<br />
typhicola was traditionally assigned to Massariosphaeria (as<br />
M. typhicola), and Massariosphaeria is characterized by<br />
staining the woody substrate purple (Crivelli 1983;<br />
Leuchtmann 1984). Eriksson (1981 p. 135) had pointed out<br />
that “Purple-staining species of Pleospora, treated by Webster<br />
(1957), are not congeneric with P. herbarum (Eriksson 1967b:<br />
13), and certainly do not even belong to the Pleosporaceae”.<br />
This is mirrored in Murispora rubicunda, a<br />
previous Pleospora species (as P. rubicunda) staining the<br />
woody substrate purple, closely related to the Amniculicolaceae<br />
in a subsequent phylogenetic study (Zhang et al.<br />
2009a). The anamorphs of this family are possibly<br />
Anguillospora longissima, Spirosphaera cupreorufescens<br />
and Repetophragma ontariense (Zhang et al. 2009a).<br />
?Arthopyreniaceae (or Massariaceae) W. Watson 1929<br />
The Arthopyreniaceae was introduced as a lichenized<br />
family of Pyrenocarpales, which comprises Acrocordia,<br />
Arthopyrenia, Athrismidium, Bottaria, Celothelium, Laurera,<br />
Leptorhaphis, Microthelia, Microtheliopsis, Polyblastiopsis,<br />
Pseudosagedia, Raciborskiella and Tomasellia<br />
(Watson 1929). Subsequently, Arthopyreniaceae was<br />
assigned under Dothideales (suborder Pseudosphaeriineae)<br />
(von Arx and Müller 1975). The generic type of Massaria<br />
(M. inquinans) and Torula herbarum and Arthopyrenia<br />
salicis together with members of Roussoella as well as<br />
Roussoellopsis form a robust clade, which makes their<br />
familial placement uncertain (Massariaceae or Arthopyreniaceae)<br />
(Schoch et al. 2009; Zhang et al. 2009a).<br />
?Cucurbitariaceae G. Winter 1885<br />
The Cucurbitariaceae is characterized by its aggregated<br />
ascomata which form from a basal stromatic structure,<br />
ostiolate, fissitunicate and cylindrical asci, and pigmented,<br />
phragmosporous or muriform ascospores (Cannon and Kirk<br />
2007). Currently, no molecular study has been able to<br />
resolve its ordinal status, but some characters are similar to<br />
Leptosphaeriaceae or Phaeosphaeriaceae (Cannon and<br />
Kirk 2007). Cucurbitaria elongata clustered within <strong>Pleosporales</strong><br />
(Schoch et al. 2006).<br />
Delitschiaceae M.E. Barr 2000<br />
The Delitschiaceae was established to accommodate<br />
some species of the Sporormiaceae, which is characterized<br />
by its ascomata with periphysate ostioles, ocular chamber<br />
surrounded by a dome and usually in having four refractive<br />
rods, ascospores with or without a septum, having a germ<br />
slit in each cell and being surrounded by a mucilaginous<br />
sheath (Barr 2000). Species of the Delitschiaceae are<br />
hypersaprotrophic on old dung or exposed wood (Barr<br />
2000). Based on a molecular phylogenetic studies, Delitschia<br />
didyma and D. winteri form a robust clade basal to<br />
other pleosporalean fungi (Schoch et al. 2009; Zhang et al.<br />
2009a). The familial status of two other genera, Ohleriella<br />
and Semidelitschia, remains undetermined.<br />
?Diademaceae Shoemaker & C.E. Babc. 1992<br />
The Diademaceae was introduced by Shoemaker and<br />
Babcock (1992) based on its ascomata opening as a flat<br />
circular lid and bitunicate asci, ascospores are fusiform,<br />
brown, mostly applanate, and having three or more<br />
transverse septate and with or lacking longitudinal septa<br />
and usually having a sheath. Five genera had been included<br />
viz. Clathrospora, Comoclathris, Diadema, Diademosa and<br />
Macrospora (Shoemaker and Babcock 1992).<br />
Didymellaceae Gruyter, Aveskamp & Verkley 2009<br />
The generic type of Didymella (D. exigua) together with<br />
some Phoma or Phoma-related species form a robust<br />
familial clade on the phylogenetic tree, thus the Didymellaceae<br />
was introduced to accommodate them (de Gruyter et<br />
al. 2009). Subsequently, Didymellaceae was assigned to
