Pleosporales - CBS - KNAW

Fungal Diversity 

DOI 10.1007/s13225-011-0117-x 

Pleosporales 

Ying Zhang & Pedro W. Crous & Conrad L. Schoch & 

Kevin D. Hyde 

Received: 31 March 2011 /Accepted: 13 June 2011 

# The Author(s) 2011. This article is published with open access at Springerlink.com 

Abstract One hundred and five generic types of Pleosporales 

are described and illustrated. A brief introduction and detailed 

history with short notes on morphology, molecular phylogeny 

as well as a general conclusion of each genus are provided. 

For those genera where the type or a representative specimen 

is unavailable, a brief note is given. Altogether 174 genera of 

Pleosporales are treated. Phaeotrichaceae as well as Kriegeriella, 

Zeuctomorpha and Muroia are excluded from 

Pleosporales. Based on the multigene phylogenetic analysis, 

the suborder Massarineae is emended to accommodate five 

families, viz. Lentitheciaceae, Massarinaceae, Montagnulaceae, 

Morosphaeriaceae and Trematosphaeriaceae. 

Y. Zhang 

Division of Microbiology, School of Biological Sciences, 

The University of Hong Kong, 

Pokfulam Road, 

Hong Kong, SAR, People’s Republic of China 

P. W. Crous 

CBS-KNAW Fungal Biodiversity Centre, 

P.O. Box 85167, 3508 AD Utrecht, The Netherlands 

C. L. Schoch 

National Center for Biotechnology Information, 

National Library of Medicine, National Institutes of Health, 

45 Center Drive, MSC 6510, 

Bethesda, MD 20892-6510, USA 

K. D. Hyde 

School of Science, Mae Fah Luang University, 

Tasud, Muang, 

Chiang Rai 57100, Thailand 

K. D. Hyde (*) 

Botany and Microbiology Department, College of Science, 

King Saud University, 

Riyadh 11442, Saudi Arabia 

e-mail: kdhyde3@gmail.com 

Keywords Generic type . Massarineae . Molecular 

phylogeny. Morphology . Pleosporales . Taxonomy 

Introduction 

Historic overview of Pleosporales 

Pleosporales is the largest order in the Dothideomycetes, 

comprising a quarter of all dothideomycetous species (Kirk 

et al. 2008). Species in this order occur in various habitats, 

and can be epiphytes, endophytes or parasites of living 

leaves or stems, hyperparasites on fungi or insects, lichenized, 

or are saprobes of dead plant stems, leaves or bark 

(Kruys et al. 2006; Ramesh 2003). 

The Pleosporaceae was introduced by Nitschke (1869), 

and was assigned to Sphaeriales based on immersed 

ascomata and presence of pseudoparaphyses (Ellis and 

Everhart 1892; Lindau 1897; Wehmeyer 1975; Winter 

1887). Taxa in this family were then assigned to Pseudosphaeriaceae 

(Theissen and Sydow 1918; Wehmeyer 1975). 

Pseudosphaeriales, represented by Pseudosphaeriaceae, 

was introduced by Theissen and Sydow (1918), and was 

distinguished from Dothideales by its uniloculate, perithecioid 

ascostromata. Subsequently, the uni- or pluri-loculate 

ascostromata was reported to be an invalid character to 

separate members of Dothideomycetes into different orders 

(Luttrell 1955). In addition, the familial type of Pseudosphaeriales 

together with its type genus, Pseudosphaeria,was 

transferred to Dothideales, thusPseudosphaeriales became a 

synonym of Dothideales. The name “Pseudosphaeriales” 

has been applied in different senses, thus Pleosporales (as an 

invalid name due to the absence of a Latin diagnosis) was 

proposed by Luttrell (1955) to replace the confusing name, 

Pseudosphaeriales, which included seven families, i.e.


Botryosphaeriaceae, Didymosphaeriaceae, Herpotrichiellaceae, 

Lophiostomataceae, Mesnieraceae, Pleosporaceae 

and Venturiaceae. Müller and von Arx (1962) however, 

reused Pseudosphaeriales with 12 families included, viz. 

Capnodiaceae, Chaetothyriaceae, Dimeriaceae, Lophiostomataceae, 

Mesnieraceae, Micropeltaceae, Microthyriaceae, 

Mycosphaerellaceae, Pleosporaceae, Sporormiaceae, Trichothyriaceae 

and Venturiaceae. 

Familial circumscriptions of the Pleosporales were based 

on characters of ascomata, morphology of asci and their 

arrangement in locules, presence and type of hamathecium, 

shape of papilla or ostioles, morphology of ascospores and 

type of habitats (Luttrell 1973) (Table1). Based on these 

characters, Luttrell (1973) included eight families, i.e. 

Botryosphaeriaceae, Dimeriaceae, Lophiostomataceae, Mesnieraceae, 

Mycoporaceae, Pleosporaceae, Sporormiaceae 

and Venturiaceae in Pleosporales. In their review of bitunicate 

ascomycetes, von Arx and Müller (1975) accepted only a 

single order, Dothideales, with two suborders, i.e. Dothideineae 

(including Atichiales, Dothiorales, Hysteriales and 

Myriangiales) and Pseudosphaeriineae (including Capnodiales, 

Chaetothyriales, Hemisphaeriales, Lophiostomatales, 

Microthyriales, Perisporiales, Pleosporales, Pseudosphaeriales 

and Trichothyriales). This proposal has however, rarely 

been followed. Three existing families, i.e. Lophiostomataceae, 

Pleosporaceae and Venturiaceae plus 11 other families 

were accepted in Pleosporales as arranged by Barr (1979a) 

(largely using Luttrell’s concepts, Table 1), and she assigned 

these families to six suborders. The morphology of pseudoparaphyses 

was given much prominence at the ordinal level in 

this classification (Barr 1983). In particular the Melanommatales 

was introduced to accommodate taxa with trabeculate 

pseudoparaphyses (Sporormia-type centrum development) 

(Barr 1983), distinguished from cellular pseudoparaphyses 

(Pleospora-type centrum development) possessed by members 

of Pleosporales sensu Barr. The order Melanommatales 

included Didymosphaeriaceae, Fenestellaceae, Massariaceae, 

Melanommataceae, Microthyriaceae, Mytilinidiaceae, 

Platystomaceae and Requienellaceae (Barr 1990a). 

Pleosporales was formally established by Luttrell and Barr 

(in Barr 1987b), characterised by perithecioid ascomata, 

usually with a papillate apex, ostioles with or without 

periphyses, presence of cellular pseudoparaphyses, bitunicate 

asci, and ascospores of various shapes, pigmentation and 

septation (Table 1). Eighteen families were included, i.e. 

Arthopyreniaceae, Botryosphaeriaceae, Cucurbitariaceae, 

Dacampiaceae, Dimeriaceae, Hysteriaceae, Leptosphaeriaceae, 

Lophiostomataceae, Parodiellaceae, Phaeosphaeriaceae, 

Phaeotrichaceae, Pleomassariaceae, Pleosporaceae, 

Polystomellaceae, Pyrenophoraceae, Micropeltidaceae, 

Tubeufiaceae and Venturiaceae. Recent phylogenetic analysis 

based on DNA sequence comparisons, however, indicated 

that separation of the orders (Pleosporales and Melanommatales) 

based on the Pleospora or Sporormia centrum type, is 

not a natural grouping, and Melanommatales has therefore 

been combined under Pleosporales (Liew et al. 2000; 

Lumbsch and Lindemuth 2001; Reynolds 1991). Six more 

families, i.e. Cucurbitariaceae, Diademaceae, Didymosphaeriaceae, 

Mytilinidiaceae, Testudinaceae and Zopfiaceae, 

were subsequently added to Pleosporales (Lumbsch and 

Huhndorf 2007). After intensive sampling and multigene 

phylogenetic studies, 20 families were accepted in Pleosporales, 

namely Aigialaceae, Amniculicolaceae, Delitschiaceae, 

Didymellaceae, Didymosphaeriaceae, Hypsostromataceae, 

Lentitheciaceae, Leptosphaeriaceae, Lindgomycetaceae, 

Lophiostomataceae, Massarinaceae, Melanommataceae, 

Montagnulaceae, Morosphaeriaceae, Phaeosphaeriaceae, 

Pleosporaceae, Pleomassariaceae, Sporormiaceae, Tetraplosphaeriaceae 

and Trematosphaeriaceae (Boehm et al. 

2009a, b; Mugambi and Huhndorf 2009b;Schochetal.2009; 

Shearer et al. 2009; Suetrongetal.2009; Tanakaetal.2009; 

Zhang et al. 2009a) (Table 1). In addition, another five 

families, i.e. Arthopyreniaceae, Cucurbitariaceae, Diademaceae, 

Teichosporaceae and Zopfiaceae are tentatively included 

(Kruys et al. 2006; Plate 1). In the most recent issue of 

Myconet, 28 families were included in Pleosporales 

(Lumbsch and Huhndorf 2010). 

Species included in Pleosporales have different ecological 

or morphological characters. For instance, members of 

the Leptosphaeriaceae have saprobic or parasitic lifestyles 

and lightly pigmented, multi-septate ascospores. Members 

of the Lophiostomataceae are mostly saprobic with ascomata 

that usually possess a compressed apex. Members of 

Sporormiaceae are coprophilous, and are characterized by 

heavily pigmented, multi-septate ascospores with germ 

slits, and with or without non-periphysate ostioles. The 

lack of DNA sequence data for representatives of numerous 

families means that their inter-relationships are unclear and 

many genera or species are artificially placed based on 

morphological classification. The most recent study on 

Venturiaceae indicated that this group had a set of unique 

morphological and ecological characters, which is distinct 

and distantly related to other members of Pleosporales 

(Kruys et al. 2006; Zhang et al. unpublished). Molecular 

phylogenetic results indicated that members of Venturiaceae 

form a robust clade separate from the core members of 

Pleosporales, and the clade of Venturiaceae was uncertainly 

placed but outside of the two currently designated 

dothideomycetous subclasses, i.e. Pleosporomycetidae and 

Dothideomycetidae (Schoch et al. 2009). In addition, 

phylogenetic analysis of rDNA sequence data indicates 

that members of Zopfiaceae (as Testudinaceae) seem to 

lack affinity with Pleosporales (Kodsueb et al. 2006b). 

Thus, 26 families are temporarily accepted in Pleosporales 

in this study, although some such as Zopfiaceae, still 

require extensive DNA sequence sampling (Table 4).


Table 1 Major circumscription changes of Pleosporales from 1955 to 2011 

References 

Luttrell 1955 

Müller and von Arx 1962 

Luttrell 1973 

Barr 1979a 

Barr 1987b 

Kirk et al. 2001, 2008 

Boehm et al. 2009a, b; Mugambi 

and Huhndorf 2009b; Schoch et 

al. 2009; Shearer et al. 2009; 

Suetrong et al. 2009; Tanaka et 

al. 2009; Zhang et al. 2009a 

Circumscriptions distinguishing Pleosporales from other orders of Dothideomycetes 

Pleospora-type centrum development. 

Ascomata perithecoid, with rounded or slit-like ostiole; asci produced within a locule, arranged regularly 

in a single layer or irregularly scattered, surrounded with filiform pseudoparaphyses, cylindrical, 

ellipsoidal or sac-like. 

Ascocarps perithecioid, immersed, erumpent to superficial on various substrates, asci ovoid to mostly 

clavate or cylindrical, interspersed with pseudoparaphyses (sometimes form an epithecium) in mostly 

medium- to large-sized locules. 

Saprobic, parasitic, lichenized or hypersaprobic. Ascomata perithecioid, rarely cleistothecioid or 

hysterothecioid, peridium pseudoparenchymatous, pseudoparaphyses cellular, narrow or broad, 

deliquescing early at times, not forming an epithecium, asci oblong, clavate or cylindrical, interspersed 

with pseudoparaphyses, ascospores mostly asymmetric. 

Saprobic, biotrophic or hemibiotrophic. Ascomata globose, subglobose or conical, asci bitunicate, oblong, 

clavate or cylindrical, cellular pseudoparaphyses, ascospores hyaline or pigmented, asymmetric or 

symmetric, with or without septa. 

Ascomata perithecioid or rarely cleistothecioid, sometimes clypeate, mostly globose, thick-walled, 

immersed or erumpent, black, sometimes setose, peridium composed of pseudoparenchymatous cells, 

pseudoparaphyses trabeculate or cellular, asci cylindrical, fissitunicate, with a well-developed ocular 

chamber, rarely with a poorly defined ring (J-), ascospores hyaline to brown, septate, thin or thickwalled, 

sometimes muriform, usually with sheath, anamorphs hyphomycetous or coelomycetous. 

Hemibiotrophic, saprobic, hypersaprobic, or lichenized. Habitats in freshwater, marine or terrestrial 

environment. Ascomata perithecioid, rarely cleistothecioid, immersed, erumpent to superficial, globose 

to subglobose, or lenticular to irregular, with or without conspicuous papilla or ostioles. Ostioles with or 

without periphyses. Peridium usually composed of a few layers of cells with various shapes and 

structures. Hamathecium persistent, filamentous, very rarely decomposing. Asci bitunicate, fissitunicate, 

cylindrical, clavate to obclavate, with or without pedicel. Ascospores hyaline or pigmented, ellipsoidal, 

broadly to narrowly fusoid or filiform, mostly septate. 

Morpho-characters used in taxonomy of Pleosporales 

Sexual characters 

According to the Linnean classification system, reproductive 

structures are the most important criteria in plant 

taxonomy, and this proposal is widely applied in fungal 

taxonomy (Gäumann 1952). In the classification of Dothideomycetes, 

reproductive characters such as the uni- or 

multilocular nature and shape of ascomata, presence and 

shape of ostioles/papillae, shape and apical structures of 

asci and shape, pigmentation and septation of ascospores 

play important roles at different ranks (Clements and Shear 

1931; Luttrell 1951, 1955, 1973). Besides the common 

morphological characters possessed by Dothideomycetes 

(bitunicate and fissitunicate asci as well as the perithecioidlike 

ascostromata), most pleosporalean fungi also have 

pseudoparaphyses among their well-arranged asci (Zhang et 

al. 2009a). Currently, classification of Pleosporales at the 

family level focuses mostly on morphological characters of 

ascomata (such as size, shape of ostiole or papilla), presence 

or absence of periphyses, characters of centrum (such as asci, 

pseudoparaphyses and ascospores) as well as on lifestyle or 

habitat (Barr 1990a; Shearer et al. 2009; Suetrong et al. 

2009; Tanaka et al. 2009; Zhang et al. 2009a), whilst relying 

extensively on DNA sequence comparisons. 

Ascomata 

Most species of Pleosporales have uniloculate ascomata. 

The presence (or absence) and forms of papilla and ostiole 

are the pitoval character of ascomata, which serve as 

important characteristics in generic or higher rank classification 

(Clements and Shear 1931). The vertically flattened papilla has 

recently been shown as an effective criterion for familial level 

classification, e.g. in the Amniculicolaceae and the Lophiostomataceae 

(Zhang et al. 2009a). Papillae and ostioles are 

present in most species of Pleosporales, except in the 

Diademaceae and Sporormiaceae. Members of Diademaceae 

have apothecial ascomata, and some genera of Sporormiaceae 

have cleistothecioid ascomata. Another coprophilous pleosporalean 

family, Delitschiaceae, can be distinguished from 

Sporormiaceae by the presence of periphysate ostioles. 

Pseudoparaphyses 

Presence of pseudoparaphyses is a characteristic of 

Pleosporales (Kirk et al. 2008; Liew et al. 2000). Although 

pseudoparaphyses may be deliquescing in some families 

when the ascomata mature (e.g. in Didymellaceae), they are 

persistent in most of other pleosporalean members. According 

to the thickness, with or without branching and density 

of septa, pseudoparaphyses were roughly divided into two 

types: trabeculate and cellular, and their taxonomic significance 

need to be re-evaluated (Liew et al. 2000).


Asci 

The asci of Pleosporales are bitunicate, usually fissitunicate, 

mostly cylindrical, clavate or cylindro-clavate, and 

rarely somewhat obclavate or sphaerical (e.g. Macroventuria 

anomochaeta Aa and Westerdykella dispersa). There are 

ocular chambers in some genera (e.g. Amniculicola and 

Asteromassaria), or sometimes with a large apical ring (J-) 

(e.g. Massaria). 

Ascospores 

Ascospores of Pleosporales canbehyalineorcolored 

to varying degrees. They may be amerosporous (e.g. 

species of Semidelitschia), phragmosporous (e.g. Phaeosphaeria 

and Massariosphaeria), dictyosporous (e.g. most 

species of Pleospora and Bimuria), or scolecosporous 

(e.g. type species of Cochliobolus, Entodesmium or 

Lophionema). Although ascospore morphology had been 

regarded as a key factor in differentiating genera under 

some families, e.g. Arthopyreniaceae (Watson 1929) and 

Testudinaceae (Hawksworth 1979), it has been proven 

variable even within a single species. For instance, two 

types of ascospores are produced by Mamillisphaeria 

dimorphospora, i.e. one type is large and hyaline, and the 

other is comparatively smaller and brown. Numerous 

studies have shown the unreliability of ascospore characters 

above genus level classification (e.g. Phillips et al. 

2008; Zhang et al. 2009a). 

Asexual states of Pleosporales 

Anamorphs of pleosporalean families 

Anamorphs of Pleosporales are mostly coelomycetous, 

but may also be hyphomycetous. Phoma or Phoma-like 

anamorphic stages and its relatives are most common 

anamorphs of Pleosporales (Aveskamp et al. 2010; de 

Gruyter et al. 2009, 2010; Hyde et al. 2011). Some of the 

reported teleomorph and anamorph connections (including 

some listed below) are, however, based on the association 

rather than single ascospore isolation followed by induction 

of the other stage in culture (Hyde et al. 2011). 

Pleosporales suborder Pleosporineae 

Pleosporineae is a phylogenetically well supported 

suborder of Pleosporales, which temporarily includes 

seven families, namely Cucurbitariaceae, Didymellaceae, 

Didymosphaeriaceae, Dothidotthiaceae, Leptosphaeriaceae, 

Phaeosphaeriaceae and Pleosporaceae, and 

contains many important plant pathogens (de Gruyter et 

al. 2010; Zhangetal.2009a). De Gruyter et al. (2009, 

2010) systematically analyzed the phylogeny of Phoma 

and its closely related genera, and indicated that their 

representative species cluster in different subclades of 

Pleosporineae. 

Plate 1 The best scoring likelihood tree of representative Pleosporales 

obtained with RAxML v. 7.2.7 for a concatenated set of 

nucleotides from LSU, SSU, RPB2 and TEF1. Family and suborder 

names are indicated where possible. The percentages of nodes present 

in 250 bootstrap pseudo replicates are shown above branches. Culture 

and voucher numbers are indicated after species names and the 

presence of the genes used in the analysis are indicated by pluses in 

this order: LSU, SSU, RPB2, TEF1 

Cucurbitariaceae 

Based on the molecular phylogenetic analysis, some 

species of Coniothyrium, Pyrenochaeta, Phoma, Phialophorophoma 

and Pleurophoma belong to Cucurbitariaceae (de 

Gruyter et al. 2010; Hyde et al. 2011). Other reported 

anamorphs of Cucurbitaria are Camarosporium, Diplodialike 

and Pleurostromella (Hyde et al. 2011; Sivanesan 1984). 

The generic type of Cucurbitaria (C. berberidis Fuckel) is 

linked to Pyrenochaeta berberidis (Farr et al. 1989). Curreya 

has a Coniothyrium-like anamorphic stage (von Arx and van 

der Aa 1983; Marincowitz et al. 2008). The generic type of 

Curreya is C. conorum (Fuckel) Sacc., which is reported to 

be linked with Coniothyrium glomerulatum Sacc. (von Arx 

and van der Aa 1983). The generic type of Rhytidiella (R. 

moriformis, Cucurbitariaceae) can cause rough-bark of 

Populus balsamifera, andhasaPhaeoseptoria anamorphic 

stage (Zalasky 1968). Rhytidiella baranyayi Funk & Zalasky, 

another species of Rhytidiella associated with the cork-bark 

disease of aspen is linked with Pseudosporella-like anamorphs 

(Funk and Zalasky 1975; Sivanesan 1984). 

Didymellaceae, Didymosphaeriaceae and Dothidotthiaceae 

As has been mentioned before, Phoma sensu lato species 

have been proved to be highly polyphyletic, and they cluster 

in six distinct familial clades within the Pleosporales 

(Aveskamp et al. 2010). Most Phoma species, including the 

generic type (P. herbarum), clustered in Didymellaceae 

(Aveskamp et al. 2010). The clade of Didymellaceae also 

comprises other sections, such as Ampelomyces, Boeremia, 

Chaetasbolisia, Dactuliochaeta, Epicoccum, Peyronellaea, 

Phoma-like, Piggotia, Pithoascus, as well as the type species 

of Ascochyta and Microsphaeropsis (Aveskamp et al. 2010; 

de Gruyter et al. 2009; Kirketal.2008; Sivanesan 1984). 

Leptosphaerulina is another genus of Didymellaceae, which 

has hyphomycetous anamorphs with pigmented and 

muriform conidia, such as Pithomyces (Roux 1986). 

The other reported anamorphs of Didymosphaeria are 

Fusicladiella-like, Dendrophoma, Phoma-like (Hyde et al. 

2011). Hyphomycetous Thyrostroma links to Dothidotthiaceae 

(Phillips et al. 2008). 

Some important plant pathogens are included within 

Didymellaceae, such as Phoma medicaginis Malbr. & 

Roum., which is a necrotrophic pathogen on Medicago 

truncatula (Ellwood et al. 2006). Phoma herbarum is 

another plant pathogen, which has potential as a

Fungal Diversity

Plate 1 (continued) 

Fungal Diversity


biocontrol agent of weeds (Neumann and Boland 2002). 

Ascochyta rabiei is a devastating disease of chickpea in 

most of the chickpea producing countries (Saxena and 

Singh 1987). 

Leptosphaeriaceae 

The anamorphic stages of Leptosphaeriaceae can be 

Coniothyrium, Phoma, Plenodomus and Pyrenochaeta. All 

are coelomycetous anamorphs, and they may have phialidic 

or annellidic conidiogenous cells. Phoma heteromorphospora 

Aa & Kesteren, the type species of Phoma sect. 

Heterospora and Coniothyrium palmarum, the generic type 

of Coniothyrium, reside in Leptosphaeriaceae (de Gruyter 

et al. 2009). 

Pleosporaceae 

Various anamorphic types can occur in Pleosporaceae, 

which can be coelomycetous or hyphomycetous, and the 

ontogeny of conidiogenous cells can be phialidic, annellidic 

or sympodial blastic. Both Ascochyta caulina and Phoma 

betae belong to Pleosporaceae (de Gruyter et al. 2009). 

Some species of Bipolaris and Curvularia are anamorphs 

of Cochliobolus. Many species of these two 

genera cause plant disease or even infect human beings 

(Khan et al. 2000). They are hyphomycetous anamorphs 

with sympodial proliferating conidiogenous cells, and 

pigmented phragmosporous poroconidia. The generic type 

of Lewia (L. scrophulariae) is linked with Alternaria 

conjuncta E.G. Simmons (Simmons 1986), and the 

generic type of Pleospora (P. herbarum) is linked with 

Stemphylium botryosum Sacc. (Sivanesan 1984). Both 

Alternaria and Stemphylium are hyphomycetous anamorphs 

characterized by pigmented, muriform conidia that develop 

at a very restricted site in the apex of distinctive conidiophores 

(Simmons 2007). 

The generic type of Pleoseptum (P. yuccaesedum) is 

linked with Camarosporium yuccaesedum (Ramaley and 

Barr 1995), the generic type of Macrospora (M. scirpicola) 

with Nimbya scirpicola (Fuckel) E.G. Simmons (Simmons 

1989), and the generic type of Setosphaeria (S. turcica) 

with Drechslera turcica (Pass.) Subram. & B.L. Jain 

(Sivanesan 1984). Pyrenophora has the anamorphic stages 

of Drechslera, and the anamorphic stage of Wettsteinina 

can be species of Stagonospora (Farr et al. 1989). 

Most common anamorphs in Pleosporaceae are Alternaria, 

Bipolaris, Phoma-like and Stemphylium, and they 

can be saprobic or parasitic on various hosts. Phoma betae 

A.B. Frank is a notorious pathogen on sugar beet, which 

causes zonate leaf spot or Phomopsis of sugar beet. 

Alternaria porri (Ellis) Cif., Stemphylium solani G.F. 

Weber, S. botryosum and S. vesicarium (Wallr.) E.G. 

Simmons can cause leaf blight of garlic (Zheng et al. 

2009). Phoma incompta Sacc. & Martelli is a pathogen on 

olive, and Stemphylium botryosum, the anamorph of 

Pleospora herbarum, causes leaf disease of olive trees 

(Malathrakis 1979). 

Phaeosphaeriaceae 

The type species of Phoma sect. Paraphoma (Phoma 

radicina (McAlpine) Boerema) as well as several pathogens 

on Gramineae, i.e. Stagonospora foliicola (Bres.) Bubák, S. 

neglecta var. colorata and Wojnowicia hirta Sacc.belongto 

Phaeosphaeriaceae (de Gruyter et al. 2009). Other anamorphs 

reported for Phaeosphaeriaceae are Amarenographium, 

Ampelomyces, Chaetosphaeronema, Coniothyrium, 

Hendersonia, Neosetophoma, ?Parahendersonia, Paraphoma, 

Phaeoseptoria, Rhabdospora, Scolecosporiella, Setophoma, 

Sphaerellopsis and Tiarospora. 

These anamorphic fungi can be saprobic, but mostly 

pathogenic on herbaceous plants. For instance, Stagonospora 

foliicola and Coniothyrium concentricum (Desm.) 

Sacc. can cause leaf spots on herbaceous plants (Zeiders 

1975), and Ampelomyces quisqualis Ces. is a hyperparasite 

of powdery mildews. 

Pleosporales suborder Massarineae 

Massarineae species are mostly saprobic in terrestrial or 

aquatic environments. Five families are currently included 

within Massarineae, viz. Lentitheciaceae, Massarinaceae, 

Montagnulaceae, Morosphaeriaceae and Trematosphaeriaceae. 

Anamorphs of the five families are summarized as 

follows. 

Lentitheciaceae 

Stagonospora macropycnidia Cunnell nests within the 

clade of Lentitheciaceae (Plate 1). A relatively broad genus 

concept of Stagonospora is currently accepted, which 

comprises parasitic or saprobic taxa. Keissleriella cladophila 

(Niessl) Corbaz is another species nesting within 

Lentitheciaceae (Zhang et al. 2009a), and is linked with 

Dendrophoma sp., which has branching conidiogenous 

cells, and 1-celled, hyaline conidia (Bose 1961; Sivanesan 

1984). 

Massarinaceae 

A relatively narrow concept tends to be accepted for 

Massarinaceae, which seems only to comprise limited 

species such as Byssothecium circinans, Massarina eburnea, 

M. cisti S.K. Bose, M. igniaria (C. Booth) Aptroot 

(anamorph: Periconia igniaria E.W. Mason & M.B. Ellis) 

and Neottiosporina paspali (G.F. Atk.) B. Sutton & Alcorn 

(Zhang et al. 2009a; Plate 1). Similarly, a relatively narrow 

generic concept of Massarina was accepted, containing 

only M. eburnea and M. cisti (Zhang et al. 2009b), and both 

species have been linked with species of Ceratophoma 

(Sivanesan 1984).


Montagnulaceae 

Montagnula has an Aschersonia anamorph, and Kalmusia 

and Paraphaeosphaeria have Coniothyrium-like, Cytoplea, 

Microsphaeropsis and Paraconiothyrium anamorphs. 

The generic type of Paraphaeosphaeria (P. michotii) is 

linked with Coniothyrium scirpi Trail (Webster 1955). The 

Coniothyrium complex is highly polyphyletic, and was 

subdivided into four groups by Sutton (1980), viz. 

Coniothyrium, Microsphaeropsis, Cyclothyrium and Cytoplea. 

Paraconiothyrium was introduced to accommodate 

Coniothyrium minitans W.A. Campb. and C. sporulosum 

(W. Gams & Domsch) Aa, which are closely related to 

Paraphaeosphaeria based on 18S rDNA sequences phylogeny 

(Verkley et al. 2004). 

Morosphaeriaceae 

Based on the multigene phylogenetic analysis in this 

study, Asteromassaria is tentatively included in Morosphaeriaceae. 

Asteromassaria macrospora is linked with 

Scolicosporium macrosporium (Berk.) B. Sutton, which is 

hyphomycetous. No anamorphic stages have been reported 

for other species of Morosphaeriaceae. 

Trematosphaeriaceae 

Three species from three different genera were included 

in Trematosphaeriaceae, i.e. Falciformispora lignatilis, 

Halomassarina thalassiae and Trematosphaeria pertusa 

(Suetrong et al. data unpublished; Plate 1). Of these, only 

Trematosphaeria pertusa, the generic type of Trematosphaeria, 

produces hyphopodia-like structures on agar 

(Zhang et al. 2008a). 

Other families of Pleosporales 

Amniculicolaceae 

Three anamorphic species nested within the clade of 

Amniculicolaceae, i.e. Anguillospora longissima (Sacc. & 

P. Syd.) Ingold, Repetophragma ontariense (Matsush.) W. 

P. Wu and Spirosphaera cupreorufescens Voglmayr 

(Zhang et al. 2009a). Sivanesan (1984, p.500)described 

the teleomorphic stage of Anguillospora longissima as 

Massarina sp. II, which fits the diagnostic characters of 

Amniculicola well. Thus this taxon may be another species of 

Amniculicola. 

Hypsostromataceae 

A Pleurophomopsis-like anamorph is reported in the 

subiculum of the generic type of Hypsostroma (H. saxicola 

Huhndorf) (Huhndorf 1992). 

Lophiostomataceae 

The concept of Lophiostomataceae was also narrowed, 

and presently contains only Lophiostoma (Zhang et al. 

2009a). Leuchtmann (1985) studied cultures of some 

Lophiostoma species, and noticed that L. caulium (Fr.) 

Ces. & De Not., L. macrostomum, L. semiliberum (Desm.) 

Ces. & De Not., Lophiostoma sp. and Lophiotrema nucula 

produced Pleurophomopsis anamorphic stages, which are 

similar to those now in Melanomma (Chesters 1938), but 

Lophiostoma and Melanomma has no proven phylogenetic 

relationship (Zhang et al. 2009a, b; Plate 1). Species of 

Aposphaeria have also been reported in Massariosphaeria 

(Farr et al. 1989; Leuchtmann 1984), but the polyphyletic 

nature of Massariosphaeria is well documented (Wang et 

al. 2007). 

Melanommataceae 

The anamorphs of the Melanommataceae are mostly 

coelomycetous and rarely hyphomycetous with various 

ontogenic structures, such as annellidic or sympodial for 

hyphomycetes (Exosporiella and Pseudospiropes) and 

coelomycetes (Aposphaeria-like and Pyrenochaeta). 

Herpotrichia is reported as having a Pyrenochaeta 

anamorphic stage with or without seta on the surface of 

pycnidia (Sivanesan 1984). Aposphaeria and Phoma-like 

have been reported in Melanomma species (Chesters 

1938; Sivanesan 1984). Similarly, the anamorphs of 

Karstenula are reported as coelomycetous, i.e. Microdiplodia 

(Constantinescu 1993). The anamorphic stage of 

Anomalemma is Exosporiella (Sivanesan 1983), and that 

of Byssosphaeria is Pyrenochaeta (Barr 1984). Ohleria 

brasiliensis Starbäck has been linked with Monodictys 

putredinis (Wallr.) S. Hughes (Samuels 1980). Astrosphaeriella 

is a contentious genus as its familial status is 

not determined yet. Here we temporarily assigned it under 

Melanommataceae, which is linked with the anamorph 

genus Pleurophomopsis. 

Pleomassariaceae 

Shearia and Prosthemium are all anamorphs of Pleomassaria, 

andProsthemium betulinum is linked with the generic 

type of Pleomassaria (P. siparia) (Barr1982b; Sivanesan 

1984; Sutton 1980; Tanaka et al. 2010). Splanchnonema is a 

genus of Pleomassariaceae, the teleomorphic morphology of 

which is difficult to distinguish from two other genera, i.e. 

Asteromassaria and Pleomassaria, and the reported anamorphs 

of Splanchnonema are Ceuthodiplospora, Myxocyclus 

and Stegonsporium, which are comparable with those of 

Asteromassaria and Pleomassaria. 

Tetraplosphaeriaceae 

Tetraplosphaeriaceae was introduced to accommodate 

the Massarina-like bambusicolous fungi that produce 

Tetraploa sensu stricto anamorphs (Tanaka et al. 2009). 

Tetraploa aristata Berk. & Broome, the generic type of 

Tetraploa is widely distributed, associated with various


substrates and many occur in freshwater or has been 

isolated from air. The polyphyletic nature of T. aristata 

has been well documented (Tanaka et al. 2009). Anamorphic 

stages can serve as a diagnostic character for this 

family. 

Diademaceae, Massariaceae, Sporormiaceae 

and Teichosporaceae 

The Sporormiaceae is coprophilous having Phoma or 

Phoma-related anamorphic states (Cannon and Kirk 2007). 

Comoclathris (Diademaceae) is linked with Alternaria-like 

anamorphs (Simmons 1952). Myxocyclus links to Massaria 

(Massariaceae) (Hyde et al. 2011). The anamorphic stage of 

Chaetomastia (Teichosporaceae) isAposphaeria- orConiothyrium-like 

(Barr 1989c). 

Generally speaking, the morphologically simple conidiophores 

are usually considered phylogenetically uninformative 

(Seifert and Samuels 2000). Phoma-like anamorphs commonly 

occur in Pleosporales, while their colorless and unicellular 

conidia are also not phylogenetically informative (Seifert and 

Samuels 2000). 

All of the above mentioned anamorphic taxa of 

Pleosporales have phialidic, annellidic or sympodial conidiogenous 

cells, representing apical wall-building type 

(compared to ring wall-building and diffused wallbuilding) 

(Nag Raj 1993), which may indicate that the 

wall-building type probably has phylogenetic significance. 

Molecular phylogeny of Pleosporales 

Numerous genes have been applied in phylogenetic 

studies of Pleosporales, mostly including LSU, SSU, 

mtSSU and ITS as well as the protein genes, such as 

RPB1, RPB2, TEF1, β-tubulin (TUB1) and actin (ACT1). 

A single gene such as ITS or LSU, has been used to study 

phylogenetic relationships between Leptosphaeria and 

Phaeosphaeria (Câmara et al. 2002) or Pleosporaceae 

and Tubeufiaceae (Kodsueb et al. 2006a, b) (Table2). The 

use of these phylogenetic markers, although making 

important contributions, has not been successful in 

resolving numerous relationships in single gene dendrograms. 

One exception is the use of SSU sequences to 

demonstrate the phylogenetic significance of pseudoparaphyses 

(Liew et al. 2000) whilst rejecting the phylogenetic 

utility of pseudoparaphyses morphology (cellular or 

trabeculate). Analyses with combined genes have had 

more success. For instance combined analyses with LSU 

and SSU sequence data could be used to define family 

level classification in a few cases (Dong et al. 1998; de 

Gruyter et al. 2009; Lumbsch and Lindemuth 2001; Pinnoi 

et al. 2007; Zhang et al. 2009b) (Table2). The addition of 

more than two genes has been used to determine relationships 

between orders. For instance, genes such as LSU, 

SSU and mtSSU have been used to analyze ordinal 

relationships in Loculoascomycetes (Lindemuth et al. 

2001), and to analyze phylogenetic relationships of 

coprophilous families in Pleosporales (Kruys et al. 

2006). Phaeocryptopus gaeumannii (T. Rohde) Petr. was 

showntobelonginDothideales based on LSU, SSU and 

ITS sequence analysis (Winton et al. 2007), while Schoch 

et al. (2006) used four genes, i.e. LSU, SSU, RPB2 and 

TEF1 to evaluate the phylogenetic relationships among 

different orders of the Dothideomycetes. Fivegenes,viz. 

LSU, SSU, TEF1, RPB1 andRPB2, were used to study the 

phylogenetic relationships of different orders within 

Dothideomycetes (Schoch et al. 2009) and of different 

families within Pleosporales (Zhang et al. 2009a) 

(Table 2). It is clear that even more genes will be required 

to address the remaining issues and the promise of genome 

analyses is within reach (www.jgi.doe.gov/sequencing/ 

why/dothideomycetes.html) forDothideomycetes. 

The importance of generic type specimens 

The type specimen (collection type) is a fundamental 

element in the current Code of Botanical Nomenclature at 

familial or lower ranks (Moore 1998). A type specimen 

fixes the name to an exact specimen at family, genera, 

species and variety/subspecies rank and is ultimately based 

on this single specimen, i.e. a family name is based on a 

genus, the genus name is based on a species, and the 

species name is based on a specimen (Kirk et al. 2008). 

The generic type is of great importance in defining 

generic circumscriptions in fungal taxonomy. The generic 

types of Pleosporales have been studied previously by 

many mycologists. For instance, Müller and von Arx 

(1962) studied the generic types of “Pyrenomycetes”, and 

described and illustrated them in detail. Sivanesan (1984) 

described and illustrated the generic representatives of 

Loculoascomycetes for both their teleomorphs and anamorphs, 

and their links were emphasized. A large number 

of pleosporalean genera have been studied by Barr (1990a, 

b). Almost all of the previous work was conducted more 

than 20 years ago, when no molecular phylogenetic studies 

could be carried out and thus had been carried out in a 

systematic fashion. 

Aim and outline of present study 

The present study had two principal objectives: 

1. To explore genera under Pleosporales based on the 

generic types and provide a detailed description and 

illustration for the type species of selected genera, 

discuss the study history of those genera, and explore 

their ordinal, familial, and generic relationships;


Table 2 List of phylogenetic studies on Pleosporales 

Year Author(s) Loci used Target fungi General conclusion 

1998 Dong et al. LSU, SSU Leptosphaeriaceae, Pleosporaceae 

and three other families 

Leptosphaeriaceae is paraphyletic and 

Pleosporaceae is monophyletic. 

2000 Liew et al. SSU Pleosporales and Melanommatales Pleosporales and Melanommatales are not 

naturial groups. 

2001 Lindemuth et al. LSU, SSU, mtSSU loculoascomycetes Loculoascomycetes are not monophyletic. 

2001 Lumbsch and 

Lindemuth 

LSU, SSU Dothideomycetes Presence of pseudoparaphyses is a major 

character at order level classification 

2002 Câmara et al. ITS Leptosphaeria and Phaeosphaeria Accepted Leptosphaeria sensu stricto. 

2006 Kodsueb et al. LSU Pleosporaceae Wettsteinina should be excluded from the 

Pleosporaceae. 

2006 Kodsueb et al. LSU Tubeufiaceae Tubeufiaceae is more closely related to the 

Venturiaceae. 

2006 Kruys et al. LSU, SSU, mtSSU coprophilous familes of 


2006 Schoch et al. LSU, SSU, TEF1, 

RPB2 

Dothideomycetes 

coprophilous familes of Pleosporales form 

phylogenetic monophyletic groups 

respectively 

Proposed the subclasses 

Pleosporomycetidae 

2007 Pinnoi et al. LSU, SSU Pleosporales phylogenetic relationships of different 

families of Pleosporales, introduced a 

new fungus–– Berkleasmium crunisia 

2007 Wang et al. LSU, SSU, RPB2 Massariosphaeria Massariosphaeria is not monophyletic 

2007 Winton et al. LSU, SSU, ITS Phaeocryptopus gaeumannii Phaeocryptopus gaeumannii nested in 

Dothideales. 

2008a Zhang et al. LSU, SSU Melanomma and Trematosphaeria Melanomma and Trematosphaeria belong 

to different families 

2009 de Gruyter et al. LSU, SSU; Phoma and related genera They are closely related with 

Didymellaceae, Leptosphaeriaceae, 

Phaeosphaeriaceae and Pleosporaceae 

2009a Zhang et al. LSU, SSU, TEF1, 

RPB1, RPB2 

2009 Mugambi and 

Huhndorf 

LSU, TEF1 


Melanommataceae, 


Amniculicolaceae and Lentitheciaceae were 

introduced, and Pleosporineae 

recircumscribed. 

Recircumscribed Melanommataceae and 

Lophiostomataceae, and reinstated 

Hypsostromataceae. 

2009 Nelsen et al. LSU and mtSSU lichenized Dothideomycetes Pyrenocarpous lichens with bitunicate asci 

are not monophyletic, but belong to at 

least two classes (Dothideomycetes and 

Erotiomycetes). 

2009 Suetrong et al. LSU, SSU, TEF1, RPB1 marine Dothideomycetes Two new families are introduced 

Aigialaceae and Morosphaeriaceae. 

2009 Shearer et al. LSU, SSU freshwater Dothideomycetes Freshwater Dothideomycetes are related to 

terrestrial taxa and have adapted to 

freshwater habitats numerous times. 

2009 Tanaka et al. LSU, SSU, TEF1, ITS, BT bambusicolous Pleosporales Introduced Tetraplosphaeriaceae with 

Tetraploa-like anamorphs. 

2009 Kruys and Wedin ITS-nLSU, mtSSU rDNA 

and β-tubulin 

Sporormiaceae 

Analyzed the inter-generic relationships as 

well as evaluated the morphological significance 

used in this family. 

2010 Hirayama et al. LSU, SSU Massarina ingoldiana sensu lato Massarina ingoldiana sensu lato is 

polyphyletic, and separated into two 

clades within Pleosporales. 

2010 Aveskamp et al. LSU, SSU, ITS and β-tubulin Phoma and related genera within Rejected current Boeremaean subdivision. 

Didymellaceae 

2010 de Gruyter et al. LSU, SSU Phoma and related genera within 

Pleosporineae 

Introduced Pyrenochaetopsis, Setophoma 

and Neosetophoma and reinstated 

Cucurbitariaceae within Pleosporineae


2. To investigate the phylogeny of Pleosporales, its interfamilial 

relationships, and the morphological circumscription 

of each family; 

In order to clarify morphological characters, the generic 

types of the majority of teleomorphic pleosporalean genera 

(> 60%) were studied. Most of them are from the “core 

families” of Pleosporales, i.e.Delitschiaceae, Lophiostomataceae, 

Massariaceae, Massarinaceae, Melanommataceae, 

Montagnulaceae, Phaeosphaeriaceae, Phaeotrichaceae, Pleomassariaceae, 

Pleosporaceae, Sporormiaceae and Teichosporaceae. 

Notes are given for those where type 

specimens could not be obtained during the timeframe 

of this study. A detailed description and illustration of 

each generic type is provided. Comments, notes and 

problems that need to be addressed are provided for each 

genus. Phylogenetic investigation based on five nuclear loci, 

viz. LSU, SSU, RPB1, RPB2 andTEF1 was carried out using 

available strains from numerous genera in Pleosporales. In 

total, 278 pleosporalean taxa are included in the phylogenetic 

analysis, which form 25 familial clades on the dendrogram 

(Plate 1). The suborder, Massarineae, is emended to 

accommodate Lentitheciaceae, Massarinaceae, Montagnulaceae, 

Morosphaeriaceae and Trematosphaeriaceae. 

Materials and methods 

Molecular phylogeny 

Four genes were used in this analysis, the large and small 

subunits of the nuclear ribosomal RNA genes (LSU, SSU) 

and two protein coding genes, namely the second largest 

subunit of RNA polymerase II (RPB2) and translation 

elongation factor-1 alpha (TEF1). All sequences were 

downloaded from GenBank as listed in Table 3. Eachof 

the individual ribosomal genes was aligned in SATé under 

default settings with at least 20 iterations. The protein coding 

genes were aligned in BioEdit (Hall 2004) and completed by 

manual adjustment. Introns were removed and all genes were 

concatenated in a single nucleotide alignment with 43% 

missing and gap characters out of a total set of 5081. The 

alignment had 100% representation for LSU, 75% for SSU, 

48% for RPB2 and65%forTEF1. The final data matrix had 

280 taxa including outgroups (Table 3). 

Previous results indicated no clear conflict amongst the 

majority of the data used (Schoch et al. 2009). A phylogenetic 

analysis of the concatenated alignment was performed on 

CIPRES webportal (Miller et al. 2009) using RAxML v. 7.2.7 

(Stamatakis 2006; Stamatakis et al. 2008) applying unique 

model parameters for each gene and codon (8 partitions). A 

general time reversible model (GTR) was applied with a 

discrete gamma distribution and four rate classes. Fifty 

thorough maximum likelihood (ML) tree searches were done 

in RAxML v. 7.2.7 under the same model, each one starting 

from a separate randomized tree and the best scoring tree 

selected with a final likelihood value of −95238.628839. Two 

isolates of Hysterium angustatum (Hysteriales, Pleosporomycetidae) 

were used as outgroups based on earlier work 

(Boehm et al. 2009a). Bootstrap pseudo-replicates were run 

with the GTRCAT model approximation, allowing the 

program to halt bootstraps automatically under the majority 

rule criterion (Pattengale et al. 2010). The resulting 250 

replicates were plotted on to the best scoring tree obtained 

previously. The phylogram with bootstrap values on the 

branches is presented in Plate 1 by using graphical options 

available in TreeDyn v. 198.3 (Chevenet et al. 2006). 

Morphology 

Type specimens as well as some other specimens were loaned 

from the following herbaria: BAFC, BISH, BPI, BR, BRIP, 

CBS, E, ETH, FFE, FH, G, H, Herb. J. Kohlmeyer, HHUF, 

IFRD, ILLS, IMI, K(M), L, LPS, M, MA, NY, PAD, PC, PH, 

RO, S, TNS, TRTC, UB, UBC, UPS and ZT. Attempts were 

made to trace and borrow all the type specimens from herbaria 

worldwide, but only some of them could be obtained. Some of 

the type specimens are in such bad condition that little 

information could be obtained. In order to obtain the location 

of specimens, original publications were searched. 

Ascostroma and ascomata were examined under an 

Olympus SZ H10 dissecting microscope. Section of the 

fruiting structures was carried out by cryotome or by handcutting. 

Measurements and descriptions of sections of the 

ascomata, hamathecium, asci and ascospores were carried out 

by immersing ascomata in water or in 10% lactic acid. 

Microphotography was taken with material mounted in water, 

cotton blue, Melzer’s reagent or 10–100% lactic acid. 

Terminologies are as in Ulloa and Hanlin (2000). In 

addition, ascomata size is defined as: small-sized: < 300 μm 

diam., medium-sized: from 300 μm to 600 μm diam., largesized: 

> 600 μm diam. 

Question mark (“?”) before family (or genus) name 

means its familial (or generic) status within Pleosporales 

(or some particular family) is uncertain. Other question 

marks after habitats, latin names or other substantives mean 

the correctness of their usages need verification. 

Results 

Molecular phylogeny 

In total, 278 pleosporalean taxa are included in the phylogenetic 

analysis. These form 25 familial clades in the dendrogram, 

i.e. Aigialaceae, Amniculicolaceae, Arthopyreniaceae,


Table 3 Taxa used in the phylogenetic analysis and their corresponding GenBank numbers. Culture and voucher abbreviations are indicated were 

available 

Species Culture/voucher 1 LSU SSU RPB2 TEF1 

Acrocordiopsis patilii BCC 28166 GU479772 GU479736 GU479811 

Acrocordiopsis patilii BCC 28167 GU479773 GU479737 GU479812 

Aigialus grandis BCC 18419 GU479774 GU479738 GU479813 GU479838 

Aigialus grandis JK 5244A GU301793 GU296131 GU371762 

Aigialus mangrovis BCC 33563 GU479776 GU479741 GU479815 GU479840 

Aigialus mangrovis BCC 33564 GU479777 GU479742 GU479816 GU479841 

Aigialus parvus A6 GU301795 GU296133 GU371771 GU349064 

Aigialus parvus BCC 32558 GU479779 GU479743 GU479818 GU479843 

Aigialus rhizophorae BCC 33572 GU479780 GU479745 GU479819 GU479844 

Aigialus rhizophorae BCC 33573 GU479781 GU479746 GU479820 GU479845 

Alternaria alternata CBS 916.96 DQ678082 DQ678031 DQ677980 DQ677927 

Amniculicola immersa CBS 123083 FJ795498 GU456295 GU456358 GU456273 

Amniculicola parva CBS 123092 FJ795497 GU296134 GU349065 

Anteaglonium abbreviatum ANM 925.1 GQ221877 GQ221924 

Anteaglonium abbreviatum GKM 1029 GQ221878 GQ221915 

Anteaglonium globosum ANM 925.2 GQ221879 GQ221925 

Anteaglonium latirostrum L100N 2 GQ221876 GQ221938 

Arthopyrenia salicis 1994 Coppins AY607730 

Arthopyrenia salicis CBS 368.94 AY538339 AY538333 

Ascochyta pisi CBS 126.54 DQ678070 DQ678018 DQ677967 DQ677913 

Ascocratera manglicola BCC 09270 GU479782 GU479747 GU479821 GU479846 

Ascocratera manglicola JK 5262 C GU301799 GU296136 GU371763 

Asteromassaria pulchra CBS 124082 GU301800 GU296137 GU371772 GU349066 

Astrosphaeriella aggregata MAFF 239485 AB524590 AB524449 

Astrosphaeriella aggregata MAFF 239486 AB524591 AB524450 AB539105 AB539092 

Astrosphaeriella bakeriana CBS 115556 GU301801 GU349015 

Astrosphaeriella stellata MAFF 239487 AB524592 AB524451 

Beverwykella pulmonaria CBS 283.53 GU301804 GU371768 

Biatriospora marina CY 1228 GQ925848 GQ925835 GU479823 GU479848 

Bimuria novae-zelandiae CBS 107.79 AY016356 AY016338 DQ470917 DQ471087 

Byssolophis sphaerioides IFRDCC2053 GU301805 GU296140 GU456348 GU456263 

Byssosphaeria jamaicana SMH1403 GU385152 GU327746 

Byssosphaeria rhodomphala GKM L153N GU385157 GU327747 

Byssosphaeria salebrosa SMH2387 GU385162 GU327748 

Byssosphaeria schiedermayeriana GKM1197 GU385161 GU327750 

Byssosphaeria schiedermayeriana GKM152N GU385168 GU327749 

Byssosphaeria villosa GKM204N GU385151 GU327751 

Byssothecium circinans CBS 675.92 AY016357 AY016339 DQ767646 GU349061 

Chaetosphaeronema hispidulum CBS 216.75 EU754144 EU754045 GU371777 

Cochliobolus heterostrophus CBS 134.39 AY544645 AY544727 DQ247790 DQ497603 

Cochliobolus sativus DAOM 226212 DQ678045 DQ677995 DQ677939 

Corynespora cassiicola CBS 100822 GU301808 GU296144 GU371742 GU349052 

Corynespora olivacea CBS 114450 GU301809 GU349014 

Corynespora smithii CABI 5649b GU323201 GU371783 GU349018 

Cucurbitaria berberidis CBS 394.84 GQ387605 GQ387544 

Decaisnella formosa BCC 25616 GQ925846 GQ925833 GU479825 GU479851 

Decaisnella formosa BCC 25617 GQ925847 GQ925834 GU479824 GU479850 

Decorospora gaudefroyi CBS 332.63 EF177849 AF394542


Table 3 (continued) 


Delitschia cf. chaetomioides GKM 1283 GU385172 

Delitschia cf. chaetomioides GKM 3253.2 GU390656 

Delitschia chaetomioides GKM1283 GU385172 GU327752 

Delitschia chaetomioides SMH3253.2 GU390656 GU327753 

Delitschia winteri CBS 225.62 DQ678077 DQ678026 DQ677975 DQ677922 

Didymella exigua CBS 183.55 EU754155 EU754056 

Didymocrea sadasivanii CBS 438 65 DQ384103 DQ384066 

Didymosphaeria futilis CMW 22186 EU552123 

Didymosphaeria futilis HKUCC 5834 GU205219 GU205236 

Dothidotthia aspera CPC 12933 EU673276 EU673228 

Dothidotthia symphoricarpi CBS119687 EU673273 EU673224 

Entodesmium rude CBS 650.86 GU301812 GU349012 

Falciformispora lignatilis BCC 21117 GU371826 GU371834 GU371819 

Falciformispora lignatilis BCC 21118 GU371827 GU371835 GU371820 

Floricola striata JK 5603 K GU479785 GU479751 

Floricola striata JK 5678I GU301813 GU296149 GU371758 

Halomassarina thalassiae BCC 17055 GQ925850 GQ925843 

Halomassarina thalassiae JK 5262D GU301816 GU349011 

Halotthia posidoniae BBH 22481 GU479786 GU479752 

Helicascus nypae BCC 36751 GU479788 GU479754 GU479826 GU479854 

Helicascus nypae BCC 36752 GU479789 GU479755 GU479827 GU479855 

Herpotrichia diffusa CBS 250.62 DQ678071 DQ678019 DQ677968 DQ677915 

Herpotrichia juniperi CBS 200.31 DQ678080 DQ678029 DQ677978 DQ677925 

Herpotrichia macrotricha GKM196N GU385176 GU327755 

Herpotrichia macrotricha SMH269 GU385177 GU327756 

Hypsostroma caimitalense GKM 1165 GU385180 

Hypsostroma saxicola SMH 5005 GU385181 

Hysterium angustatum CBS 123334 FJ161207 FJ161167 FJ161129 FJ161111 

Hysterium angustatum CBS 236.34 FJ161180 GU397359 FJ161117 FJ161096 

Julella avicenniae BCC 18422 GU371823 GU371831 GU371787 GU371816 

Julella avicenniae BCC 20173 GU371822 GU371830 GU371786 GU371815 

Julella avicenniae JK 5326A GU479790 GU479756 

Kalmusia scabrispora MAFF 239517 AB524593 AB524452 AB539093 AB539106 

Kalmusia scabrispora NBRC 106237 AB524594 AB524453 AB539094 AB539107 

Karstenula rhodostoma CBS 690.94 GU301821 GU296154 GU371788 GU349067 

Katumotoa bambusicola MAFF 239641 AB524595 AB524454 AB539095 AB539108 

Keissleriella cladophila CBS 104.55 GU301822 GU296155 GU371735 GU349043 

Keissleriella rara CBS 118429 GU479791 GU479757 

Kirschsteiniothelia elaterascus A22-5A/HKUCC7769 AY787934 AF053727 

Lentithecium aquaticum CBS 123099 GU301823 GU296156 GU371789 GU349068 

Lentithecium arundinaceum CBS 123131 GU456320 GU456298 GU456281 

Lentithecium arundinaceum CBS 619.86 GU301824 GU296157 FJ795473 

Lentithecium fluviatile CBS 122367 GU301825 GU296158 GU349074 

Lepidosphaeria nicotiae CBS 101341 DQ678067 DQ677963 DQ677910 

Leptosphaeria biglobosa CBS 298.36 GU237980 GU238207 

Leptosphaeria biglobosa CBS 303.51 GU301826 GU349010 

Leptosphaeria doliolum CBS 505.75 GU301827 GU296159 GU349069 

Leptosphaeria dryadis CBS 643.86 GU301828 GU371733 GU349009 

Leptosphaerulina argentinensis CBS 569.94 GU301829 GU349008




Leptosphaerulina australis CBS 311.51-T FJ795500 GU456357 GU456272 

Leptosphaerulina australis CBS 317.83 GU301830 GU296160 GU371790 GU349070 

Leptosphearia maculans DAOM 229267 DQ470946 DQ470993 DQ470894 DQ471062 

Letendraea helminthicola CBS 884.85 AY016362 AY016345 

Letendraea padouk CBS 485.70 AY849951 GU296162 

Lindgomyces breviappendiculatus KT 1399 AB521749 AB521734 

Lindgomyces cinctosporae R56-1 AB522431 AB522430 

Lindgomyces cinctosporae R56-3 GU266245 GU266238 

Lindgomyces ingoldianus KH 100 JCM 16479 AB521737 AB521720 

Lindgomyces rotundatus KH 114 JCM 16484 AB521742 AB521725 

Lophiostoma alpigenum GKM 1091b GU385193 

Lophiostoma arundinis CBS 621.86 DQ782384 DQ782383 DQ782386 DQ782387 

Lophiostoma caulium CBS 623.86 GU301833 GU296163 GU371791 

Lophiostoma compressum IFRD 2014 GU301834 GU296164 FJ795457 

Lophiostoma crenatum CBS 629.86 DQ678069 DQ678017 DQ677965 DQ677912 

Lophiostoma fuckelii CBS 101952 DQ399531 

Lophiostoma fuckelii CBS 113432 EU552139 

Lophiostoma fuckelii GKM 1063 GU385192 

Lophiostoma macrostomum CBS 122681 EU552141 

Lophiostoma macrostomum HHUF 27293 AB433274 

Lophiostoma macrostomum KT 635 AB433273 AB521731 

Lophiostoma quadrinucleatum GKM1233 GU385184 GU327760 

Lophiostoma sagittiforme HHUF 29754 AB369267 

Lophiotrema brunneosporum CBS 123095 GU301835 GU296165 GU349071 

Lophiotrema lignicola CBS 122364 GU301836 GU296166 GU349072 

Massarina arundinariae MAFF 239461 AB524596 AB524455 AB539096 AB524817 

Massarina arundinariae NBRC 106238 AB524597 AB524456 AB539097 AB524818 

Lophiotrema nucula CBS 627.86 GU301837 GU296167 GU371792 GU349073 

Loratospora aestuarii JK 5535B GU301838 GU296168 GU371760 

Macroventuria anomochaeta CBS 525.71 GU456315 GU456346 GU456262 

Massaria anomia CBS 123109 GU301792 GU296130 GU349062 

Massaria anomia CBS 591.78 GU301839 GU296169 GU371769 

Massaria ariae M52 HQ599382 HQ599456 HQ599322 

Massaria aucupariae M49 HQ599384 HQ599455 HQ599324 

Massaria campestris M28 HQ599385 HQ599449 HQ599459 HQ599325 

Massaria conspurcata M14 HQ599393 HQ599441 HQ599333 

Massaria gigantispora M26 HQ599397 HQ599447 HQ599337 

Massaria inquinans M19 HQ599402 HQ599444 HQ599460 HQ599342 

Massaria lantanae M18 HQ599406 HQ599443 HQ599346 

Massaria macra M3 HQ599408 HQ599450 HQ599348 

Massaria mediterranea M45 HQ599417 HQ599452 HQ599357 

Massaria platanoidea M7 HQ599420 HQ599457 HQ599462 HQ599359 

Massaria pyri M21 HQ599424 HQ599445 HQ599363 

Massaria vindobonensis M27 HQ599429 HQ599448 HQ599464 HQ599368 

Massaria vomitoria M13 HQ599437 HQ599440 HQ599466 HQ599375 

Massarina cisti CBS 266.62 FJ795447 FJ795490 FJ795464 

Massarina eburnea CBS 473.64 GU301840 GU296170 GU371732 GU349040 

Massarina igniaria CBS 845.96 GU301841 GU296171 GU371793 

Massarina ricifera JK 5535 F GU479793 GU479759




Massariosphaeria phaeospora CBS 611.86 GU301843 GU296173 GU371794 

Mauritiana rhizophorae BCC 28866 GU371824 GU371832 GU371796 GU371817 

Mauritiana rhizophorae BCC 28867 GU371825 GU371833 GU371797 GU371818 

Melanomma pulvis-pyrius CBS 124080 GU456323 GU456302 GU456350 GU456265 

Melanomma pulvis-pyrius CBS 371.75 GU301845 GU371798 GU349019 

Melanomma pulvis-pyrius SMH 3291 GU385197 

Melanomma rhododendri ANM 73 GU385198 

Misturatosphaeria aurantonotata GKM1238 GU385173 GU327761 

Misturatosphaeria aurantonotata GKM1280 GU385174 GU327762 

Misturatosphaeria claviformis GKM1210 GU385212 GU327763 

Misturatosphaeria kenyensis GKM1195 GU385194 GU327767 

Misturatosphaeria kenyensis GKM L100Na GU385189 GU327766 

Misturatosphaeria minima GKM169N GU385165 GU327768 

Misturatosphaeria tennesseensis ANM911 GU385207 GU327769 

Misturatosphaeria uniseptata SMH4330 GU385167 GU327770 

Monascostroma innumerosum CBS 345.50 GU301850 GU296179 GU349033 

Monotosporella tuberculata CBS 256.84 GU301851 GU349006 

Montagnula anthostomoides CBS 615.86 GU205223 GU205246 

Montagnula opulenta CBS 168.34 DQ678086 AF164370 DQ677984 

Morosphaeria ramunculicola BCC 18405 GQ925854 GQ925839 

Morosphaeria ramunculicola JK 5304B GU479794 GU479760 GU479831 

Morosphaeria velataspora BCC 17059 GQ925852 GQ925841 

Morosphaeria velataspora BCC 17058 GQ925851 GQ925840 

Massariosphaeria grandispora CBS 613 86 GU301842 GU296172 GU371725 GU349036 

Massariosphaeria typhicola CBS 123126 GU301844 GU296174 GU371795 

Neophaeosphaeria filamentosa CBS 102202 GQ387577 GQ387516 GU371773 GU349084 

Neotestudina rosatii CBS 690.82 DQ384069 

Neottiosporina paspali CBS 331.37 EU754172 EU754073 GU371779 GU349079 

Ophiosphaerella herpotricha CBS 240.31 DQ767656 DQ767650 DQ767645 DQ767639 

Ophiosphaerella herpotricha CBS 620.86 DQ678062 DQ678010 DQ677958 DQ677905 

Ophiosphaerella sasicola MAFF 239644 AB524599 AB524458 AB539098 AB539111 

Paraconiothyrium minitans CBS 122788 EU754173 EU754074 GU371776 GU349083 

Paraphaeosphaeria michotii CBS 591.73 GU456326 GU456305 GU456352 GU456267 

Paraphaeosphaeria michotii CBS 652.86 GU456325 GU456304 GU456351 GU456266 

Phaeosphaeria ammophilae CBS 114595 GU301859 GU296185 GU371724 GU349035 

Phaeosphaeria avenaria CBS 602.86 AY544684 AY544725 DQ677941 DQ677885 

Phaeosphaeria avenaria DAOM 226215 AY544684 AY544725 DQ677941 DQ677885 

Phaeosphaeria brevispora MAFF 239276 AB524600 AB524459 AB539099 AB539112 

Phaeosphaeria brevispora NBRC 106240 AB524601 AB524460 AB539100 AB539113 

Phaeosphaeria caricis CBS 120249 GU301860 GU349005 

Phaeosphaeria elongata CBS 120250 GU456327 GU456306 GU456345 GU456261 

Phaeosphaeria eustoma CBS 573.86 DQ678063 DQ678011 DQ677959 DQ677906 

Phaeosphaeria luctuosa CBS 308.79 GU301861 GU349004 

Phaeosphaeria nigrans CBS 576.86 GU456331 GU456356 GU456271 

Phaeosphaeria nodorum CBS 259.49 GU456332 GU456285 

Phaeosphaeria oryzae CBS 110110 GQ387591 GQ387530 

Phaeosphaeriopsis musae CBS 120026 GU301862 GU296186 GU349037 

Phoma apiicola CBS 285.72 GU238040 GU238211 

Phoma betae CBS 109410 EU754178 EU754079 GU371774 GU349075




Phoma complanata CBS 268.92 EU754180 EU754081 GU371778 GU349078 

Phoma cucurbitacearum CBS 133.96 GU301863 GU371767 

Phoma exigua CBS 431.74 EU754183 EU754084 GU371780 GU349080 

Phoma glomerata CBS 528.66 EU754184 EU754085 GU371781 GU349081 

Phoma herbarum CBS 276.37 DQ678066 DQ678014 DQ677962 DQ677909 

Phoma radicina CBS 111.79 EU754191 EU754092 GU349076 

Phoma valerianae CBS 630.68 GU238150 GU238229 

Phoma vasinfecta CBS 539.63 GU238151 GU238230 

Phoma violicola CBS 306.68 GU238156 GU238231 

Phoma zeae-maydis CBS 588.69 EU754192 EU754093 GU371782 GU349082 

Platychora ulmi CBS 361.52 EF114702 EF114726 

Lophiostoma compressum GKM1048 GU385204 GU327772 

Lophiostoma scabridisporum BCC 22836 GQ925845 GQ925832 GU479829 GU479856 

Lophiostoma scabridisporum BCC 22835 GQ925844 GQ925831 GU479830 GU479857 

Pleomassaria siparia CBS 279.74 DQ678078 DQ678027 DQ677976 DQ677923 

Pleospora ambigua CBS 113979 AY787937 

Pleospora herbarum CBS 191.86 DQ247804 DQ247812 DQ247794 DQ471090 

Polyplosphaeria fusca CBS 125425 AB524607 AB524466 AB524822 

Polyplosphaeria fusca MAFF 239687 AB524606 AB524465 

Preussia funiculata CBS 659.74 GU301864 GU296187 GU371799 GU349032 

Preussia lignicola CBS 264.69 GU301872 GU296197 GU371765 GU349027 

Preussia terricola DAOM 230091 AY544686 AY544726 DQ470895 DQ471063 

Prosthemium betulinum CBS 127468 AB553754 AB553644 

Prosthemium canba JCM 16966 AB553760 AB553646 

Prosthemium orientale JCM 12841 AB553748 AB553641 

Prosthemium stellare CBS 126964 AB553781 AB553650 

Pseudotetraploa curviappendiculata CBS 125426 AB524610 AB524469 AB524825 

Pseudotetraploa curviappendiculata MAFF 239495 AB524608 AB524467 

Pseudotetraploa javanica MAFF 239498 AB524611 AB524470 AB524826 

Pseudotetraploa longissima MAFF 239497 AB524612 AB524471 AB524827 

Pseudotrichia guatopoensis SMH4535 GU385202 GU327774 

Pyrenochaeta acicola CBS 812.95 GQ387602 GQ387541 

Pleurophoma cava CBS 257.68 EU754199 EU754100 

Pyrenochaeta corn CBS 248.79 GQ387608 GQ387547 

Pyrenochaeta nobilis CBS 292.74 GQ387615 GQ387554 

Pyrenochaeta nobilis CBS 407.76 DQ678096 DQ677991 DQ677936 

Pyrenochaeta quercina CBS 115095 GQ387619 GQ387558 

Pyrenochaeta unguis-hominis CBS 378.92 GQ387621 GQ387560 

Pyrenochaetopsis decipiens CBS 343.85 GQ387624 GQ387563 

Pyrenophora phaeocomes DAOM 222769 DQ499596 DQ499595 DQ497614 DQ497607 

Pyrenophora tritici-repentis OSC 100066 AY544672 DQ677882 

Quadricrura bicornis CBS 125427 AB524613 AB524472 AB524828 

Quadricrura meridionalis CBS 125684 AB524614 AB524473 AB524829 

Quadricrura septentrionalis CBS 125428 AB524617 AB524476 AB524832 

Quintaria lignatilis BCC 17444 GU479797 GU479764 GU479832 GU479859 

Quintaria lignatilis CBS 117700 GU301865 GU296188 GU371761 

Quintaria submersa CBS 115553 GU301866 GU349003 

Repetophragma ontariense HKUCC 10830 DQ408575 DQ435077 

Rimora mangrovei JK 5246A GU301868 GU296193 GU371759




Rimora mangrovei JK 5437B GU479798 GU479765 

Roussoella hysterioides CBS 125434 AB524622 AB524481 AB539102 AB539115 

Roussoella hysterioides MAFF 239636 AB524621 AB524480 AB539101 AB539114 

Roussoella pustulans MAFF 239637 AB524623 AB524482 AB539103 AB539116 

Roussoellopsis tosaensis MAFF 239638 AB524625 AB539104 AB539117 

Saccothecium sepincola CBS 278.32 GU301870 GU296195 GU371745 GU349029 

Salsuginea ramicola KT 2597.1 GU479800 GU479767 GU479833 GU479861 

Salsuginea ramicola KT 2597.2 GU479801 GU479768 GU479834 GU479862 

Setomelanomma holmii CBS 110217 GU301871 GU296196 GU371800 GU349028 

Setosphaeria monoceras AY016368 AY016368 

Massaria platani CBS 221.37 DQ678065 DQ678013 DQ677961 DQ677908 

Sporormiella minima CBS 524.50 DQ678056 DQ678003 DQ677950 DQ677897 

Stagonospora macropycnidia CBS 114202 GU301873 GU296198 GU349026 

Tetraploa aristata CBS 996.70 AB524627 AB524486 AB524836 

Tetraplosphaeria nagasakiensis MAFF 239678 AB524630 AB524489 AB524837 

Lophiostoma macrostomoides GKM1033 GU385190 GU327776 

Lophiostoma macrostomoides GKM1159 GU385185 GU327778 

Thyridaria rubronotata CBS 419.85 GU301875 GU371728 GU349002 

Tingoldiago graminicola KH 68 AB521743 AB521726 

Trematosphaeria pertusa CBS 122368 FJ201990 FJ201991 FJ795476 GU456276 

Trematosphaeria pertusa CBS 122371 GU301876 GU348999 GU371801 GU349085 

Trematosphaeria pertusa SMH 1448 GU385213 

Triplosphaeria cylindrica MAFF 239679 AB524634 AB524493 

Triplosphaeria maxima MAFF 239682 AB524637 AB524496 

Triplosphaeria yezoensis CBS 125436 AB524638 AB524497 AB524844 

Ulospora bilgramii CBS 110020 DQ678076 DQ678025 DQ677974 DQ677921 

Verruculina enalia BCC 18401 GU479802 GU479770 GU479835 GU479863 

Verruculina enalia BCC 18402 GU479803 GU479771 GU479836 GU479864 

Westerdykella cylindrica CBS 454.72 AY004343 AY016355 DQ470925 DQ497610 

Westerdykella dispersa CBS 508.75 DQ468050 U42488 

Westerdykella ornata CBS 379.55 GU301880 GU296208 GU371803 GU349021 

Wicklowia aquatica AF289-1 GU045446 

Wicklowia aquatica CBS 125634 GU045445 GU266232 

Xenolophium applanatum CBS 123123 GU456329 GU456312 GU456354 GU456269 

Xenolophium applanatum CBS 123127 GU456330 GU456313 GU456355 GU456270 

Zopfia rhizophila CBS 207.26 DQ384104 L76622 

1 BCC Belgian Coordinated Collections of Microorganisms; CABI International Mycological Institute, CABI-Bioscience, Egham, Bakeham Lane, 

U.K.; CBS Centraalbureau voor Schimmelcultures, Utrecht, The Netherlands; DAOM Plant Research Institute, Department of Agriculture 

(Mycology), Ottawa, Canada; DUKE Duke University Herbarium, Durham, North Carolina, U.S.A.; HHUF Herbarium of Hirosaki University, 

Japan; IFRDCC Culture Collection, International Fungal Research & Development Centre, Chinese Academy of Forestry, Kunming, China; 

MAFF Ministry of Agriculture, Forestry and Fisheries, Japan; NBRC NITE Biological Resource Centre, Japan; OSC Oregon State University 

Herbarium, U.S.A.; UAMH University of Alberta Microfungus Collection and Herbarium, Edmonton, Alberta, Canada; UME Herbarium of the 

University of Umeå, Umeå, Sweden; Culture and specimen abbreviations: ANM A.N. Miller; CPC; P.W. Crous; EB E.W.A. Boehm; EG E.B.G. 

Jones; GKM G.K. Mugambi; JK J. Kohlmeyer; KT K. Tanaka; SMH S.M. Huhndorf 

Cucurbitariaceae/Didymosphaeriaceae, Delitschiaceae, 

Didymellaceae, Dothidotthiaceae, Hypsostromataceae, Lentitheciaceae, 

Leptosphaeriaceae, Lindgomycetaceae, 

Lophiostomataceae, Massariaceae, Massarinaceae, Melanommataceae, 

Montagnulaceae, Morosphaeriaceae, 

Phaeosphaeriaceae, Pleomassariaceae, Pleosporaceae, 

Sporormiaceae, Testudinaceae/Platystomaceae, Tetraplosphaeriaceae, 

Trematosphaeriaceae and Zopfiaceae (Plate 1). 

Of these, Lentitheciaceae, Massarinaceae, Montagnulaceae, 

Morosphaeriaceae and Trematosphaeriaceae form a robust


clade in the present study and in previous studies (Schoch et 

al. 2009; Zhang et al. 2009a, b). We thus emended the 

suborder, Massarineae, to accommodate them. 

Pleosporales suborder Massarineae Barr, Mycologia 71: 

948. (1979a). emend. 

Habitat freshwater, marine or terrestrial environment, 

saprobic. Ascomata solitary, scattered or gregarious, globose, 

subglobose, conical to lenticular, immersed, erumpent 

to superficial, papillate, ostiolate. Hamathecium of dense or 

rarely few, filliform pseudoparaphyses. Asci bitunicate, 

fissitunicate, cylindrical, clavate or broadly clavate, pedicellate. 

Ascospores hyaline, pale brown or brown, 1 to 3 or 

more transverse septa, rarely muriform, narrowly fusoid, 

fusoid, broadly fusoid, symmetrical or asymmetrical, with 

or without sheath. 

Accepted genera of Pleosporales 

Acrocordiopsis Borse & K.D. Hyde, Mycotaxon 34: 535 

(1989). (Pleosporales, genera incertae sedis) 

Generic description 

Habitat marine, saprobic. Ascomata seated in blackish 

stroma, scattered or gregarious, superficial, conical to 

semiglobose, ostiolate, carbonaceous. Hamathecium of 

dense, long trabeculate pseudoparaphyses. Asci 8-spored, 

cylindrical with pedicels and conspicuous ocular chambers. 

Ascospores hyaline, 1-septate, obovoid to broadly fusoid. 

Anamorphs reported for genus: none. 

Literature: Alias et al. 1999; Barr 1987a; Borse and Hyde 

1989. 

Type species 

Acrocordiopsis patilii Borse & K.D. Hyde, Mycotaxon 34: 

536 (1989). (Fig. 1) 

Ascomata 1–2 mm high×1.8–3 mmdiam.,scatteredor 

gregarious, superficial, conical or semiglobose, with a 

flattened base not easily removed from the substrate, ostiolate, 

black, very brittle and carbonaceous and extremely difficult to 

cut (Fig. 1a and b). Peridium 250–310 μm thick, to 600 μm 

thick near the apex, thinner at the base, comprising three 

types of cells; outer cells pseudoparenchymatous, small 

heavily pigmented thick-walled cells of textura epidermoidea, 

cells 0.6–1×6–10 μm diam., cell wall 5–9 μm thick; 

cells near the substrate less pigmented, composed of cells of 

textura prismatica, cellwalls1–3(−5) μm thick; inner cells 

less pigmented, comprised of hyaline to pale brown thinwalled 

cells, merging with pseudoparaphyses (Fig. 1c, d and 

e). Hamathecium of dense, long trabeculate pseudoparaphyses, 

ca. 1 μm broad, embedded in mucilage, hyaline, 

Fig. 1 Acrocordiopsis patilii (from IMI 297769, holotype). a 

Ascomata on the host surface. b Section of an ascoma. c Section of 

lateral peridium. d Section of the apical peridium. e Section of the 

basal peridium. Note the paler cells of textura prismatica. f 

Cylindrical ascus. g Cylindrical ascus in pseudoparaphyses. h, i 

One-septate ascospores. Scale bars: a=3 mm, b=0.5 mm, c=200 μm, 

d, e =50 μm, f–i=20 μm 

anastomosing and sparsely septate. Asci 140–220×13– 

17 μm (x ¼ 165:3 15:6mm, n=10), 8-spored, bitunicate, 

fissitunicate, cylindrical, with short pedicels, 15–25(−40) μm 

long, with a large and conspicuous ocular chamber (Fig. 1f 

and g). Ascospores 17.5–25×12.5–15(−20) μm 

(x ¼ 21:5 13:6mm, n=10), uniseriate to partially overlapping, 

ovoid or ellipsoidal, hyaline, 1-septate, not constricted 

at the septum, smooth-walled (Fig. 1h and i). 

Anamorph: none reported. 

Material examined: INDIA, Indian Ocean, Malvan 

(Maharashtra), on intertidal wood of Avicennia alba Bl., 

30 Oct. 1981 (IMI 297769, holotype). 

Notes 

Morphology 

Acrocordiopsis was formally established by Borse and 

Hyde (1989) as a monotypic genus represented by A. patilii 

based on its “conical or semiglobose superficial carbonaceous 

ascomata, trabeculate pseudoparaphyses, cylindrical, 

bitunicate, 8-spored asci, and hyaline, 1-septate, obovoid or 

ellipsoid ascospores”. Acrocordiopsis patilii was first 

collected from mangrove wood (Indian Ocean) as a marine 

fungus, and a second marine Acrocordiopsis species 

was reported subsequently from Philippines (Alias et al. 

1999). Acrocordiopsis is assigned to Melanommataceae 

(Melanommatales sensu Barr 1983) based on its ostiolate 

ascomata and trabeculate pseudoparaphyses (Borse and 

Hyde 1989). Morphologically, Acrocordiopsis is similar to 

Astrosphaeriella sensu stricto based on the conical ascomata 

and the brittle, carbonaceous peridium composed of thickwalled 

black cells with rows of palisade-like parallel cells at 

the rim area. Ascospores of Astrosphaeriella are, however, 

elongate-fusoid, usually brown or reddish brown and 

surrounded by a gelatinous sheath when young; as such 

they are readily distinguishable from those of Acrocordiopsis. 

A new family (Acrocordiaceae) was introduced by Barr 

(1987a) to accommodate Acrocordiopsis. This proposal, 

however, has been rarely followed and Jones et al. (2009) 

assigned Acrocordiopsis to Melanommataceae. 

Phylogenetic study 

Acrocordiopsis patilii nested within an unresolved clade 

within Pleosporales (Suetrong et al. 2009). Thus its familial 

placement is unresolved, but use of the Acrocordiaceae 

could be reconsidered with more data. 

b

Fungal Diversity


Concluding remarks 

Acrocordiopsis, Astrosphaeriella sensu stricto, Mamillisphaeria, 

Caryospora and Caryosporella are morphologically 

similar as all have very thick-walled carbonaceous ascomata, 

narrow pseudoparaphyses in a gelatinous matrix (trabeculae) 

and bitunicate, fissitunicate asci. Despite their similarities, the 

shape of asci and ascospores differs (e.g. Mamillisphaeria has 

sac-like asci and two types of ascospores, brown or hyaline, 

Astrosphaeriella has cylindro-clavate asci and narrowly fusoid 

ascospores, both Acrocordiopsis and Caryosporella has 

cylindrical asci, but ascospores of Caryosporella are reddish 

brown). Therefore, the current familial placement of Acrocordiopsis 

cannot be determined. All generic types of Astrosphaeriella 

sensu stricto, Mamillisphaeria and Caryospora 

should be recollected and isolated for phylogenetic study. 

Aigialus Kohlm. & S. Schatz, Trans. Br. Mycol. Soc. 85: 

699 (1985). (Aigialaceae) 


Habitat marine, saprobic. Ascomata mostly subglobose in 

front view, fusoid in sagittal section, rarely subglobose, 

scattered, immersed to erumpent, papillate, ostiolate, ostiole 

rounded or slit-like, periphysate. Peridium 2-layered. Hamathecium 

of trabeculate pseudoparaphyses. Asci 8-spored, 

cylindrical, pedicellate, with an ocular chamber and conspicuous 

apical ring. Ascospores ellipsoidal to fusoid, muriform, 

yellow brown to brown, with terminal appendages. 


Literature: Eriksson 2006; Jones et al. 2009; Kohlmeyer 

and Schatz 1985; Lumbsch and Huhndorf 2007. 

Type species 

Aigialus grandis Kohlm. & S. Schatz, Trans. Br. Mycol. 

Soc. 85: 699 (1985). (Fig. 2) 

Ascomata 1–1.25 mm high×1–1.3 mm diam. in front view, 

250–400 μm broad in sagittal section, vertically flattened 

subglobose, laterally compressed, scattered, immersed to semiimmersed, 

papillate, with an elongated furrow at the top of the 

papilla, wall black, carbonaceous, ostiolate, ostiole filled with 

branched or forked septate periphyses (Fig. 2a). Peridium 70– 

100 μm thick laterally, up to 150 μm thick at the apex, thinner 

at the base, comprising two cell types, outer layer composed of 

small heavily pigmented thick-walled pseudoparenchymatous 

cells, cells 1–2 μm diam., cell wall 2–5 μm thick, inner layer 

thin, composed of small hyaline cells (Fig. 2b). Hamathecium 

of dense, very long trabeculate pseudoparaphyses, 0.8–1.2 μm 

broad, embedded in mucilage, anastomosing and branching 

above the asci. Asci 450–640×22–35 μm (x ¼ 505 30mm, 

n=10), 8-spored, bitunicate, fissitunicate, cylindrical to 

cylindro-clavate, with a long furcate pedicel, 90–180 μm long, 

with a low truncate ocular chamber and a refractive apical 

apparatus (to 12 μm wide×4 μm high) (Fig. 2e and f). 

Ascospores 75–95×15–26 μm (x ¼ 84:3 17:5mm, n=10), 

obliquely uniseriate and partially overlapping, broadly fusoid 

to fusoid with narrowly rounded ends in front view, flat on one 

side from side view (14–20 μm thick), yellowish brown, apical 

cells usually hyaline, muriform, with 14–17(−18) transversal 

septa, 1–3 longitudinal septa in most cells, slightly constricted 

at the septa, with a gelatinous cap at each end (Fig. 2c and d). 


Material examined: BELIZE, Wee-Wee Cay, on submerged 

wood of roots and branches of Rhizophora mangle 

L., Mar. 1983, leg. J. Kohlmeyer (NY, J.K. 4332b, isotype). 

Notes 

Morphology 

Aigialus was formally established by Kohlmeyer and Schatz 

(1985) based on its immersed or semi-immersed ascomata 

with periphysate ostiole, trabeculate pseudoparaphyses, cylindrical 

and fissitunicate asci, and distinctive muriform ascospores 

with gelatinous sheath or caps. There are five accepted 

species in the genus, namely A. grandis, A. mangrovei Borse, 

A. parvus S. Schatz & Kohlm., A. rhizophorae Borse and A. 

striatispora K.D. Hyde (Jones et al. 2009). Aigialus was first 

assigned to the Melanommatales, but its familial status was 

uncertain (Kohlmeyer and Schatz 1985). Barr (1990b) 

included Aigialus in Massariaceae based on its conspicuous 

apical ring in the asci and ascospore characters, and this has 

subsequently been widely followed (Eriksson 2006; Hawksworth 

et al. 1995; Kirk et al. 2001; Lumbsch and Huhndorf 2007). 


The generic type of Aigialus (A. grandis) together with 

other three marine species, i.e. A. mangrovei, A. parvus as well 

as A. rhizophorae form a robust clade on the phylogenetic 

tree. Thus a new family, Aigialaceae, was introduced to 

accommodate Aigialus together with Ascocratera and Rimora 

(Suetrong et al. 2009). 


The pleosporalean status of Aigialus has been phylogenetically 

verified, and the single branch containing Aigialus, 

Ascocratera and Rimora represents a familial rank of 

Aigialaceae (Suetrong et al. 2009). 