Fungal Diversity<br />
Pleosporineae (suborder of <strong>Pleosporales</strong>) (Zhang et al.<br />
2009a). A detailed study was conducted on the Didymellaceae<br />
based on LSU, SSU rDNA, ITS as well as β-tubulin,<br />
which indicated that many Phoma or Phoma-related<br />
species/fungi reside in this clade of the Didymellaceae<br />
(Aveskamp et al. 2010).<br />
Didymosphaeriaceae Munk 1953<br />
The Didymosphaeriaceae was introduced by Munk (1953),<br />
and was revived by Aptroot (1995) based on its distoseptate<br />
ascospores and trabeculate pseudoparaphyses, mainly anastomosing<br />
above the asci. The familial status of the Didymosphaeriaceae<br />
is debatable, and Lumbsch and Huhndorf (2007)<br />
assigned it to the Montagnulaceae, while von Arx and Müller<br />
(1975) treated it as a synonym of the Pleosporaceae. Inthis<br />
study, Didymosphaeria futilis (the generic type of Didymosphaeria)<br />
is closely related to the Cucurbitariaceae (Plate 1).<br />
Herein, we accept it as a separate family containing three genera,<br />
namely Appendispora, Didymosphaeria and Phaeodothis. More<br />
information could only be obtained by further molecular work<br />
based on correctly identified strains.<br />
Dothidotthiaceae Crous & A.J.L. Phillips 2008<br />
Dothidotthiaceae was introduced to accommodate the<br />
single genus Dothidotthia, which is characterized by gregarious,<br />
erumpent, globose ascomata, hyaline, septate pseudoparaphyses,<br />
8-spored, bitunicate, clavate asci, ellipsoid, 1-<br />
septate ascospores, and has anamorphic Thyrostroma<br />
(Phillips et al. 2008). In this study, Dothidotthiaceae is<br />
closely related to Didymellaceae, but it is still treated as a<br />
separate family (Plate 1).<br />
Hypsostromataceae Huhndorf 1994<br />
Hypsostromataceae was introduced based on two tropical<br />
genera (i.e. Hypsostroma and Manglicola), which have superficial,<br />
large, elongate ascomata with a soft-textured, pseudoparenchymatic<br />
wall, trabeculate pseudoparaphyses and stipitate asci<br />
attached in a basal arrangement in the centrum; asci with an<br />
apical chamber and fluorescing ring; and fusiform, septate<br />
ascospores (Huhndorf 1994). Hypsostromataceae was assigned<br />
to Melanommatales sensu Barr (Huhndorf 1994). In a<br />
subsequent phylogenetic study, Hypsostromataceae was recovered<br />
as a strongly supported monophyletic group nested within<br />
<strong>Pleosporales</strong> (Mugambi and Huhndorf 2009b).<br />
Lentitheciaceae Yin. Zhang, C.L. Schoch, J. Fourn., Crous<br />
& K.D. Hyde 2009<br />
Phylogenetic analysis based on multi-genes indicate that<br />
freshwater taxa, e.g. Lentithecium fluviatile, L. arundinaceum,<br />
Stagonospora macropycnidia, Wettsteinina lacustris,<br />
Keissleriella cladophila, Katumotoa bambusicola and<br />
Ophiosphaerella sasicola form a well supported clade, which<br />
most likely represent a familial rank (Zhang et al. 2009a).<br />
Their morphology, however, varies widely, e.g. ascomata<br />
small- to medium-sized, ascospores fusoid to filliform,<br />
hyaline to pale yellow, 1- to multi-septate (Zhang et al.<br />
2009a). In particular, they are saprobic on monocotyledons<br />
or dicotyledons. Currently, no conspicuous, unique morphological<br />
character has been noted in Lentitheciaceae, which<br />
makes it difficult to recognize based on morphology.<br />
Leptosphaeriaceae M.E. Barr 1987a<br />
The Leptosphaeriaceae was introduced by Barr (1987a)<br />
based on Leptosphaeria. The familial status of the Leptosphaeriaceae<br />
is subsequently supported by molecular phylogenetic<br />
studies, in which members of the Leptosphaeriaceae<br />
form a paraphyletic clade with moderate bootstrap support<br />
(Dong et al. 1998; de Gruyter et al. 2009;Schochetal.2009;<br />
Zhang et al. 2009a). Coniothyrium palmarum, the generic<br />
type of Coniothyrium nested within this family (de Gruyter<br />
et al. 2009). Further molecular phylogenetic study is needed,<br />