Amniculicola Yin. Zhang & K.D. Hyde, Mycol. Res. 112: 

1189 (2008). (Amniculicolaceae) 


Habitat freshwater, saprobic. Ascomata solitary, scattered, 

or in small groups, initially immersed, becoming erumpent,


Fig. 2 Aigialus grandis (from NY, J.K. 4332b, isotype). a Ascomata 

on the host surface. Note the longitudinal slit-like furrow which is the 

ostiole. b Section of the peridium. c, d Released ascospores. e 

Ascospores in ascus. Note the conspicuous apical ring. f Cylindrical 

ascus with a long pedicel. Scale bars: a=1 mm, b=200 μm, c–f= 

20 μm


to nearly superficial, globose, subglobose to conical, wall 

black, roughened; apex well differentiated into two tuberculate 

flared lips surrounding a slit-like ostiole. Peridium thin, 2- 

layered, outer layer composed of small heavily pigmented 

thick-walled cells of textura angularis, inner layer composed 

of hyaline thin-walled cells of textura angularis. Hamathecium 

of dense, long trabeculate pseudoparaphyses, embedded 

in mucilage, anastomosing between and above the asci. Asci 

8-spored, bitunicate, fissitunicate, cylindrical to narrowly 

fusoid, short pedicellate, with an ocular chamber and a small 

apical apparatus. Ascospores fusoid, hyaline, 1-septate, 

constricted at the septum, surrounded by an irregular hyaline 

gelatinous sheath. 

Anamorphs reported for genus: Anguillospora longissima, 

Spirosphaera cupreorufescens and Repetophragma 

ontariense (Zhang et al. 2008c, 2009c). 

Literature: Zhang et al. 2008c, 2009a, c. 

Type species 

Amniculicola lignicola Ying Zhang & K.D. Hyde, Mycol. 

Res. 112: 1189 (2008). (Fig. 3) 

Ascomata 350–450 μm high×300–500 μm diam., 

solitary, scattered, or in small groups of 2–3, initially 

immersed, becoming erumpent, to nearly superficial, with 

basal wall remaining immersed in host tissue, globose, 

subglobose, broadly or narrowly conical, often laterally 

flattened, with a flattened base not easily removed from the 

substrate, wall black, roughened, often bearing remnants of 

wood fibers; apex well differentiated into two tuberculate 

flared lips surrounding a slit-like ostiole, 150–250 μm long, 

filled with a purplish amorphous matter, oriented in the axis 

of the wood fibers; underlying wood stained pale purple 

(Fig. 3a and b). Peridium 40–55 μm thick laterally, up to 

120 μm thick at the apex, thinner at the base, coriaceous, 

2-layered, outer layer composed of small heavily pigmented 

thick-walled cells of textura angularis, cells4–9 μm diam., 

cell wall 2–3 μm thick, apex cells smaller and walls thicker, 

inner layer composed of hyaline thin-walled cells of textura 

angularis, 8–16 μm diam., in places with columns of textura 

prismatica, oriented perpendicular to the ascomatal surface, 

and larger, paler cells of textura prismatica towards the 

interior and at the base, 10–25 μm (Fig. 3c, d and e). 

Hamathecium of dense, long trabeculate pseudoparaphyses 


Fig. 3 Amniculicola lignicola (from PC 0092661, holotype). a 

Superficial ascomata gregarious on the host surface. b An erumpent 

ascoma with elongated papilla and slit-like ostiole. c Habitat section of 

a superficial ascoma. d, e Section of an ascoma and the partial 

peridium. f Cylindrical 8-spored ascus with a short pedicel. g Hyaline, 

1-septate broadly fusoid ascospores. Scale bars: a=1 mm, b–d= 

100 μm, e=50 μm, f, g=20 μm 

1- (rarely 2- to 3-) septate, fusoid, reddish brown, constricted 

at the main septum. 

Anamorphs reported for genus: Exosporiella (= Phanerocorynella) 

(Sivanesan 1983).


Fig. 4 Anomalemma epochnii (from K(M):143936, syntype). a 

Gregarious ascomata on the host surface. b, c Bitunicate asci. Note the 

wide pseudoparaphyses. d Section of the apical peridium comprising 

thick-walled cells of textura angularis. e–h Fusoid to broadly fusoid 

ascospores. Scale bars: a=0.5 mm, b–h=20 μm


Literature: Berkeley and Broome 1866; Keissler 1922;Massee 

1887; Saccardo 1878a; Sivanesan 1983. 

Type species 

Anomalemma epochnii (Berk. & Broome) Sivan., Trans. 

Br. Mycol. Soc. 81: 328 (1983). (Fig. 4) 

≡ Sphaeria epochnii Berk. & Broome, Ann. Mag. nat. 

Hist., Ser. 3 18: 128 (1866). 

Ascomata 340–500 μm high×170–286 μm diam., gregarious 

on the intertwined hyphae, superficial, papillate, wall 

black, coriaceous, roughened (Fig. 4a). Peridium composed of 

two types of cells, outer layer 17–22 μm wide, composed of 

heavily pigmented thick-walled cells of textura angularis, 

cells up to 8×13 μm diam., cell wall 1–1.5 μm thick, inner 

layer 30–34 μm thick, composed of hyaline thin-walled cells 

(Fig. 4d). Hamathecium of dense, long cellular pseudoparaphyses, 

2–4 μm broad, septate. Asci 75–108×9.5–12.5 μm 

(x ¼ 92:8 11:1mm, n=10), 8-spored, bitunicate, fissitunicate, 

dehiscence not observed, cylindro-clavate to clavate, with 

a furcate pedicel up to 6–25 μm long, with a small ocular 

chamber best seen in immature asci (ca. 2μm wide×1 μm 

high) (Fig. 4b and c). Ascospores 20–25(−30)×5–7.5 μm 

(x ¼ 23:1 6:3mm, n=10), obliquely uniseriate and partially 

overlapping to biseriate, fusoid to narrowly fusoid with narrowly 

rounded ends, brown, 1-septate, rarely 2- to 3-septate, deeply 

constricted at the median septum, smooth (Fig. 4e, f, g and h). 

Anamorph: Exosporiella fungorum (Fr.) P. Karst. 

(Sivanesan 1983). 

= Epochnium fungorum Fr., Syst. mycol. 3: 449 (1832). 

Mycelium composed of branched, septate, pale brown 

hyphae. Stroma none. Conidiophores macronematous or 

semi-macronematous, mononematous, hyaline, smooth, 

branched towards the apex. Conidiogenous cells monoblastic, 

cylindrical or doliform. Conidia cylindrical or 

ellipsoidal, dry, 3-4-septate, smooth, hyaline or pale brown. 

Material examined: UK, England, Warleigh near Bath, 

on fungus on bark (Epochnium sp.), Mar. 1866, leg. 

Warbright? (K(M):143936, syntype, ex herb. C.E. 

Broome). 

Notes 

Morphology 

Sphaeria epochnii was first described and illustrated by 

Berkeley and Broome (1866) from Britain and the anamorphic 

stage is the hyphomycetous Epochniella fungorum. Sphaeria 

epochnii has subsequently been transferred to Melanomma (as 

M. epochnii (Berk. & Broome) Sacc.; Saccardo 1878a), 

Byssosphaeria (as B. epochnii (Berk. & Broome) Cooke; 

Massee 1887) andChaetosphaeria (as C. epochnii (Berk. & 

Broome) Keissl.; Keissler 1922). The deposition of Sphaeria 

epochnii in Chaetosphaeria is obviously unacceptable, as 

Chaetosphaeria has unitunicate asci. Melanomma has been 

reported having Aposphaeria or Pseudospiropes anamorphs, 

which differs from Exosporiella (Sivanesan 1983). In addition, 

the presence of well developed prosenchymatous stroma 

in Sphaeria epochnii can also readily distinguish it from 

Melanomma (Sivanesan 1983). 

The gregarious ascomata and formation of prosenchymatous 

stroma of Anomalemma resembles those of Cucurbitaria, 

but the pleosporaceous dictyosporous ascospores of Cucurbitaria 

readily distinguish it from Anomalemma epochnii. In 

addition, the pseudoparenchymatous peridium, fungicolous 

habitat and brown 1-septate ascospores, which later becoming 

3-septate differ from any other pleosporalean genus. Thus a 

new genus, Anomalemma, was introduced to accommodate it 

(Sivanesan 1983). Anomalemma is presently monotypic. 


None. 


Anomalemma epochnii certainly resembles Byssosphaeria 

in its ascomata clustering together in groups on closely 

intertwined hyphae and brown ascospores, and may well be 

included in this genus. Its fungicolous habitat, however, 

distinguishes it from Byssosphaeria. 

Appendispora K.D. Hyde, Sydowia 46: 29 (1994a). 

(?Didymellaceae) 


Habitat terrestrial, saprobic. Ascomata small, clustered, 

immersed, subglobose or irregularly pyriform. Peridium 

thin. Hamathecium of dense, long trabeculate pseudoparaphyses. 

Asci 8-spored, bitunicate, fissitunicate, cylindrical, 

apical rounded with ocular chamber and faint ring, with 

short pedicels. Ascospores uniseriate to partially overlapping, 

fusoid, brown, 1-septate, slightly constricted at the septum. 


Literature: Hyde 1994a. 

Type species 

Appendispora frondicola K.D. Hyde, Sydowia 46: 30 

(1994a). (Fig. 5) 

Ascomata 120–280 μm high×180–280 μm diam., clustered, 

immersed with minute ostioles visible through cracks 

or blackened dots on the host surface, subglobose or 

irregularly pyriform (Fig. 5a and b). Peridium 40 μm thick, 

comprising two types of cells; outer cells, small heavily 

pigmented thick-walled cells of textura angularis, innercells 

compressed, hyaline. Hamathecium of dense, very long 

trabeculate pseudoparaphyses, ca. 1μm broad, embedded in 

mucilage, hyaline, anastomosing (Fig. 5e). Asci 130–144×11–


Fig. 5 Appendispora frondicola 

(from BRIP 21354, holotype). 

a Immersed ascomata on 

host surface. b Valsoid configuration 

of the ascomata. c Cylindrical 

ascus. d Squash showing 

asci and numerous pseudoparaphyses. 

e Thin strands of 

anastomosing pseudoparaphyses. 

f, g Ascospores with one or two 

appendages. Scale bars: 

a, b=0.5 mm, c–g=30 μm 

13 μm, 8-spored, bitunicate, fissitunicate, cylindrical, with an 

ocular chamber and faint ring, with short pedicels (Fig. 5c and 

d). Ascospores 21–30×7–9 μm, uniseriate to partially overlapping, 

fusoid, brown, 1-septate, slightly constricted at the 

septum, with an irregular ridged ornamentation and 3–5 

narrow appendages at each end (Fig. 5f and g). 


Material examined: BRUNEL, Jalan, Muara, Simpang 

835, on dead rachis of Oncosperma horridum on forest floor, 

Nov. 1992, K.D. Hyde 1652 (BRIP 21354, holotype). 

Notes 

Morphology 

Appendispora was described as a saprobe of palm, and is 

characterized by small, immersed ascomata, bitunicate, 

fissitunicate asci, trabeculate pseudoparaphyses, brown, 1- 

septate, appendaged ascospores with irregular wall striations 

(Hyde 1994a).Basedonitstrabeculatepseudoparaphyses 

embedded within gel matrix and its brown 

ascospores, Appendispora was assigned to Didymosphaeriaceae 

(Barr 1987b; Hyde1994a). 


None. 


The saprobic habitat and association with monocots, 

cylindrical asci, trabeculate pseudoparaphyses as 

well as its brown, 1-septate ascospores make it 

difficult to determine a better phylogenetic position than 

Didymellaceae.


Ascorhombispora L. Cai & K.D. Hyde, Cryptog. Mycol. 

28: 294 (2007). (Pleosporales, genera incertae sedis) 


Habitat freshwater, saprobic. Ascomata solitary or gregarious, 

superficial, globose to subglobose, dark brown to black, short 

papillate, ostiolate, coriaceous. Peridium relatively thin, 

textura angularis in longitudinal section, 2-layered. Hamathecium 

not observed. Asci 8-spored, obpyriform, broadly clavate 

to saccate, pedicellate, bitunicate, apex rounded, persistent. 

Ascospores overlapping 2-3-seriate, broadly fusoid to rhomboid, 

thick-walled, surrounded by mucilaginous sheath, 3- 

euseptate, not constricted at septa, median septum wide, 

forming a darker band, central cells large, trapezoid, dark 

brown to black, verruculose, polar end cells small and paler. 


Literature: Cai and Hyde 2007. 

Type species 

Ascorhombispora aquatica L. Cai & K.D. Hyde, Cryptog. 

Mycol. 28: 295 (2007). (Fig. 6) 

Ascomata 140–170 μm high×150–185 μm diam., solitary 

or gregarious, superficial, globose to subglobose, dark brown 

to black, short papillate, ostiolate, ostioles rounded, small, 

coriaceous. Peridium relatively thin, 10–18 μm wide, textura 

angularis in longitudinal section, composed of two layers of 

angular cells, outer later dark brown to black, relatively thickwalled, 

inner layer hyaline, relatively thin-walled (Fig. 6a and 

b). Hamathecium not observed. Asci 100–198×72–102 μm 

(x ¼ 186 88mm, n=15), 8-spored, obpyriform, broadly 

clavate to saccate, pedicellate, bitunicate, apex rounded, 

deliquescent (Fig. 6c, d and e). Ascospores 30.5–45×16– 

26.5 μm (x ¼ 38:5 21mm, n=25), overlapping 2-3-seriate, 

broadly fusoid to rhomboid, thick-walled, surrounded by 

mucilaginous sheath, 3-euseptate, not constricted at septa, 

median septum wide, forming a darker band, central cells 

large, trapezoid, 11–18 μm long, dark brown to black, 

verruculose, polar end cells small, hemispherical, 3.5–4 μm 

long, subhyaline to pale brown, smooth (Fig. 6f). 


Material examined: CHINA, Yunnan, Jinghong, on 

submerged bamboo in a small forest stream, 26 Jan. 2003, 

leg. det. L. Cai, CAI-1H31 (HKU(M) 10859, holotype). 

Notes 

Morphology 

Ascorhombispora was introduced as a monotypic 

genus from freshwater by Cai and Hyde (2007), and is 

characterized by superficial, coriaceous, non-stromatic 

ascomata, large, saccate asci; lack of interascal filaments 

and trapezoid (rhombic), 3-septate, dark brown to black 

ascospores with smaller end cells which are subhyaline to 

pale brown. Ascorhombispora is most comparable with 

Caryospora and Zopfia. But the globose to subglobose 

ascomata and thin peridium, saccate asci lacking interascal 

pseudoparaphyses, and the 3-septate, rhomboid ascospores 

with the paler end cells of Ascorhombispora differs from 

those of Caryospora (Cai and Hyde 2007). 


Phylogenetic analysis based on either SSU or LSU 

rDNA sequences indicated that Ascorhombispora aquatica 

belongs to Pleosporales, but its familial placement was left 

undetermined (Cai and Hyde 2007). 


The sac-shaped asci and absence of pseudoparaphyses 

are uncommon in Pleosporales, especially among those 

from freshwater. 

Asteromassaria Höhn., Sber. Akad. Wiss. Wien, Math.- 

naturw. Kl., Abt. I 126: 368 (1917). (?Morosphaeriaceae) 


Habitat terrestrial, saprobic. Ascomata medium-sized, 

clustered, at first immersed and then breaking through 

the host surface and becoming superficial, globose, 

subglobose, coriaceous. Peridium 2-layered, thicker near 

the base. Hamathecium of dense, septate, cellular pseudoparaphyses 

which branch and anastomosing frequently 

between and above asci. Asci (4-)8-spored, bitunicate, 

cylindro-clavate to clavate, with a short truncated pedicel 

and a small ocular chamber. Ascospores obliquely uniseriate 

and partially overlapping to biseriate, fusoid to 

fusoid-ellipsoidal, pale brown when mature, 1-septate, 

some becoming 3-septate when old, constricted at the 

median septum. 

Anamorphs reported for genus: Scolicosporium (Sivanesan 

1984). 

Literature: Barr 1982a; b; 1993a; Boise 1985; Shoemaker 

and LeClair 1975; Sivanesan 1987; Tanaka et al. 2005. 

Type species 

Asteromassaria macrospora (Desm.) Höhn., F. von, Sber. 

Akad. Wiss. Wien, Math.-naturw. Kl., Abt. I 126: 368 

(1917). (Fig. 7) 

≡ Sphaeria macrospora Desm., Ann. Sci. Nat. Bot. 10: 

351 (1849). 

Ascomata 400–600 μm high×450–650 μm diam., 4–20 

clustered together, at first immersed and then breaking 

through the host surface and becoming superficial, globose,


Fig. 6 Ascorhombispora aquatica (from HKU(M) 10859, holotype). 

a Section of an ascoma. b Section of a partial peridium. c Immature 

ascus. d–f Mature asci with ascospores. Note the deliquescent ascal 

wall in f. Note the wide, dark band in the medium septum of 

ascospores in d and e and the mucilaginous sheath and paler end cells 

in e and f. Scale bars: a=20 μm, b–f=15 μm (figures referred to Cai 

and Hyde 2007) 

subglobose, not easily removed from the substrate, wall black, 

coriaceous, roughened, apex usually widely porate, with or 

without papilla (Fig. 7a). Peridium 70–90 μm wide, thicker 

near the base where it is up to 180 μm wide, comprising two 

cell types, outer cells composed of heavily pigmented 

small cells, cells 3–5 μm diam., inner layer composed of 

less pigmented cells of textura angularis, 10–20 μm 

diam. (Fig. 7b and c). Hamathecium of dense, septate, 2– 

3 μm broad, pseudoparaphyses which branch and anastomosing 

frequently between and above asci (Fig. 7d). Asci 

(180-)200–280×28–43 μm (x ¼ 230 35mm, n=10), 8- 

Fig. 7 Asteromassaria macrospora (from L, 1004). a Ascomata 

clustered in a group breaking through the host surface. b Section of an 

ascoma. c Section of a partial peridium. Note the cells of textura 

angularis. d Pseudoparaphyses. Note the branches. e Upper part of the 

ascus illustrating the ocular chamber. f Ascus with a short pedicel. g–j 

Ascospores. Note the mucilaginous sheath in G and minutely verruculose 

ornamentation in J. Scale bars: a=0.5 mm, b, c=100 μm, d–j=10 μm 

b

Fungal Diversity


spored (sometimes 4-spored), bitunicate, fissitunicate dehiscence 

not observed, cylindro-clavate to clavate, with a short 

truncated pedicel up to 30 μm, with a small ocular chamber 

(ca. 3μm wide×3 μm high) (Fig. 7e and f). Ascospores 50– 

58×(14-)18–21 μm (x ¼ 55:3 18:2mm, n=10), obliquely 

uniseriate and partially overlappingtobiseriate,fusoidto 

fusoid-ellipsoidal, with narrowly rounded ends, lightly 

brown when mature, 1-septate, some becoming 3-septate 

when old, constricted at the median septum, the upper 

cell often broader and longer than the lower one, 

minutely verrucose (Fig. 7g, h, i and j). 

Anamorph: Scolicosporium macrosporium (Berk.) B. 

Sutton. 

Acervuli immersed in bark, brown, discrete, up to 

250 μm diam., opening by irregular rupture of the 

overlaying tissues. Peridium of thin-walled angular cells. 

Conidiophores cylindrical, 1-2-septate, up to 30 μm long 

and 3–5 μm wide. Conidiogenous cells holoblastic, 1-2- 

annellate, cylindrical, hyaline. Conidia 100–190×12– 

15 μm, fusoid, pale brown with paler or hyaline ends, 7– 

17 transverse septate, smooth-walled, with a tapered apex 

and truncate base (adapted from Sivanesan 1984). 

Material examined: CZECH REPUBLIC, Mährisch- 

Welвkirchen (Hranice), Wsetin (Vsetin), Berg Čap., on 

Fagus sylvatica L., Aug. 1938, F. Petrak (L, 1004). 

Notes 

Morphology 

In this study we were unable to obtain the holotype, 

so we used a collection of Petrak’s. The main morphological 

characters of Asteromassaria are the medium- to 

large-sized, globose to depressed ascomata opening with a 

pore, clavate to oblong asci, narrowly cellular pseudoparaphyses, 

pale to dark brown, bipolar symmetric, mostly 

fusoid, distoseptate or euseptate ascospores (Barr 1993a). 

The bipolar symmetric ascospores of Asteromassaria can 

readily be distinguished from other genera of this family 

(Barr 1993a; Tanaka et al. 2005). Currently, it comprises 

12 species (Tanaka et al. 2005; http://www.mycobank.org, 

28-02-2009). 


Asteromassaria pulchra (Harkn.) Shoemaker & P.M. 

LeClair is basal to Morosphaeriaceae in the phylogenetic 

tree based on four genes, but its placement is 

influenced by taxon sampling that was different in 

several analyses. 


Asteromassaria can be distinguished from other comparable 

genera, i.e. Pleomassaria and Splanchnonema by 1- 

septate and pale brown ascospores, thick-walled textura 

angularis peridium and Scolicosporium anamorphic stage 

(see under Pleomassaria). 

Astrosphaeriella Syd. & P. Syd., Annls mycol. 11: 260 

(1913). (?Melanommataceae) 


Habitat terrestrial, saprobic. Ascomata densely scattered or 

in small groups, erumpent through the outer layers of the 

host tissues to nearly superficial, reflexed pieces of the 

ruptured host tissue usually persisting around the base of 

the ascomata, often star-like, conical to semiglobose, with a 

central papilla. Peridium upper wall usually comprising a 

thick dark brittle pseudoparenchymatous layer, base usually 

flattened and thin-walled. Hamathecium of dense, filliform, 

trabeculate pseudoparaphyses, embedded in mucilage. Asci 

8-spored, bitunicate, fissitunicate, cylindro-clavate to narrowly 

fusoid. Ascospores narrowly fusoid with acute ends, 

hyaline, pale brown or brown, 1-3-septate. 

Anamorphs reported for genus: Pleurophomopsis (Hyde 

et al. 2011). 

Literature: von Arx and Müller 1975; Barr 1990a; Chen 

and Hsieh 2004; Hawksworth 1981; Hawksworth and 

Boise 1985; Hyde and Fröhlich 1998; Hyde et al. 2000; 

Kirk et al. 2001; Sydow and Sydow 1913; Tanaka and 

Harada 2005a; b; Tanaka et al. 2009. 

Type species 

Astrosphaeriella stellata Syd. & P. Syd., Annls mycol. 11: 

260 (1913). (Fig. 8) 


densely scattered or in small groups, erumpent through the 

outer layers of the host tissues to nearly superficial, reflexed 

pieces of the ruptured host tissue usually persisting around 

the base of the ascomata, forming star-like flanges around the 

ascomata from the surface view; ascomata broadly conical, 

with a flattened base not easily removed from the substrate, 

wall black; apex with a central papilla which is black and 

shiny at maturity, scarcely projecting (Fig. 8a). Peridium 40– 

70 μm thick, carbonaceous and crisp, 1-layered, composed of 

very small dark brown thick-walled pseudoparenchymatous 

cells, cells 2–5 μm diam., cell wall 2–6 μm thick, in places at 

the base composed of hyaline cells of textura prismatica, 

cells 5×8 μm diam. (Fig. 8b). Hamathecium of dense, very 

long trabeculate pseudoparaphyses,


Fig. 8 Astrosphaeriella fusispora 

(BISH 145726). a Ascomata 

forming a small group on 

host surface. Note the remains 

of the host forming flanges 

around the ascomata. b Section 

of the partial peridium. Note the 

black peridium and wedge of 

palisade cells between the lateral 

and basal walls. c Asci in trabeculate 

pseudoparaphyses. d–f 

Narrowly fusoid ascospores. 

Scale bars: a=1 mm, b= 

100 μm, c=50 μm, d=15 μm, 

f=10 μm 

ends, hyaline, turning pale brown when mature, 1(−3)- 

septate, constricted at the median septum (Fig. 8d,e and f). 


Material examined: USA, Hawaii, Kapano Gulch, in 

bamboo culms, 5 Jun. 1947, leg. Kopf & Rogers, det. Miller 

(BISH 145726, as Astrosphaeriella fusispora Syd. & P. Syd.). 

Notes 

Morphology 

Astrosphaeriella has been treated as a synonym of Microthelia 

(von Arx and Müller 1975), but the large conical 

ascomata, numerous trabeculate pseudoparaphyses and 1- 

septate and elongated ascospores of Astrosphaeriella all 

disagree with those of Microthelia (Hawksworth 1981). It 

was assigned to Platystomaceae by Barr (1990a) inPleosporales 

or Melanommataceae by Kirk et al. (2001). 

Following a systematic study of Astrosphaeriella, only four 

species were accepted, i.e. A. aosimensis I. Hino & Katum., 

A. stellata, A. trochus (Penz. & Sacc.) D. Hawksw. and A. 

venezuelensis M.E. Barr & D. Hawksw. (Hawksworth 1981), 

and it was defined as a tropical genus, occurring exclusively 

on palms or bamboo. Astrosphaeriella stellata was selected 

as the type of Astrosphaeriella,andA. fusispora was regarded


as a synonym of A. stellata (Hawksworth 1981). More taxa 

were subsequently added (Barr 1990a; Hawksworth and 

Boise 1985; Hyde and Fröhlich 1998), and the generic 

concept extended to include three elements: 1. typical semiimmersed 

to superficial ascomata with flattened base, 

cylindro-clavate asci with fusoid ascospores and trabeculate 

pseudoparaphyses, i.e. Astrosphaeriella sensu stricto (e.g. 

A. fusispora and A. vesuvius (Berk. & Broome) D. Hawksw. 

& Boise); 2. Trematosphaeria-like with rounded ascomata (e. 

g. A. africana D. Hawksw.); and 3. Massarina-like species 

with immersed ascomata (e.g. A. bakeriana (Sacc.) K.D. 

Hyde & J. Fröhl.) (Chen and Hsieh 2004; Tanaka and Harada 

2005a; b). Currently, a broad generic concept of Astrosphaeriella 

is accepted, and 47 taxa are included in Astrosphaeriella. 


Phylogenetic analysis based on LSU and SSU nurDNA 

sequence data indicates that Astrosphaeriella is polyphyletic, 

and located in the basal region of the Pleosporales between 

Testudinaceae and Zopfiaceae/Delitschiaceae (Tanaka et al. 

2009), or basal to Aigialaceae (Schoch et al. 2009). The 

genus is, however, clearly not related to Trematosphaeria as 

previously understood (Boise 1985). 


Astrosphaeriella is currently polyphyletic and new 

collections of the different elements listed above are needed 

in order to understand the placement of various species. We 

suggest that some immersed bambusicolous species may 

belong in Tetraplospheariaceae. 

Asymmetricospora J. Fröhl. & K.D. Hyde, Sydowia 50: 

183 (1998). (?Melanommataceae) 


Habitat terrestrial, saprobic. Ascomata solitary or in small 

groups, immersed, black, lenticular in section, uni- or often 

multi-locular, with a central ostiole without tissue differentiation. 

Upper peridium carbonaceous, thicker at sides and apex. 

Lower peridium composed of irregular-shaped, hyaline cells. 

Hamathecium of trabeculate pseudoparaphyses, branching and 

anastomosing between and above asci, embedded in mucilage. 

Asci 8-spored, bitunicate, fissitunicate unknown, clavate, short 

pedicellate. Ascospores 1-septate, hyaline, constricted at the 

septum, with a broad, spreading mucilaginous sheath. 


Literature: Fröhlich and Hyde 1998. 

Type species 

Asymmetricospora calamicola J. Fröhl. & K.D. Hyde, 

Sydowia 50: 184 (1998). (Fig. 9) 


solitary or in small groups of 2–10, immersed and forming 

slightly protruding domes on the substrate surface, with nearwhite 

rim around the central ostiole; in vertical view lenticular, 

multi- or rarely unilocular, individual locules 175–270 μm 

high×320–400 μm diam., with a flattened base, ostiole a 

central opening without tissue differentiation (Fig. 9a). Upper 

peridium 32–70 μm wide, carbonaceous, composed of a few 

layers of black walled cells of textura angularis. Lower 

peridium thinner, composed of hyaline cells of textura 

globulosa or textura prismatica (Fig. 9b). Hamathecium of 

long trabeculate pseudoparaphyses, 1.2–1.6(−2) μm wide, 

branching and anastomosing between and above asci, 

embedded in mucilage. Asci 137.5–207.5×26–35 μm 

(x ¼ 172:8 31:5mm, n=20), 8-spored, bitunicate, fissitunicate 

dehiscence not observed, clavate, with short pedicel (to 

25 μm), with ocular chambers (ca. 3μm wide×4 μm high) 

(Fig. 9c and d). Ascospores 35–55×10.5–15 μm 

(x ¼ 44:7 12:4mm, n=50), biseriate, navicular to obovoid, 

hyaline, becoming pale brown when senescent, straight or 

usually curved, smooth, asymmetric, 1-septate, the upper cell 

larger with a rounded end, basal cell with a tapering end, 

constricted at the septum, with spreading mucilaginous sheath 

(Fig. 9e, f and g) (data from Fröhlich and Hyde 1998). 


Material examined: AUSTRALIA, North Queensland, 

Palmerston, Palmerston National Park, on dead rattan of 

Calamus caryotoides A.Cunn. ex Mart., Mar. 1994, J. 

Fröhlich (HKU(M) 7794, holotype). 

Notes 

Morphology 

Asymmetricospora was introduced as a monotypic genus 

represented by A. calamicola based on its “absence of a 

subiculum, the absence of short dark setae around the papilla 

and its asymmetric ascospores” (Fröhlich and Hyde 1998). 

Because of the immersed ascomata, ostiole and peridium 

morphology, fissitunicate asci and trabeculate pseudoparaphyses, 

Asymmetricospora was assigned to Melanommataceae 

(sensu Barr 1990a; Fröhlich and Hyde 1998). 

Morphologically Asymmetricospora can be distinguished 

from its most comparable genus, Astrosphaeriella, by its 

ostiole, which is a simple opening without tissue differentiation, 

asymmetric ascospores, and the usually multiloculate 

fruiting body (Fröhlich and Hyde 1998). 


None. 


The placement of Asymmetricospora under Melanommataceae 

remains to be confirmed.


Fig. 9 Asymmetricospora calamicola (from HKU(M) 7794, holotype). 

a Ascomata immersed in the substrate. b Section of the 

peridium. c Mature and immature asci in pseudoparaphyses (in cotton 

Barria Z.Q. Yuan, Mycotaxon 51: 313 (1994). 

(Phaeosphaeriaceae) 


Habitat terrestrial, parasitic. Ascomata small- to medium-sized, 

solitary or scattered, immersed, globose, subglobose, ostiolate, 

coriaceous. Apex with or without papilla and with a pore-like 

ostiole. Peridium 2-layered. Hamathecium of dense, long 

cellular pseudoparaphyses, septate, embedded in mucilage. 

blue). d Clavate ascus with a small ocular chamber. e–g Ascospores 

with sheath. Scale bars: a, b=0.5 mm, c=50 μm, d–g=20 μm 

Asci bitunicate, fissitunicate, cylindrical to clavate, with a short, 

furcate pedicel. Ascospores ellipsoid, hyaline at first, turning 

brown at maturity, 1-septate, strongly constricted at the septum. 


Literature: Yuan 1994. 

Type species 

Barria piceae Z.Q. Yuan, Mycotaxon 51: 314 (1994). 

(Fig. 10)


Fig. 10 Barria piceae (from 

NY 92003, isotype). a Ascoma 

on the host surface. Note the 

wide opening ostiole. b Section 

of the partial peridium with two 

types of cells. c, d Asci with 

ocular chambers and short pedicels. 

e, f Ellipsoid ascospores 

which are turning brown with 

thin sheath around them. Scale 

bars: a=0.5 mm, b=50 μm, c, 

d=20 μm, e, f=10 μm 


solitary, scattered, immersed, globose, subglobose, coriaceous, 

apex with or without papilla and with a pore-like 

ostiole (Fig. 10a). Peridium 20–35 μm thick, comprising 

two cell types, the outer cells comprising 3–4 layers of 

brown pseudoparenchymatous cells, cells 4–5 μm diam., 

cell wall 2–3 μm thick, inner cells comprising 3–4 layers of 

pale brown compressed cells, cells 2×16 μm diam., cell 

wall 0.5–1.5 μm thick (Fig. 10b). Hamathecium of dense, 

long cellular pseudoparaphyses, 2–3 μm broad, septate. 

Asci 135–200(−220)×14–20 μm (x ¼ 156 16:6mm, n= 

10), 8-spored, bitunicate, fissitunicate, cylindrical to clavate, 

with a short, furcate pedicel, up to 22 μm long, with a 

large ocular chamber (ca. 4μm wide×3 μm high) (Fig. 10c 

and d). Ascospores 19–21.5×10–12 μm (x ¼ 20:4 11mm, 

n=10), uniseriate to partially overlapping, ellipsoid, hyaline 

or greenish with numerous small guttules at first and olive 

green to smoky brown at maturity, 1-septate, strongly 

constricted at the septum, foveolate, surrounded with sheath 

(Fig. 10e and f). 


Material examined: CHINA, Xinjiang Province, Uygur, 

Urumqi, Tianshan Mountain, on needles of Picea schrenkiana, 

1 Jul. 1992, Z.Q. Yuan (NY 92003, isotype).


Notes 

Morphology 

Barria was established by Yuan (1994) as a monotypic 

genus represented by B. piceae according to its “two-celled, 

pigmented ascospores, pseudoparenchymatous peridium and 

narrowly cellular pseudoparaphyses” thus differing in its 

combination of characters from all of the morphologically 

related dothideomycetous genera, such as Didymosphaeria, 

Didymopleella or Stegasphaeria. The taxon was considered 

to belong in Phaeosphaeriaceae. Ascomata and 

colour or shape of ascospores, however, readily distinguish 

it from other 1-septate Phaeosphaeriaceae genera, 

i.e. Didymella, Lautitia and Metameris (Yuan 1994). 

Barria piceae causes blight of spruce needles. 


None. 


The status of Barria with its unusual verrucose ascospores 

and thick gel coating is uncertain. In many ways it 

resembles Belizeana, with its cylindrical asci, 1-septate, 

ellipsoid ascospores with sheath and verruculose surface 

(Kohlmeyer and Volkmann-Kohlmeyer 1987). However, 

the latter is a marine genus while Barria causes leaf blight 

of terrestrial Picea (Yuan 1994). The placement in 

Phaeosphaeriaceae seems logical based on the parasitic 

life style, thin and simple peridium, wide cellular pseudoparaphyses 

and brown ascospores. However, molecular 

data are needed to confirm this. 

Belizeana Kohlm. & Volkm.-Kohlm., Bot. Mar. 30: 195 



Habitat marine, saprobic. Ascomata solitary, scattered, or 

in small groups, medium-sized, immersed to semiimmersed, 

subglobose to broadly ampulliform, black, 

ostiolate, carbonaceous. Peridium thin, comprising several 

layers of brown thin-walled cells of textura angularis. 

Hamathecium of dense, filliform pseudoparaphyses, rarely 

branched. Asci 8-spored, bitunicate, fissitunicate, broadly 

cylindrical to clavate, with a short pedicel and an ocular 

chamber. Ascospores uniseriate, broadly ellipsoidal, hyaline, 

turn pale brown when senescent, 1-septate, constricted 

at the septum, thick-walled, 2-layered, mature 

spores with tuberculate ornamentation between the two 

layers. 

Anamorphs reported for genus: Phoma-like (Kohlmeyer 

and Volkmann-Kohlmeyer 1987). 

Literature: Kohlmeyer and Volkmann-Kohlmeyer 1987. 

Type species 

Belizeana tuberculata Kohlm. & Volkm.-Kohlm., Bot. 

Mar. 30: 196 (1987). (Fig. 11) 

Ascomata 170–300 μm high×160–290 μm diam., solitary, 

scattered, or in small groups of 2–3, immersed to semiimmersed, 

subglobose to broadly ampulliform, carbonaceous, 

black, pale brown on the sides, ostiolate, epapillate or shortly 

papillate, ostiolar canal filled with a tissue of hyaline cells 

(Fig. 11a). Peridium 25–35 μm wide, comprising several 

layers thin-walled cells of textura angularis, which are 

hyaline inwardly, near the base composed of a hyaline 

hyphal mass producing asci, up to 20 μm thick (Fig. 11b, c 

and e). Hamathecium of dense, ca. 2 μm broad, filliform 

pseudoparaphyses, rarely branched, embedded in mucilage 

(Fig. 11g). Asci 145–170×20–30 μm (x ¼ 163 25mm, n= 

10), 8-spored, bitunicate, fissitunicate, broadly cylindrical to 

clavate with a short pedicel, thick-walled, with a small ocular 

chamber (Fig. 11d, f and h). Ascospores 21–26 × 13–18 μm 

(x ¼ 22 15mm, n = 10), uniseriate, broadly ellipsoidal, 

hyaline, turn pale brown when senescent, 1-septate, constricted 

at the septum, thick-walled, 2-layered, mature spores 

with tuberculate ornamentation between the two layers 

(Fig. 11i and j). 

Anamorph: Phoma-like (Kohlmeyer and Volkmann- 

Kohlmeyer 1987). 

Material examined: BELIZE, Twin Cays, on Laguncularia 

sp., 7 Apr. 1983, leg. & det. J. Kohlmeyer (Herb. J. 

Kohlmeyer No. 4398, holotype); AUSTRALIA, Towra 

Point, New South Wales, trunk of eroded tree with oysters 

and shipworms, intertidal zone, Botany Bay, 23 Aug. 1981 

(Herb. J. Kohlmeyer No. 4209, paratype). 

Notes 

Morphology 

Belizeana was formally established to accommodate B. 

tuberculata, an obligate marine fungus, which is characterized 

by verrucose ascospores (Kohlmeyer and Volkmann- 

Kohlmeyer 1987). Belizeana tuberculata canbeassignedto 

Pleosporaceae (Pleosporales) according to Luttrell’s (1973) 

treatment and keys of von Arx and Müller (1975), but cannot 

resolve a proper family based on Barr (1979a, 1983). The 

unique morphology together with obligate marine habitat 

makes B. tuberculata readily distinguishable from all other 

taxa of Pleosporaceae. 


None. 


The ascospores of Belizeana tuberculata are most comparable 

with those of Acrocordiopsis patilii, but the superficial


Fig. 11 Belizeana tuberculata 

(from Herb. J. Kohlmeyer No. 

4398, holotype). a Immersed to 

semi-immersed ascomata. b, e 

Vertical section of an ascoma. c 

Section of a partial peridium. d 

Squash mounts with a large 

number of asci. f Broadly 

cylindrical ascus with a large 

ocular chamber. g Filliform 

pseudoparaphyses. h Apical part 

of an ascus. Note the large 

ocular chamber. i, j One-septate 

ascospores. Scale bars: 

a=0.3 mm, b=100 μm, 

c=20 μm, d, e=50 μm, 

f–j=10 μm 

conical ascomata of A. patilii are distinct from B. tuberculata. 

Thus, the familial placement of Belizeana is still undetermined. 

Biatriospora K.D. Hyde & Borse, Mycotaxon 26: 263 

(1986). (Pleosporales, genera incertae sedis)



Habitat marine, saprobic. Ascomata large, solitary or gregarious, 

immersed, subglobose to pyriform, ostiolate, papillate, 

periphysate, black, branching, carbonaceous. Hamathecium 

of dense, long trabeculate pseudoparaphyses, embedded in 

mucilage. Asci 8-spored, bitunicate, fissitunicate, cylindrical, 

with apical apparatus. Ascospores uniseriate to partially 

overlapping, fusoid, hyaline when young, becoming brown 

to dark brown at maturity, multi-septate towards each end, 

with a hyaline, globose refractive chamber or appendage at 

each end, not constricted at the septum. 


Literature: Hyde and Borse 1986; Suetrong et al. 2009. 

Type species 

Biatriospora marina K.D. Hyde & Borse, Mycotaxon 26: 

264 (1986). (Fig. 12) 


or gregarious, immersed, subglobose to pyriform, ostiolate, 

papillate, periphysate, black, carbonaceous (Fig. 12.2a). Hamathecium 

of dense, long trabeculate pseudoparaphyses, 1– 

1.5 μm broad, branching, embedded in mucilage. Asci 175– 

Fig. 12 1 Biatriospora marina 

(from IMI 297768, holotype). a, 

b Cylindrical asci. Note the 

mucilage pseudoparaphyses in 

(a) and the conspicuous ocular 

chamber in (b). c, d Ascospores 

with hyaline end chambers 

(arrowed). Scale bars: a, b= 

50 μm, c, d=20 μm. 2 Line 

drawings of Biatriospora marina 

(based on holotype). a Section 

through ascocarp showing 

asci and pseudoparaphyses. b 

Asci and pseudoparaphyses. c 

Ascospores. Scale bars: a= 

200 μm, b=40 μm, c=30 μm 

(figure with permission from 

Hyde and Borse 1986)


Fig. 12 

2 (continued) 

400×22–40 μm, 8-spored, bitunicate, fissitunicate, cylindrical, 

with long pedicels and apical apparatus (Fig. 12.1a, b, 2b). 

Ascospores 55–82×16–25 μm, uniseriate to partially overlapping, 

fusoid, hyaline when young, becoming brown to 

dark brown at maturity, 2-4-septate towards each end, and 

with a hyaline, globose refractive chamber or appendage 

at each end, 6–8×4–6 μm diam., not constricted at the 

septum (Fig. 12.1c, d, 2c).



Material examined: SEYCHELLES, 2 Jan. 1984 (Herb. 

IMI 297768 holotype). 

Notes 

Morphology 

Biatriospora was introduced to accommodate a 

marine fungus B. marina, which is characterized by 

horizontal ascomata and ascospores with polar, globose 

refractive chambers and polar septa (Hyde and Borse 

1986). Polar refractive chambers can also occur in other 

marine fungi, such as Lulworthia and Aigialus. The 

chambers have been proposed as important for spore 

attachment to substrates in a liquid environment (Hyde 

and Borse 1986). 


Multigene phylogenetic analysis indicated that Biatriospora 

marina forms a separate branch, sister to other families of 

Pleosporales (Suetrong et al. 2009), and maybe related to 

species in Roussoella (Plate 1). 


The familial status of Biatriospora can not be determined. 

Bicrouania Kohlm. & Volkm.-Kohlm., Mycol. Res. 94: 



Habitat marine, saprobic. Ascomata immersed gregarious, 

erumpent to superficial, globose to subglobose, black, 

periphysate, coriaceous, epapillate or papillate, ostiolate. 

Peridium thin, 2-layered. Hamathecium of dense, long 

trabeculate pseudoparaphyses, branching and anastomosing 

between and above the asci. Asci 8-spored, bitunicate, 

fissitunicate, cylindrical, with a thick, furcate pedicel 

lacking ocular chamber. Ascospores obliquely uniseriate 

and partially overlapping, ellipsoidal with broadly rounded 

ends, reddish brown, 1-septate, thick-walled, constricted at the 

septum. 


Literature: Jones et al. 2009; Kohlmeyer and Volkmann- 

Kohlmeyer 1990. 

Type species 

Bicrouania maritima (P. Crouan & H. Crouan) Kohlm. & 

Volkm.-Kohlm., Mycol. Res. 94: 685 (1990). (Fig. 13) 

≡ Sphaeria maritima P. Crouan & H. Crouan, Florule du 

Finistére, Paris: 27 (1867) non Sphaeria maritima Cooke & 

Plowright, Grevillia 5: 120 (1877). 


gregarious, immersed, mostly erumpent to superficial, 

globose to subglobose, black, coriaceous, with a rough 

surface, papillate or epapillate, ostiolate, periphysate 

(Fig. 13a). Peridium 40–50 μm thick laterally, up to 

75 μm thick at the apex, thinner at the base, 2-layered, 

outer layer composed of small heavily pigmented 

pseudoparenchymatous cells, inner layer very thin, 

composed of hyaline thin-walled small cells, merging into 

pseudoparaphyses (Fig. 13a and b). Hamathecium of 

dense, very long trabeculate pseudoparaphyses, 0.8– 

1.2 μm broad, branching and anastomosing between and 

above the asci. Asci 170–225×17.5–22.5 μm 

(x ¼ 199:6 20mm, n=10), 8-spored, bitunicate, fissitunicate, 

cylindrical, with a thick, furcate pedicel which is up 

to 70 μm long, lacking ocular chamber (Fig. 13c, d and e). 

Ascospores 22–26×12–15 μm (x ¼ 24:5 13:3mm, n= 

10), obliquely uniseriate and partially overlapping, ellipsoidal 

with broadly rounded ends, reddish brown, 1- 

septate, slightly constricted at the septum, thick-walled, 

with a thick darkened band around the septum, smooth 

(Fig. 13c, d and e). 


Material examined: FRANCE, Finistère, on Halimone 

portulacoides (IMI 330806, isotype, as Sphaeria 

maritima). 

Notes 

Morphology 

When Kohlmeyer and Volkmann-Kohlmeyer (1990) studied 

the four marine Didymosphaeria species, the monotypic 

Bicrouania was established to accommodate B. maritima (as 

Didymosphaeria maritima (P. Crouan & H. Crouan) Sacc.), 

which could be distinguished from Didymosphaeria by its 

superficial ascomata lacking a clypeus, thick-walled asci and 

its association with algae (Kohlmeyer and Volkmann- 

Kohlmeyer 1990). Jones et al. (2009) agreed that it cannot 

be placed in Didymosphaeria based on its superficial 

ascomata, but that it does have many similarities with 

Didymosphaeria. Molecular data are required to determine 

its relationship with Didymosphaeria and to resolve its 

higher level placement. 


None. 


Besides the morphological differences, its marine and 

substrate habitats also differ from Didymosphaeria. 

Bimuria D. Hawksw., Chea & Sheridan, N. Z. J. Bot. 17: 

268 (1979). (Montagnulaceae)


Fig. 13 Bicrouania maritima 

(from IMI 330806, isotype). a 

Section of an ascoma. b Section 

of papilla. Note the periphyses. 

c–e Eight-spored asci. Note the 

furcated pedicel. Scale bars: a, 

b=100 μm, c–e=20 μm 


Habitat terrestrial, saprobic. Ascomata solitary, superficial, 

globose, dark brown, epapillate, ostiolate. Peridium thin, 

pseudoparenchymatous. Hamathecium of few, cellular 

pseudoparaphyses, embedded in mucilage, rarely anastomosing 

and branching. Asci bitunicate, fissitunicate, broadly 

clavate with short pedicels, 2-3-spored. Ascospores muriform, 

broadly ellipsoid, dark brown with subhyaline end 

cells, verrucose. 


Literature: Barr 1987b; Hawksworth et al. 1979; 

Lumbsch and Huhndorf 2007. 

Type species 

Bimuria novae-zelandiae Hawksworth, Chea & Sheridan, 

N. Z. J. Bot. 17: 268 (1979). (Fig. 14) 

Ascomata (185-)200×310(-330) μm diam., solitary, scattered, 

semi-immersed or superficial, globose, hyaline when


Fig. 14 Bimuria novae-zelandiae (from CBS 107.79, isotype). a–c 

Asci with a short pedicel and small ocular chamber. d Immature ascus 

(in cotton blue). e Partial ascospore. Note the convex verrucae on the 

ascospore surface. f Released ascospores. Note the lighter end cells, 

germ pore and the longiseptum (arrowed). g Fissitunicate ascus 

dehiscent. Scale bars: a–g=20 μm 

young, turning dark brown to black when mature, ostiolate, 

the ostiole more or less sessile or raised into a very short neck. 

Peridium 5–8(-12) μm thick, comprising 2–3 layers of 

radically compressed pseudoparenchymatous cells, cells 10– 

15 μm diam. in surface view, cell wall 2–3 μm thick. 

Hamathecium consisting of few, 2.5–4 μm broad cellular 

pseudoparaphyses, embedded in mucilage, rarely anastomosing 

and branching, septate, 7–13 μm long between 

two septa. Asci (65-)80–95×20–32.5 μm (x ¼ 75:6 

29:4mm, n=10), (1-)2(-3)-spored, bitunicate, fissitunicate, 

broadly clavate, with a short and small knob-like pedicel 

which is up to 13 μm long, ocular chamber best seen in 

immature asci (Fig. 14a, b, c, d and g). Ascospores 

accumulating in a subglobose black shiny mass adhering 

together outside the ostiole, 55–68×25–28 μm (x ¼ 59 

26mm, n=10), broadly ellipsoid but becoming narrowed 

towards the poles, muriform with (5-)7 transverse septa, cells 

with (0-)l(-2) longitudinal septa in each cell, no constriction


at the septa, dark brown, the apical cells paler with no 

longitudinal septa, verruculose (Fig. 14e and f). 


Material examined: NEW ZEALAND, North Island, 

Wairarapa District, Nutty Farm, isolated from soil, 3 Mar. 

1978, Chea Chark Yen & J.E. Sheridan (CBS 107.79, 

isotype). 

Notes 

Morphology 

Bimuria novae-zelandiae was first isolated from soil of a 

barley field in New Zealand (Hawksworth et al. 1979). Based 

on B. novae-zelandiae, the genus is characterized by a very 

thin peridium, mostly 2-spored and fissitunicate asci as well as 

the muriform, dark brown, verrucose ascospores (Hawksworth 

et al. 1979). Because of its unique morphological characters, 

the familial placement of this genus has been debatable and it 

has been placed in Pleosporaceae (Hawksworth et al. 1979), 

in Phaeosphaeriaceae (Barr 1987b)andinMelanommataceae 

(Lumbsch and Huhndorf 2007). 

Morphologically, Bimuria is most comparable with 

some superficially similar or allied genera, in particular 

Montagnula (Hawksworth et al. 1979). However, the thick 

carbonaceous peridium distinguishes Montagnula from 

that of Bimuria (Hawksworth et al. 1979). In addition, the 

ascospores of Montagnula are discharged forcibly through 

the ostiole instead of forming a mass outside of the ostiole 

as in Bimuria (Hawksworth et al. 1979). Ascomauritiana 

lignicola V.M. Ranghoo & K.D. Hyde has somewhat 

similar ascospores in 4-spored asci, but this taxon has 

unitunicate asci (Ranghoo and Hyde 1999). The morphological 

characters of Bimuria, such as ascospore release 

and large, thick-walled ascospores may be an adaptation to 

its soil-borne habitat (Hawksworth et al. 1979). 


Bimuria novae-zelandiae was found to be closely related 

to Phaeodothis winteri (Niessl) Aptroot (syn. Didymosphaerella 

opulenta (De Not.) Checa & M.E. Barr) and 

Montagnula opulenta (De Not.) Aptroot in analysis of 

combined sequences, i.e. SSU rDNA, LSU rDNA, RPB2 

and TEF1 sequences (Schoch et al. 2006, 2009). These two 

species had been included by Barr (2001) in her new family 

Montagnulaceae. 


We agree with Barr (2001) and include the genus in 

Montagnulaceae based on both morphological and phylogenetic 

characters. 

Bricookea M.E. Barr, Mycotaxon 15: 346 (1982). 

(?Phaeosphaeriaceae) 


Habitat terrestrial, saprobic (or parasitic?). Ascomata small- to 

medium-sized, solitary, scattered, or in small groups, immersed, 

erumpent to superficial, depressed globose, papillate, 

ostiolate. Peridium thin. Hamathecium filliform, cellular 

pseudoparaphyses, embedded in mucilage, anastomosing, 

septate. Asci bitunicate, fissitunicate, cylindrical, cylindroclavate 

or slightly obclavate, with a short knob-like pedicel, 

with an ocular chamber. Ascospores hyaline, ellipsoid to 

narrowly obovoid, 3-septate, constricted at each septum. 


Literature:Barr1982a;Berlese1896;Holm1957; Shoemaker 

and Babcock 1989a. 

Type species 

Bricookea sepalorum (Vleugel) M.E. Barr, Mycotaxon 15: 

346 (1982). (Fig. 15). 

≡ Metasphaeria sepalorum Vleugel, Svensk bot. Tidskr. 

2: 369 (1908). 


scattered, or in small groups, or forming locules in massive 

stromatic tissues, initially immersed, becoming erumpent, to 

nearly superficial, depressed globose, black, membraneous, 

roughened; apex rounded, sometimes very short and almost 

inconspicuous, with a somewhat slit-like or Y-shaped ostiole 

(Fig. 15a). Peridium 16–30 μm wide, comprising two types of 

cells, outer cells heavily pigmented thick-walled textura 

angularis, cells 4.5–8 μm diam., cell wall 1–1.5 μm thick, 

inner cells of subhyaline thin-walled textura angularis, cells 

larger than outer cells (Fig. 15b). Hamathecium of long 

cellular pseudoparaphyses, 1.5–2 μm broad, embedded in 

mucilage, anastomosing, septate. Asci 63–83×9.5–11 μm 


oblong, cylindro-clavate or slightly obclavate, with a short 

knob-like pedicel which is 5–13 μm long, with an ocular 

chamber (Fig. 15c, d and e). Ascospores (14-)15.5–19×5– 

7 μm (x ¼ 16:9 5:9mm, n=10), obliquely uniseriate and 

partially overlapping to biseriate, ellipsoid to narrowly 

obovoid, hyaline, 3-septate, constricted at each septum, the 

cells above central septum often broader than the lower ones, 

smooth (Fig. 15f, g, h, i and j). 


Material examined: SWEDEN, on Juncus filliformis, 

Stockholm, J. Vleugel. Jul. 1907 (S, type as Metasphaeria 

sepalorum Vleugel). 

Notes 

Morphology 

Bricookea was formally established by Barr (1982a) asa 

monotypic genus represented by B. sepalorum basedonits


Fig. 15 Bricookea sepalorum (from S, type). a Ascomata on host 

surface (arrowed). b Section of partial peridium. Note thick-walled 

out layer and thin-walled inner layer. c–e Cylindrical to slightly 

obclavate asci with short knob-like pedicels. f–j Hyaline, 3-septate 

smooth-walled ascospores. Scale bars: a=0.5 mm, b=50 μm, c–j= 

10 μm


“globose to depressed ascomata, slit-like ostiole with labial 

cells, bitunicate asci, cellular pseudoparaphyses, and hyaline 

septate ascospores”. Bricookea was morphologically assigned 

to Phaeosphaeriaceae. Holm(1957) checked the authentic 

collections from North America and type material from 

Europe, and observed that the ascospores of collections from 

North America were significantly larger than those from the 

type material from Sweden. Thus, Shoemaker and Babcock 

(1989a) considered that the collections from North America 

represented a new species, which they introduced as B. barrae 

Shoemaker & C.E. Babc. Although the short slit-like 

ostiole has previously been reported (Shoemaker and 

Babcock 1989a), it is inconspicuous in the type specimen 

from Sweden. Currently, only two species are accommodated 

in this genus. 


None. 


The knob-shaped pedicel, slit-like ostiole, hyaline 

ascospores as well as the herbaceous substrate all disagree 

with any current pleosporalean family. Thus, we temporarily 

retain this genus under Phaeosphaeriaceae until DNA 

sequence comparisons can be carried out. 

Byssolophis Clem., in Clements & Shear, Gen. fung., Edn 2 

(Minneapolis): 286 (1931). (Pleosporales, genera incertae 

sedis) 


Habitat terrestrial, saprobic. Ascomata medium-sized, gregarious, 

semi-immersed to erumpent, coriaceous, ovoid, with a 

conspicuous elongate slit-like ostiole on the top. Peridium not 

observed. Hamathecium of dense, long pseudoparaphyses, 

anastomosing and branching between and above the asci. Asci 

8-spored, bitunicate, fissitunicate, cylindrical or cylindroclavate, 

with a furcate pedicel. Ascospores fusoid, hyaline, 

turning faintly brown when old, 1-septate, with a short 

terminal appendage at each end. 


Literature: Clements and Shear 1931; Holm 1986; Müller 

and von Arx 1962. 

Type species 

Byssolophis byssiseda (Flageolet & Chenant.) Clem., Gen. 

Fung. (Minneapolis): 286 (1931). (Fig. 16) 

≡ Schizostoma byssisedum Flageolet & Chenant., in 

Chenantaise, Bull. Soc. mycol. Fr. 35: 125 (1919). 

Ascomata 300–450 μm high×600–750 μm long× 

350–420 μm broad, gregarious, semi-immersed to 

erumpent, coriaceous, ovoid with a flattened base and 

apex with a elongate slit-like ostiole, up to 700 μm long and 

200 μm wide (Fig. 16a). Peridium not observed. Hamathecium 

of dense, long pseudoparaphyses, up to 1.5–2.5 μm 

broad, anastomosing and branching between and above the 

asci (Fig. 16b). Asci 80–105×(5-)7.5–10 μm 

(x ¼ 91 8mm; n=10), 8-spored, bitunicate, fissitunicate, 

cylindrical or cylindro-clavate, with a furcate pedicel and a 

small ocular chamber (J-) (Fig. 16d). Ascospores 18–20 

(−28)×4.5–6(−7.5) μm (x ¼ 20:8 5:7mm, n=10), uniseriate 

to biseriate, fusoid, hyaline, turning faintly brown when 

old, 1-septate, with 1–2 distinct oil drops in each cell and 

usually with a short terminal appendage at each end 

(Fig. 16c). 


Material examined: on decaying wood (K(M):164030, 

isotype). 

Notes 

Morphology 

Byssolophis was introduced as a monotypic genus based 

on B. byssiseda, which is characterized by its semiimmersed, 

gregarious, ovoid ascomata, with a conspicuous 

central apical ostiolar slit (Holm 1986). Subsequently, two 

more species were introduced, viz. B. ampla (Berk. & 

Broome) L. Holm and B. sphaerioides (P. Karst.) E. Müll. 

(Holm 1986; Müller and von Arx 1962). 


The current phylogeny places Byssolophis sphaerioides 

in proximity of Hypsostromataceae without resolving any 

sister taxa (Plate 1). 


The slit-like ostiole, cylindrical asci, hyaline and 1- 

septate ascospores as well as the form of pseudoparaphyses 

are similar to species in Lophiostoma. Thus, Byssolophis 

may be a synonym of Lophiostoma. 

Byssosphaeria Cooke, Grevillea 7: 84 (1879). 

(Melanommataceae) 


Habitat terrestrial, saprobic. Ascomata medium-sized, 

scattered to gregarious, superficial, globose, subglobose 

to turbinate, non papillate with white, orange, red or green 

ostiolar region, wall black. Hamathecium of dense, 

long trabeculate pseudoparaphyses, embedded in mucilage, 

anastomosing between and above the asci. Asci 

bitunicate, fissitunicate, clavate to nearly cylindrical, 

with a furcate pedicel. Ascospores fusoid with narrow


Fig. 16 Byssolophis byssiseda 

(from K(M):164030, isotype). a 

Ascomata gregarious on the host 

surface. b Numerous pseudoparaphyses. 

c Fusoid ascospores 

with or without terminal appendages. 

d Clavate ascus with a 

short furcate pedicel. Scale bars: 

a=1 mm, b–d=10 μm 

ends, straight or slightly curved, brown, 1-septate when 

young. 

Anamorphs reported for genus: Pyrenochaeta or Chaetophoma-like 

(Barr 1984; Hawksworth et al. 1995; Samuels 

and Müller 1978). 

Literature: von Arx and Müller 1975; Barr 1984; Boise 

1984; Bose 1961; Chen and Hsieh 2004; Cooke and 

Plowright 1879; Hyde et al. 2000; Luttrell 1973; Mugambi 

and Huhndorf 2009b; Müller and von Arx 1962; Samuels 

and Müller 1978. 

Type species 

Byssosphaeria keitii (Berk. & Broome) Cooke [as ‘Byssosphaeria 

keithii’], (1879). (Fig. 17) 

≡ Sphaeria keitii Berk. & Broome [as ‘Sphaeria keithii’], 

Ann. Mag. Nat. Hist., IV 17: 144 (1876). 

Ascomata 360–500(−600) μm high×420–640 μm 

diam., scattered or in small groups, superficial with 

basal subiculum anchoring on the substrate, globose, 

subglobose to turbinate, non-papillate with pore-like


Fig. 17 Byssosphaeria schiedermayriana (from K(M):108784, 

holotype). a Superficial ascomata on the host surface. b Brown, 1- 

septate ascospores. c Section of the lateral peridium. Note the outer 

textura angularis and inner textura epidermoidea cells. d, e Furcate 

asci with a long pedicel. f Dehiscent ascus. Scale bars: a=0.5 mm, c= 

50 μm, b, d–f=15 μm 

ostiole, ostiolar region sometimes with orange and 

greenish tint, wall black, roughened, coriaceous 

(Fig. 17a). Peridium 55–85 μm thick, peridium outside 

of the substrate comprising two cell types, outer layer 

composed of brown thick-walled cells of textura epidermoidea, 

cells 1–3 μm diam., inner layer composed of 

small hyaline cells, cells 3–5 μm diam., merging into 

pseudoparaphyses; peridium inside the substrate one 

layer, composed of large pale brown cells of textura 

angularis, cells6–13 μm diam. (Fig. 17c). Hamathecium 

of dense, long trabeculate pseudoparaphyses, 1–2 μm 

broad, embedded in mucilage, anastomosing between and 

above the asci. Asci 90–120(−148)×10–14 μm, 8-spored, 

bitunicate, fissitunicate, cylindro-clavate to clavate, 

biseriate above and uniseriate below, pedicel 15–20 

(−53) μm long, the immature asci usually with longer 

and furcate pedicel (−68 μm) (Fig. 17d,e and f). 

Ascospores 29–34(−38)×5.5–8(−10) μm, fusoid with 

narrow ends, mostly straight, sometimes slightly curved, 

smooth, pale brown, 1-septate, becoming 3-septate after 

discharge, with hyaline appendages at each acute to 

subacute end; in some mature spores the appendage may 

be absent (Fig. 17b). 

Anamorph: Pyrenochaeta sp. (Barr 1984; Samuels and 

Müller 1978). 

Pycnidia 70–500 μm diam. Conidiogenous cells phialidic, 

lining cavity, 5–8×4–6 μm to 5–10×3–6 μm. Conidia 

2.5–3.5(−4)×1.5–2(−3) μm, hyaline, ellipsoid or subglobose 

(Barr 1984). 

Material examined: ERIE, Dublin, Glasnevin Botanic 

Garden, on old rope, Jun. 1872, W. Keit (K(M):108784, 

holotype, asSphaeria keitii Berk. & Broome).


Notes 

Morphology 

Byssosphaeria was introduced by Cooke and Plowright 

(1879) based on its superficial ascomata seated on a “tomentose 

subiculum of interwoven threads”, which includes various 

species in Sphaeria and Byssisedae, and was validly typified by 

B. keitii (Cooke 1878). Byssosphaeria keitii was treated as a 

synonym of B. schiedermayeriana (Fuckel) M.E. Barr by 

Sivanesan (1971), and B. schiedermayeriana exclusively occurs 

in tropical regions or greenhouse environments in temperate 

regions (Barr 1984). Morphologically, B. keitii is characterized 

by its large ascomata with orange to reddish plain apices, and is 

closely related to B. rhodomphala (Berk.) Cooke (Barr 1984). 

For a long time, Byssosphaeria was assigned to 

Herpotrichia sensu lato, and Byssosphaeria schiedermayeriana 

was renamed as H. schiedermayeriana Fuckel 

(von Arx and Müller 1975; Bose 1961; Luttrell 1973; 

Müller and von Arx 1962; Sivanesan 1971). After studying 

Herpotrichia in North America, Barr (1984) accepted a 

relatively narrow generic concept, Herpotrichia sensu 

stricto, and revived Byssosphaeria; this proposal is 

supported by phylogenetic study (Mugambi and Huhndorf 

2009b). Currently Byssosphaeria comprises 32 species 

(http://www.mycobank.org, 08-01-2009). 


The monophyletic nature of Byssosphaeria is well 

demonstrated, as well as its familial status in Melanommataceae 

(Mugambi and Huhndorf 2009b). 


Orange and greenish plain apices exist in the specimen 

we examined, which is different from records as “orange, 

bright or dull reddish plain apices” by Barr (1984). This 

might be because different specimens have different 

colours, or there may be a variation of apical colour within 

a single species, as both orange and green can coexist on 

the same ascoma (see Fig. 17a). The coloured apical rim, 

together with the trabeculate pseudoparaphyses as well as 

the presence of subiculum make Byssosphaeria readily 

distinguishable from other morphologically comparable 

genera, e.g. Herpotrichia and Keissleriella (Hyde et al. 2000). 

Calyptronectria Speg., Anal. Mus. nac. Hist. nat. B. Aires 

19: 412 (1909). (Melanommataceae) 


Habitat terrestrial, saprobic. Ascomata small- to mediumsized, 

solitary, scattered, or in small groups, immersed, 

lenticular to subglobose, papillate, ostiolate. Hamathecium 

of long, filliform pseudoparaphyses, branching and anastomosing, 

embedded in mucilage. Asci 4- to 8-spored, 

bitunicate, fissitunicate, cylindrical to cylindro-clavate, with 

a short, furcate pedicel. Ascospores muriform, broadly 

fusoid to fusoid with broadly to narrowly rounded ends, 

hyaline. 


Literature: Barr 1983; Rossman et al. 1999; Spegazzini 

1909. 

Type species 

Calyptronectria platensis Speg., Anal. Mus. nac. Hist. nat. 

B. Aires 19: 412 (1909). (Fig. 18) 


scattered, immersed, lenticular to subglobose, papillate, 

ostiolate (Fig. 18a and b). Apex with a small and slightly 

protruding papilla. Peridium 18–30 μm wide, comprising two 

types of cells, outer layer composed of pseudoparenchymatous 

cells, cells 3–6 μm diam., cell wall 1–2 μm thick, inner 

layer comprising less pigmented cells, merging with pseudoparaphyses 

(Fig. 18b and c). Hamathecium of long, filliform 

pseudoparaphyses, 1–2 μm broad, branching and anastomosing, 

embedded in mucilage. Asci 98–140×12.5–20 μm 

(x ¼ 107 15:4mm, n=10), 8-spored, sometimes 4-spored, 

bitunicate, fissitunicate, cylindrical to cylindro-clavate, with 

a short, furcate pedicel, 12–20 μm long,withanocular 

chamber (to 4 μm wide×3 μm high) (Fig. 18e and f). 

Ascospores 17– 22.5 μ m ×(6.3-)7.5– 10 μ m 

(x ¼ 19:8 7:6mm, n=10), biseriate above and uniseriate 

below, ellipsoid to broadly fusoid with broadly to narrowly 

rounded ends, hyaline, usually with (3-)5 transverse septa, 

with or without 1–3 longitudinal septa in the central cells, 

constricted at the median septum, the upper cell often broader 

than the lower one, smooth, surrounded by an irregular 

hyaline gelatinous sheath up to 3 μm thick (in dry specimen) 

(Fig. 18d). 


Material examined: ARGENTINA, La Plata, on decaying 

branches of Manihot carthaginensis (Jacq.) Müll., Sept. 

1906, Spegazzini (LPS 1209, holotype). 

Notes 

Morphology 

Calyptronectria is a relatively poorly studied genus, 

which was formally established based on C. argentinensis 

Speg. and C. platensis, with C. platensis being chosen as 

the generic type (Spegazzini 1909). Morphologically, 

Calyptronectria is characterized by its immersed ascomata,


Fig. 18 Calyptronectria platensis (from LPS 1209, holotype). a 

Appearance of ascomata scattered in the substrate (after removing the 

out layer of the substrate). Note the protruding papilla. b Section of an 

ascoma. c Section of the partial peridium. Note the lightly pigmented 

trabeculate pseudoparaphyses and hyaline, muriform ascospores 

as well as its peridium that turns reddish brown in KOH 

(Rossman et al. 1999) (not shown here). Subsequently, C. 

indica Dhaware was introduced from India, and Barr (1983) 

transferred Teichospora ohiensis Ellis & Everh. to Calyptronectria 

as C. ohiensis (Ellis & Everh.) M.E. Barr. 

However, this proposal is inappropriate as the type specimen 

of T. ohiensis is “unitunicate” (Barr 1983; Rossman et al. 

1999). Subsequently, Rossman et al. (1999) transferred 

pseudoparenchymatous cells. d Released ascospores with mucilaginous 

sheath. e Eight-spored asci in hamathecium and embedded in gel 

matrix. f Ascus with a short pedicel. Scale bars: a=0.5 mm, b= 

100 μm, c=50 μm, d–f=10 μm 

Calyptronectria ohiensis to Thyridium (as T. ohiense (Ellis 

& Everh.) Rossman & Samuels). 


None. 


The immersed ascomata, trabeculate pseudoparaphyses, 

bitunicate asci, hyaline and muriform ascospores as well as


the reaction of peridium to KOH (turns reddish brown) 

make it distinguishable from all other reported genera 

(Rossman et al. 1999). Thus Calyptronectria is a morphologically 

well defined genus. 

Carinispora K.D. Hyde, J. Linn. Soc., Bot. 110: 97 (1992). 

(Pleosporales, genera incertae sedis) 


Habitat marine, saprobic. One or two ascomata per 

stroma. Ascomata scattered or in small groups, developing 

beneath the host epidermis, erumpent, lenticular, 

ostiolate, lacking periphyses. Peridium pale brown, 

composed of thin-walled elongated cells at the sides 

and thick-walled cells of textura epidermoidea at the 

base. Hamathecium of dense, long filliform pseudoparaphyses, 

embedded in mucilage, anastomosing between 

and above the asci, rarely septate. Asci 8-spored, 

bitunicate, fissitunicate, clavate to cylindrical, with a 

short furcate pedicel, apex with an ocular chamber and 

apical ring. Ascospores biseriate, narrowly fusoid, yellow 

to pale brown, multi-septate, constricted at the septa, the 

two central cells being the largest, surrounded by a 

gelatinous sheath. 


Literature: Hyde 1992a, 1994b. 

Type species 

Carinispora nypae K.D. Hyde, J. Linn. Soc., Bot. 110: 99 

(1992). (Fig. 19) 

One or two ascomata per stroma. Ascomata up to 0.8 mm 

diam., scattered or in small groups, developing beneath the 

host epidermis, crust-like, as circular spots, wall brown, with 

a small central ostiole, in section 225–285 μm high×510– 

750 μm diam., lenticular, ostiolar canal lacking periphyses 

(Fig. 19a and b). Peridium 35–45 μm wide at sides, pale 

brown, at sides composed of a thin layer of thin-walled 

elongate cells, fusing with the stromatic tissue and host cells, 

at the base composed of thick-walled cells, forming a textura 

epidermoidea and fusing with host cells. A wedge of pale 

brown hyphae forming a textura porrecta is present at the 

rim (Fig. 19c). Hamathecium of dense, long filliform 

pseudoparaphyses 1–3 μm broad, embedded in mucilage, 

anastomosing between and above the asci, rarely septate. 

Asci 142–207×14.2–19.8 μm, 8-spored, bitunicate, fissitunicate, 

clavate to cylindrical, with a furcate pedicel, up to 

40 μm long, apex with an ocular chamber and apical ring (to 

2 μm wide×3 μm high, J-), developing from ascogenous 

tissue at the base of the ascocarp (Fig. 19d, e, f, g and h). 

Ascospores 42–66×7–10.6 μm, biseriate, narrowly fusoid 

with broadly to narrowly rounded ends, somewhat curved, 

yellow to pale brown, yellow in mass, 7-8-septate, constricted 

at the septa, the two central cells being the largest, 

surrounded by a gelatinous sheath; the sheath has a central 

“spine” and curved polar extrusions (Fig. 19i and j). 


Material examined: BRUNEI DARUSSALAM, Tungit 

Api Api mangrove, from decaying intertidal fronds of Nypa 

fruticans Wurmb., 14 Apr. 1987, K.D. Hyde (BRIP 17106, 

holotype). 

Notes 

Morphology 

Carinispora is distinguished from Phaeosphaeria by its 

saprobic life style and lenticular ascomata formed under the 

host epidermis, peridium structure and sheath surrounding the 

ascospores (Hyde 1992a, 1994b). Two species were reported, 

i.e. C. nypae and C. velatispora K.D. Hyde. 


Suetrong et al. (2009) could not resolve Carinispora 

nypae in a phylogeny based on four genes. 


Both Carinispora nypae and C. velatispora are reported 

as marine fungi, which should be taken into consideration 

for their familial placement. 

Caryosporella Kohlm., Proc. Indian Acad. Sci., Pl. Sci. 94: 



Habitat marine, saprobic. Ascomata densely scattered or 

gregarious, superficial, subglobose, black, papillate, ostiolate, 

periphysate, carbonaceous. Peridium carbonaceous. Hamathecium 

of dense, long trabeculate pseudoparaphyses, anastomosing 

and branching above the asci. Asci 8-spored, bitunicate, 

fissitunicate, cylindrical. Ascospores ellipsoidal to broadly fusoid 

with narrowly hyaline rounded ends, deep reddish brown, thickwalled, 

1-septate with hyaline germ pore at each end. 

Anamorphs reported for genus: suspected spermatia 

(Kohlmeyer 1985). 

Literature: Eriksson 2006; Kohlmeyer 1985; Lumbsch and 

Huhndorf 2007. 

Type species 

Caryosporella rhizophorae Kohlm., Proc. Indian Acad. 

Sci., Pl. Sci. 94: 356 (1985). (Fig. 20) 

Ascomata 0.8–1.1 mm high×0.9–1.2 mm diam., densely 

scattered or gregarious, superficial with a flattened base, not 

easily removed from the host surface, subglobose, black, short


papillate, ostiolate, periphysate, carbonaceous (Fig. 20a 

and b). Peridium 120–150 μm thick at sides, up to 

200 μm thick at the apex, thinner at the base, 3-layered, 

outer layer composed of golden-yellow, very thickwalled 

cells of textura epidermoidea, mixed with subglobose, 

large cells near the surface, cells 7–15 μm 

diam., middle layer composed of deep brown, very thickwalled 

cells of textura epidermoidea, inner layer composed 

of hyaline, thin-walled cells of textura prismatica, 

up to 50×5 μm diam., merging with pseudoparaphyses 

(Fig. 20b, c and d). Hamathecium of dense, long 

trabeculate pseudoparaphyses, 1.5-2 μm wide, anastomosing 

and branching above the asci. Asci 225–250 

(−275)×14–17 μm (x ¼ 137 16:3mm, n=10), 8-spored, 

bitunicate, fissitunicate, cylindrical, with a long, narrowed, 

pedicel which is up to 75 μm long, apical 

characters not observed (Fig. 20e). Ascospores 25–28 

(−30)×9–13 μm (x ¼ 26:8 11mm, n=10), uniseriate to 

partially overlapping, ellipsoidal to broadly fusoid with 

narrow hyaline rounded ends, deep reddish brown, thickwalled, 

1-septate with hyaline germ pore at each end, slightly 

constricted at the septum, verruculose, sometimes with “net”- 

like ridged ornamentations (Fig. 20f and g). 

Anamorph: suspected spermatia (Kohlmeyer 1985). 

Material examined: BELIZE, Twin Cays, tip of prop 

root of Rhizophora mangle, 18 Mar. 1984, J. Kohlmeyer 

(NY. Herb. J. Kohlmeyer No. 4532a, holotype). 

Notes 

Morphology 

Caryosporella was formally established by Kohlmeyer 

(1985) based on the obligate marine fungus, C. rhizophorae, 

which is characterized by its superficial ascomata, 

3-layered peridium, filliform trabeculate pseudoparaphyses, 

and brown, 1-septate ascospores. Caryosporella was 

originally assigned to Massariaceae despite several major 

differences, such as the superficial ascomata, reddish 

brown ascospores (Kohlmeyer 1985). Subsequently, 

Caryosporella was assigned to Melanommataceae 

(Eriksson 2006; Lumbsch and Huhndorf 2007). 


Suetrong et al. (2009) showed that a single isolate of 

Caryosporella rhizophorae does not reside in Pleosporales, 

but is related to Lineolata rhizophorae (Kohlm. & E. 

Kohlm.) Kohlm. & Volkm.-Kohlm. and placed in Dothideomycetidae 

incertae sedis. 


As an obligate marine fungus, the familial placement of 

Caryosporella rhizophorae is uncertain but it may not 

belong to Pleosporales. 

Fig. 19 Carinispora nypae (from BRIP 17106, holotype). a b 

Ascomata on the host surface. b Section of an ascoma. c Section of 

a partial peridium. d, e, g, h Asci with ocular chambers and short 

pedicels. f The ocular chamber and apical ring of ascus. i–j Narrowly 

fusoid ascospores. Scale bars: a=1 mm, b, c=100 μm, d, g, h=50 μm, 

e, f, i, j =10 μm 

Chaetomastia (Sacc.) Berl., Icon. fung. (Abellini) 1: 38 

(1890). (Teichosporaceae) 

≡ Melanomma subgen. Chaetomastia Sacc., Syll. fung. 

(Abellini) 2: 113 (1883). 


Habitat terrestrial, saprobic. Ascomata relatively small, 

scattered, or in small groups, superficial, globose or 

subglobose, black, papillate, ostiolate, coriaceous. Peridium 

relatively thin, 1-layered, composed of heavily 

pigmented cells of textura angularis. Hamathecium of 

dense, long cellular pseudoparaphyses, embedded in 

mucilage. Asci mostly 4-spored, bitunicate, fissitunicate, 

broadly cylindrical with a furcate pedicel, with a large 

ocular chamber, especially apparent in immature asci. 

Ascospores ellipsoid to broadly fusoid with broadly to 

narrowly rounded ends, brown, 3-septate, constricted at all 

septa. 

Anamorphs reported for genus: coelomycetous where 

known: conidia hyaline or brown, aseptate or 1-septate 

(Aposphaeria- orConiothyrium-like) (Barr 1989c). 

Literature: Barr 1987b, 1989c; 1993a; b; 2002; Berlese 

1890; Clements and Shear 1931; Eriksson 1999; Eriksson 

and Hawksworth 1987, 1998; Holm 1957; Leuchtmann 

1985; Saccardo 1883. 

Type species 

Chaetomastia hirtula (P. Karst.) Berl., Icon. fung. (Abellini) 

1: 38 (1890). (Fig. 21) 

≡ Sphaeria hirtula P. Karst., Fungi Fenn. Exs. N. 825 (1869). 


scattered or in groups, superficial, globose, wall black; 

apex often opening with a broad pore within slightly 

raised papilla, up to 30 μm diam., coriaceous (Fig. 21a). 

Peridium 20–26 μm thick, 1-layered, composed of 

heavily pigmented cells of textura angularis, cells up to 

5×15 μm diam., cell wall up to 3.5 μm thick (Fig. 21b). 

Hamathecium of dense, long cellular pseudoparaphyses, 

embedded in mucilage. Asci 90–130×12.5–17.5(−22.5) 

μm (x ¼ 111 16:3mm, n=10), mostly 4-spored, bitunicate, 

fissitunicate, broadly cylindrical, with a furcate 

pedicel, 18–48 μm long, with a large ocular chamber 

best seen in immature asci (to 3 μm wide×3 μm high) 

(Fig. 21c and d). Ascospores 20.5–27×7–10 μm

Fungal Diversity


Fig. 20 Caryosporella rhizophoriae (from NY. Herb. J. Kohlmeyer 

No. 4532a, holotype). a Gregarious ascomata on host surface. b 

Section of an ascoma. c, d Section of partial peridium at sides (c) and 

base (d). Note the three layers. e Asci with long peduncles in 

pseudoparaphyses. f, g Ascospores. Note the “net”-like ridged 

ornamentation of spore surface and hyaline germ pores. Scale bars: 

a=1 mm, b=200 μm, c–e=100 μm, f, g=10 μm 

(x ¼ 23:5 8:2mm, n=10), uniseriate to partially overlapping, 

ellipsoid to broadly fusoid with broadly to 

narrowly rounded ends, brown, 3-septate, verruculose, 

constricted at all septa, constricted at the median septum, 

the cell above the central septum often broader than the 

others (Fig. 21e and f).


Fig. 21 Chaetomastia hirtula (from H, FFE 825, kleptotype). a 

Superficial ascomata gregarious on the host surface. b Section of a 

partial peridium. Note the cells of textura angularis with relatively 

thick wall. c, d Cylindrical asci with long and furcate pedicels. e, f 

Brown, 3-septate ascospores. Scale bars: a=0.5 mm, b=50 μm, c–f= 

15 μm 


Material examined: FINLAND, ETELÄ-HÄME (EH/ 

Ta), Tammela, Mustiala, På Rub. id., 8 May 1866. P.A. 

Karsten (H, FFE 825, kleptotype). 

Notes 

Morphology 

Chaetomastia was introduced by Saccardo (1883) asa 

subgenus of Melanomma, and five species were included, 

i.e. M. canescens Speg., M. cucurbitarioides Speg., 

M. hirtulum (P. Karst.) Sacc., M. hispidulum Sacc. and 

M. pilosellum P. Karst. Berlese (1890) promoteditto 

genus rank. Subsequently, Chaetomastia hirtula (P. 

Karst.) Berl. was selected as the lectotype species of 

the genus (Clements and Shear 1931). Chaetomastia has 

been regarded as having unitunicate asci (Eriksson and 

Hawksworth 1986, 1998; Eriksson 1999). However its 

bitunicate status was confirmed by Holm (1957). Holm 

(1957) treated C. hirtula as Melanomma hirtulum (P.


Karst.) Sacc., and Leuchtmann (1985) transferred this 

species to Montagnula sensu lato based on the ascospore 

morphology and the hyphae surrounding the ascomata. 

Barr (1987b) suggested that ascoma, peridium structure 

and ascospore characters pointed Montagnula sensu 

stricto to Phaeosphaeriaceae, while the characters of 

ascomata and peridium structure of Chaetomastia were 

thought to fit the definition of Dacampiaceae (Barr 

1987b). In particular, the peridium and ascospore characters 

of C. hirtula are comparable with those of the generic 

type of Massariosphaeria (M. phaeospora). Thus, Barr 

(1989c) accepted Massariosphaeria sensu stricto and 

assigned the phragmosporous species of Massariosphaeria 

sensu lato to Chaetomastia. 

Barr (2002) later assigned Chaetomastia to Teichosporaceae 

based on its saprobic or hypersaprobic lifestyle, 

occurring on woody stems and peridium structure, and this 

is widely followed (Eriksson 2006; Lumbsch and Huhndorf 

2007). Currently, 11 species are accepted in this genus 

(http://www.indexfungorum.org/). 


None. 


Familial placement of Chaetomastia is undetermined 

currently but has been included in the Teichosporaceae 

by authoritative sources (Eriksson 2006; Lumbsch and 

Huhndorf 2007) or the Dacampiaceae (http://www. 

indexfungorum.org/). 

Chaetoplea (Sacc.) Clem., Gen. Fung. (Minneapolis): 275 

(1931). (?Phaeosphaeriaceae) 

≡ Pyrenophora subgen. Chaetoplea Sacc., Syll. fung. 

(Abellini) 2: 279 (1883). 


Habitat terrestrial, saprobic. Ascomata small to medium, 

immersed, erumpent to superficial, globose to subglobose, 

papillate, ostiolate. Peridium not examined. Hamathecium 

of dense, long, narrowly cellular pseudoparaphyses. Asci 8- 

spored or 4-spored, bitunicate, fissitunicate, cylindroclavate, 

with a thick, furcate pedicel. Ascospores ellipsoid 

or fusoid, pale brown to brown, phragmosporous or 

muriform. 

Anamorphs reported for genus: Microdiplodia-like (Barr 

1990b). 

Literature: Barr 1981; 1987a; b; 1990b; Clements and 

Shear 1931; Ramaley and Barr 1995; Yuan and Barr 

1994. 