in which more related taxa are included.<br />
Lindgomycetaceae K. Hirayama, Kaz. Tanaka & Shearer 2010<br />
Lindgomycetaceae was introduced as a monotypic<br />
family represented by Lindgomyces (Hirayama et al.<br />
2010). Lindgomycetaceae is another freshwater family in<br />
<strong>Pleosporales</strong>, which is characterized by its subglobose to<br />
globose, ostiolate and papillate ascomata, numerous, septate,<br />
branching and anastomosing pseudoparaphyses, fissitunicate,<br />
cylindrical to clavate, 8-spored asci, fusiform to<br />
cylindrical, uni- to multiseptate, hyaline to brown ascospores<br />
usually covered with an entire sheath and/or bipolar<br />
mucilaginous appendages (Hirayama et al. 2010).<br />
Lophiostomataceae Sacc. 1883<br />
The Lophiostomataceae had been characterized by its<br />
slot-like ostiole on the top of a flattened neck (Holm and<br />
Holm 1988). Based on this, 11 genera were assigned under<br />
the Lophiostomataceae, viz. Byssolophis, Cilioplea, Entodesmium,<br />
Herpotrichia, Lophiella, Lophionema, Lophiostoma,<br />
Lophiotrema, Massariosphaeria, Muroia and<br />
Quintaria (Holm and Holm 1988). The Lophiostomataceae<br />
was thought to be heterogeneous, as the “papilla form is an<br />
unstable and highly adaptive character” (Holm and Holm<br />
1988). Most recent phylogenetic analysis support the<br />
monophyletic status of the Lophiostomataceae sensu stricto<br />
(which tends to comprise a single genus of Lophiostoma)<br />
(Zhang et al. 2009a, b). The familial placement of other<br />
genera, however, remains unresolved.<br />
Massarinaceae Munk 1956<br />
The Massarinaceae was established based on Keissleriella,<br />
Massarina, Metasphaeria, Pseudotrichia and Trichometasphaeria<br />
(Munk 1956). Subsequently, the Massarinaceae is<br />
sometimes treated as a synonym of Lophiostomataceae (Barr
Fungal Diversity<br />
1987b). Based on a multigene phylogenetic study, the generic<br />
type of Massarina (M. eburnea) together with M. cisti,<br />
Neottiosporina paspali and Byssothecium circinans form a<br />
well supported clade (Zhang et al. 2009a, b). It seems that a<br />
relatively narrow familial concept should be accepted.<br />
Melanommataceae G. Winter 1885<br />
The traditional circumscription of the Melanommataceae<br />
was based on its globose or depressed perithecial ascomata,<br />
bitunicate and fissitunicate asci, pigmented phragmosporous<br />
ascospores as well as the trabeculate pseudoparaphyses<br />
(Barr 1990a; Sivanesan 1984). However, the family has<br />
recently proved polyphyletic (Liew et al. 2000; Kodsueb et<br />
al. 2006a; Kruys et al. 2006; Wang et al. 2007). Bimuria,<br />
Ostropella, Trematosphaeria and Xenolophium occur outside<br />
Melanommataceae (Mugambi and Huhndorf 2009b;<br />
Zhang et al. 2009a). Species of Byssosphaeria, Bertiella,<br />
Herpotrichia, Pseudotrichia, Pleomassaria as well as<br />
Melanomma resided in the clade of Melanommataceae<br />
(Mugambi and Huhndorf 2009b; Schoch et al. 2009; Zhang<br />
et al. 2009a). The familial status of many genera previously<br />
listed under this family remains to be sorted out (Lumbsch<br />
and Huhndorf 2007).<br />
Montagnulaceae M.E. Barr 2001<br />
The Montagnulaceae was introduced to accommodate<br />
some pleosporalean genera with ascomata immersed under<br />
a clypeus, a pseudoparenchymatous peridium with small<br />
cells, cylindric or oblong asci with pedicels and brown<br />
ascospores (Barr 2001). Three genera were included, i.e.<br />
phragmosporous Kalmusia, dictyosporous Montagnula and<br />
didymosporous Didymosphaerella (Barr 2001). Our molecular<br />
phylogenetic analysis based on multi-genes indicated<br />
that species from Kalmusia, Phaeosphaeria, Bimuria,<br />
Didymocrea, Paraphaeosphaeria, Karstenula, Letendraea<br />
as well as Montagnula resided in the monophylogenetic<br />
clade of the Montagnulaceae (Schoch et al. 2009; Zhang et<br />