Fig. 22 Chaetoplea calvescens (from FH-81113, isotype). a, b Fourspored 

and 8-spored asci. c Released ascospores. Scale bars: a–c= 

10 μm 

Type species 

Chaetoplea calvescens (Fr.) Clem., Gen. Fung. (Minneapolis): 

275 (1931). (Fig. 22) 

≡ Sphaeria calvescens Fr. Scleromyc. Sueciae 401. 

Ascomata not examined. Peridium not examined. 

Hamathecium of dense, long, narrow cellular pseudoparaphyses, 

2–3 μm broad, septate, branching and anastomosing. 

Asci 90–110×10–12 μm, 8-spored, rarely 4-spored, 

bitunicate, fissitunicate, cylindro-clavate, with a thick, 

furcate pedicel which is up to 30 μm long (Fig. 22a 

and b). Ascospores 13–18×5.5–7 μm, obliquely uniseriate 

and partially overlapping, broadly fusoid to oblong 

with broadly rounded ends, pale brown, 2-3-septate, 

constricted at the septa, containing four refractive globules 

(Fig. 22c). 

Note: The specimen is only a slide, and no peridium or 

ascomata information could be obtained. 

Anamorph: coelomycetous, conidia yellowish, 1- 

septate, 9–13×4–5(−8) μm (Webster and Lucas 1959); 

Microdiplodia henningsii Staritz=Chaetodiplodia caudina 

Karst. (Sutton 1980) (referred to Barr 1990b (p50)). 

Material examined: SWEDEN, sub-collection: Curtis 

Herbarium, verified by R.A. Shoemaker, leg. E.M. Fries 

401 (FH-81113, isotype, microscope slide). 

Notes 

Morphology 

Chaetoplea was introduced based on C. calvescens, 

which has been regarded as similar to Pleospora or 

Leptosphaeria (Eriksson and Hawksworth 1987; 

Wehmeyer 1961; von Arx and Müller 1975). Based on 

the differences in ascomata, peridium structure, pseudoparaphyses 

as well as its anamorphic stage, Chaetoplea 

was maintained as a separate genus (Barr 1990b; 

Yuan and Barr 1994). Chaetoplea sensu lato was 

accepted by Barr (1990b), which included some species 

of Teichospora as well as the subgenus Pleospora subg. 

Cylindrosporeae. 

The following is from the label of specimen. 

“Sphaeria calvescens, Scler. Suecicae (Ed. 2) 401. No 

specimen of Scler. Suecicae 401 is now at Uppsala 

according to R. Santesson 1966. This Curtis Herbarium 

specimen in the Farlow Herbarium is isotype. 

Wehmeyer (1961) in his Pleospora monograph did 

not study any portion of the Scler. Suecicae exsiccatus 

b

Fungal Diversity


401, nor did Webster & Lucas in the taxonomic and 

life-history study (Trans. Brit. Myc. Soc. 42, 332– 

342. 1959) of this species. 

The specimen has most of the features described by 

Webster & Lucas including the presence of the 

conidial state Microdiplodia henningsii Staritz. I did 

not see vertical septa in the ascospores. Webster & 

Lucas note that vertical septa may be occasionally be 

lacking. The fungus is otherwise as they describe it 

although some perithecia collapse and appear cupulate.”—by 

R.A. Shoemaker. 


None. 


The substrate of Chaetoplea sensu Barr (1990b) can 

be herbaceous stalks, decorticated wood or periderm, or 

old cotton cloth and string, which may indicate its 

heterogeneous nature. The ascospores seem very much 

like Phaeosphaeria which may be an earlier name; more 

details concerning the ascomatal, peridial and hamathecial 

structures are needed to make any conclusion. 

Cilioplea Munk, Dansk botanisk Arkiv 15: 113 (1953). 



Habitat terrestrial, saprobic. Ascomata small- to medium-sized, 

solitary, scattered or in small groups, immersed, globose or 

subglobose, papilla covered with short and blackish setae, 

coriaceous. Peridium thin, comprising small heavily pigmented 

thick-walled cells of textura angularis. Hamathecium 

of cellular pseudoparaphyses. Asci 8-spored, bitunicate, 

fissitunicate, broadly clavate, with a short, furcate pedicel, and 

small ocular chamber. Ascospores fusoid to narrowly fusoid 

with narrowly rounded ends, pale brown to reddish brown, 

multi-transverse septa, usually with one longitudinal septum in 

some central cells, constricted at the primary septum. 


Literature: Barr 1990b, 1992b; Crivelli 1983; Lumbsch 

and Huhndorf 2007; Müller 1951; Munk 1953, 1957. 

Type species 

Cilioplea coronata (Niessl) Munk, Dansk botanisk Arkiv 

15: 113 (1953). (Fig. 23) 

≡ Pleospora coronata Niessl, Notiz. Pyr.: 16 (1876). 


scattered, or in small groups, immersed, globose or subglobose, 

wall black, papilla raised, 50–80 μm high, with short 

and blackish setae, coriaceous (Fig. 23a). Peridium 9–15 μm 

thick laterally, up to 28 μm thick at the apex, thinner at the 

base, 1-layered, composed of small heavily pigmented thickwalled 

cells of textura angularis, cells up to 4×2.5 μm diam., 

cell wall 2–3 μm thick, apex cells smaller and walls thicker 

(Fig. 23b). Hamathecium of long cellular pseudoparaphyses, 

2–3 μm broad. Asci (60-)80–108×10–15 μm 


broadly clavate, with a short, thick, furcate pedicel, 5– 

15 μm long, and a small ocular chamber (to 3 μm wide× 

2 μm high) (Fig. 23c and d). Ascospores 21–27.5×5.5– 

7.5 μm (x ¼ 24 6:7mm, n=10), biseriate to uniseriate at 

base, fusoid to narrowly fusoid with narrowly rounded ends, 

pale reddish brown, 5–7 transverse septa (mostly 5), usually 

with one longitudinal septum in some central cells, deeply 

constricted at the median septum, the part above the primary 

septum shorter and broader, smooth-walled. 


Material examined: GERMANY, Hadiberg. on Reseda 

lutea Hadiberg, 20 Sept. 1875, Niessl (M 175-89-290, 

lectotype; M 175-89-291, type). 

Notes 

Morphology 

Cilioplea was introduced by Müller (1951) as a 

subgenus of Pleospora, and this was followed by Munk 

(1957), who had earlier proposed it as a separate genus 

typified by C. coronata based on its hairy papilla, clavate 

asci as well as its “perfectly paraphysoid” (see Munk 

1953). A relatively narrow concept of Pleospora was 

accepted by Crivelli (1983), and four species was 

assigned under the separate genus Cilioplea, viz. C. 

coronata, C. genisticola (Fautrey & Lambotte) Crivelli, 

C. kansensis (Ellis & Everh.) Crivelli and C. nivalis 

(Niessl) Crivelli. Subsequently, another six species were 

added (Barr 1990b, 1992b). Currently, ten species are 

included under Cilioplea. 


None. 


The most striking character of Cilioplea is its setose 

papilla, which has been shown to have no phylogenetic 

significance in Lentitheciaceae (Zhang et al. 2009a). 

Cilioplea was assigned under Lophiostomataceae (Lumbsch 

and Huhndorf 2007), but there is little morphological 

similarity with the Lophiostomataceae sensu stricto (Zhang 

et al. 2009a). Thus its familial placement needs further study. 

Crivellia Shoemaker & Inderb., in Inderbitzin, Shoemaker, 

O’Neill, Turgeon & Berbee, Can. J. Bot. 84: 1308 (2006). 

(Pleosporaceae)


Fig. 23 Cilioplea coronata (M 175-89-290, lectotype). a Immersed 

ascomata in small groups on the host surface (the covering host tissue 

was removed). b Section of a partial ascoma. Note the thin peridium. c 

Clavate asci within pseudoparaphyses. d Ascus with a small ocular 

chamber. Scale bars: a=0.5 mm, b=100 μm, c=50 μm, d=10 μm 


Habitat terrestrial, hemibiotrophic or parasitic. Ascomata 

small- to medium-sized, scattered, immersed, erumpent to 

nearly superficial, papillate, ostiolate. Peridium thin, composed 

of two cells types, outer cells of thick walled and 

textura angularis, inner cells thin-walled, yellow. Hamathecium 

of dense, long and thin pseudoparaphyses. Asci (4-) 

8-spored, bitunicate, fissitunicate dehiscence not observed, 

broadly cylindrical to cylindrical, with a short, 

furcate pedicel and an ocular chamber. Ascospores 

fusoid to broadly fusoid, pale brown, septate, sometimes 

with one or two vertical septa in the middle cells, constricted 

at the septa. 

Anamorphs reported for genus: Brachycladium (Inderbitzin 

et al. 2006). 

Literature: Inderbitzin et al. 2006. 

Type species 

Crivellia papaveracea (De Not.) Shoemaker & Inderb., 

Can. J. Bot. 84: 1308 (2006). (Fig. 24) 

≡ Cucurbitaria papaveracea De Not., Sfer. Ital.: 62 

(1863). 


densely scattered, immersed, erumpent to nearly superficial, 

flattened globose, dark brown, papillate, ostiolate 

(Fig. 24a). Peridium 25–30 μm thick, thicker near the apex 

and thinner at the base, composed of two cell types, outer 

cells of thick-walled and textura angularis, cells up to 10× 

5 μm diam., cell wall 2–4 μm thick, inner cells thin-walled, 

yellow (Fig. 24b). Hamathecium of dense, long, 1–2 μm


broad, rarely septate pseudoparaphyses. Asci 85–125×10– 

13 μm (x ¼ 106 11mm, n=10), (4-)8-spored, bitunicate, 

fissitunicate dehiscence not observed, broadly cylindrical to 

cylindrical, with a short, furcate pedicel, with a relatively large 

ocular chamber (Fig. 24c and d). Ascospores (16-)19–24×5– 

7.5 μm (x ¼ 20:4 6:3mm, n=10), overlapping uniseriate to 

rarely biseriate, fusoid to broadly fusoid, pale brown, 3- 

septate, sometimes with one or two vertical septa in the 

middle cells, constricted at the septa, the upper cell often 

broader than the lower one, smooth-walled. 

Anamorph: Brachycladium penicillatum (Corda) Fr. 

(Inderbitzin et al. 2006). 

Material examined: AUSTRIA, Vienna, on decaying 

stems of Papaver rhoeas L., 28 Oct. 2001, W. Jaklitsch 

(UBC F14995, epitype). 

Notes 

Morphology 

Crivellia was separated from Pleospora and introduced 

as a new genus by Inderbitzin et al. (2006) based on their 

differences in ascospore morphology and anamorphic 

stages. Crivellia is characterized by having small- to 

medium-sized ascomata, and yellow, 3-septate ascospores 

Fig. 24 Crivellia papareracea 

(from UBC F14995, epitype). 

a Gregarious ascomata immersed 

within the host surface. 

b Section of an ascoma. c Asci 

within pseudoparaphyses. d 

Cylindrical ascus with a 

short pedicel. Scale bars: 

a=1 mm, b=100 μm, c, 

d=30 μm


with one or two vertical septa in central cells. Its 

Brachycladium anamorphic stage with phragmosporous 

conidia also differs from that of Stemphylium, which is 

the anamorphic stage of Pleospora (Inderbitzin et al. 2006). 

Currently, two species are included within Crivellia, i.e. C. 

homothallica Inderb. & Shoemaker and C. papaveracea. 


Crivellia papaveracea was shown to be closely related to 

some species of Alternaria, and its pleosporaceous status was 

confirmed following molecular studies (Inderbitzin et al. 2006). 


Crivellia seems to belong to Pleosporaceae, and may be 

closely related to Pleospora. 

Decaisnella Fabre, Annls Sci. Nat., Bot., sér. 6 9:112 



Habitat terrestrial, saprobic. Ascomata medium to large, 

immersed to erumpent, clypeate, papillate, ostiolate. Hamathecium 

of dense, long, cellular pseudoparaphyses, rarely 

septate, embedded in mucilage. Asci mostly 4- or 8-spored, 

rarely 2-spored, cylindrical to cylindro-clavate, with a 

furcate pedicel. Ascospores muriform, dark brown, oblong 

with broadly rounded ends. 


Literature: Barr 1986; 1990a; b; Fabre 1878; Saccardo 

1883. 

Type species 

Decaisnella spectabilis Fabre, Annls Sci. Nat., Bot., sér. 6 

9: 112 (1879). (Fig. 25) 


solitary, scattered, or in small groups of 2–3, immersed 

to erumpent, clypeate, globose or subglobose, black, 

roughened, with a blunt papilla up to 170 μm high, 

apex with a round ostiole, coriaceous (Fig. 25a). 

Peridium 70–90 μm thick at sides, thicker near the apex, 

comprising two types of cells; part immersed in host 

tissue, outer layer pseudoparenchymatous, 55–65 μm 

thick, pigmented, inner layer composed of lightly pigmented 

to hyaline thin-walled compressed cells, 15– 

23 μm thick, cells 3.5–7 μm diam., part above host 

tissue heavily pigmented covered by clypeus tissues 

(Fig. 25b). Hamathecium of dense, long, cellular pseudoparaphyses, 

1.5–3 μm broad, rarely septate, embedded in 

mucilage. Asci 150– 200×15– 25(− 33) μ m 

(x ¼ 181 20:6mm, n=10), (2-)4-spored, bitunicate, fissitunicate, 

broadly cylindrical, with a short, thick, furcate 

pedicel which is 20–40 μm long, no apical apparatus 

observed (Fig. 25e). Ascospores 37–45×12–17 μm 

(x ¼ 43 15mm, n=10), uniseriate and sometimes slightly 

overlapping, oblong with broadly rounded ends, dark 

brown, verrucose or smooth, 7–9 transverse septa and 1– 

3 longitudinal septa in some of the cells, no constriction at 

the septa (Fig. 25c and d). 


Material examined: GERMANY, Valsalpe in der Ramsau, 

Bayer, Alpen, on Rhamnus pumila Turra., Jul. 1913, Karl Arnold 

(NY2082, syntype as Teichospora megalocarpa Rehm). 

Notes 

Morphology 

Decaisnella was formally established by Fabre (1879), 

but was treated as a synonym of Teichospora by 

Saccardo (1883). This was followed by several mycologists 

over a long time. The main morphological differences 

between Decaisnella and Teichospora include the 

size and septation of ascospores, shape of ascomata, 

structure of peridium and type of pseudoparaphyses (Barr 

1986). Thus Barr (1986) revivedDecaisnella and assigned it 

to Massariaceae based on the shape of ascomata and large, 

distoseptate ascospores. Currently, 15 species are accepted 

under Decaisnella (http://www.mycobank.org/MycoTaxo. 

aspx). Neither the size of ascomata nor the ascospore 

characters have proven sufficient to place taxa at the family 

level in Pleosporales (Zhang et al. 2009a), and therefore 

familial placement of Decaisnella remains uncertain. 


Decaisnella formosa resided in the clade of Lophiostomataceae 

and in proximity to Lophiostoma macrostomoides 

De Not. (Plate 1). 


The muriform ascospores, saprobic life style and 4- 

spored asci point Decaisnella spectabilis to Montagnulaceae, 

but this can only be confirmed following a molecular 

phylogenetic study. 

Delitschia Auersw., Hedwigia 5: 49 (1866). (Delitschiaceae) 


Habitat terrestrial, saprobic (coprophilous). Ascomata 

medium- to large-sized, solitary or scattered, immersed to 

erumpent, globose or subglobose, apex with or without 

papilla, ostiolate. Peridium thin, composed of compressed 

cells. Hamathecium of dense, long pseudoparaphyses, 

anastomosing and branching. Asci 8-spored, cylindrical to 

cylindro-clavate, with short pedicel. Ascospores uni- to


Fig. 25 Decaisnella spectabilis (NY2082, syntype). a Appearance of 

ascomata on the host surface. b Section of a partial peridium 

(immersed in the substrate). Note the pseudoparenchymatous out 

triseriate, pale to dark brown, ellipsoid, 1-septate, usually 

constricted at the septum, smooth, with a full length germ 

slit in each cell. 


layer. c, d Muriform ascospores. Note the minuitely verrucose 

ornamentation. e Ascus with a short pedicel. Scale bars: a=0.5 mm, 

b=100 μm, c–e=20 μm 

Literature: Auerswald 1866; Barr 2000; Cain 1934; 

Dennis 1968; Eriksson 2006; Griffiths 1901; Hyde and 

Steinke 1996; Kirschstein 1911; Kruys et al. 2006; Luck- 

Allen and Cain 1975; Lumbsch and Huhndorf 2007;


Moreau 1953; Munk 1957; Romero and Samuels 1991; 

Schoch et al. 2006; Winter 1887. 

Type species 

Delitschia didyma Auersw., Hedwigia 5: 49 (1866). 

(Fig. 26) 

Ascomata 400–800 μm diam., solitary or scattered, 

immersed, globose or subglobose,black,papillashort,70– 

130 μm broad, central, with a wide opening, coriaceous 

(Fig. 26a). Peridium ca. 15μm thick laterally, up to 35 μm 

thick at the apex, up to 30 μm at the base, comprising a 

single layer of small lightly pigmented thin-walled cells of 

textura angularis, cells 4–10 μm diam., cell wall


Fig. 26 Delitschia didyma 

(from L, 1950). a Ascomata on 

the substrate surface. Note the 

ostiolar opening. b Section of 

peridium. Note the small cells of 

textura angularis. c Released 

and unreleased ascospores. Note 

the germ slit in each cell. d, e 

Asci with ascospores and short 

pedicels with rounded ends. 

Scale bars: a=0.5 mm, 

b =30 μm, c–e=70 μm 

Type species 

Didymosphaeria futilis (Berk. & Broome) Rehm, Hedwigia 

18: 167 (1879). (Fig. 27) 

≡ Sphaeria futilis Berk. & Broome, Ann. Mag. nat. 

Hist., Ser. 2 9: 326 (1852). 

Ascomata 190–230 μm high×240–340 μm diam., scattered, 

or in small groups, immersed to slightly erumpent, 

subglobose to ovoid, membraneous, near-hyaline, under 

clypeus, papillate, periphysate (Fig. 27a and c). Papilla 

central, up to 100 μm high, black, with a pore-like ostiole 

(Fig. 27a and c). Peridium 30–40 μm wide upper part, 6– 

23 μm wide near the base, 1-layered, composed of brown 

pseudoparenchymatous cells of textura angularis, cell wall 2– 

3 μm thick (Fig. 27b). Hamathecium of dense, long 

trabeculate pseudoparaphyses, 0.8–1.5 μm broad, anastomosing 

mostly above the asci, embedded in mucilage (Fig. 27d). 

Asci 90–110×7.5–10 μm (x ¼ 97 9mm, n=10), 2–4-spored, 

rarely 8-spored, bitunicate, fissitunicate, cylindrical, with a 

furcate pedicel, 17.5–27.5 μm long, with a large ocular (to 

2.5 μm wide×4 μm high) (Fig. 27d, e and f). Ascospores 14– 

15.5×(5.5-) 6–7.5 μm (x ¼ 14:8 6:9mm, n=10), uniseriate, 

ellipsoid with obtuse ends, brown, 1-septate, distoseptate, 

slightly to not constricted, capitate (Fig. 27g). 

Anamorph: Dendrophoma sp., Fusicladiella sp. vel aff. 

(Sivanesan 1984).


Material examined: UK, England, Norfolk, King’s 

Cliffe; on dead stem (in ramis emortuis) Rosa sp., Mar. 

1850, M.J. Berkeley (K(M): 147683, holotype). 

Notes 

Morphology 

Didymosphaeria is a widely distributed genus with wide 

host range (Aptroot 1995). Didymosphaeria was formally 

established by Fuckel (1870) based on six ascomycetous 

species, and D. epidermidis (Fries) Fuckel (or D. peltigerae 

Fuckel) has been chosen as the lectotype species (see 

comments by Aptroot 1995). Hawksworth and David 

(1989: 494) proposed to conserve the genus with a lectotype 

specimen, Fungi Rhenani 1770. The genus had been 

considered as a depository to accommodate all types of 

didymosporous pyrenocarpous ascomycetes. Many workers 

have tried to redefine the genus and excluded some species. 

Saccardo (1882) restricted the genus to brown-spored species, 

and about 100 species have been excluded subsequently (Barr 

1989a, b, 1990a, 1992a, b, 1993b; Hawksworth1985a, b; 

Hawksworth and Boise 1985; Hawksworth and Diederich 

Fig. 27 Didymosphaeria futilis 

(from K(M): 147683, holotype). 

a Two immersed ascomata on 

the host surface (one of them is 

cut horizontally). b Section of 

an ascoma. Note the thin peridium. 

c Hand cut portion of 

ascoma showing habitat in 

wood. d Asci in pseudoparaphyses. 

Note the trabeculate pseudonparaphyses 

anastomosing 

above the asci. e, f Four-spored 

asci with long pedicels which 

are rounded at their bases. g 

Brown, 1-septate ascospores 

with spinulose ornamentation. 

Scale bars: a=0.3 mm, b, c= 

100 μm, d–g=10 μm


1988; Scheinpflug 1958). Over 400 epithets of Didymosphaeria 

were included until the monograph of Aptroot (1995). 

Aptroot (1995) examined more than 3000 specimens under 

the name Didymosphaeria. The type specimen of Didymosphaeria 

(Fungi Rhenani 1770) represents the widespread and 

common D. futilis (Aptroot 1995). In this study, we did not 

get the lectotype specimen, but described the type of D. futilis 

(Sphaeria futilis). Using a narrow concept (ignoring differences 

of host or country of origin), Aptroot (1995) accepted 

only seven species, which were closely related with the 

generic type of Didymosphaeria with over 100 synonyms 

distributed among them. Many taxa were found to belong to 

other groups, i.e. Aaosphaeria, Amphisphaeria, Astrosphaeriella, 

Dothidotthia, Flagellosphaeria, Kirschsteiniothelia, Megalotremis, 

Montagnula, Munkovalsaria, Mycomicrothelia, 

Parapyrenis or Phaeodothis. Didymosphaeria is mainly characterized 

by a peridium consisting of flattened or irregular cells 

or completely hyphae; a hamathecium consisting of narrow, 

trabeculate paraphysoids or paraphyses, richly anastomosing 

above the asci; and brown thinly distoseptate ascospores. 

Didymosphaeriaceae was maintained as a separated family 

within Pleosporales by Aptroot (1995) because of the 

distoseptate ascospores and trabeculate pseudoparaphyses 

mainly anastomosing above the asci. This proposal, however, 

has not received much support (Lumbsch and Huhndorf 2007). 


There have been few molecular investigations of 

Didymosphaeria when compared to the morphological 

studies. Didymosphaeria futilis resided in the clade of 

Cucurbitariaceae (or Didymosphaeriaceae) (Plate 1). The 

correct identification of the Didymosphaeria strain used for 

sequencing, however, has not been verified. 


Didymosphaeria is a well established genus represented 

by D. futilis. Of particular significance are the narrow 

pseudoparaphyses which anastomose above the asci and 

brown 1-septate ascospores with indistinct distosepta. Familial 

placement of Didymosphaeria is unclear yet because 

of insufficient molecular data. 

Dothidotthia Höhn., Ber. Deutsch. Bot. Ges. 36: 312 

(1918). (Didymellaceae) 


Habitat terrestrial, saprobic. Ascomata medium-sized, solitary, 

clustered or somewhat gregarious, erumpent, subglobose, 

apex somewhat papillate to depressed, coriaceous. Peridium 

composed of a few layers of dark brown cells of textura 

angularis, and giving rise dark brown, thick-walled hyphae in 

the basal region, 2-layered. Hamathecium septate pseudoparaphyses 

branched in upper part above asci. Asci 8-spored, 

bitunicate, clavate, straight to curved. Ascospores biseriate to 

obliquely uniseriate, ellipsoid, pale brown, 1-septate. 

Anamorphs reported for genus: Dothiorella and Thyrostroma 

(Hyde et al. 2011; Phillips et al. 2008). 

Literature: Barr1989b; Phillips et al. 2008. 

Type species 

Dothidotthia symphoricarpi (Rehm) Höhn., Ber. Deutsch. 

Bot. Ges. 36: 312 (1918). (Fig. 28) 

≡ Pseudotthia symphoricarpi Rehm, Ann. Mycol. 11: 

169 (1913). 

Ascomata up to 500 μm high×550 μm diam., gregarious 

clustered, rarely solitary, erumpent, subglobose, apex somewhat 

papillate to depressed, coriaceous (Fig. 28a). Peridium 20– 

80 μm thick, composed of 3–6 layers of dark brown cells of 

textura angularis, giving rise dark brown, thick-walled hyphae 

in the basal region, 2-layered, outer layer wall thicker and inner 

layer wall thinner (Fig. 28b). Hamathecium hyaline, septate 

pseudoparaphyses, 2–3 μm wide, branched in upper part above 

asci. Asci 70–120×15–22 μm, 8-spored, bitunicate, clavate, 

straight to curved (Fig. 28c, d and e). Ascospores (20-)22–23 

(−26)×(8-)9–10(−11) μm, biseriate to obliquely uniseriate and 

partially overlapping, ellipsoid tapering towards subacutely 

rounded ends, pale brown, 1-septate, constricted at the septum, 

smooth (Fig. 28f) (description referred to Phillips et al. 2008). 

Anamorph: Thyrostroma negundinis (Phillips et al. 2008). 

Material examined: USA, North Dakota, on branches 

of Symphoricarpos occidentalis Hook. (NY, holotype); 

Colorado, San Juan Co, c. 0.5 mile up Engineer Mountain 

Trail from turnoff at mile 52.5, Hwy 550, dead twigs of 

Symphoricarpos rotundifolius A. Gray, 24 Jun. 2004, A.W. 

Ramaley 0410 (BPI 871823, epitype). 

Notes 

Morphology 

Dothidotthia was formally established to accommodate 

Pseudotthia symphoricarpi (Montagnellaceae, Dothideales) 

(von Höhnel 1918a). Many mycologists considered Dothidotthia 

closely related to a genus of Venturiaceae such as 

Dibotryon by Petrak (1927), or Gibbera by von Arx and 

Müller (1954) and Müller and von Arx (1962). Dothidotthia 

had been treated as a synonym of Gibbera (von Arx 1954; 

Müller and von Arx 1962), which was followed by 

Shoemaker (1963) and Eriksson and Hawksworth (1987). 

Based on the coelomycetous anamorphic stage and peridium 

structure, shape of asci, as well as morphology of 

pseudoparaphyses, Barr (1987b, 1989b) retrieved Dothidotthia, 

and considered it closely related to Botryosphaeria 

(Botryosphaeriaceae). Currently, 11 species are included 

within Dothidotthia (http://www.mycobank.org, 01–2011).


Fig. 28 Dothidotthia symphoricarpi 

(from NY, holotype). a 

Clustered ascomata on the host 

stubstrate. b Longitudinal section 

through an ascoma. c, d 

Asci with pale brown, 1-septate 

ascospores. e Immature asci. f 

Pale brown, 1-septate ascospores 

within asci. g Conidia of 

Thyrostroma anamorph in association 

with ascomata. Scale 

bars: a=0.5 mm, b=100 μm, 

c–g=10 μm. (figure with permission 

from Phillips et al. 

2008) 


Based on a multi-gene phylogenetic analysis, Dothidotthia 

formed a separate familial clade (Phillips et al. 2008). 

Thus Dothidotthiaceae was introduced to accommodate it 

(Phillips et al. 2008). 


By comparing the morphological characters and phylogenetic 

dendrograms by Phillips et al. (2008) and de Gruyter et al. 

(2009), Dothidotthia seems closely related to Didymellaceae, 

but Dothidotthiaceae should still be treated as a separate family.


Dubitatio Speg., Anal. Soc. cient. argent. 12: 212 

(1881). (Arthopyreniaceae (or Massariaceae)) 



densely scattered, or in small groups of 2–4, 

immersed, covered with white crystaline rim, papillate, 

ostiolate. Hamathecium of dense pseudoparaphyses, long, 

2–3 μm broad, branching and anastomosing. Asci cylindrical, 

pedicellate, with furcate pedicel. Ascospores 1-septate, 

asymmetrical, reddish to dark brown. 

Anamorphs reported for genus: Aplosporella-like (Rossman 

et al. 1999). 

Literature: Barr 1979b, 1987b; Müller and von Arx 1962; 

Rossman et al. 1999; Spegazzini 1881. 

Type species 

Dubitatio dubitationum Speg., Anal. Soc. cient. argent. 12: 

212 (1881). (Fig. 29) 


solitary, densely scattered, or in small groups of 2–4, 

immersed, with a protruding papilla, 110–160 μm high, 

160–250 μm diam., globose or subglobose, black, 

covered with white crystalline material which becomes 

hyaline and gel-like in water, ostiolate (Fig. 29a and b). 

Peridium 18–25 μm thick laterally (excluding the rim), up 

to 35 μm thick at the apex, thinner at the base, 1-layered, 

composed of small pale brown thin-walled cells of textura 

prismatica, cells 5–12×3–5 μm diam., cell wall up to 

1 μm thick, apex cells smaller and walls thicker 

(Fig. 29b). Hamathecium of dense, long pseudoparaphyses, 

2–3 μm broad, branching and anastomosing between 

and above the asci. Asci 150–190(−230)×12.5–15 μm 

(x ¼ 172:5 13:4mm, n =10), (6-)8-spored, rarely 4- 

spored, bitunicate, fissitunicate, cylindrical, with a furcate 

pedicel which is up to 40 μm long, ocular chamber not 

observed (Fig. 29c, d and e). Ascospores 19–22.5×10– 

12 μm (x ¼ 20:2 11:4mm, n=10), uniseriate to obliquely 

uniseriate and partially overlapping, broadly ellipsoid 

with broadly to narrowly rounded ends, reddish brown, 

1-septate, constricted at septum, asymmetric with a 

larger upper cell, thick-walled, possibly distoseptate 

(Fig. 29f, g and h). 

Anamorph: Aplosporella-like (for detailed description 

see Rossman et al. 1999). 

Conidiomata globose, ca. 300 μm diam. Conidia 

holoblastic, broadly fusoid, 13–15×7–10 μm, dark brown, 

finely spinulose (Rossman et al. 1999). 

Material examined: ARGENTINA, Buenos Aires, 

Tuyu, on Celtis tala Gill., Jan. 1881, leg. det. C. Spegazzini 

(NY, isotype; LPS, holotype). 

Notes 

Morphology 

When established Dubitatio, Spegazzini (1881) considered 

it as intermediate between Sphaeriaceae and Nectriaceae 

as has been mentioned by Rossman et al. (1999). Müller 

and von Arx (1962) treated Dubitatio as a synonym of 

Passerinula, while the differences of ascomata and ascospores 

could easily distinguish these two genera (Rossman et al. 

1999). After checking the type specimen, Dubitatio was assigned 

to Dothideomycetes, and considered closely related to Dothivalsaria 

in the Massariaceae (Barr 1979b, 1987b). Dubitatio 

chondrospora was assigned to Pseudomassaria (as P. chondrospora 

(Ces.) Jacz.) (Barr 1964; Müller and von Arx 1962). 


None. 


The black ascomata with white crystalline covering and 

central white ostiolar region as well as the asymmetrical 

reddish brown ascospores are striking characters of Dubitatio 

dubitationum. The genus cannot be assigned to any 

family with certainty based on morphological characters 

and fresh collections are needed for sequencing. 

Entodesmium Reiss, Hedwigia 1: 28 (1854). 



Habitat terrestrial, saprobic (or parasitic?). Ascomata 

scattered or in small groups, immersed, papillate, ostiolate, 

periphysate. Peridium thin, comprising one cell type of 

pigmented pseudoparenchymatous cells. Hamathecium of 

dense, long pseudoparaphyses, septate, embedded in mucilage. 

Asci 8-spored, bitunicate, fissitunicate, cylindrical, with 

furcate pedicel. Ascospores ellipsoid to filliform, multiseptate, 

deeply constricted at the primary septum (usually 

near apex), breaking into partspores. 


Literature: von Arx and Müller 1975; Barr1992b; Eriksson 

1967a; b; Holm1957; Liewetal.2000; Shoemaker 1984a, b. 

Type species 

Entodesmium rude Reiss, Hedwigia 1: 28 (1854). (Fig. 30) 

Ascomata 160–250 μm high×150–300 μm diam., in groups, 

immersed with long and protruding cylindrical papilla, 

globose to subglobose, black, coriaceous (Fig. 30a). Papilla 

100–220 μm long, 70–120 μm broad, cylindrical, with 

periphysate ostiole. Peridium 25–33 μm wide, comprising 

pseudoparenchymatous cells, cells up to 10×7.5 μm diam., cell 

wall up to 2 μm thick, beak cells smaller and wall thicker


Fig. 29 Dubitatio dubitationum (from NY, isotype; LPS, holotype). 

a Appearance of ascomata scattered on the host surface. Note the 

exposed white covering around the ostioles. b, c Section of an ascoma. 

(Fig. 30b and c). Hamathecium of dense, long pseudoparaphyses, 

septate, 2–3 μm wide, embedded in mucilage. Asci 

100–175×8–13 μm (x ¼ 147:5 11:3mm, n =10), 8- 

spored, bitunicate, fissitunicate, cylindrical, with a furcate 

pedicel which is 18–50 μm long, and with a low ocular 

chamber (ca. 1μm wide×1 μm high) (Fig. 30e,f, g and h). 

Note the white covering (see arrow). d–f Cylindrical asci with short 

furcate pedicels. g, h Asymmetrical, 1-septate reddish-brown ascospores. 

Scale bars: a=1 mm, b=100 μm, c-e=50 μm, f–h=20 μm 

Ascospores 108–138×3–3.5 μm (x ¼ 123 3:2mm, n=10), 

filliform, brown, multi-septate, breaking into 22–28 partspores, 

5–7×3–3.5 μm diam. (Fig. 30d). 


Material examined: GERMANY, Königstein, on stems of 

Coronilla varia L., 20 May 1895, W. Krieger (H, Krieger 1070).


Fig. 30 Entodesmium rude (from H, Krieger 1070). a Ascomata in 

groups on the host surface. Note the erumpent papilla which is 

cylindrical and has an inconspicuous ostiole. b Section of part of an 

ascoma. Note the arrangement of asci and pseudoparaphyses. c Section 

of the peridium comprising cells of textura angularis. d Part-spores 

inside the ascus. e Relatively immature ascus with filliform ascospores 

and low ocular chamber. f–h Mature and immature asci with pedicels. 

Scale bars: a=0.5 mm, b, c=50 μm, d–h=10 μm 

Notes 

Morphology 

Entodesmium is characterized by having immersed 

ascomata dark cylindrical, periphysate papillae, numerous 

clavate to cylindrical asci surrounded by narrowly 

cellular pseudoparaphyses, and ellipsoidal to filliform 

multi-septate ascospores (Barr 1992b; Shoemaker 

1984b). Currently, five species, viz. Entodesmium eliassonii 

L. Holm, E. lapponicum (L. Holm) L. Holm, E. 

mayorii (E.Müll.)L.Holm,E. niessleanum (Rabenh. ex 

Niessl) L. Holm and E. rude are accepted in this genus 

(Holm 1957; Shoemaker 1984b). Von Arx and Müller 

(1975) assignedEntodesmium to the Pleosporaceae sensu 

lato, and Shoemaker (1976) assigned E. rude (as 

Ophiobolus rudis) to Ophiobolus sensu lato based on 

the fragmenting filliform ascospores. According to the 

short, blackish beak and periphysate ostiole, Barr (1992b)


assigned Entodesmium to Lophiostomataceae. Thehosts 

of Entodesmium are restricted to stems of legumes (Barr 

1992b; Shoemaker 1984b). 


Limited phylogenetic studies indicate that Entodesmium 

rude may have affinities to Phaeosphaeriaceae (Liew et al. 

2000; Plate 1). 


Species of Entodesmium share several morphological 

characters, such as immersed, papillate ascomata, periphysate 

ostioles, pale yellow to light yellowish brown, multiseptate 

(≥ 3), narrowly fusoid to filliform ascospores, and 

are specific to legumes. All of the above similarities 

indicate a close relationship among members of Entodesmium. 

We do not agree with Barr (1992b) who assigned 

Entodesmium to Lophiostomataceae because the ascomata 

are immersed, the papilla are not laterally compressed 

and the peridium comprises a single type of cells of 

textura angularis. These characters plus multi-septate, 

lightly pigmented ascospores, which break up into 

partspores and host specificity to legumes support inclusion 

in Phaeosphaeriaceae. Entodesmium multiseptatum 

(G. Winter) L. Holm and E. niessleanum were originally 

described as Leptosphaeria species (Shoemaker 1984b) 

indicating their similarity to Phaeosphaeria with which 

Leptosphaeria is commonly confused (Shoemaker 1984a; 

Shoemaker and Babcock 1989b). Phylogenetic study has 

also shown that Entodesmium rude is related to members 

of Phaeosphaeriaceae (Liew et al. 2000). Thus we assign 

Entodesmium to Phaeosphaeriaceae as a separate genus 

until further phylogenetic analysis is carried out on 

verified specimens. 

Eudarluca Speg., Revta Mus. La Plata 15: 22 (1908). 



Habitat terrestrial, parasitic. Ascomata small, solitary, 

scattered, immersed to erumpent, subglobose, ostiolate, 

papillate. Peridium thin, composed of a few layers cells of 

textura prismatica. Hamathecium of dense, cellular pseudoparaphyses, 

septate. Asci 8-spored, bitunicate, fissitunicate, 

cylindrical to fusoid, with a furcate pedicel. 

Ascospores broadly fusoid to fusoid, hyaline to pale yellow, 

rarely 1- or 3- septate, mostly 2-septate, constricted at the 

primary septum. 

Anamorphs reported for genus: Sphaerellopsis (Sivanesan 

1984). 

Literature: Bayon et al. 2006; Eriksson 1966; Katumoto 

1986; Ramakrishnan 1951; Spegazzini 1908. 

Type species 

Eudarluca australis Speg., Revta Mus. La Plata 15: 22 

(1908). (Fig. 31) 


scattered, or in small groups, semi-immersed to erumpent, 

subglobose to broadly ellipsoid, wall black, ostiolate, apex 

with a short papilla, 40–70 μm broad (Fig. 31a and b). 

Peridium


Fig. 31 Eudarluca australis (from LPS 5.415, type). a Ascomata on 

the host surface. b Section of an ascoma. c Section of a partial 

peridium. Note the thin peridium with cells of textura angularis. d–g 

Asci with short pedicels. h Ascospores. Note the 2-septate hyaline 

ascospore. Scale bars: a, b=100 μm, c=50 μm, d–h=10 μm 

Falciformispora K.D. Hyde, Mycol. Res. 96: 26 (1992). 

(Trematosphaeriaceae) 


Habitat freshwater, saprobic. Ascomata small, scattered to 

gregarious, erumpent to nearly superficial, depressed 

globose to ovoid, black, ostiolate, epapillate, coriaceous. 

Peridium thin, comprising two cells types, outer layer 

composed of thick-walled cells of textura angularis, inner 

layer composed of hyaline compressed cells. Hamathecium 

long and cellular pseudoparaphyses, septate, embedded in 

mucilage. Asci 8-spored, bitunicate, fissitunicate, broadly 

clavate to fusoid, with a short, thick pedicel. Ascospores


fusoid to somewhat clavate, hyaline, usually slightly 

curved, multi-septate. 


Literature: Hyde 1992b; Raja and Shearer 2008. 

Type species 

Falciformispora lignatilis K.D. Hyde, Mycol. Res. 96: 27 

(1992). (Fig. 32) 

Ascomata 180–270 μm high×250–340 μm diam., scattered 

to gregarious, erumpent and eventually superficial, depressed 

globose to ovoid, black, ostiolate, epapillate, coriaceous 

(Fig. 32a). Peridium up to 35 μm wide, comprising two cell 

types, outer layer composed of thick-walled cells of textura 

angularis, upto8μm diam., cell wall up to 5 μm thick, inner 

layer composed of hyaline compressed cells, cells 12×3 μm 

diam., cell wall 1–1.5 μm thick (Fig. 32a). Hamathecium long 

and cellular pseudoparaphyses, 2–3 μm broad, septate, 

embedded in mucilage. Asci 115–130×23–31 μm, 8-spored, 

bitunicate, fissitunicate, broadly clavate to fusoid, with a short, 

thick pedicel, 8–15 μm long,withanocularchamber(to5μm 

wide×3 μm high) (Fig. 32b and c). Ascospores 42–50×8– 

10 μm, 2–3 seriate, fusoid to somewhat clavate, hyaline, 

usually slightly curved, 6–8-septate, mostly 7-septate, slightly 

constricted at all septa, smooth-walled, surrounded by a thin 

mucilaginous sheath which is longer at the base (up to 20– 

30 μm) (Fig.32d, e and f). 


Material examined: MEXICO, Nova Hispania, mangrove 

near Boca de Pascuales, saprobic on immersed 

intertidal mangrove wood, Mar. 1988, K.D. Hyde (BRIP 

16972, holotype). 

Notes 

Morphology 

Falciformispora was formally established by Hyde (1992b) 

as a monotypic genus and was assigned to Pleosporaceae by 

comparing with Setosphaeria, but Setosphaeria has the 

anamorphic stage of Exserohilum and is exclusively parasitic 

on Gramineae unlike Falciformispora. The setae on the 

ascomata of Setosphaeria could also serve as a distinguishing 

character from Falciformispora. Raja and Shearer (2008) 

also collected this species from freshwater in Florida. They 

considered that the species was more closely related to 

Chaetomastia than Setosphaeria, but that Falciformispora 

differed in having hyaline ascospores. 


Phylogenetic analyses in Schoch et al. (2009) and Suetrong et 

al. (2009) placed Falciformispora lignatilis in Trematosphaeriaceae 

in proximity to another marine species associated 

with mangroves, Halomassarina thalassiae. 


Phylogenetic work confirmed that the saprobic habitat of 

Falciformispora is inconsistent with most other members of 

Pleosporaceae. The hyaline multi-septate ascospores with a 

mucilaginous sheath indicate affinities to Lophiostomataceae 

but this is not supported in DNA sequence comparisons. 