al. 2009a).<br />
Morosphaeriaceae Suetrong, Sakay., E.B.G. Jones & C.L.<br />
Schoch 2009<br />
Four marine species, viz. Massarina ramunculicola<br />
(as Morosphaeria ramunculicola), Massarina velataspora<br />
(Morosphaeria velataspora), Helicascus kanaloanus and<br />
H. nypae together with the freshwater species Kirschsteiniothelia<br />
elaterascus form a well supported clade, which<br />
most likely represent a familial rank (Suetrong et al.<br />
2009). Thus, Morosphaeriaceae was introduced to accommodate<br />
these taxa (Suetrong et al. 2009). In<br />
this study, Asteromassaria pulchra is basal to other<br />
species of Morosphaeriaceae, and gets well support<br />
(Plate 1). Thus we tentatively assign Asteromassaria under<br />
Morosphaeriaceae.<br />
Phaeosphaeriaceae M.E. Barr 1979a<br />
The Phaeosphaeriaceae was introduced to accommodate<br />
some pleosporalean genera which have saprobic, parasitic or<br />
hyperparasitic lifestyles and have small- to medium-sized,<br />
subglobose or conical ascomata, bitunicate asci and hyaline or<br />
pigmented ascospores with or without septation (Barr 1979a).<br />
Fourteen genera were included, viz. Comoclathris, Didymella,<br />
Eudarluca, Heptameria, Leptosphaeria, Loculohypoxylon,<br />
Metameris, Microthelia, Nodulosphaeria, Ophiobolus, Paraphaeosphaeria,<br />
Rhopographus, Scirrhodothis and Teichospora<br />
(Barr 1979a), which were subsequently assigned to<br />
various families, such as Loculohypoxylon and Teichospora to<br />
the Teichosporaceae, Paraphaeosphaeria to the Montagnulaceae,<br />
Leptosphaeria to the Leptosphaeriaceae, Comoclathris<br />
to the Diademaceae, Didymella to the Didymellaceae and<br />
Heptameria and Rhopographus to genera incertae sedis of<br />
Dothideomycetes (Aveskamp et al. 2010; de Gruyter et al.<br />
2009; Lumbsch and Huhndorf 2007; Zhang et al. 2009a).<br />
Based on multi-gene phylogenetic analysis, a relatively narrow<br />
familial concept is accepted, which is mostly associated with<br />
monocotyledons, with perithecoid, small- to medium-sized<br />
ascomata, and septate ascospores which are fusiform to<br />
filliform (Zhang et al. 2009a). Four genera were accepted,<br />
Ophiosphaerella, Phaeosphaeria, Entodesmium and Setomelanomma<br />
(Zhang et al. 2009a). Together with Cucurbitariaceae,<br />
Didymellaceae, Didymosphaeriaceae, Dothidotthiaceae, Leptosphaeriaceae<br />
and Pleosporaceae, thePhaeosphaeriaceae is<br />
assigned under Pleosporineae (Zhang et al. 2009a).<br />
Pleomassariaceae M.E. Barr 1979a<br />
Both Asteromassaria and Splanchnonema were designated<br />
as representative genera of Pleomassariaceae (Barr<br />
1979a). Currently, four genera are included in Pleomassariaceae,<br />
viz. ?Lichenopyrenis, ?Splanchnonema, ?Peridiothelia<br />
and Pleomassaria (Table 4). The generic type of<br />
Pleomassaria (P. siparia) clustered with species of Melanommataceae<br />
in previous and present studies (Schoch et al.<br />
2009; Zhang et al. 2009a; Plate 1). Zhang et al. (2009a) has<br />
attempted to assign Pleomassariaceae to Melanommataceae<br />
(Zhang et al. 2009a). Based on the distinct morphology and<br />
anamorphic stage of Pleomassaria siparia as well as the<br />
divergence of dendrogram, we hesitantely reinstate Pleomassariaceae<br />
as a separate family in this study.<br />
Pleosporaceae Nitschke 1869<br />
The Pleosporaceae is one of the earliest introduced<br />
families in Dothideomycetes. The Pleosporaceae was<br />
originally assigned under Sphaeriales, which accommodated<br />
species with paraphyses and immersed perithecia (Ellis<br />
and Everhart 1892; Lindau 1897; Winter 1887). Subsequently,<br />
many of the Pleosporaceae species were transferred<br />
to the Pseudosphaeriaceae, which was subsequently<br />
elevated to ordinal rank as Pseudosphaeriales (Theissen
Fungal Diversity<br />
and Sydow 1918). Luttrell (1955) introduced the <strong>Pleosporales</strong><br />