Carinispora is also similar and may be related. 

Hadrospora Boise, Mem. N. Y. bot. Gdn 49: 310 (1989). 



Habitat terrestrial (or freshwater?), saprobic. Ascomata 

small- to medium-sized, solitary, scattered, or in groups, 

immersed to nearly superficial, globose to subglobose, 

papillate. Peridium thin, comprising pseudoparenchymatous 

cells. Hamathecium dense, narrowly cellular, embedded 

in mucilage. Asci bitunicate, fissitunicate, oblong to 

ovoid, with a short pedicel. Ascospores ellipsoid to broadly 

fusoid with narrow ends, reddish brown, multi-septate, 

constricted at the primary septum. 

Anamorphs reported for genus: Zalerion (Tanaka and 

Harada 2003a). 

Literature: Boise1984, 1989; Fisher and Webster 1992;Shearer 

and Crane 1971; Tanaka and Harada 2003a; Webster1993. 

Type species 

Hadrospora fallax (Mouton) Boise, Mem. N. Y. bot. Gdn 

49: 310 (1989). (Fig. 33) 

≡ Trematosphaeria fallax Mouton, Bull. Soc. R. Bot. 

Belg. 25: 155, (1886). 


scattered or in groups, initially immersed, becoming erumpent to 

nearly superficial, with basal wall remaining immersed in host 

tissue, not easily removed from the substrate, globose or 

subglobose, roughened, papillate, coriaceous (Fig. 33a). Peridium 

30–45 μm wide, comprising cells of pseudoparenchymatous,upto12.5×9μm 

diam. (Fig. 33b and c). Hamathecium of 

dense, narrowly cellular pseudoparaphyses, 1–2 μm broad, 

embedded in mucilage. Asci 150–200×40–60 μm 


oblong to ovoid, with a short pedicel, 10–24 μm long, 

with a ocular chamber (to 5 μm wide×6 μm high) (Fig. 33d). 


biseriate to 4-seriate, ellipsoid to broadly fusoid with narrow 

ends, reddish brown with paler end cells, 8-septate, 

constricted at the primary septum, smooth-walled (Fig. 33e, 

f, g, h and i). 

Anamorph: Zalerion sp. (Tanaka and Harada 2003a). 

Material examined: BELGIUM, Beaufays, on cut off, 

still hard wood. Oct. 1922, V. Mouton (BR, Capsa: K 7534,


Fig. 32 Falciformispora lignatilis 


a Section of a superficial 

ascoma. The peridium comprises 

two layers. b, c Squash 

mounts showing asci with wide 

pseudoparaphyses. The asci are 

cylindro-clavate with very short 

pedicels. d–f Hyaline multiseptate 

ascospores. Note the elongated 

appendage at the base 

(arrow head). Scale bars: 

a, b=100 μm, c=50 μm, 

d–f=10 μm 

holotype). (Note: The specimen is not in good condition, 

only a few ascomata left). 

Notes 

Morphology 

Boise (1989) formallyestablishedHadrospora to accommodate 

Trematosphaeria fallax and T. clarkia (Sivan.) Boise, 

and Hadrospora fallax (syn. T. fallax) was selected as the 

generic type. Hadrospora is a widely distributed species that 

has been reported from Belgium, China, Italy, Japan, 

Switzerland and the United States (Boise 1989; Fisher and 

Webster 1992; Shearer and Crane 1971; Tanaka and Harada 

2003a; Webster 1993). Hadrospora was temporarily 

assigned to Phaeosphaeriaceae based on its “small, thinwalled 

ascomata and narrow cellular pseudoparaphyses” 

(Boise 1989), which is distinguished from other genera of 

Phaeosphaeriaceae by its “large, stout ascospores that form 

within oblong-ovoid asci” (Boise 1989). Currently, Hadrospora 

includes two species, i.e. H. fallax and H. clarkii 

(Sivan.) Boise differentiated by ascospore size. 


None. 


Hadrospora seems not closely related to Phaeosphaeriaceae. 

Halotthia Kohlm., Nova Hedwigia 6: 9 (1963). (?Zopfiaceae) 


Habitat marine, saprobic. Ascomata large, solitary, gregarious 

or confluent, broadly conical to subglobose, 

flattened at the base, carbonaceous, immersed to erumpent, 

ostiolate, epapillate. Peridium plectenchymatous. 

Hamathecium of dense, long, cellular pseudoparaphyses, 

septate, branching. Asci 8-spored, bitunicate, cylindrical, 

with a short pedicel. Ascospores uniseriate, ellipsoidal, 

subcylindrical or obtuse-fusoid, dark brown, 1-septate, 

constricted at the septum. 


Literature: Kohlmeyer 1963; Suetrong et al. 2009.


Fig. 33 Hadrospora fallax 

(from BR, Capsa: K 7534, 

holotype). a Ascomata forming 

a cluster on the host surface. b 

Section of an ascoma. Note the 

peridium structure. c Section of 

a partial peridium. Note the 

pseudoparenchymatous cells. d 

Asci in pseudoparaphyses. e–i 

Reddish brown multiseptate 

ascospores. Scale bars: a= 

0.5 mm, b=100 μm, c, d= 

50 μm, e–i=20 μm 

Type species 

Halotthia posidoniae (Durieu & Mont.) Kohlm., Nova 

Hedwigia 6: 9 (1963). (Fig. 34) 

≡ Sphaeria posidoniae Durieu & Mont. Exploration 

scientifique de l’Algérie, pp. 502–503, Taf. 25, Abb. 8a-i, 1849. 

Ascomata 0.8–1.1 mm high×1.5–2.1 mm diam., solitary, 

gregarious or confluent, broadly conical to subglobose, 

flattened at the base, carbonaceous, immersed to 

erumpent, ostiolate, epapillate (Fig. 34a). Peridium 165– 

275 μm thick at sides, thicker near the apex, plectenchymatous. 

Hamathecium of dense, long cellular pseudo-


paraphyses, 1.5–2 μm broad, septate, branching. Asci 

275–290×25–35 μm, 8-spored, bitunicate, cylindrical, 

with a short pedicel (Fig. 34b, c and d). Ascospores 37– 

60.5×16.5–26 μm, uniseriate, ellipsoidal, subcylindrical 

or obtuse-fusoid, dark brown, 1-septate, constricted at the 

septum (Fig. 34e, f, g and h) (adapted from Kohlmeyer 

and Kohlmeyer 1979). 


Material examined: ITALY, in rhizomes of Posidonia 

oceanica (Posidoniaceae), 1861, Caldesi (S, isotype of 

Sphaeria posidoniae) 

Notes 

Morphology 

Halotthia was introduced to accommodate the marine 

fungus, H. posidoniae (as Sphaeria posidoniae), which is 

characterized by immersed to erumpent, large, carbonaceous 

ascomata, thick peridium, bitunicate, 8-spored, 

cylindrical asci, ellipsoidal, 1-septate, and dark brown 

ascospores (Kohlmeyer 1963). Morphologically, Halotthia 

is most comparable with Bicrouania maritima, but the 

conical ascomata with flattened base of H. posidoniae can 

be readily distinguished from B. maritima. 


Phylogenetically, Halotthia posidoniae, Pontoporeia 

biturbinata and Mauritiana rhizophorae form a robust 

clade, which may represent a potential family (Suetrong et 

al. 2009). 


Currently the familial status of Halotthia is unresolved 

(Suetrong et al. 2009). 

Helicascus Kohlm., Can. J. Bot. 47: 1471 (1969). 

(Morosphaeriaceae) 


Habitat marine, saprobic. Ascostromata lenticular, immersed, 

black, carbonaceous, enclosing several loculi, 

pseudoclypeus composed of host cells enclosed in black 

stromatic fungus material. Ascomata depressed ampulliform, 

horizontally arranged under a black pseudoclypeus, 

ostiolate, torsellioid ostioles, papillate. Peridium absent, 

partitions between loculi formed of brown, isodiametric or 

elongated cells of the stroma. Hamathecium of dense, long 

pseudoparaphyses. Asci 8-spored, bitunicate, subcylindrical 

to oblong clavate, with a short pedicel and conspicuous 

apical ring. Ascospores uniseriate, obovoid, brown, 1- 

septate, at each end with a germ pore, surrounded with 

dissolving sheath. 


Literature: Kohlmeyer1969; Kohlmeyer and Kohlmeyer 

1979; Suetrongetal.2009. 

Type species 

Helicascus kanaloanus Kohlm., Can. J. Bot. 47: 1471 

(1969). (Fig. 35) 

Ascostromata 0.6–0.78 mm high×1.25–2.75 mm diam., 

lenticular, immersed, black, carbonaceous, enclosing 3–4(−5) 

loculi, pseudoclypeus composed of host cells enclosed in 

black stromatic fungus material (Fig. 35a). Ascomata 235– 

370 μm high×440–800 μm diam., depressed ampulliform, 

horizontally arranged under a black pseudoclypeus, ostioles 

70–170 μm diam., torsellioid ostiole (Adams et al. 2005), 

papilla slightly rising over the surface of the pseudoclypeus, 

subconical,canal filled with thick, bright orange to yellowish 

periphyses, 270–435 μm high, 255–300 μm diam. Peridium 

absent, partitions between loculi formed of brown, isodiametric 

or elongated cells of the stroma. Hamathecium of 

dense, very long pseudoparaphyses. Asci 250–335×25– 

30 μm, 8-spored, subcylindrical, finally oblong-clavate 

(400–480 μm long), with a short pedicel, bitunicate, 

thick-walled, physoclastic, apically multi-layered and 

annulate, ectoascus forming a third, thin permeable outer 

layer around the base, endoascus swelling in water and 

becoming coiled at maturity, finally stretching and 

pushing the ascus into the ostiolar canal (Fig. 35b). 

Ascospores 36.5–48.5×18–22.5 μm, uniseriate, obovoid, 

brown, 1-septate, at each end with a germ pore, 

surrounded with dissolving sheath, 2.7–5.4 μm thick, 

with funnel-shaped, apical indentations (Fig. 35c and d) 

(adapted from Kohlmeyer and Kohlmeyer 1979). 


Material examined: USA, Hawaii, Oahu, Kaneohe Bay, 

Heeia Swamp, on Rhizophora mangle, 4 Jun. 1968 (Herb. J. 

Kohlmeyer No. 2566, holotype; No. 2565, 2567, paratype). 

Notes 

Morphology 

Helicascus is another marine genus, which is characterized 

by its thin additional sheath around the base of the asci, 

the coiling and stretching mechanism of the basal part of the 

endoascus and its conspicuous apical apparatus which is not 

that common in bitunicate asci (Kohlmeyer 1969). The 

immersed stroma comprising several loculi sharing one 

common ostiole is another striking character of Helicascus. 


Multigene phylogenetic analysis indicated that both 

Helicascus kanaloanus and H. nypae K.D. Hyde nested 

within Morosphaeriaceae (Suetrong et al. 2009).


Fig. 34 Halotthia posidoniae (from S, isotype of Sphaeria posidoniae). a Ascomata gregarious on the host surface. b–d Mature or immature 

cylindrical asci. e–h Ellipsoidal, dark-brown, 1-septate ascospores. Scale bars: a=1 mm, b–d=50 μm, e–h=5 μm 


Helicascus is a well defined marine genus. 

Herpotrichia Fuckel, Fungi rhenani exsic.: no. 2171 

(1868). (Melanommataceae) 


Habitat terrestrial, parasitic, hyperparasitic or saprobic. 

Ascomata medium-sized, immersed, erumpent to nearly 

superficial, scattered to gregarious, globose to subglobose


Fig. 35 Helicascus kanaloanus 

(from Herb. J. Kohlmeyer No. 

2566, holotype). a Section of 

ascostroma immersed in the host 

tissue. Note the torsellioid ostiole. 

b One-septate, brown, 

asymmetrical ascospores within 

the asci. c, d Released thickwalled 

ascospores. Note the 

germ pore at the lower end of 

the ascospores. Scale bars: a= 

0.5 mm, b–d=20 μm 

with a broad pore. Peridium composed of pseudoparenchymatous 


embedded in mucilage, septate, branching. Asci 

cylindrical to cylindro-clavate, with a furcated pedicel. 

Ascospores fusoid, ellipsoid or oblong with broadly to 

narrowly round ends, 1-septate, constricted at the septum, 

uni- to biseriate. 

Anamorphs reported for genus: Pyrenochaeta or Pyrenochaeta-like 

(Sivanesan 1984). 

Literature: von Arx and Müller 1975; Barr1984; Cannon 

1982; Freyer and van der Aa 1975; Mugambi and 

Huhndorf 2009b; Samuels 1973; Samuels and Müller 

1978; Sivanesan 1971, 1984. 

Type species 

Herpotrichia rubi Fuckel, Fungi rhenani exsic 2171. 

(1868). (Fig. 36) 

Ascomata 220–430 μm high×240–390(-530) μm 

diam., scattered to gregarious, immersed to erumpent, 

rarely superficial, globose to subglobose, wall black, 

coriaceous, apex with a small sometimes inconspicuous 

papilla, usually with a pore, lacking periphyses (Fig. 36a 

and b). Peridium 32–45 μm wideatthesides,upto60μm 

wide at the apex, basal wall thinner, all walls comprising 

cells of textura angularis, cells 2.5–4 μm diam., cell wall 

2–4(−7) μm thick, exterior cells more thick-walled and 

pigmented, inner cells thin-walled and less pigmented, 

comprising thin-walled cells up to 9 μm diam., apex cells 

smaller and walls thicker (Fig. 36b and c). Hamathecium 

of dense, long pseudoparaphyses, 2–3 μm broad, embedded 

in mucilage, septate, branching (Fig. 36e). Asci 105– 

150×12.5–15 μm (x ¼ 137:5 13:8mm, n=10), 8-spored, 

bitunicate, fissitunicate, cylindrical to cylindro-clavate, 

with a furcate pedicel that is 20–42.5 μm long, and 

ocular chamber up to 2.5 μm wide×2.5 μm high 

(Fig. 36d and f). Ascospores 17.5–25×(5.5-)6.3–9 μm 

(x ¼ 20:5 7:3mm, n=10), biseriate to partially overlapping 

uniseriate near the base, fusoid with narrowly 

rounded ends, hyaline when immature and becoming pale 

brown, 1-septate, deeply constricted at the septum, the 

upper cell often broader than the lower one, verruculose 

(Fig. 36g and h). 

Anamorph: Pyrenochaeta rhenana Sacc. (Sivanesan 

1984). 

Material examined: AUSTRIA, on Rubus idaeus L., 

very rarely in the spring, in the Oestreicher meadow forest 

(G, F. rh. 2171, type).


Notes 

Morphology 

Herpotrichia was established by Fuckel (1868) comprising 

two species H. rhenana Fuckel and H. rubi Fuckel, but 

no generic type was assigned. Bose (1961) designatedH. 

rhenana as the lectotype species with H. rubi as a synonym. 

This proposal was followed by Müller and von Arx (1962) 

and Sivanesan (1971). Herpotrichia rubi was later assigned 

as the generic type (Holm 1979) as it was found to be validly 

published 2 years earlier than H. rhenana, thus having 

priority (Cannon 1982). However, Cannon (1982) reported 

that Sphaeria herpotrichoides Fuckel (1864, cited as a 

synonym of H. rhenana) was the earliest name. Thus he 

made a new combination as H. herpotrichoides (Fuckel) P.F. 

Cannon and cited H. rubi as the synonym. Herpotrichia rubi 

is maintained as the type of the genus (Holm 1979; Cannon 

1982), but the current name is H. herpotrichoides. 

Herpotrichia is a morphologically well studied genus 

(Barr 1984; Bose 1961; Müller and von Arx 1962; 

Pirozynski 1972; Samuels and Müller 1978; Sivanesan 

1971, 1984), and Herpotrichia sensu lato is characterized 

by having subglobose, pyriform to obpyriform ascomata 

and a peridium of textura angularis or comprising thickwalled 

polygonal cells with thin-walled hyaline cells 

towards the centre. Asci are clavate to cylindrical, 4–8- 

spored and ascospores are hyaline at first, becoming pale to 

dark brown, one to many septate, constricted or not at the 

septa and often surrounded by a mucilaginous sheath. 

Several morphologically distinct genera were synonymized 

under Herpotrichia using the above broad circumscription 

(Barr 1984; Müller and von Arx 1962; Sivanesan 1984). In 

particular, Barr kept Lojkania as a separate genus after 

studying its type material (Barr 1984, 1990a). Sivanesan 

(1984) was also of the opinion that Lojkania and Neopeckia 

were distinct genera as several of their characters differed. 

Byssosphaeria and Pseudotrichia have subsequently been 

assigned to Melanommataceae, Lojkania to Fenestellaceae 

and Neopeckia to Coccoideaceae (Barr 1984). Herpotrichia 

sensu stricto is represented by H. rubi and has erumpent to 

superficial ascomata or immersed in a subiculum, clavate to 

cylindrical, 4–8-spored, stalked asci with a conspicuous 

apical “nasse”, hyaline, 1-septate ascospores, usually becoming 

pale brown and several septate, constricted or not 

constricted at septa, usually surrounded by sheath (Sivanesan 

1984). Currently, about 90 species are included in this genus 

(http://www.indexfungorum.org/, 12/01/2009). 


Herpotrichia diffusa (Schwein.) Ellis & Everh., H. 

juniperi (Duby) Petr., H. herpotrichoides and H. macrotricha 

have been shown to have phylogenetic affinity with 

the generic types of Byssosphaeria schiedermayeriana, 

Melanomma pulvis-pyrius and Pleomassaria siparia, which 

had been assigned under Melanommataceae (Kruys et al. 

2006; Mugambi and Huhndorf 2009b; Schoch et al. 2006, 

2009; Zhang et al. 2009a). In this study, Pleomassaria 

siparia together with its closely related species of Prosthemium 

is kept in a separate family, viz Pleomassariaceae. 


Even species under Herpotrichia sensu stricto (according 

to Sivanesan 1984) have diverse hosts (such as 

gymnosperms (H. coulteri (Peck) S.K. Bose and H. parasitica 

(R. Hartig) Rostr.) and angiosperms (H. diffusa and 

H. villosa Samuels & E. Müll.)) or substrates (like dead or 

living leaves, bark or decorticated wood) (Sivanesan 1984). 

Species of Herpotrichia sensu stricto are also reported from 

various locations such as Europe, Asia or America, and 

they have various life styles, e.g. parasitic, hyperparasitic or 

saprobic (Sivanesan 1984). Additional factors (like hosts or 

locations) may need to be considered in order to get a 

natural concept for Herpotrichia. 

Immotthia M.E. Barr, Mycotaxon 29: 504 (1987). 

(Teichosporaceae) 


Habitat terrestrial, hyperparasitic. Ascomata gregarious, 

globose, superficial, ostiolate, periphysate. Hamathecium 

of cellular pseudoparaphyses. Asci 8-spored, bitunicate, 

cylindrical, with a short pedicel. Ascospores 1-seriate, 

ellipsoidal, brown to reddish brown, 1-septate, constricted 

at the septum, smooth. 


Literature: Barr 1987a, 2002; Wang et al. 2004. 

Type species 

Immotthia hypoxylon (Ellis & Everh.) M.E. Barr, Mycotaxon 

29: 504 (1987). (Fig. 37) 

≡ Amphisphaeria hypoxylon Ellis & Everh., J. Mycol. 2: 

41 (1886). 

Ascomata gregarious, globose, superficial, ostiolate, 

periphysate, papillate (Fig. 37a). Hamathecium of cellular 

pseudoparaphyses, 2–2.5 μm broad, septate. Asci 60–82× 

7–9 μm, 8-spored, bitunicate, cylindrical, with a short 

pedicel (Fig. 37b, c and d). Ascospores 10–13×4.4–5.4 μm, 

1-seriate, ellipsoidal, brown to reddish brown, 1-septate, 

constricted at the septum, smooth (Fig. 37f, g and h) 

(adapted from Wang et al. 2004). 


Material examined: USA, Louisiana, Pointe a la 

Hache, on decaying wood, a branch of Carya oliviformis 

(Juglandaceae) lying on the ground in grass (parasitic on


Fig. 36 Herpotrichia rubi (from g, f. rh. 2171, type). a Numerous 

ascomata gregariously immersed in the host tissue. b Section of an 

ascoma. Note the central ostiole and peridium structure and also note 

the arrangement of asci and pseudoparaphyses. c Section of partial 

lateral peridium which comprises cells of textura angularis. d Part of a 

mature squashed ascus. e Relatively wide, septate pseudoparaphyses. f 

Immature ascus. Note the furcate pedicel. g–h One-septate ascospores. 

Note the verruculose ornamentation which is visible at the sides. Scale 

bars: a=0.5 mm, b=100 μm, c=50 μm, d=20 μm, e–h=10 μm


some effused Hypoxylon), 30 Dec. 1885, A.B. Langlois, No. 138 

(NY, holotype of Amphisphaeria hypoxylon Ellis & Everh.). 

Notes 

Morphology 

Immotthia was introduced to accommodate a species of 

Amphisphaeria (A. hypoxylon), which has bitunicate asci, 

and is characterized by superficial, ostiolate, periphysate, 

papillate ascomata, cellular pseudoparaphyses, bitunicate, 8- 

spored, cylindrical asci, ellipsoid, smooth, brown to reddish 

brown, 1-septate ascospores (Barr 1987a; Wangetal.2004). 


None. 


It seems that those Amphisphaeria species with bitunicate 

asci should be assigned to Pleosporales. Morphologically, 

Immotthia is somewhat comparable with Herpotrichia. 

Isthmosporella Shearer & J.L. Crane, Mycologia 91: 141 



Habitat freshwater, saprobic. Ascomata small- to mediumsized, 

scattered, immersed, erumpent to superficial, globose, 

papillate, ostiolate, periphysate, membranous. Peridium 2- 

layered, outer layer composed of brown, pseudoparenchymatic, 

fusoid-cylindric cells, inner layer composed of fusoid, 

subhyaline to pale brown, compressed cells. Hamathecium of 

rare, broad, septate, interascal pseudoparaphyses. Asci 8- 

spored, bitunicate, fissitunicate, oblong to clavate, with a 

short pedicel, ocular chamber not observed. Ascospores 3–4 

seriate, cylindrical to fusoid, isthmoid at centre, constricted at 

septa, isthmus 1-septate, surrounded by a gelatinous sheath. 


Literature: Shearer and Crane 1999. 

Type species 

Isthmosporella pulchra Shearer & J.L. Crane, Mycologia 

91: 142 (1999). (Fig. 38) 

Ascomata 240–330 μm diam., scattered on decorticated 

wood, immersed, erumpent to superficial, globose, black, 

papillate, papilla short, cylindrical, 60 μm long×55 μm 

wide, ostiolate, periphysate, membranous (Fig. 38a). Peridium 

2-layered, outer 3–4 cell layers composed of brown, 

pseudoparenchymatic, fusoid-cylindric cells, 2–6.5 μm 

long; inner layer composed of 5–7 rows of fusoid, subhyaline 

to pale brown compressed cells, 11–20×2–3.5 μm 

diam. (Fig. 38a and b). Hamathecium of rare, broad, 

septate, interascal pseudoparaphyses (Fig. 38f). Asci (95-) 

135–160(−175)×(25-)30–45(−60) μm, 8-spored, bitunicate, 

fissitunicate, oblong to clavate, with a short pedicel, ocular 

chamber not observed (Fig. 38c, d and e). Ascospores 80– 

105(−110)×(7-)8–10 μm, 3–4-seriate, cylindrical to fusoid, 

isthmoid at centre, sometimes bent at isthmus and becoming 

u- or v- shaped, end cells tapering, 12–17-phragmoseptate, 

constricted at septa, isthmus 1-septate, 2–5.5×2– 

4.5 μm diam., hyaline, frequently fragmenting to form 

partspores; filled with lipid droplets that merge to form 

large guttules; surrounded by a gelatinous sheath with a 

dense region near the isthmus, sheath greatly enlarging in 

water (Fig. 38g, h, i and j). 


Colonies on yeast soluble starch agar containing balsa 

wood sticks effuse, white. Hyphae hyaline, septate. 

Material examined: USA, New York, Adirondack Park. 

Piercefield. Tupper Lake at public boat launch from Rt. 30, 

UTM Zone 18, 539840 mE, 4892100mN; 44°10″59″N, 80° 

31′6″W, on submerged, decorticated wood, 7 Jul. 1994, J.L. 

Crane & C.A. Shearer A-254-1 (ILLS 53086, holotype). 

Notes 

Morphology 

Isthmosporella was described as a freshwater genus 

typified by I. pulchra, and is characterized by globose, 

pseudoparenchymatous ascomata, sparse, septate pseudoparaphyses, 

fissitunicate asci and hyaline, cylindrical to 

fusoid, phragmoseptate, isthmoid ascospores surrounded 

with a gelatinous sheath (Shearer and Crane 1999). Based 

on the morphological characters, i.e. small, globose 

ascomata, peridium with small pseudoparenchymatous cells 

and sparse pseudoparaphyses, Isthmosporella was assigned 

to the Phaeosphaeriaceae (Shearer and Crane 1999). The 

aquatic habitat of Isthmosporella, however, disagree with 

the Phaeosphaeriaceae. Isthmosporella seems less likely to 

belong to Pleosporineae. 


None. 


Molecular phylogenetic studies should be conducted to 

explore its familial placement within Pleosporales. 

Kalmusia Niessl, Verh. nat. Ver. Brünn 10: 204 (1872). 

(Montagnulaceae) 



solitary, scattered or in small groups, immersed to


Fig. 37 Immotthia hypoxylon (from holotype of Amphisphaeria hypoxylon). a Ascomata gregarious on host surface. b–d Bitunicate asci. e– 

h Released 1-septate ascospores. Scale bars: a=0.5 mm; b–h=10 μm 

erumpent, globose or subglobose, often laterally flattened, 

coriaceous, wall black, with or without papilla. 

Hamathecium of dense, filliform, delicate, septate pseudoparaphyses, 

branching and anastomosing between and 

above asci, embedded in mucilage. Asci bitunicate, 

fissitunicate unknown, clavate, with a long, furcate 

pedicel. Ascospores narrowly ovoid to clavate, pale 

brown, 3-septate, distoseptate. 

Anamorphs reported for genus: Cytoplea (Petrak and 

Sydow 1926).


Fig. 38 Isthmosporella pulchra 

(from ILLS 53086, holotype). 

a Section of an ascoma. b 

Section of a partial peridium. c– 

e Broadly clavate asci with short 

pedicels. f Pseudoparaphyses. 

g–j Ascospores. Note the 2- 

celled isthmus in J and mucilaginous 

sheath in G and H. 

Scale bars: a=50 μm, b–j= 

20 μm (figure with permission 

from http://fungi.life.illinois. 

edu/search/fwa_lit_refs) 

Literature: Barr 1987b, 1990a, 1992a; Lindau 1897; von 

Niessl 1872. 

Type species 

Kalmusia ebuli Niessl, Verh. nat. Ver. Brünn 10: 204 

(1872). (Fig. 39) 


scattered, or in small groups, immersed to erumpent, 

globose or subglobose, coriaceous, wall black, with or 

without papilla, ostiolate (Fig. 39a). Papilla small, up to 

100 μm high, with small ostioles (Fig. 39a). Peridium 15– 

40 μm wide, comprising one cell type of small, pigmented, 

thick-walled cells of textura prismatica to textura angularis, 

cells ca. 5×3μm diam., cell wall 2–3 μm thick (Fig. 39b 

and c). Hamathecium of dense, delicate pseudoparaphyses, 1– 

1.5 μm broad, septate, branching and anastomosing between 

and above asci, embedded in mucilage. Asci 75–125×10– 

15 μm (x ¼ 90:5 12mm, n=10), 8-spored, bitunicate, 

fissitunicate unknown, clavate, with a long, narrowed, furcate


pedicel which is up to 45 μm long, and a low ocular chamber 

(ca. 2μm wide×1 μm high) (Fig. 39d, e and f). Ascospores 

15–18×5.5–6.5 μm (x ¼ 16:3 5:8mm, n=10), biseriate, 

narrowly ovoid to clavate, pale brown, 3-distoseptate, without 

constriction, smooth-walled (Fig. 39g, h and i). 


Material examined: BELGIUM, Dolembreux, on branchlets 

and pieces of stumps of Sarothamnus scoparius from 

woodland, Oct. 1922, V. Mouton (BR 101525–63, holotype). 

Notes 

Morphology 

Kalmusia was formally established by von Niessl (1872), 

and is mainly characterized as “immersed, sphaeroid ascoma 

with central, stout papilla, surrounded by hyphae in the 

substrate, stipitate asci with septate pseudoparaphyses, and 

brown, 3-septate, inequilateral ascospores” (Barr 1992a). 

The most morphologically comparable genus to Kalmusia 

is Thyridaria, which had been treated as a subgenus 

under Kalmusia (Lindau 1897), and was subsequently 

transferred to Platystomaceae in Melanommatales (Barr 

1987b, 1990a). Compared to Thyridaria, Kalmusia has 

sphaeroid ascomata, a peridium of small pseudoparenchymatous 

cells, basal asci and very thin pseudoparaphyses, 

thus it was assigned to Phaeosphaeriaceae of the Pleosporales 

by Barr (1990a), and the genus is utilized to 

accommodate both K. ebuli and K. clivensis (Berk. & 

Broome) M.E. Barr, as well as closely related species, i.e. 

K. utahensis (Ellis & Everh.) Huhndorf & M.E. Barr and K. 

coniothyrium (Fuckel) Huhndorf (Barr 1992a). But this 

proposal is questionable, as the clavate, distoseptate ascospores, 

as well as the clavate asci with very long pedicels are 

uncommon in Phaeosphaeriaceae, and most recent phylogenetic 

study indicated that some species of Kalmusia reside 

outside of Phaeosphaeriaceae (Zhang et al. 2009a). 


Both Kalmusia scabrispora Teng Kaz. Tanaka, Y. 

Harada & M.E. Barr and K. brevispora (Nagas. & Y. 

Otani) Yin. Zhang, Kaz. Tanaka & C.L. Schoch reside in 

the clade of Montagnulaceae (Zhang et al. 2009a). Familial 

placement of Kalmusia can only be verified after the DNA 

sequences of the generic type (K. ebuli) are obtained. 


Kalmusia is distinct amongst the Pleosporales as it has 

pale brown ascospores with indistinct distosepta and clavate 

asci with long pedicels. Although both K. scabrispora and 

K. brevispora reside in the clade of Montagnulaceae, they 

both lack the distoseptate ascospores that are possessed by 

the generic type (K. ebuli). Thus, the familial placement of 

Kalmusia is still undetermined. 

Karstenula Speg., Decades Mycologicae Italicae ad no. 94 

(in sched.) (1879). (Montagnulaceae) 


Habitat terrestrial, saprobic. Ascomata rarely small-, 

usually medium-sized, immersed usually under thin 

clypeus, scattered to gregarious, with flattened top and 

rounded pore-like ostiole, coriaceous. Peridium 2-layered, 

outer layer composed of reddish brown to dark brown 

small cells, inner layer of pale compressed cells. Hamathecium 

of dense, cellular pseudoparaphyses. Asci cylindrical 

to cylindro-clavate with short furcate pedicel. 

Ascospores muriform, ellipsoid to fusoid, reddish brown 

to dark brown. 

Anamorphs reported for the genus: Microdiplodia 

(Constantinescu 1993). 

Literature: Barr 1990a; Eriksson and Hawksworth 1991; 

Kodsueb et al. 2006a; Munk 1957; Zhang et al. 2009a. 

Type species 

Karstenula rhodostoma (Alb. & Schwein.) Speg., Decades 

Mycologicae Italicae no. 94. (1879). (Fig. 40) 

≡ Sphaeria rhodostoma Alb. & Schwein., Consp. fung. 

(Leipzig): 43 (1805). 


or gregarious, immersed in the subiculum which sometimes 

sloths off, globose or subglobose, black, flattened top often 

white or reddish and sometimes slightly protruding out of the 

substrate surface, usually with a wide opening ostiole after 

removing the cover, coriaceous (Fig. 40a and b). Peridium 

30–40 μm wide, comprising two cell types, outer region 1- 

layered, composed of relatively small heavily pigmented 

thick-walled compressed cells, cells 2–4×5–10 μm diam., 

cell wall 2–4 μm thick, inner layer cells larger and wall 

thinner, comprising cells of textura angularis, merging with 

pseudoparaphyses (Fig. 40c and d). Hamathecium of dense, 

long cellular pseudoparaphyses 2–3.5 μm broad, septate, 

branching or anastomosing not observed. Asci 150–210× 

12.5–15 μm (x ¼ 182 13:1mm, n=10), 8-spored, bitunicate, 

fissitunicate, cylindrical, with a broad, furcate pedicel which is 

12–35 μm long, and with an ocular chamber (to 4 μm wide× 

3 μm high) (Fig. 40e and f). Ascospores 20–26×7.5–10 μm 

(x ¼ 22:4 8mm, n=10), obliquely uniseriate and partially 

overlapping, ellipsoid, reddish brown, with 3 transverse septa 

and a vertical septum in one or two central cells, constricted at 

the septa, verruculose (Fig. 40g, h and i). 

Anamorph: Microdiplodia frangulae Allesch. (Constantinescu 

1993). 

Conidiomata globose to subglobose, 330–495 μm diam., 

in subiculum. Conidia 9–13×4–5 μm, reddish brown, 1- 

septate (information obtained from Barr 1990a).


Fig. 39 Kalmusia ebuli (from BR 101525–63, holotype). a Immersed 

to erumpent ascomata scattered on the host surface. b Section 

of a partial peridium. Note the compressed peridium cells. c Section of 

an ascoma. d–f Eight-spored asci with long pedicels. g Partial ascus in 

pseudoparaphyses. h, i Ascospores with 3 thick-walled septa. Scale 

bars: a=0.5 mm, b=50 μm, c=100 μm, d–g=20 μm, h, i=10 μm 

Material examined: Fries, Suecia (received by herbarium 

in 1834) (PH 01048835, type, as Sphaeria rhodostoma 

Alb. & Schwein.). 

Notes 

Morphology 

Karstenula is an ambiguous genus, which has been 

synonymized under Pleomassaria (Lindau 1897; Winter 

1885). Some of the ascomata characters are even comparable 

with those of Didymosphaeria, such as ascomata 

seated in subiculum or beneath a clypeal thickening, the 

development of apex vary in a large degree, even to the 

occasional formation of a blackened internal clypeus, and 

sometimes apical cells become reddish or orange-brown 

(Barr 1990a). Barr (1990a) redefined the concept of 

Karstenula (sensu lato), which encompasses some species 

of Thyridium. In her concept, however, Barr (1990a) 

treated Karstenula as having trabeculate pseudoparaphyses 

and this is clearly not the case. In most cases the


Fig. 40 Karstenula rhodostoma (from PH 01048835, type). a Line 

of ascomata on host surface (after remove the decaying cover). Note 

the wide ostiolar opening and light colored region around the ostiole. 

b Immersed ascoma under the decaying cover (see arrow). c, d 

Section of the peridium. The peridium comprises small thick-walled 

cells in the outer layer. The outside comprises defuse hyphae which is 

probably part of the subiculum. e Ascus with a short furcate pedicel. f 

Partial ascus showing arrangement of ascospores. g–i Released 

ascospores. Note the transverse and rarely vertical septa. Scale bars: 

a, b=0.5 mm, c=50 μm, d–f=20 μm, g–i=10 μm


ascospores were brown with transverse septa and sparse 

longitudinal septa. 

The ascomata of this species are similar to those found in 

Byssosphaeria and Herpotrichia, especially in the paler area 

around the ostiole and even in peridial structure and 

development under a subiculum. The numerous wide cellular 

pseudoparaphyses and cylindrical asci (in Herpotrichia) are 

also similar. The main difference of Karstenula from other 

two genera are the 3-septate ascospores with rare longitudinal 

septa (1-septate in Byssosphaeria and Herpotrichia). 


Karstenula forms a robust phylogenetic clade with 

Phaeodothis winteri (Niessl) Aptroot, Didymocrea sadasivanii, 

Bimuria novae-zelandiae, Montagnula opulenta, Curreya 

pityophila (J.C. Schmidt & Kunze) Arx & E. Müll. and 

some species of Letendraea and Paraphaeosphaeria (Kodsueb 

et al. 2006a; Zhang et al. 2009a). Consequently, 

Karstenula might be included in Montagnulaceae. 


The description of the type of Karstenula here clearly 

excludes it from Melanommataceae as it has wide 

pseudoparaphyses. But its Montagnulaceae status can only 

be confirmed by more phylogenetic work including 

sequencing the generic type of Karstenula (K. rhodostoma). 

Katumotoa Kaz. Tanaka & Y. Harada, Mycoscience 46: 

313 (2005). (Lentitheciaceae) 


Habitat terrestrial or freshwater, saprobic. Ascomata smallto 

medium-sized, scattered or in small groups, immersed to 

erumpent, with a central protruding hairy papilla, subglobose. 

Peridium thin, comprising several layers of thinwalled 

compressed cells. Hamathecium of dense, cellular, 

filliform, embedded in mucilage, branching and anastomosing. 

Asci 8-spored, bitunicate, fissitunicate, clavate with 

short furcate pedicels. Ascospores apiosporous and hyaline 

when young, becoming 2-septate with reddish brown 

echinate central cell at maturity, with long gelatinous 

terminal appendages. 


Literature: Tanaka and Harada 2005b; Tanaka et al. 2009; 

Zhang et al. 2009a. 

Type species 

Katumotoa bambusicola Kaz. Tanaka & Y. Harada, 

Mycoscience 46: 313 (2005). (Fig. 41) 

Some information for the following description is from 

Tanaka and Harada (2005). 


scattered or in small groups, immersed, becoming erumpent, 

with a slightly protruding papilla covered with brown 

hyphae, subglobose (Fig. 41a). Peridium 13–30 μm thick, 

composed of a few layers of lightly pigmented, depressed 

cells. Hamathecium of dense, long cellular pseudoparaphyses, 

1.5–3 μm broad, embedded in mucilage, branching and 

anastomosing. Asci 110–160×17.5–24 μm (x ¼ 139 21mm, 

n=10), 8-spored, bitunicate, fissitunicate, cylindro-clavate 

withashortfurcatepedicelwhichisupto25μm long 

(Fig. 41b and d). Ascospores 39–50(−57)×7–10 μm 

(x ¼ 45:8 8:2mm, n=10), biseriate, fusoid to narrowly 

fusoid with acute ends, usually curved, apiosporus and 

hyaline when young, constricted at the primary septum, the 

upper cell longer and broader than the lower one, smooth, 

surrounded by a bipolar sheath which is up to 15 μm long, 

best seen in India ink, senescent ascospores yellowish 

brown, 2–4-septate (Fig. 41c). 


Material examined: JAPAN, Mt. Iwate, near Yakebashiri, 

Hirakasa, Nishine, Iwate, on culms of Oryza sativa L., 19 

Oct. 2003, K. Tanaka (HHUF 28663, holotype). 

Notes 

Morphology 

Katumotoa was formally established by Tanaka and 

Harada (2005b) to accommodate the monotypic species, 

K. bambusicola, which is characterized by immersed 

ascomata with a thin peridium comprising thin-walled 

compressed cells, cellular pseudoparaphyses, cylindroclavate 

and fissitunicate asci and fusoid ascospores with 

an elongated bipolar mucilaginous sheath. Based on its 

immersed ascomata, psuedoparenchymatous peridium 

cells and cellular pseudoparaphyses, Katumotoa was 

assigned to Phaeosphaeriaceae (Tanaka and Harada 

2005b; Tanaka et al. 2009), but this classification has 

been shown to be incorrect in subsequent phylogenetic 

studies (Tanaka et al. 2009; Zhangetal.2009a). 


Phylogenetic analysis based on five genes (LSU, SSU, 

RPB1, RPB2 and EF1) indicates that Katumotoa bambusicola 

resides in Lentitheciaceae, and this receives high 

bootstrap support (Zhang et al. 2009a). In particular, K. 

bambusicola forms a robust clade with Ophiosphaerella 

sasicola (Nagas. & Y. Otani) Shoemaker & C.E. Babc., 

which has filliform ascospores (Shoemaker and Babcock 

1989b). 


The hyaline, apiosporous ascospores which become 

2–4-celled with central reddish brown cells and large


Fig. 41 Katumotoa bambusicola (from HHUF 28663, holotype). a Ascomata scattered on the host surface. b Asci in pseudoparaphyses. c 

Hyaline ascospore with long terminal appendages. d Clavate ascus with a short pedicel. Scale bars: a=0.5 mm. b–d=20 μm 

unraveling appendages are the most striking features of 

this species and readily distinguish it from other 

pleosporalean taxa. Both Katumotoa bambusicola and 

Ophiosphaerella sasicola are associated with bambusicolous 

hosts, which might indicate that host spectrum in 

this case, has greater phylogenetic significance than some 

morphological characters (Zhang et al. 2009a). 

Keissleriella Höhn., Sber. Akad. Wiss. Wien, Math.-naturw. 

Kl., Abt. 1 128: 582 (1919). (Lentitheciaceae) 


Habitat terrestrial or freshwater, saprobic. Ascomata smallto 

medium-sized, immersed, erumpent to nearly superficial, 

globose, papillate, ostiolate. Papilla covered by dark setae 

or small blackened cells. Peridium thick, composed of cells 

of pseudoparenchymatous and inner layer composed of pale 


rarely septate, anastomosing and branching. Asci 4- or 8- 

spored, bitunicate, fissitunicate, cylindro-clavate, with a


furcate pedicel and a small ocular chamber. Ascospores 

hyaline to pale brown, ellipsoid to fusoid, 1-septate, 

constricted at the septum (Barr 1990a). 

Anamorphs reported for genus: Dendrophoma (Bose 1961). 

Literature: von Arx and Müller 1975; Bose 1961; Barr 

1990a; Dennis 1978; Eriksson 1967a; von Höhnel 1919; 

Luttrell 1973; Munk 1957; Zhang et al. 2009a. 

Type species 

Keissleriella aesculi (Höhn.) Höhn., Sber. Akad. Wiss. 

Wien, Math.-naturw. Kl., Abt. 1 128: 582 (1919). (Fig. 42) 

≡ Pyrenochaeta aesculi Höhn., Ber. dt. bot. Ges. 35: 249 

(1917). 

Ascomata ca. 250μm high×450 μm diam., gregarious, 

immersed to erumpent, globose or subglobose, with a 

small black papilla, ca. 75μm high and 110 μm broad, 

with short black external setae (Fig. 42a). Peridium ca. 

25–40 μm wide laterally, up to 70 μm near the apex, 

thinner at the base, comprising two types of cells which 

merge in the middle; outer cells composed of small 

heavily pigmented thick-walled cells, cells ca. 4 μm 

diam., cell wall up to 4 μm thick, and thick near the apex 

and thinner laterally and absent in the immersed part of 

the ascoma, inner cells less pigmented, comprising 

lightly pigmented to hyaline cells, 5–7 μm thick 

(Fig. 42a). Hamathecium of dense, long pseudoparaphyses, 

0.8–1.2 μm broad, rarely septate, anastomosing and 

branching, thicker near the base, ca. 2μm, constricted 

near the septum (Fig. 42b). Asci 80–120×6–11 μm (x ¼ 

101 8:5mm, n=10), 4- or 8-spored, bitunicate, fissitunicate, 

cylindro-clavate, with a furcate pedicel which is 

up to 20–40 μm long, with a small ocular chamber 

(Fig. 42e and f). Ascospores 13–18×4–5.5 μm 


overlapping, fusoid with narrowly rounded ends, 

hyaline, 1-septate, constricted at the septum, smooth 

(Fig. 42c and d). 


Material examined: AUSTRIA, Brentenmaistal in the 

Viennese forest, Aesculus hippocastanum L., 1916, Höhnel 

(FH, holotype of Otthiella aesculi). (Note: only two slides; 

setae cannot be seen from the slides but could be seen from 

the drawings on the cover). 

Notes 

Morphology 

Keissleriella is characterized by ascomata with setae in 

and over the papilla, asci are cylindrical and ascospores are 

hyaline, 1-septate. Based on the morphological characters, 

K. aesculi was regarded as conspecific with K. sambucina; 

as an earlier epithet, K. sambusina typifies the genus (see 

comments by Barr 1990a). Munk (1957) placed Trichometasphaeria 

and Keissleriella in Massarinaceae, and distinguished 

them by their substrates (Trichometasphaeria 

occurs on herbaceous plants and Keissleriella on woody 

substrates). Bose (1961) combined Trichometasphaeria 

under Keissleriella, which was followed by some workers 

(von Arx and Müller 1975; Dennis 1978; Eriksson 1967a; 

Luttrell 1973). Barr (1990a), however, maintained these as 

distinct genera based on the differences of peridium 

structure and pseudoparaphyses. 


The phylogeny of Keissleriella is poorly studied. 

Limited phylogenetic information indicates that K. cladophila 

forms a robust clade with other species of Lentitheciaceae 



The presence of black setae on the surface of papilla is a 

striking character of Keissleriella, but phylogenetic significance 

of setae is undetermined yet. 

Lentithecium K.D. Hyde, J. Fourn. & Yin. Zhang, Fungal 

Divers. 38: 234 (2009). (Lentitheciaceae) 

= Tingoldiago K. Hirayama & Kaz. Tanaka, Mycologia 

102: 740 (2010) syn. nov. 


Habitat freshwater, saprobic. Ascomata small, scattered 

or gregarious, immersed, slightly erumpent, depressed 

spherical to lenticular, ostiolate, papillate or epapillate. 

Peridium thin. Hamathecium of cellular pseudoparaphyses. 

Asci 8-ascospored, bitunicate, fissitunicate, clavate, 

short-stipitate. Ascospores broadly fusoid with broadly 

rounded ends, 1-septate, constricted, hyaline, usually 

with sheath. 


Literature: Shearer et al. 2009; Zhang et al. 2009a, b. 

Type species 

Lentithecium fluviatile (Aptroot & Van Ryck.) K.D. Hyde, J. 

Fourn. & Yin. Zhang, Fungal Divers. 38: 234 (2009). (Fig. 43) 

≡ Massarina fluviatilis Aptroot & Van Ryck., Nova 

Hedwigia 73: 162 (2001). 


or gregarious, immersed, slightly erumpent, subglobose to 

depressed spherical, under a small black pseudostroma 

originating from the apical part of the peridium, apex slightly 

papillate, ostiole rounded, 60–70 μm diam. (Fig. 43a and b). 

Peridium 15–20 μm thick at sides and at base, comprising 4–5 

layers of angular cells more thick-walled outwards, 50–55 μm


Fig. 42 Keissleriella sambucina (from FH, holotype of Otthiella 

aesculi). a Section of an ascoma. b Pseudoparaphyses which are 

narrow (less than 1.5 μm) and branch and anastomosing as 

trabeculate. c, d Hyaline ascospores with distinct constrictions at the 

septa. e Asci amongst narrow pseudoparaphyses. F. Ascus with a 

pedicel and ocular chamber. Scale bars: a=100 μm, b–f=10 μm 

thick at apex, of small very thick-walled cells. Hamathecium 

of cellular pseudoparaphyses, 2–2.5 μm broad (Fig. 43c and 

d). Asci 89–100×19–21 μm, 8-spored, bitunicate, fissitunicate, 

clavate, bumpy, short-stipitate, apex without obvious 

apical chamber (Fig. 43e). Ascospores 27–35×8.5–9.4 μm,, 2- 

3-seriate, broadly fusoid with broadly rounded ends, straight to 

slightly curved, 1-septate, slightly constricted, with four large 

guttules, hyaline, smooth-walled, a very thin mucilaginous 

sheath can be occasionally observed in India ink but in most 

cases no sheath can be observed (Fig. 43f and g).



Material examined: FRANCE, Haute Garonne: Avignonet, 

Lac de Rosel, artificial lake, on bark and wood of a 

submerged branch Populus sp., 23 Nov. 2006, leg. Michel 

Delpont, det. Jacques Fournier (IFRD 2039, holotype). 

Notes 

Morphology 

Lentithecium was introduced to accommodate some freshwater 

fungi previous assigned under Massarina, suchasM. 

arundinacea (Sowerby) Leuchtm. and M. fluviatilis (Zhang et 

al. 2009a). It is characterized by its immersed and 

lenticular ascomata, thin peridium which is almost equal 

in thickness, short pedicellate asci and fusoid or filliform, 

hyaline or rarely lightly pigmented, 1- to multi-septate 

ascospores (Zhang et al. 2009b). Lentitheciaceae was 

introduced to accommodate Lentithecium and some other 

related taxa (Zhang et al. 2009a). 


The clade of Lentitheciaceae comprises the generic type 

Lentithecium fluviatile, as well as L. arundinaceum 

(Sowerby) K.D. Hyde, J. Fourn. & Yin. Zhang, Stagonospora 

macropycnidia, Wettsteinina lacustris (Fuckel) Shoemaker 

& C.E. Babc., Keissleriella cladophila, and the 

bambusicolous species Katumotoa bambusicola and 

Ophiosphaerella sasicola, which receive high bootstrap 

support (Zhang et al. 2009a). 


Tingoldiago graminicola K. Hirayama & Kaz. Tanaka 

form a robust clade with species of Lentithecium (Shearer et 

al. 2009). Tingoldiago has lenticular immersed to erumpent 

ascomata, numerous and septate pseudoparaphyses, 

cylindro-clavate asci and hyaline, 1-septate ascospores with 

sheath. All of these characters fit Lentithecium well. We 

treat Tingoldiago as a synonym of Lentithecium. 

Leptosphaeria Ces. & De Not., Comm. Soc. crittog. Ital. 1: 

234 (1863). (Leptosphaeriaceae) 


Habitat terrestrial, saprobic or parasitic. Ascomata small- to 

medium-sized, solitary, scattered or in small groups, 

erumpent to superficial, subglobose, broadly or narrowly 

conical, papillate, ostiolate. Peridium thick, comprising 

layers of cells of textura angularis. Hamathecium of dense 

cellular pseudoparaphyses, embedded in mucilage, anastomosing 

and branching. Asci 8-spored, bitunicate, fissitunicate 

unknown, cylindrical with a furcate pedicel and a large 

ocular chamber. Ascospores fusoid or narrowly fusoid, 

brown or reddish brown, 3-septate, constricted at each 

septum. 

Anamorphs reported for genus: Coniothyrium and 

Phoma (Hyde et al. 2011; Sivanesan 1984). 

Literature: von Arx and Müller 1975; Barr1987a, b; Cesati 

and de Notaris 1863; Crane and Shearer 1991; Dongetal. 

1998; Eriksson 1967a; Eriksson and Hawksworth 1986, 

1991; de Greuter et al. 1988; Hedjaroude1969; von Höhnel 

1907;Holm1957, 1975;Huhndorfetal.1990; Luttrell 1973; 

Müller 1950; Munk 1957; Saccardo 1878b, 1883, 1891, 

1895; Schochetal.2009; Shearer 1993; Shearer et al. 1990; 

Shoemaker 1984a; Sivanesan1984; Zhang et al. 2009a. 

Type species 

Leptosphaeria doliolum Ces. & De Not., Comm. Soc. 

crittog. Ital. 1: 234 (1863). (Fig. 44) 

≡ Sphaeria doliolum Pers., Icon. Desc. Fung. Min. 

Cognit. (Leipzig) 2: 39 (1800). 


scattered or in small groups, superficial, subglobose, broadly 

or narrowly conical, with a flattened base on the host surface, 

black, usually with 2–4 ring-like ridges surrounding the 

ascomata surface, apex with a conical, usually shiny papilla 

(Fig. 44a). Peridium 85–110 μm wide at sides, thinner at the 

apex, comprising two types of cells, outer layer composed of 

small thick-walled cells of textura angularis, cells


lectotype species (de Greuter et al. 1988; Holm1975; 

Shearer et al. 1990). Leptosphaeria was originally 

defined based mainly on the characters of ascospores 

being ellipsoid or fusoid, one to many septa, hyaline to 

dark brown. These few common characters meant that 

Leptosphaeria comprised many species, and some of 

them should be assigned to either Euascomycetes or 

Loculoascomycetes (Crane and Shearer 1991). Leptosphaeria 

had been divided based on host and habitat 

(Saccardo 1878b, 1891, 1895) as well as the pseudothecium 

(glabrous, hairy, setose) and ascospore septation 

(see comments by Crane and Shearer 1991). von Höhnel 

(1907) used centrum structure in the classification of 

Leptosphaeria, and divided Leptosphaeria into three 

genera, viz. Leptosphaeria, Scleropleella and Nodulosphaeria. 

Müller (1950) subdivided Leptosphaeria into 

four sections based on pseudothecial and centrum 

structure as well as ascospore characters. This classification 

was modified by Munk (1957), who named these 

four sections as section I (Eu-Leptosphaeria), section II 

(Para-Leptosphaeria), section III (Scleropleella) and 

section IV (Nodulosphaeria). Holm (1957) used a 

relatively narrow concept for Leptosphaeria, which 

included species closely related to the generic type, L. 

doliolum. This viewpoint was accepted by some workers 

(Eriksson 1967a; Hedjaroude 1969; Shoemaker 1984a). 

Nevertheless, it still seems a heterogeneous group of 

fungi (see comments by Crane and Shearer 1991). Its position 

among the Loculoascomycetes is also debated. It has been 

placed in the Pleosporaceae (von Arx and Müller 1975; 

Luttrell 1973; Sivanesan 1984) orLeptosphaeriaceae (Barr 

1987a, b; Eriksson and Hawksworth 1991) orPhaeosphaeriaceae 

(Eriksson and Hawksworth 1986). 


Molecular phylogenetic analysis based on multigenes 

indicated that species of Leptosphaeria (including the 

generic type L. doliolum) and Neophaeosphaeria form a 

paraphyletic clade with moderate bootstrap support (Dong 

et al. 1998; Schoch et al. 2009; Zhang et al. 2009a), which 

is sister to other families of Pleosporales (Zhang et al. 

2009a). Thus the familial rank of the Leptosphaeriaceae 

could be temporarily verified, but further molecular 

phylogenetic study is needed in which more related taxa 

should be included. 


Morphologically, Leptosphaeria is mostly comparable 

with Amarenomyces, Bricookea, Diapleella, Entodesmium, 

Melanomma, Nodulosphaeria, Paraphaeosphaeria, Passeriniella, 

Phaeosphaeria and Trematosphaeria. While it 

prefers non-woody parts of dicotyledonous hosts, its 

cylindrical ascus with short pedicel and smooth, fusoid 

Fig. 43 Lentithecium fluviatile (from IFRD 2039). a Erumpent b 

ascomata scattering on the host surface. b Habitat section of the 

immersed ascomata. c, d Section of an ascoma and a partical peridium. 

Note the peridium cells of textura angularis. e Clavate 8-spored ascus 

with a short pedicel. f, g Hyaline, 1-septate broadly fusoid ascospores. 

Scale bars: a, b=0.5 mm, c=100 μm, d=50 μm, e–g=20 μm 

and multi-septate ascospores make it readily distinguishable 

from all other genera (Shoemaker 1984a). 

Leptosphaerulina McAlpine, Fungus diseases of stonefruit 

trees in Australia and their treatment: 103 (1902). 

(Didymellaceae) 


Habitat terrestrial, parasitic or saprobic. Ascomata small, 

scattered, immersed, globose to subglobose, with a small, 

slightly protruding papilla, ostiolate. Peridium thin. Hamathecium 

of rare or decomposing cellular pseudoparaphyses. 

Asci bitunicate, obpyriform. Ascospores broadly clavate or 

cylindrical, hyaline, turning pale brown when old, asymmetrical, 

multi-septate, smooth-walled. 

Anamorphs reported for genus: Pithoascus and Pithomyces 

(Hyde et al. 2011). 

Literature: Barr 1972; Chlebicki 2002; Crivelli 1983; 

Kodsueb et al. 2006a; Zhang et al. 2009a. 

Type species 

Leptosphaerulina australis McAlpine, Fungus diseases of 

stone-fruit trees in Australia and their treatment: 103 (1902). 

(Fig. 45) 

Ascomata 140–170 μm diam., scattered, immersed, 

globose to subglobose, with a small slightly protruding 

papilla, ostiolate (Fig. 45a). Peridium thin, composed of 

one or two layers of large cells of textura angularis, pale 

brown (Fig. 45a). Hamathecium of rare or decomposing 

cellular pseudoparaphyses, up to 5 μm broad, filling 

the gaps between the asci. Asci 38–53×55–75 μm 

(x ¼ 67:5 43:3mm, n=10), 8-spored, without pedicel, bitunicate, 

fissitunicate dehiscence not observed, obpyriform, with a 

large ocular chamber and apical ring (Fig. 45b and c). 

Ascospores 30–40(-47)×11–14 μm (x ¼ 36:5 13mm, n= 

10), broadly clavate, hyaline, turning pale brown when old, 

asymmetrical, upper hemisphere usually with one transverse 

septum and with a somewhat narrowly rounded 

end, lower hemisphere usually with two transverse septa 

and with broadly rounded ends, slighted constricted at 

the primary septum, mostly with one vertical septum in 

each central cell, smooth, with thin gelatinous sheath 

when young, 2–3 μm thick (Fig. 45d and e). 

Anamorph: none reported.

Fungal Diversity


Fig. 44 Leptosphaeria doliolum (from L, lectotype). a Ascomata on the host surface. Note the shiny black surface. b Section of the partial peridium. 

Note the uneven thickness. c–e Asci with a short pedicel. f Three ascospores in ascus. Scale bars: a=0.5 mm, b=100 μm, c–f=20 μm


Material examined: USA, Kansas, Kansas State College, 

on Poa pratensis L. Grass plots, 2 Jul. 1953, leg. T. 

Rogerson, det. L.E. Wehmeyer (NY, C.T. Rogerson 3836). 

Notes 

Morphology 

Leptosphaerulina, introduced by McAlpine (1902), is 

characterized by small immersed ascomata, obpyriform asci 

with a large ocular chamber and apical ring as well as 

muriformly septate ascospores which may be hyaline or 

pigmented. Species of Leptosphaerulina may occur on 

monocotyledons or dicotyledons. Leptosphaerulina is most 

comparable with Pleospora, and the only difference between 

them is that Leptosphaerulina has smaller ascomata and 

hyaline ascospores that only become pigmented after 

discharge, whereas the ascospores of Pleospora become 

brown within the asci. Currently, about 60 names are 

accepted in this genus, and some even reported from marine 

environments, e.g. L. mangrovei (Inderbitzin et al. 2000). 


Based on multigene phylogenetic analysis, two putative 

strains of Leptosphaerulina australis, the generic type of 

Leptosphaerulina, from Switzerland (CBS 311.51) and 

Indonesia (CBS 317.83) resided within Didymellaceae (de 

Gruyter et al. 2009; Zhang et al. 2009a). 


Because of its morphological confusion with Pleospora 

and the diversity of habitats within the genus, Leptosphaerulina 

sensu lato is likely to be polyphyletic. Fresh 

collections of this species are needed from Australia to 

epitypify this taxon and define the genus in a strict sense. 

The specimen described here is a collection from USA and 

therefore may not represent the type. 

Lewia M.E. Barr & E.G. Simmons, Mycotaxon 25: 289 

(1986). (Pleosporaceae) 


Habitat terrestrial, parasitic or saprobic? Ascomata 

small, scattered, erumpent to nearly superficial at maturity, 

subglobose to globose, black, smooth, papillate, 

ostiolate. Papilla short, blunt. Peridium thin. Hamathecium 

of pseudoparaphyses. Asci (4–6-)8-spored, bitunicate, 

fissitunicate, cylindrical to cylindro-clavate, with a 

short, furcate pedicel. Ascospores muriform, ellipsoid to 

fusoid. 

Anamorphs reported for genus: Alternaria (Simmons 1986). 

Literature: Kwasna and Kosiak 2003; Kwasnaetal.2006; 

Simmons 1986, 2007; Vieira and Barreto 2006. 

Type species 

Lewia scrophulariae (Desm.) M.E. Barr & E.G. Simmons, 

Mycotaxon 25: 294 (1986). (Fig. 46) 

≡ Sphaeria scrophulariae Desm., Plantes cryptogames 

du Nord de la France, ed. 1 fasc. 15:no. 718 

(1834). 

Ascomata ca. 150–200 μm diam., scattered, erumpent to 

nearly superficial at maturity, subglobose to globose, black, 

smooth, papillate. Papilla short, blunt. Peridium thin. 

Hamathecium of septate pseudoparaphyses, ca. 2–2.5 μm 

broad, anastomosing or branching not observed. Asci 100– 

140×13–17 μm, (4–6-)8-spored, bitunicate, fissitunicate, 

cylindrical to cylindro-clavate, with a short, furcate pedicel, 

ocular chamber unknown (Fig. 46a). Ascospores ellipsoid, 

5 (rarely 6 or 7) transversal septa and one longitudinal 

septum mostly through the central cells, yellowish brown to 

gold-brown, 20–24×8–10 μm (x ¼ 21:5 9:1mm, n=10), 

constricted at median septum, smooth or verruculose 

(Fig. 46b, e and f). 

Anamorph: Alternaria conjuncta (Simmons 1986). 

Primary conidiophore simple with a single conidiogenous 

locus; conidia produced in chains, the first conidia 

in chain is larger, 30–45×10–12 μm, 7 transverse septa, 

1–2 longitudinal or oblique septa in lower cells. 

Secondary conidiophore with 5–7 conidiogenous loci, 

sometimes branched; sporulation in chains, rarely 

branched. 

Material examined: (FH, slide from lectotype). 

Note: The specimen contains only a slide, so limited 

structures could be observed e.g. ascospores. The information 

about ascomata, peridium and whole asci is referred to 

Simmons (1986). 

Notes 

Morphology 

Lewia has “Pleospora-like” teleomorphs, while it has 

Alternaria anamorphs, which are characterized by the 

beakless conidia connected together with secondary 

conidiophore (Simmons 1986). Based on these characters, 

more species under this genus were subsequently reported, 

i.e. Lewia avenicola Kosiak & Kwaśna (Kwasna and 

Kosiak 2003); L. chlamidosporiformans B.S. Vieira & R. 

W. Barreto (Vieira and Barreto 2006); L. alternarina (M. 

D. Whitehead & J.G. Dicks.) E.G. Simmons and L. 

daucicaulis E.G. Simmons (Simmons 2007). Currently 

Lewia comprises 15 species (http://www.mycobank.org, 

24-02-2009). 


Phylogenetic analysis based either on SSU rDNA 

sequences or on multigenes indicated that Lewia species


Fig. 45 Leptosphaerulina australis (from NY, C.T. Rogerson 3836). 

a. Compressed ascoma. Note the obpyriform asci within the ascoma 

and the thin peridium. b, c. Eight-spored asci released from the 

ascomata. Note the apical apparatus (arrowed). d. Ascospores with 

thin sheath. e. An old pale brown ascospore. Scale bars: a-c=50 μm, 

d, e=10 μm 

(Allewia eureka (E.G. Simmons) E.G. Simmons=L. 

eureka) form a robust clade with other members of 

Pleosporaceae (Schoch et al. 2006; Schochetal.2009; 

Zhang et al. 2009a). 


Its position in Pleosporaceae is confirmed. 

Lichenopyrenis Calat., Sanz & Aptroot, Mycol. Res. 105: 

634 (2001). (?Pleomassariaceae) 


Habitat terrestrial, parasitic on lichens. Ascomata mediumsized, 

globose or subglobose. Hamathecium of dense, 

filliform, branching, septate pseudoparaphyses. Asci bitunicate, 

fissitunicate, clavate, with a short sometimes furcate 

pedicel. Ascospores ellipsoidal with broadly rounded ends, 

pale orange-brown, 1-distoseptate. 

Anamorphs reported for genus: see below. 

Literature: Calatayud et al. 2001.


Fig. 46 Lewia scrophulariae 

(from FH, slide from lectotype). 

a Cylindrical ascus with a 

short pedicel. b Ascospores in 

asci. c–f Released muriform 

brown ascospores. Scale bars: 

a=20 μm, b–f=10 μm 

Type species 

Lichenopyrenis galligena Calat., Sanz & Aptroot, Mycol. 

Res. 105: 636 (2001). (Fig. 47) 

Ascomata 140–260 μm high×140–250 μm diam., gregarious, 

initially immersed in galls, later becoming erumpent, 

globose or subglobose, black, roughened (Fig. 47a and b). 

Peridium 18–25 μm wide, composed of 2–5 layers of heavily 

pigmented cells of textura angularis to compressed, cells 

6–11 μm diam., cell wall 1–3 μm thick (Fig. 47c, d and e). 

Hamathecium of dense, long filamentous pseudoparaphyses, 

2.5–4 μm broad, branching, septate. Asci 65–85× 

15–20 μm (x ¼ 74 18mm, n=10), 8-spored, bitunicate, 

fissitunicate, broadly clavate, with a short, thick, some-


times furcate pedicel, up to 13 μm long, ocular chamber 

not observed (Fig. 47f, g, h and k). Ascospores 16–20×9– 

11 μm (x ¼ 18 10mm, n=10), biseriate, ellipsoidal, pale 

orange-brown, 1-distoseptate, with prominent swelling at 

the septum, containing refractive globules, smooth (Fig. 47i, 

jandl). 

Anamorph: The following description is from Calatayud et 

al. (2001). 

Conidiomata pycnidial, arising in galls together with the 

ascomata, immersed, ca. 100–200 μm diam.; wall dark 

brown throughout, composed of 2–5 layers of angular to 

laterally compressed cells; cells relatively large, ca. 8– 

16 μm diam. in superficial view. Conidiophores formed by 

1–3 cells, frequently branched and with the uppermost cells 

bearing 1–4 conidiogenous cells; cells±cylindrical, hyaline 

except at the base, which are sometimes pale brown, 7–15× 

3–4 μm. Conidiogenous cells tapered towards the apex, 14– 

18×3–4 μm. Conidia 5–7×1.5–2 μm. Vegetative hyphae 

hyaline. 

Material examined: SPAIN, Andalucía, Province, Jaén, 

Andújar, lichenicolous on Leptochidium albociliatum 

(Desm.) M. Choisy on acid volcanic rock, 19 Apr. 2000, 

V. Calatayud (MA-Lichen 12715, holotype). 

Notes 

Morphology 

Lichenopyrenis was formally established by Calatayud et 

al. (2001) based on its “perithecioid ascomata with peridium 

comprising compressed cells, fissitunicate and J- asci, wide 

hamathecium filaments, and 1-septate pale orange-brown 

ascospores with distoseptate thickenings at maturity”, andis 

monotypic with L. galligena. The genus was temporarily 

assigned to Pleomassariaceae. Lichenopyrenis galligena is a 

parasite of lichens, occurring in galls in the thallus of the host 

(Calatayud et al. 2001). 


None. 


This is one of the few species that are parasitic on 

lichens. The most comparable species are Parapyrenis 

lichenicola Aptroot & Diederich and Lacrymospora parasitica 

Aptroot (both in Requienellaceae, Pyrenulales) as 

well as some species from Dacampiaceae. The peridium 

structure, cellular pseudoparaphyses, distoseptate and 

smooth, orange-brown ascospores as well as the anamorphic 

stage of Lichenopyrenis can easily distinguish from all 

of them (Calatayud et al. 2001). 

Lineolata Kohlm. & Volkm.-Kohlm., Mycol. Res. 94: 687 



Habitat marine, saprobic (or perthophytic?). Ascomata 

medium-sized, gregarious, immersed to erumpent, obpyriform, 

ostiolate, papillate. Peridium thin, comprising two 

types of cells; outer cells thick stratum pseudostromatic, 

inner stratum thin, composed of a few layers of hyaline 

cells of textura angularis. Hamathecium of dense, long 

trabeculate pseudoparaphyses, embedded in mucilage, 

anastomosing and septate. Asci 8-spored, bitunicate, cylindrical, 

with short pedicels, with an ocular chamber. 

Ascospores uniseriate to partially overlapping, ellipsoidal, 

dark brown, 1-septate. 


Literature: Kohlmeyer and Kohlmeyer 1966; Kohlmeyer 

and Volkmann-Kohlmeyer 1990. 

Type species 

Lineolata rhizophorae (Kohlm. & E. Kohlm.) Kohlm. & 

Volkm.-Kohlm., Mycol. Res. 94: 688 (1990). (Fig. 48) 

≡ Didymosphaeria rhizophorae Kohlm. & E. Kohlm. 

Icones Fungorum Maris (Lehre) 4 & 5: tab. 62a (1967). 


gregarious, immersed to erumpent, obpyriform, ostiolate, 

papillate, subcarbonaceous to subcoriaceous, blackish 

brown (Fig. 48a). Peridium 37–45 μm thick, comprising 

two types of cells; outer cells thick stratum pseudostromatic, 

composed of irregular or roundish, dark brown cells, 

on the outside with a more or less recognizable hyphal 

structure, enclosing some decaying cells of the host, inner 

stratum thin, composed of four or five layers of hyaline, 

polygonal, elongate, thin-walled cells with large lumina, 

merging into the pseudoparaphyses. Hamathecium of 

dense, long trabeculate pseudoparaphyses, 1–1.5 μm broad, 

embedded in mucilage, anastomosing and septate. Asci 

150–175×14–17.5 μm, 8-spored, bitunicate, cylindrical, 

with short pedicels, with an ocular chamber (Fig. 48b). 

Ascospores 23–32(−33)×9–12 μm, uniseriate to partially 

overlapping, ellipsoid, dark brown, 1-septate, not or slightly 

constricted at the septum, striate by delicate costae that run 

parallel or in a slight angle to the longitudinal axis of the 

ascospore (Fig. 48c, d, e and f) (adapted from Kohlmeyer 

and Kohlmeyer 1979). 


Material examined: US, Florida, Middle Torch Key, on 

Rhizophora mangle, 21 Nov. 1965, J. Kohlmeyer (Herb. J. 

Kohlmeyer No. 2390b, isotype); Pirate Grove Key, on R. 

mangle, 5 Jan. 1964 (Herb. J. Kohlmeyer No. 1721 paratype); 

Florida, Virginia Key, on R. mangle, 1 Jan. 1964, leg. 

E. Kohlmeyer (Herb. J. Kohlmeyer No. 1751 paratype); 

Florida, Torch Key, on R. mangle, 20 Nov. 1965, leg. J. 

Kohlmeyer (Herb. J. Kohlmeyer No. 2423 paratype).


Fig. 47 Lichenopyrenis galligena (from MA-Lichen 12715, holotype). 

a, b Ascomata forming in the host tissues. c, d Sections of 

ascomata. e Section of a partial peridium. f–h, k Broadly clavate asci. 

Note the short rounded pedicel. i, j, l Ascospores. Note the small 

swellings at the septa. Scale bars: a, b=0.5 mm, c, d=100 μm, e= 

50 μm, f–h, k=20 μm, i, j, l=10 μm


Notes 

Morphology 

Lineolata was monotypified by L. rhizophorae, whichwas 

originally introduced by Kohlmeyer and Kohlmeyer (1966)asa 

species of Didymosphaeria (as D. rhizophorae).Basedonthe 

morphology of ascomata and asci, Barr (1990a) assignedit 

under Lojkania (as L. rhizophorae). Kohlmeyer and Volkmann- 

Kohlmeyer (1990) restudied this species and noticed that the 

absence of clypeus, almost superficial ascomata, coloured 

peridium, a hamathecium with gelatinous matrix, asci with 

apical ring-like structure and the ornamented ascospores are 

quite different from the modified concept of Didymosphaeria. 

Thus they introduced Lineolata to accommodate D. rhizophorae 

(Kohlmeyer and Volkmann-Kohlmeyer 1990). 


Three isolates of Lineolata rhizophorae from varied geographic 

localities were analyzed by Suetrong et al. (2009)andshowntobe 

related to Caryosporella rhizophorae in Dothideomycetidae and 

excluded from Pleosporomycetidae and Pleosporales. 


Based on initial molecular work it is likely that this species 

does not belong to Pleosporales in spite of its dense 

pseudoparaphyses and other characters shared with the order. 

Fig. 48 Lineolata rhizophorae 

(from Herb. J. Kolmeyer No. 

2390b, isotype of Didymosphaeria 

rhizophorae). a Ascomata 

immersed in the host 

substrate with protruding papilla. 

b Ascospores within an ascus. 

Note the ascospore 

arrangement. c–f One-septate 

ascospores. Note the striate ornamentation 

in (c). Scale bars: 

a=100 μm, a, b=20 μm, c–f= 

10 μm


Loculohypoxylon M.E. Barr, Mycotaxon 3: 326 (1976). 

(Teichosporaceae) 


Habitat terrestrial, saprobic. Ascomata relatively small, 

gregarious, immersed to erumpent, globose or subglobose, 

forming under a clypeus, papillate, ostiolate. 

Peridium thin, a single layer comprising hyaline thinwalled 

cells of textura angularis or textura prismatica. 

Hamathecium of septate pseudoparaphyses. Asci (2–4-)8- 

spored, bitunicate, cylindrical to cylindro-clavate, with a 

short, furcate pedicel, and wide ocular chamber. Ascospores 

broadly elliptic to subglobose, often apiculate at 

both ends, pale to dark brown, aseptate, with a germ 

slit. 


Literature: von Arx and Müller 1975; Barr 1976. 

Type species 

Loculohypoxylon grandineum (Berk. & Rav.) Barr, Mycotaxon 

3: 326 (1976). (Fig. 49) 

≡ Diatrype grandinea Berk. & Rav., in Berkeley, 

Grevillea 4: 95 (1876). 

Ascomata 85–130 μm high×75–145 μm diam., gregarious, 

immersed to widely erumpent, globose or subglobose, 

under a reddish brown to black clypeus, papillate, 

ostiolate (Fig. 49a and b). Peridium 18–30 μm thick 

laterally, 1-layered, composed of hyaline thin-walled cells 

of textura angularis to prismatica, cells up to 5×9 μm 

diam., cell wall 0.5–1 μm thick, apex cells smaller and 

walls thicker (Fig. 49c). Hamathecium comprising 2–3 μm 

broad, septate pseudoparaphyses. Asci 70–90×10– 

12.5 μm (x ¼ 76:5 10:9mm, n=10), (2–4-)8-spored, 

bitunicate, cylindrical to cylindro-clavate, with a short, 

furcate pedicel, up to 25 μm long, with a wide ocular chamber 

(Fig. 49f, g, and h). Ascospores 7.5–10×5–7 μm 

(x ¼ 8:3 5:9mm, n=10), uniseriate to partially overlapping 

at the upper part, broadly elliptic to subglobose, often 

apiculate at both ends, pale to dark brown, aseptate, with a 

germ slit (Fig. 49d and e). 


Material examined: USA, New Jersey, Newfield, on 

bark of Quercus coccinea, Sept. 1878, as Diatrype 

grandinea, Ellis N.A.F. 494 (NY, MASS); on Quercus 

sp. wood, Nov. 1893, as Anthostoma grandinea B. & Rav., 

Ellis & Everhart, N.A.F. 494 (NY); Newfield, Oct. 1881, 

as Diatrype grandinea (NY); Newfield, Jan. 1882, on 

Quercus coccinea, asDiatrype grandinea B. & Rav, Ex 

Herb Ellis (NY); Newfield, Nov. 1893, as Anthostoma 

grandinea, on bark of fallen trunks of Quercus coccinea 

(NY). 

Notes 

Morphology 

Loculohypoxylon grandineum is one of the rare pleosporalean 

species having aseptate ascospores. When emphasis 

is given to ascospore morphology, Semidelitschia 

(monotypified by S. agasmatica Cain & Luck-Allen) is the 

most comparable genus. The large ascomata and ascospores, 

the mucilaginous sheath surrounding the ascospores 

as well as the coprophilous habitat of S. agasmatica differ 

from L. grandineum greatly. Thus Loculohypoxylon was 

introduced as a new genus. 


None. 


Aseptate ascospores are rare in Pleosporales, and the 

position of this fungus needs further verification. The familial 

status of Loculohypoxylon in Teichosporaceae is questionable, 

as it is simply based on the similarity of living habitat, ascomata 

and asci with Immotthia and Teichospora (Barr 2002). 

Lophionema Sacc., Syll. fung. (Abellini) 2: 717 (1883). 



Habitat terrestrial, saprobic? Ascomata solitary, scattered or 

in small groups, immersed to erumpent, globose to subglobose, 

with a flattened base, wall black, papillate, ostiolate. 

Peridium comprising two types of cells which merge in the 

middle. Hamathecium of trabeculate pseudoparaphyses, 

septate, rarely anastomosing and branching. Asci 8-spored, 

bitunicate, fissitunicate unknown, clavate to cylindro-clavate, 

with a short and furcate pedicel and a small inconspicuous 

ocular chamber. Ascospores filliform, hyaline to pale yellow, 

multi-septate, slightly constricted at each septum. 


Literature: Barr1992b; ChestersandBell1970; Ellis and 

Everhart 1892; Höhnel 1909; Solheim 1949. 

Type species 

Lophionema vermisporum (Ellis) Sacc., Syll. fung. (Abellini) 

2: 717 (1883). (Fig. 50) 

≡ Lophiostoma vermispora Ellis, Bull. Torrey bot. Club 

9: 19 (1882). 


solitary, scattered or in small groups of 2–3, immersed to 

erumpent, globose to subglobose, black, papillate, ostiolate. 

Papilla 80–120 μm high, up to 150 μm broad, cylindrical to 

somewhat vertically flattened neck; mostly with a short


Fig. 49 Loculohypoxylon grandineum (from N.A.F. 494). a Appearance 

of ascomata on the host surface. b Habitat section of ascomata. c 

Section of an ascoma. Note the pale brown thin-walled peridium cells. 

slot-like ostiole, periphysate (Fig. 50a). Peridium 30–45 μm 

wide at the sides and slightly thicker at the apex, 

d, e Uniseriate ascospores in asci. f–f Cylindro-clavate asci with 

ascospores. Note the ocular chamber in (g). Scale bars: a=100 μm, 

b=200 μm, c=50 μm, d–h=10 μm 

2-layered, lateral walls and wall adjacent to neck comprising 

two types of cells which merge in the middle; outer


cells small heavily pigmented thick-walled cells of textura 

angularis, cells 4–7 μm diam., cell wall 3.5–5 μm thick, 

inner cells less pigmented, comprising thin-walled compressed 

cells; apical wall cells smaller and walls thicker, 

basal wall thinner (ca. 15μm wide), composed of lightly 

pigmented thin-walled compressed cells (Fig. 50b and c). 

Hamathecium of trabeculate pseudoparaphyses, 1–2 μm 

broad, septate, anastomosing and branching rarely between 

and mostly above the asci. Asci 105–130(−150)×10–15 μm 

(x ¼ 123 12mm, n=10), 8-spored, bitunicate, fissitunicate 

dehiscence not observed, clavate to cylindro-clavate, with a 

short, narrow, furcate pedicel which is 10–25 μm long, and 

a small inconspicuous ocular chamber (to 1.5 μm wide× 

1 μm high) (Fig. 50d and e). Ascospores (80-)90–115×3– 

5 μm (x ¼ 95 3:5mm, n=10), filliform, gradually tapering 

towards the base, hyaline to light yellow, (6-)7(−8)-septate, 

slightly constricted at each septum, smooth (Fig. 50f). 



dead stems of Oenothera biennis, Aug. 1881, Ellis (NY 

643, holotype, NY 885, isotype). 

Notes 

Morphology 

Lophionema is a relatively poorly studied genus, which 

was formally established by Saccardo (1883) as a monotypic 

genus represented by L. vermisporum based on its “globose 

ascomata, compressed ostiole, cylindrical to clavate ascus, 

and filamentous, septate, subhyaline to lightly pigmented 

ascospores”. Lophionema vermisporum was consequently 

listed as the generic type (Clements and Shear 1931). Berlese 

(1890) placed the genus in Lophiostomataceae but mentioned 

that the genus was similar to Ophiobolus according to 

the variable apex, and Shoemaker (1976) transferred Lophionema 

vermisporum to Ophiobolus sensu lato. Chesters and 

Bell (1970) however, had regarded Lophionema as related to 

Lophiostoma despite the distinct ascospore morphology. Barr 

(1992b) assignedLophionema to Entodesmium based on the 

morphology of ascomata, papilla, peridium structure, pseudoparaphyses 

as well as the hyaline or slightly yellowish 

ascospores with a terminal appendage (not observed here). 

Species of Entodesmium, however, exclusively occur on 

legumes, but Lophionema vermisporum does not. We also 

note that the filliform ascospores, bitunicate asci, pseudoparaphyses 

and nature of the peridium may also be considered 

as typical of genera in the Tubeufiaceae (Barr 1980; 

Kodsueb et al. 2006b). 


None. 


The immersed to erumpent ascomata, trabeculate pseudoparaphyses 

and laterally flattened papilla and periphysate 

ostioles indicate that this genus should be included in 

Lophiostomataceae. We do not accept the above proposals 

and, consider that Lophionema should be maintained as a 

separate genus with filliform ascospores in Lophiostomataceae 

until representative taxa can be sequenced and 

analyzed. Currently Lophionema comprises 10 species 

(http://www.mycobank.org, 08-01-2009). However, many 

of these are poorly studied and obscure. 

Lophiostoma Ces. & De Not., Comm. Soc. crittog. Ital. 1: 

219 (1863). (Lophiostomataceae) 


Habitat terrestrial, saprobic. Ascomata immersed to erumpent, 

usually with a distinct depressed papilla and a slotlike 

ostiole. Hamathecium of dense, long, septate pseudoparaphyses, 

embedded in mucilage, anastomosing and 

branching between and above the asci. Peridium unequal 

in thickness, thicker near the apex and thinner at base. Asci 

usually clavate. Ascospores 1-septate, multi-septate or even 

muriform, hyaline to deep brown, usually with terminal 

appendages. 

Anamorphs reported for genus: Pleuorphomopsis-like 

(Hyde et al. 2011). 

Literature: Barr 1990a; Chesters and Bell 1970; Holm and 

Holm 1988; Hyde and Aptroot 1998; Hyde et al. 2002; 

Tanaka and Harada 2003b; Yuan and Zhao 1994. 

Type species 

Lophiostoma macrostomum (Tode) Ces. & De Not., 

Comm. Soc. crittog. Ital. 1: 219 (1863). (Fig. 51) 

≡ Sphaeria macrostoma Tode, Fung. mecklenb. sel. 

(Lüneburg) 2: 12 (1791). 


densely scattered to gregarious, semi-immersed to erumpent, 

globose or subglobose, with a small to large 

flattenedcrest-likeraisedareaabovetheascomatawhich 

is variable in shape, up to 300 μm high and 480 μm 

wide, with a slit-like ostiole along the full length of the 

crest (Fig. 51a and b). Peridium 30–45 μm thick at the 

sides, thicker at the apex and thinner at the base, 

composed of one cell type of small lightly pigmented 

thin-walled cells of textura prismatica, cellsca. 6–9×3– 

4 μm diam., apex composed of pseudoparenchymatous 

cells (Fig. 51b). Hamathecium of dense, filliform, up to 

3 μm near the base and less than 1.5 μm broad in the


Fig. 50 Lophionema vermisporum (from NY-643, holotype). a 

Appearance of ascomata on the host surface. Note the form of the 

neck. b Section of the peridium. c Peridium comprising two types of 

cells which merge in the middle; outer cells small heavily pigmented 

thick-walled cells of textura angularis, inner cells less pigmented, and 

comprising thin-walled compressed cells. d, e Cylindro-clavate, 8- 

spored asci. f A 7-septate filliform ascospore. Scale bars: a=0.5 mm, 

b=100 μm, c=50 μm, d–f=10 μm


upper place, septate pseudoparaphyses, embedded in 

mucilage, anastomosing and branching between and 

above the asci (Fig. 51f). Asci 110–145×10–15 μm 

(x ¼ 127:5 13mm, n=10), 8-spored, bitunicate, fissitunicate 

(ectotunica no constriction), cylindro-clavate, with a 

furcate pedicel and a small ocular chamber (to 1.5 μm wide× 

2 μm high) (J-) (Fig. 51c, d and e). Ascospores 27–38(−43)× 

5–7.5 μm (x ¼ 31:2 6:4mm, n=10), biseriate, fusoid, 

curved, hyaline, usually 1-septate, with 3–5 septaandfaintly 

brown when old, with (2-)3(−4) distinct oil drops in each cell 

and short terminal appendage at ends (Fig. 51h, i and j), and 

ornamented with warts when spores are senescent (Fig. 51g). 