(lacking a Latin description), which is characterized by<br />
its Pleospora-type of centrum development. Based on this,<br />
the Pleosporaceae and the Lophiostomataceae as well as<br />
other five families were placed in <strong>Pleosporales</strong> (Luttrell<br />
1955). Pleosporaceae is the largest and most typical family<br />
in <strong>Pleosporales</strong>. Wehmeyer (1975) stated that the Pleosporatype<br />
centrum development is verified in a small number of<br />
genera, and centrum development in the majority of genera<br />
is unknown; thus the placement of families or genera is quite<br />
arbitrary. In addition, the circumscription of Pleosporaceae<br />
is not clear-cut, and “……ascostromata of many different<br />
types, which are previously placed in various other families<br />
(Trichosphaeriaceae, Melanommataceae, Cucurbitariaceae,<br />
Amphisphaeriaceae etc.) are to be found here” (Wehmeyer<br />
1975). Thus, the heterogeneous nature of <strong>Pleosporales</strong> is<br />
obvious (Eriksson 1981), and had been confirmed by<br />
subsequent molecular phylogenetic studies (e.g. Kodsueb et<br />
al. 2006a). Based on the multi-gene phylogenetic analysis,<br />
some species from Lewia, Cochliobolus, Pleospora, Pyrenophora<br />
and Setosphaeria resided in the Pleosporaceae<br />
(Zhang et al. 2009a).<br />
Sporormiaceae Munk 1957<br />
The Sporormiaceae is the largest coprophilous family in<br />
<strong>Pleosporales</strong>, which bears great morphological variation.<br />
Ascomata vary from cleistothecoid to perithecoid, asci are<br />
regularly or irregularly arranged, clavate or spherical,<br />
ascospores with or without germ slits or ornamentations.<br />
Based on phylogenetic analysis, Sporormiaceae is most<br />
likely monophyletic as currently circumscribed (Kruys et al.<br />
2006; Kruys and Wedin 2009).<br />
?Teichosporaceae M.E. Barr 2002<br />
The Teichosporaceae was introduced by segregating some<br />
non-lichenized members of the Dacampiaceae which are<br />
apostrophic on woody stems and periderm or hypersaprotrophic<br />
on other ascomycetous fungi (Barr 2002). The Dacampiaceae<br />
together with its synonym, Pyrenidiaceae was only<br />
maintained to accommodate its lichenicolous genera (Barr<br />
2002). This proposal does not have any molecular phylogenetic<br />
support.<br />
Tetraplosphaeriaceae Kaz. Tanaka & K. Hirayama 2009<br />
The Tetraplosphaeriaceae was introduced to accommodate<br />
five genera, i.e. Tetraplosphaeria, Triplosphaeria,<br />
Polyplosphaeria and the anamorphic genera Pseudotetraploa<br />
and Quadricrura (Tanaka et al. 2009). The Tetraplosphaeriaceae<br />
is characterized by its Massarina-like<br />
teleomorphs and its Tetraploa-like anamorphs with setaelike<br />
appendages, and its monophylogenetic status has been<br />
recently confirmed based on DNA phylogenetic studies<br />
(Tanaka et al. 2009).<br />
Trematosphaeriaceae<br />
Three species, viz. Falciformispora lignatilis, Halomassarina<br />
thalassiae and Trematosphaeria pertusa form a<br />
robust clade, which forms a sister group with other<br />
pleosporalean families (Schoch et al. 2009; Suetrong et al.<br />
2009). Trematosphaeriaceae is waiting to be formally<br />
proposed (Suetrong et al. data unpublished).<br />
?Zopfiaceae G. Arnaud ex D. Hawksw. 1992<br />
The Zopfiaceae was introduced by Arnaud (1913), but<br />
was invalid due to the lack of a Latin diagnosis (see<br />
comments by Eriksson and Hawksworth 1992). The Zopfiaceae<br />
was formally introduced by Eriksson and Hawksworth<br />
(1992), and is characterized by its cleistothecial ascomata,<br />
thick-walled peridium, globose or saccate asci and oneseptate,<br />
dark brown ascospores (Cannon and Kirk 2007).<br />
Currently, eleven genera are included, but the family is likely<br />
polyphyletic (Kruys et al. 2006).<br />
Excluded family<br />
Phaeotrichaceae Cain 1956<br />
The cleistothecioid ascomata, ascospores with germ pore<br />
at each end and the absence of pseudoparaphyses indicate<br />
that the Phaeotrichaceae may not be closely related to<br />
<strong>Pleosporales</strong>. This was confirmed by DNA based phylogenies<br />
(Schoch et al. 2009). Thus, we exclude it from<br />
<strong>Pleosporales</strong>.<br />
Final remarks<br />
Problems and concerns<br />
Recently, many new pleosporalean lineages from freshwater<br />
(Shearer et al. 2009; Zhang et al. 2009a), marine<br />
(Suetrong et al. 2009) or from bambusicolous hosts (Tanaka<br />
et al. 2009) have been reported. In particular, large-scale<br />
phylogenetic analysis indicate that numerous unresolved<br />
clades still exist, which may also indicate that a large<br />
number of fungal lineages are not resolved. As has been<br />
estimated, 95% of all fungi are unreported (Hawksworth<br />
1991), and a large portion of them might exist only as<br />
hyphae (or DNA-only fungi, Taylor 1993). Under the<br />
influence of human activities, environmental situations are<br />
changing quickly, which may result in numerous fungal<br />
taxa losing their habitats and/or become endangered. More<br />
field work is urgently needed.<br />
A future polyphasic approach to study <strong>Pleosporales</strong><br />
The use of DNA sequence comparisons have proved<br />
invaluable in modern concepts of fungal taxonomy. It is now<br />
clear many fungi do not produce reproductive structures or
Fungal Diversity<br />
only do so under very rare circumstances and many fungi<br />
cannot be cultured (Begerow et al. 2010). More and more<br />
morpho-species have proven to be cryptic taxa, and already a<br />
large percentage of fungal diversity is documented only by<br />
DNA sequences. DNA sequence analysis is an essential way<br />
to resolve these problems. But are they enough for fully<br />
informed fungal taxonomy? Each single morphological<br />
character may be the outcome of the expression of one to<br />
numerous genes, which might be composed of thousands of<br />
base pairs. DNA barcoding methods are “a breakthroughfor<br />
identification, but they will not supplant the need to<br />
formulate and rigorously test species hypothesis” (Wheeler<br />
et al. 2004). Thus, integration of classical morphological<br />
approaches and DNA and protein based sequence comparisons<br />
are critical to produce a modern taxonomy that reflects<br />
evolutionary similarities and differences (DeSalle et al. 2005;<br />
Godfray 2002). In particular, the advent of comparative<br />
genomics and advances in our understanding of secondary<br />
metabolites and host or habitat spectra allow the possibility<br />
to tie phylogenetic hypotheses derived from DNA and<br />
protein sequence to the biology of the organisms. (Bitzer et<br />
al. 2008; Stajich et al. 2009; Zhang et al. 2009a, b).<br />
Acknowledgement We are grateful to the Directors and Curators of<br />
the following herbaria for loan of specimens in their keeping: BAFC,<br />
BISH, BPI, BR, BRIP, <strong>CBS</strong>, E, ETH, FFE, FH, G, H, Herb. J.<br />
Kohlmeyer, HHUF, IFRD, ILLS, IMI, K(M), L, LPS, M, MA, NY,<br />
PAD, PC, PH, RO, S, TNS, TRTC, UB, UBC, UPS and ZT; to Dr. L.<br />
Cai, Dr. A.J.L. Phillips, Dr. C. Shearer and some other mycologists for<br />
their permission to use or refer to their published figures, to J.K. Liu,<br />
H. Zhang, Y.L. Yang and J. Fournier for helping me loan or collect<br />
specimens, to H. Leung for technical help. The third coauthor<br />
acknowledges the Intramural Research Program of the NIH, National<br />
Library of Medicine. The Global Research Network and King Saud<br />
University are also thanked for support.<br />
Open Access This article is distributed under the terms of the<br />
Creative Commons Attribution Noncommercial License which permits<br />
any noncommercial use, distribution, and reproduction in any<br />
medium, provided the original author(s) and source are credited.<br />
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