Material examined: SWEDEN, Smaland, Femsjö par., 

Femsjö, on Prunus, 2006, Elias Fries, det. Geir Mathiassen 

(UPS, lectotype, asSphaeria macrostoma Fr.). FRANCE, 

Ariège, Rimont, Las Muros, on dead stems of Vitis vinifera, 

2 Sept. 1996 (IFRD2005). 

Notes 

Morphology 

Lophiostoma is morphologically a well studied genus 

(Barr 1990a; Chesters and Bell 1970; Holm and Holm 

1988; Mugambi and Huhndorf 2009b; Yuan and Zhao 

1994), and currently it comprises about 30 species (Tanaka 

and Harada 2003b). The genus was characterized as 

having immersed to erumpent ascomata with a cylindrical 

or crest-like papilla and full length, slit-like ostiole; a 

peridium of unequal thickness, which was broader near the 

base (Lophiostoma-type); mostly clavate, bitunicate asci 

and 1- to several septate, hyaline to pigmented ascospores 

with terminal appendages or surrounded by a mucilaginous 

sheath (Holm and Holm 1988). This definition was 

followed by Barr (1990a), Yuan and Zhao (1994) and 

Hyde et al. (2002). 

The crest-like papilla has been regarded as a prominent 

morphological character of Lophiostoma macrostomum 

(Chesters and Bell 1970; Holm and Holm 1988). In the 

lectotype specimen, the raised area above the ascomata is 

up to 300 μm high and 480 μm long, and seen as a 

flattened or even Y-shaped crest (Fig. 51a). In Lophiostoma 

curtum (Fr.) De Not. and Lophiotrema boreale Math. the 

raised area above the ascomata varies considerably in 

height or is even lacking (Holm and Holm 1988). Thus 

the variable “crest-like raised area in Lophiostomataceae” 

was explained as an evolutionarily adaptation to the hard 

substrate within which the ascomata develop (Holm and 

Holm 1988). The ascospores of L. macrostomum usually 

turn reddish brown when mature, and minutely verrucose 

ornamentation was also found on the surface of the 

pigmented ascospores. Hyaline ascospores that became 

pigmented with age are common in Lophiostoma, such as 

in L. appendiculatum Fuckel, L. massarioides (Sacc.) L. 

Holm & K. Holm, L. semiliberum, L. subcorticale Fuckel 

and L. winteri (Holm and Holm 1988; Tanaka and Harada 

2003b). The phylogenetic significance of this character 

should be observed carefully in the future but at present its 

phylogenetic significance is unclear as this also occurs in 

some Lophiotrema species. 


Phylogenetic affinity with some Massarina species has 

been reported by Liew et al. (2002), and several Massarina 

species were transferred into Lophiostoma. Inasystematic 

study of Lophiostoma- and Massarina-related fungi conducted 

by Zhang et al. (2009b), Lophiostoma taxa clustered 

into two groups; one includes the type species L. macrostomum 

with crest-like ostioles, L. rugulosum Yin. Zhang, J. 

Fourn. & K.D. Hyde with a wide, umbilicate pore surrounded 

by 4–6 radial ridges, and L. glabro-tunicatus with small 

ostiolar pores; the other cluster comprises Lophiostoma-like 

taxa with slot-like ostioles lacking raised crests, which 

includes L. arundinis (Pers.) Ces. & De Not., L. caulium, L. 

compressum (Pers.) Ces. & De Not., L. crenatum (Pers.) 

Fuckel, L. fuckelii (Sacc.) Sacc., L. macrostomoides, L. 

semiliberum and L. viridarium Cooke, which seems to 

represent a natural group at the family level. This conclusion 

is tentative until verified sequences of L. macrostomum are 

included in analyses (see comments of Zhang et al. 2009a). 


We tend to accept a narrow concept of Lophiostomataceae, 

which only comprises species of Lophiostoma sensu 

stricto (Zhang et al. 2009a). 

Lophiotrema Sacc., Michelia 1: 338 (1878). (Pleosporales, 

genera incertae sedis) 



with or without short papilla. Hamathecium of dense, 

long, septate pseudoparaphyses, anastomosing and branching 

between and above asci. Asci cylindrical to cylindroclavate. 

Ascospores hyaline, 1–3-septate, usually with 

mucilaginous sheath. 



Holm 1988; Saccardo 1878a; Tanaka and Harada 2003c; 

Tang et al. 2003; Yuan and Zhao 1994. 

Type species 

Lophiotrema nucula (Fr.) Sacc., Michelia 1: 338 (1878). 

(Fig. 52)


≡ Sphaeria nucula Fr., Syst. mycol. (Lundae) 2: 466 (1823). 


erumpent to nearly superficial, with basal wall remaining 

immersed in host tissue, globose to subglobose, often 

laterally flattened, with a flattened base not easily removed 

from the substrate, black, roughened; with a cylindrical or 

slightly compressed papilla. Papilla to 120 μm long and 

150 μm high, protruding, with a pore-like ostiole (Fig. 52a). 

Peridium 25–30 μm wide, very thin at the base, composed of 

heavily pigmented pseudoparenchymatous cells near the apex, 

cells 2–2×6 μm diam., wall 1–3(−4) μm thick, lower sides 

composed of pigmented cells of textura angularis, 3–5 μm 

diam., wall 0.8–1.5 μm thick, ostiole wall composed of 

heavily pigmented and thick-walled small cells (Fig. 52b and 

c). Hamathecium of dense, long, septate pseudoparaphyses, 

1–2 μm broad, anastomosing and branching between and 

above asci, embedded in mucilage (Fig. 52i). Asci 90–115× 

9–11.5 μm (x ¼ 99:5 11:5mm; n=10), 8-spored, bitunicate, 

fissitunicate, cylindrical, with a short, narrowed, furcate 

pedicel which is up to 10 μm long, with a small ocular 

chamber (ca. 1.5μm wide×1 μm high) (J-) (Fig. 52d, e, f 

and h). Ascospores 17–21(−25)×(4-)5–6.5 μm 


overlapping to biseriate, broadly fusoid to fusoid, with 

narrowly rounded ends, hyaline and lightly pigmented on 

very rare occasions when senescent, 1-septate, 3-septate 

when old, constricted at the median septum, the upper cell 

often broader than the lower one (Fig. 52g). 


Material examined: on decaying wood (UPS, lectotype 

as Sphaeria nucula Fr.). 

Notes 

Morphology 

Holm and Holm (1988) provided a relatively strict 

definition for Lophiotrema after they examined several specimens 

including the type materialwhichtheylectotypified. 

Lophiotrema was mainly defined on the unique characters of 

small to medium ascomata, a “Lophiotrema-type” peridium 

and 1-septate ascospores. In Lophiotrema, Holm and Holm 

(1988) considered the ascomata to be small- to medium-sized, 

ca. pyriform but neck often reduced, even lacking and 

sometimes cylindrical. The peridium was of approximately 

equal thickness, 20–30 μm, composed of an outer textura 

angularis of uniformly pigmented cells, up to 12 μm, andan 

inner layer of very small hyaline cells, with somewhat 

thickened walls. Asci are cylindrical, spores hyaline, at first 

1-septate, becoming 3-septate, with distinct guttules, often 

with a mucilaginous sheath. Much emphasis was given to the 

1-septate ascospores by Holm and Holm (1988) when they 

described and distinguished the three Lophiotrema species: L. 

boreale, L. nucula, L. vagabundum (Sacc.) Sacc. and two 

Fig. 51 Lophiostoma macrostomum (a–h, j from UPS, leptotype; i b 

from IFRD 2005). a Appearance of ascomata on the host surface. 

Note the raised crest-like areas and full length germ slits. b Section of 

the peridium. c–e Cylindro-clavate asci with ascospores arranged in a 

2-3-seriate manner. f Hamathecium comprising branching and septate 

pseudoparaphyses. g–j Released or unreleased ascospores. Note the 

smooth young ascospores with terminal sheath, and the verrucose 

senescent ascospores. Scale bars: a=0.5 mm, b=200 μm, c–j=10 μm 

other unnamed species. This concept was widely accepted by 

later workers (Kirk et al. 2001; YuanandZhao1994). Tanaka 

and Harada (2003c) considered the peridium and asci to 

distinguish Lophiotrema from Lophiostoma, while Tang et al. 

(2003) introduced a new Lophiotrema species with elongated 

slit-like ostiole stating that the main difference between 

Lophiotrema and Lophiostoma were size of ascomata, 

structure of peridium, shape of asci and sheath of ascospores. 

This peridium concept however, is not supported by the 

lectotype specimen we examined here, which has a flattened 

thin-walled base. Thus the “Lophiotrema-like peridium” sensu 

Holm and Holm (1988) should not serve as a diagnostic 

character of Lophiotrema, while the ostiole, asci and 

ascospores might have some phylogenetic significance (Zhang 

et al. 2009b). No anamorph is yet known for Lophiotrema. 

Although the ascospores was reported by Holm and Holm 

(1988) to be verruculose this could not be observed in the 

lectotype examined under light microscope (1000 ×) in the 

present study. 


In the phylogenetic study of Lophiostoma, Massarina and 

related genera (Zhang et al. 2009b), Lophiotrema nucula 

formed a consistent and robust clade with three other 

Lophiotrema species: L. lignicola Yin. Zhang, J. Fourn. & 

K.D. Hyde, L. brunneosporum Yin. Zhang, J. Fourn. & K.D. 

Hyde and L. vagabundum, separate from other members of 

Lophiostoma and Massarina sensu stricto. This clade might 

represent Lophiotrema sensu stricto, however, the correctness 

of strains of L. vagabundum (CBS 628.86) and L. 

nucula (CBS 627.86) used in the phylogenetic study are not 

verified and warrant further study. 


Holm and Holm (1988) distinguished Lophiostoma from 

Lophiotrema based on the smaller ascomata, 1-septate 

versus multi-septate ascospores, and peridial wall structure. 

However, we doubt that these distinguishing characters 

(size of ascomata, number of septa of ascospores) can be 

confidently used to separate these genera and we could not 

establish any characters that could reliably distinguish 

between these two genera. The molecular data, however, 

does separate Lophiostoma macrostomum and Lophiotrema 

nucula into separate clades and provides some support that 

these are separate genera. Although the strain of L. nucula

Fungal Diversity


Fig. 52 Lophiotrema nucula (from UPS, lectotype). a Ascomata on 

the host surface. b Section of a partial ascoma. c Peridium structure 

near the apex. d, h Cylindrical asci in the pseudoparaphyses. e, f 

Upper part of the asci, showing the small ocular chamber near the 

apex. h Mature ascospores. i Pseudoparaphyses. Scale bars: a= 

0.5 mm, b=100 μm, c, d=30 μm, e–i=10 μm


(CBS 627.86) was isolated by K. & L. Holm, who had 

examined the type specimen of L. nucula (Holm and Holm 

1988), the culture of Lophiostoma macrostomum used in 

the analysis are unverified (see comment by Zhang et al. 

2009b). For the purpose of this monograph we tentatively 

maintain Lophiotrema as distinct from Lophiostoma. 

Macroventuria Aa, Persoonia 6: 359 (1971). 

(Didymellaceae) 


Habitat terrestrial, saprobic. Ascomata small, solitary, scattered, 

or in groups, initially immersed, becoming erumpent, to nearly 

superficial, globose to subglobose, roughened with cylindrical 

setae erect from apex. Peridium thin, membranous. Hamathecium 

of cellular pseudoparaphyses, seems to easily disappear 

when mature. Asci bitunicate, somewhat obclavate to fusoid. 

Ascospores fusoid with broadly to narrowly rounded ends, 

hyaline, 1-septate, constricted at the septum. 


Literature: van der Aa 1971; von Arx and Müller 1975; 

Barr 1987a. 

Type species 

Macroventuria wentii Aa, Persoonia 6: 361 (1971). (Fig. 53) 

Ascomata 135–180 μm diam., rarely more than 200 μm 

diam., solitary, scattered or in groups, initially immersed, 

becoming erumpent, to nearly superficial, with basal wall 

remaining immersed in host tissue, globose to subglobose, 

broadly or narrowly conical, setae erect from the apical region of 

the ascomata, cylindrical or tapering to the rounded or pointed 

tip, brown, up to 90 μm long, 5–7.5 μm thick (Fig. 53a). 

Peridium, 25–35 μm thick, 2-layered, out layer composed of 

relatively thick-walled cells of textura angularis, cell wall up to 

3 μm thick; inner layer cells with a thinner wall and subhyaline; 

near apex cells smaller (Fig. 53a). Hamathecium of cellular 

pseudoparaphyses, 1–2 μm thick, evanescing not sure. Asci 75– 

93×24–30 μm, 8-spored, without pedicel, bitunicate, somewhat 

obclavate to fusoid (Fig. 53b). Ascospores 22–32×8–14 μm, 1– 

3 seriate, fusoid with broadly to narrowly rounded ends, hyaline, 

1-septate, constricted at the septum, smooth (Fig. 53b) (description 

adapted from van der Aa 1971). 


Material referred: USA, Nevada; Death Valley, plant 

litter, F.W. Went, 229, 1970 (CBS 526.71, holotype). 

Notes 

Morphology 

Macroventuria was formally established by van der Aa 

(1971) represented by M. anomochaeta and M. wentii based 

on its “near-hyaline, 1-septate ascospores, setose ascomata, 

and saprobic life style”. Almost all of the above characters 

(except the saprobic life style) point this group of fungi to 

Venturiaceae. Thus Macroventuria was assigned to this 

family as a relatively primitive genus (van der Aa 1971). 

Subsequently, von Arx and Müller (1975) assigned Macroventuria 

to Pseudosphaeriaceae (Dothideales), and this 

proposal was followed by Barr (1987a). 


Phylogenetic analysis based on combined SSU rDNA 

and LSU rDNA sequences indicated that both of Macroventuria 

anomochaeta and M. wentii form a robust clade 

with Leptosphaerulina argentinensis (Speg.) J.H. Graham 

& Luttr., L. australis, L. trifolii (Rostr.) Petr. and 

Platychora ulmi, which appear to share phylogenetic 

affinities with the Leptosphaeriaceae and Phaeosphaeriaceae, 

but detached from other members of Venturiaceae 

and Pleosporaceae (Kodsueb et al. 2006a). In addition, 

culture characters also support the close relationship 

between Macroventuria and Leptosphaerulina (Barr 

1987a). Analysis based on five genes, i.e. SSU, LSU, 

RPB1, RPB2 andTEF1, indicated Macroventuria anomochaeta 

resides in the well supported clade of Didymellaceae 



The morphological characters, such as small ascomata 

and hyaline, 1-septate ascospores all point at Didymellaceae, 

thus the familial status of Macroventuria is verified. 

Mamillisphaeria K.D. Hyde, S.W. Wong & E.B.G. Jones, 

Nova Hedwigia 62: 514 (1996b). (?Melanommataceae) 


Habitat freshwater, saprobic. Ascomata superficial, scattered 

or gregarious, conical, carbonaceous, papillate. 

Hamathecium of dense, filliform, trabeculate pseudoparaphyses. 

Asci broadly clavate to clavate, with small ocular 

chambers and short pedicels. Ascospores of two types, (1): 

2-4-seriate, ellipsoid, hyaline, slightly constricted at the 

main septum; with apical appendages at each end and 

around the ascospore; (2) 1-2-seriate, ellipsoid to fusoid, 

brown, with mucilaginous sheath around the ascospore 

(Hyde et al. 1996b). 


Literature: Hyde et al. 1996a, b. 

Type species 

Mamillisphaeria dimorphospora K.D. Hyde, S.W. Wong 

& E.B.G. Jones, Nova Hedwigia 62: 515 (1996b). (Fig. 54)


Fig. 53 Macroventuria wenti. 

a Ascomata. Note the setae. b 

Ascus and ascospores. Scale 

bars: a=50 μm, b=10 μm 

(figures referred to van der Aa 

1971) 

Following description is adapted from Hyde et al. 

1996a, b). 


scattered or in small groups, superficial, conical, carbonaceous, 

papillate, under pseudostroma which forms a thin 

layer on the host surface, up to 50 μm thick between the 

ascomata and 125–250 μm thick on the ascomata surface 

(Fig. 54a and b). Peridium 10–25 μm thick, comprising 

several layers of compressed, densely packed, thin-walled, 

hyaline cells. A wedge-shaped area of vertically orientated 

hyaline palisade-like cells occurs at the periphery 

(Fig. 54b). Hamathecium of dense, trabeculate pseudoparaphyses, 

ca. 1μm broad, hyaline, branching and anastomosing, 

septate, embedded in mucilage (Fig. 54e). Two 

types of asci and ascospores exist in the same ascoma: 

TYPE 1: asci 185–320×40–100 μm (x ¼ 210 78mm, n= 

50), 8-spored, cylindro-clavate, bitunicate, fissitunicate, 

short-pedicellate, with an ocular chamber (to 13 μm 

wide×5 μm high) (Fig. 54c and d). Ascospores 66–84× 

20–38 μm (x ¼ 78 25mm, n=50), 2-4-seriate, hyaline, 

ellipsoidal, constricted at the central septum, with pad-like 

mucilaginous appendage at each end and with some 

mucilage associated around the spore, and TYPE 2: asci 

158–242×8–15 μm (x ¼ 182 11mm, n=50), 8-spored, 

cylindrical, bitunicate, fissitunicate, pedicellate, with an 

ocular chamber and faint apical ring, ascospores 29–42× 

6–9 μm (x ¼ 35 7mm, n =50), 1-2-seriate, brown, 

ellipsoidal-fusoid, surrounded by a thin mucilaginous 

sheath (Fig. 54f, g, h, i and j). 


Material examined: BRUNEI, on submerged wood, 

Aug. 1997, leg. K.D. Hyde (HKU(M) 7425). 

Notes 

Morphology 

Mamillisphaeria was established as a monotypic genus 

according to its bitunicate, fissitunicate asci, trabeculate 

pseudoparaphyses and dimorphic ascospores, which is 

typified by the widely distributed freshwater fungus, M. 

dimorphospora (Hyde et al. 1996a, b). The most striking 

character of this fungus is its dimorphic ascospores, i.e. 

one type is large and hyaline, and the other is comparatively 

smaller and brown. Only a few ascomycetes have 

been reported having dimorphic ascospores, such as 

Aquasphaeria dimorphospora and Nectria heterospora 

Speg. (Hyde 1995b; Spegazzini 1889). Dimorphic ascospores 

appear to have evolutionary benefits, for example 

the large ascospores with mucilaginous sheaths may 

facilitate nutrient storage for germination and enhanced 

collision and attachment to substrates. The smaller brown 

ascospores may help resist desiccation and damage by UV 

light and contribute to aerial dispersal, which might 

explain the worldwide distribution of M. dimorphospora 

(Hyde et al. 1996a, b). 


None. 


Although in the key by Barr (1990a), M. dimorphospora 

can be referred to Massariaceae, itistemporarily 

assigned to Melanommataceae here based on its 

trabeculate pseudoparaphyses embedded in mucilage 

(Hyde et al. 1996a, b).


Fig. 54 Mamillisphaeria dimorphospora (from HKU(M) 7425, 

paratype?). a Ascomata scattered on the host surface. Note the small 

papilla. b Section of an ascoma. c, d Asci (TYPE 1). e Trabeculate 

pseudoparaphyses in a gelatinous matrix. f–j Ascospores (TYPE 1). 

Scale bars: a=0.5 mm, b–d=100 μm, e=10 μm, f–j=20 μm


Massarina Sacc., Syll. fung. (Abellini) 2: 153 (1883). 

emend. (Massarinaceae) 


Habitat terrestrial, saprobic. Ascomata immersed or superficial, 

scattered or clustered, globose, conical globose to 

lenticular, papillate or epapillate, ostiolate. Hamathecium of 

dense, cellular pseudoparaphyses. Asci clavate to cylindrical, 

with short pedicels. Ascospores ellipsoid to fusoid, 

hyaline, 1- to 3-septate, with or without mucilaginous 

sheath. 

Anamorphs reported for genus: Ceratophoma (Sivanesan 

1984). 

Literature: Aptroot 1998; Barr 1990a; Bose 1961; Eriksson 

and Yue 1986; Hyde 1995a; Hyde and Aptroot 1998; 

Liew et al. 2002; Saccardo 1883; Sivanesan 1984; Tanaka 

and Harada 2003d; Zhang et al. 2009a, b. 

Type species 

Massarina eburnea (Tul. & C. Tul.) Sacc., Syll. fung. 

(Abellini) 2: 153 (1883). (Fig. 55) 

≡ Massaria eburnea Tul. & C. Tul., Sel. Fung. Carp. 2: 

239 (1863). 

Ascomata to 250 μm high×500–700 μm diam., solitary or 

in small clusters, forming under raised dome-shaped areas, 

with blackened centres, with a central ostiole, immersed 

within the cortex of thin dead branches, ellipsoidal, 

rounded from above, clypeate, neck central, short and 

barely noticeable on host surface (Fig. 55a). Clypeus ca. 

250 μm diam., 60 μm thick, brown, comprising compact 

brown-walled cells of textura angularis to globulosa 

beneath host epidermal cells (Fig. 55b). Peridium ca. 

20 μm thick comprising 3–5 layers of hyaline compressed 

cells, fusing at the outside with the host (Fig. 55e). 

Hamathecium filamentous, cellular pseudoparaphyses, ca. 

2 μm broad, septate, embedded in mucilage, without 

anastomosing (Fig. 55d). Asci 108–170×18–22 μm 

(x ¼ 144:5 18:8mm, n=10), 8-spored, cylindro-clavate, 

pedunculate, bitunicate, fissitunicate, (1-)2-seriate, apically 

rounded, with an ocular chamber and faint ring (J-) (Fig. 55c 

and f). Ascospores 30–38×8–12 μm (x ¼ 32:4 8:6mm, n= 

10), fusoid to ellipsoid, 4-celled, constricted at the septa, 

hyaline, with acute rounded ends and surrounded by (5– 

8 μm diam.) mucilaginous sheath (Fig. 55g). 

Anamorph: Ceratophoma sp. (Sivanesan 1984). 

Material examined: FRANCE, on twig of Fagus sp., 

(Desmazières 1764. P, holotype of Sphaeria pupula var 

minor), (Mycotheca universalis no. 1951 lectotype). AUS- 

TRIA, Silesia, Karlsbrunn, on dead twigs of Fagus sylvatica 

L., Aug. and Sept. 1890, Niessl., De Thümen, sub. Massarina 

eburnea, ETH. Saxonia, Königsbrunn, on twigs of Fagus 

sylvatica, Apr. 1882, W. Krieger, Rabenhorst & Winter, Fungi 

europaei no. 2767, ETH; FRANCE, on a dead twig of Fagus 

sylvatica, Deux Sèvres, Villiers en Bois, Forêt de Chizé, 

Rimbaud, 14 Apr. 2008, leg. det. Paul Leroy (IFRD 2006). 

Notes 

Morphology 

Massarina was introduced by Saccardo (1883) for 

species of pyrenocarpous ascomycetes that had previously 

been placed in Massaria, but typically had hyaline 

ascospores (Bose 1961). The family Massarinaceae was 

described by Munk (1956) to accommodate Massarina. 

This family was not commonly used and Massarina was 

later placed within the Lophiostomataceae in the Pleosporales 

(Barr 1990a; Bose 1961; Eriksson and Yue 1986). Of 

the 160 epithets listed in his monograph, Aptroot 

accepted only 43 species (Aptroot 1998). The concept 

of Massarina was widely accepted as having single or 

aggregated, immersed to erumpent, spherical to hemispherical, 

pseudothecioid ascomata; cellular pseudoparaphyses; 

bitunicate, cylindrical to clavate or obpyriform 

asci; and hyaline, 1–3(−7)-septate, fusoid to long ellipsoid 

ascospores that mostly have a mucilaginous sheath or 

appendages (Aptroot 1998; Hyde and Aptroot 1998; 

Tanaka and Harada 2003d). 

In the holotype of Sphaeria pupula var. minor (P) and 

lectotype of Massarina eburnea (ETH), ascospores are 

reported as “not constricted at the septa” (Hyde 1995a). 

However, in one of our recent collections, ascospores that 

are constricted at their septa were observed (Fig. 55g), 

which was consistent with the description by Fallah and 

Shearer (2001). This might be because this character is not 

clear in the old (over 100 years) and dry herbarium 

specimens, or it may be variable between collections. 


Recent morphological, molecular and anamorphic results 

indicate, however, that Massarina is polyphyletic (Hyde 

1995a; Kirk et al. 2001; Liew et al. 2002). Based on the 

rDNA dataset, Massarina cisti and the type of Massarina 

(M. eburnea) forms a robust clade representing Massarina 

sensu stricto (Zhang et al. 2009a, b). 


Massarina sensu stricto should be accepted, which 

seems to only include some terrestrial and saprobic species. 

Massariosphaeria (E. Müll.) Crivelli, Diss. Eidgenöss. 

Techn. Hochschule Zürich 7318: 141 (1983). 

(?Amniculicolaceae) 

≡ Leptosphaeria subgen. Massariosphaeria E. Müll., 

Sydowia 4: 206 (1950).


Fig. 55 Massarina eburnea 

(from IFRD 2006). a Ascomata 

on the host surface. b Section 

of an ascoma. c Ascus with a 

short pedicel. d Cellular pseudoparaphyses. 

e Section of the 

peridium comprising a few 

layers of compressed cells. f 

Asci in pseudoparaphyses. g 

Three-septate ascospores. Scale 

bars: a=0.5 mm, b=100 μm, 

c–g=20 μm 


Habitat terrestrial or freshwater, saprobic. Ascomata 

medium-sized, scattered, or in small groups, immersed, 

erumpent to superficial, subglobose, black; apex with a 

wide and usually somewhat compressed papilla. Peridium 

thick or thin, usually thicker near the apex, 

composed of 2–3 layers of thick walled scleroparenchymatous 

cells. Hamathecium of dense, trabeculate 

pseudoparaphyses. Asci 8-spored, bitunicate, cylindrical 

to cylindro-clavate, with a short, thick, furcate pedicel. 

Ascospores fusoid to narrowly ellipsoid, brown or dark 

brown, multi-septate. 


Literature: Barr1989c; Huhndorf et al. 1990; Kohlmeyer et al. 

1996; Müller 1950; Tanaka and Harada 2004;Tanakaetal.2005. 

Type species 

Massariosphaeria phaeospora (E. Müll.) Crivelli, Ueber 

die Heterogene Ascomycetengattung Pleospora Rabh.; 

Vorschlag für eine Aufteilung (Diss. Eid genössischen Tech 

Hochsch Zürich 7318): 141 (1983). (Fig. 56) 

≡ Leptosphaeria phaeospora E. Müll., Sydowia 4: 208 

(1950). 


scattered, or in small groups, immersed, semi-immersed, 

subglobose, black, apex wide papilla, sometimes slightly 

compressed, 40–70(−100) μm broad (Fig. 56a). Peridium


10–20 μm wide at sides, comprising one cell type of 

2–3 layers of thick walled scleroparenchymatous cells, 

cell wall 2–5 μm thick, peridium thicker near 

the apex (Fig. 56b). Hamathecium of dense, trabeculate 

pseudoparaphyses, 1–2 μm broad, septate, branching 

and anastomosing. Asci 120–173×18–25 μm 

(x ¼ 133:2 20:5mm, n=10), 8-spored, bitunicate, fissitunicate 

dehiscence not observed, broadly cylindrical to 

cylindro-clavate, with a short, thick, furcate pedicel, up to 

15 μm long. Ascospores 32.5–42×10–13 μm 

(x ¼ 36 11:2mm, n=10), narrowly ellipsoid, usually 

slightly curved, dark brown, 7–9 septa, slightly constricted 

at the median septum (Fig. 56c and d). 


Material examined: SWITZERLAND, Kt. Wallis, 

Findelen, Artemisiae campestris L., 10 Sept. 1895, H. 

Wegelin (ZT, holotype). 

Notes 

Morphology 

Massariosphaeria was established by Müller (1950) as 

a section of Leptosphaeria based on its large, thick-walled 

ascospores with a mucilaginous sheath as well as its 

ascomata with a thick apex. Massariosphaeria was 

introduced as a separate genus by Crivelli (1983), 

characterized by its wide peridial apex comprising 

thick-walled cells, compressed to round papilla, and 

relatively large, thick-walled, reddish brown to brown, 

multi-septate to dictyosporous ascospores, usually surrounded 

by a sheath (Crivelli 1983; Huhndorf et al. 1990; 

Leuchtmann 1984). In particular, Crivelli (1983) emphasized 

that species of Massariosphaeria often stain the woody 

substrate (or culture) purple, and this was accepted by 

Leuchtmann (1984). Barr (1989c) had treated Massarios- 

Fig. 56 Massariosphaeria 

phaeospora (ZT, holotype). a 

Ascomata scattering on the host 

surface. Note the immersed to 

erumpent ascomata. b Section of 

a partial peridium. Note the 

peridium structure. c, d Released 


a=1 mm, b–d=20 μm


phaeria as a synonym of Chaetomastia, but this viewpoint 

was rarely followed. 


The polyphyletic nature of Massariosphaeria is detected 

by analyzing SSU and LSU rDNA sequences (Wang et al. 

2007). The purple staining character has shown phylogenetic 

significance in Amniculicolaceae, a freshwater family from 

France (Zhang et al. 2009a). A single isolate of M. 

phaeospora was shown to be unrelated to Amniculicolaceae 

and clustered with a single isolate of Thyridaria rubronotata 

(Schoch et al. 2009; Zhang et al. 2009a). 


Based on phylogenetic analysis, staining the substrate purple 

may have more phylogenetic significance than morphological 

characters (Zhang et al. 2009a). Thus, the generic circumscription 

of Massariosphaeria should be re-evaluated by further 

phylogenetic study with more relevant taxa included. 

Mauritiana Poonyth, K.D. Hyde, Aptroot & Peerally, 

Fungal Divers. 4: 102 (2000). (?Zopfiaceae) 


Habitat terrestrial, saprobic. Ascomata medium-sized, gregarious, 

ovoid, immersed, ostiolate, ostiole rounded. Peridium 

thin, thicker near the apex. Hamathecium of dense, cellular 

pseudoparaphyses, branching. Asci 8-spored, bitunicate, cylindrical 

to cylindro-clavate, with a short pedicel and a small 

ocular chamber. Ascospores 2-3-seriate, fusoid with rounded 

ends, dark brown with paler apical cells, multi-septate, 

distoseptate, slightly constricted at the primary septum. 


Literature: Hawksworth et al. 1995; Poonyth et al. 2000; 

Suetrong et al. 2009. 

Type species 

Mauritiana rhizophorae Poonyth, K.D. Hyde, Aptroot & 

Peerally, Fungal Divers. 4: 102 (2000). (Fig. 57) 


gregarious, ovoid, immersed, ostiolate, ostiole rounded 

(Fig. 57a). Peridium 40–60 μm thick laterally, thicker near 

the apex (Fig. 57a and b). Hamathecium of dense, long 

cellular pseudoparaphyses, 1.5–2 μm broad, branching. Asci 

130–180×20–25 μm (x ¼ 156 21:8mm, n=10), 8-spored, 

bitunicate, cylindrical to cylindro-clavate, with a short pedicel, 

with a small ocular chamber (Fig. 57c, d and e). Ascospores 

29–40×9–13 μm (x ¼ 35:4 11mm, n=10), 2-3-seriate, 

fusoid with rounded ends, dark brown with paler apical cells, 

9–13-distoseptate, slightly constricted at the primary septum, 

smooth (Fig. 57f,g,handi). 


Material examined: MAURITIUS, Grand Gaube, 

Melville mangrove, on dead decorticated Rhizophora 

mucronata Lam.woodstillattachedtolivingtree,Jan. 

1995, A.D. Poonyth (HKU(M)10219, holotype). 

Notes 

Morphology 

Mauritiana was introduced to accommodate the mangrove 

fungus, M. rhizophorae, which is characterized by 

immersed ostiolate, periphysate ascomata, thin peridium, 

bitunicate, 8-spored, cylindrical to cylindro-clavate asci, 

fusoid, smooth, hyaline to pale brown, multi-septate and 

distoseptate ascospores (Poonyth et al. 2000). But after 

carefully studying the type of M. rhizophorae, notypical 

distoseptate ascospores observed. The pigmented curved 

septum of the ascospore gives a “thickened” appearance. 

Based on its immersed ascomata, presence of cellular 

pseudoparaphyses, thick-walled, fissitunicate asci and 

brown, phragmosporous ascospores constricted at the primary 

septum, Mauritiana was assigned to the Pyrenulales sensu 

stricto (Melanommatales sensu lato, Dothideales sensu lato) 

(Hawksworth et al. 1995; Poonyth et al. 2000). 


Based on a multigene phylogenetic analysis, Mauritiana 

rhizophorae resided within a paraphyletic clade (Suetrong et 

al. 2009) sister to marine fungi Halotthia posidonia and 

Pontoporeia biturbinata. In this study, the dendrogram shows 

it to be closely related to the Sporormiaceae and Lophiostomataceae, 

which may indicate an uncircumscribed familial clade 

(Plate 1). Thus, its familial placement remains undetermined. 


The “thickened” septa of ascospores of Mauritiana 

rhizophorae is quite unique in Pleosporales, which makes 

it easily distinguishable from other genera.. 

Melanomma Nitschke ex Fuckel, Jb. nassau. Ver. Naturk. 

23–24: 159 (1870). (Melanommataceae) 


Habitat terrestrial, saprobic. Ascomata immersed, erumpent 

to nearly superficial, medium- to large-sized, globose to 

subglobose, coriaceous, gregarious, short papillate. Peridium 

pseudoparenchymatous cells outside with pale compressed 

cells inside. Asci cylindric to clavate with short 

pedicels. Hamathecium of dense, filamentous, branching, 

rarely anastomosing, septate pseudoparaphyses. Ascospores 

pale brown, reddish brown to olive-brown, ellipsoid to 

fusoid, 2 to multi-septate, constricted at the main septum.


Fig. 57 Mauritiana rhizophorae 

(from HKU(M)10219, 

holotype). a Vertical section of 

an ascoma. Note the thin layer of 

fungal tissue (pseudostroma?) on 

the host surface. b Section of a 

partial peridium. c Pseudoparaphyses 

and immature ascus. d 

Fissitunicate asci. e Asci showing 

thickening of the apical wall. 

f–i Ascospores with transverse 

septa and paler polar cells. Scale 

bars: a=40 μm, b, d–i=10 μm, 

c=20 μm 

Anamorphs reported for genus: Aposphaeria, Nigrolentilocus, 

Phoma-like and Pseudospiropes (Chesters 1938; 

Sivanesan 1984). 

Literature: Barr 1990a; Chesters 1938; Fuckel 1870; 

Saccardo 1878; Zhang et al. 2008a. 

Type species 

Melanomma pulvis-pyrius (Pers.) Fuckel, Jb. nassau. Ver. 

Naturk. 23–24: 160 (1870). (Fig. 58) 

≡ Sphaeria pulvis-pyrius Pers., Syn. meth. fung. 

(Göttingen) 1: 86 (1801). 


gregarious, substrate surface covered with a thin layer of 

brown psueodstroma, superficial, globose, subglobose, 

broadly or narrowly conical, often laterally flattened, black, 

roughened and irregular, often bearing remnants of wood 

fibers; apex short papillate, often somewhat puckered or 

sulcate (Fig. 58a). Peridium 70–90 μm wide, to 180 μm 

wide at the base, coriaceous, comprising two types of cells, 

outer cells small heavily pigmented thick-walled cells of 

textura angularis, apical cells smaller and walls thicker, 

individual cell walls to 6 μm thick, inner cells lightly 

pigmented to hyaline thin-walled cells of textura angularis,


5–8 μm diam., individual cell wall to 1.5–2 μm thick, in 

places with columns of textura prismatica, and larger, paler 

cells of textura prismatica towards the interior and at the 

base (Fig. 58b). Hamathecium of dense, filamentous, 1–2 

(−2.5) μm broad, branching, rarely anastomosing, septate 

pseudoparaphyses. Asci 98–123×6.5–7.5(−9) μm 

(x ¼ 109 7:5mm, n=10), 8-spored, bitunicate, fissitunicate, 

cylindrical to fusoid, with a short, furcate pedicel, to 25 μm 

long, with an ocular chamber (Fig. 58c, d, e, f and g). 

Ascospores 14–17.5(−19)×4.5–6.5 μm (x ¼ 15:8 5:2mm, 

n=10), obliquely uniseriate and partially overlapping, broadly 

fusoid to fusoid with broadly rounded ends, straight or slightly 

curved, smooth, olive-brown, 4-celled, slightly constricted at 

the septa, the second cell from the top slightly wider than the 

others, no sheath (Fig. 58h,i,j,kandl). 

Anamorph: Aposphaeria agminalis Sacc. or Phoma 

agminalis Sacc. (Sivanesan 1984). 

Colonies (of epitype) reaching 4 cm diam. after 20 days 

growth on PDA at 25°C, depressed to raised, cottony to woolly, 

with rhizoidal margin, grey, reverse darkened. Phoma-like 

anamorph has been reported by Chesters (1938) and 

Sivanesan (1984), but no anamorphic stage was observed 

in the cultures of IFRDCC 2044, CBS 109.77 and CBS 

371.75 after culturing 3 months on PDA. 

Material examined:ondecayingwood(UPS,Scler.suec. 

n. 120, holotype,asSphaeria pulvis-pyrius Pers.); FRANCE, 

Ariège, Rimont, Saurine, on bark of Salix caprea, 10Apr. 

2008, Jacques Fournier (IFRD 2001, epitype). 

Notes 

Morphology 

Melanomma, the familial type of Melanommataceae, was 

formally established by Fuckel (1870, p 159) based on its small, 

carbonaceous ascomata, having: “sporen meist 2–3 mal septirt, 

selten ohne Scheidewand, braun oder wasscrhell.” (Chesters 

1938; Fuckel 1870). Saccardo (1878, p. 344) emended this 

genus as “Spores ovate or oblong, multi-septate, coloured.” 

Subsequently, Saccardo (1883, p. 98) extended the description 

of Melanomma as “Perithecia gregarious, seldom scattered, 

somewhat superficial, sphaerical, papillate or blunt, carbonaceous, 

smooth or somewhat hairy. Asci elongate, for the most 

part accompanied by paraphyses, 8-spored. Spores oblong or 

somewhat spindle-shaped, two to many septate, olive or dark 

brown. Species of Sphaeria belong here for the most part.” 

Melanomma pulvis-pyrius was erected as the lectotype 

species (Barr 1990a; Chesters1938). Barr (1990a) gavea 

detailed circumscription for Melanomma, under which 

Melanomma contains about 20 species (Kirk et al. 2001). 

Melanomma pulvis-pyrius is characterized by its gregarious, 

superficial ascomata with short papillate, cylindrical asci with a 

short pedicel and fusoid, olive-brown, 3-septate ascospores 

(Chesters 1938; Zhang et al. 2008a). One of the diagnostic 

characters of Melanommataceae is the trabeculate pseudoparaphyses, 

although no typical trabeculate pseudoparaphyses could 

be found in the neotype (Scler. suec. n. 120, UPS) and epitype 

(IFRD 2001) of M. pulvis-pyrius (Zhang et al. 2008a). 


Phylogenetic analysis based on five genes (LSU, SSU, 

RPB1, RPB2 andEF1) indicates that Melanomma pulvispyrius 

forms a robust clade with Byssosphaeria, Herpotrichia 

and Pleomassaria siparia (Pleomassariaceae) and 

likely represents a separate family (or families comprising 

Melanommataceae) (Zhang et al. 2008a; Mugambi and 

Huhndorf 2009b). A more recent phylogenetic analysis 

included a group of coelomycete species with stellate 

conidia, isolated from Fagales trees clustered in Melanommataceae 

(Tanaka et al. 2010; Plate 1). 


The Melanomma concept based on ascospore morphology 

appears polyphyletic. 

Metameris Theiss. & Syd., Annls mycol. 13: 342 (1915). 



Habitat terrestrial, saprobic or parasitic. Ascostromata erumpent 

through the host surface in linear rows parallel to the host fibers. 

Ascomata small, globose to subglobose, black, coriaceous. 

Peridium composed of large lightly pigmented cells of textura 

angularis. Hamathecium of rare, broad pseudoparaphyses, 

septate, constricted at the septa. Asci bitunicate, fissitunicate, 

broadly cylindrical to slightly obclavate, with a short, thick, 

knob-like pedicel. Ascospores hyaline, 1- (rarely 2-) septate. 


Literature: von Arx and Müller 1975; Barr1972; Clements 

and Shear 1931; Eriksson 2006; Lumbsch and Huhndorf 

2007; TheissenandSydow1915. 

Type species 

Metameris japonica (Syd.) Syd., Annls mycol., 13(3–4): 

342 (1915). (Fig. 59) 

≡ Monographus japonicus Syd. Annls mycol. 10: 408 

(1912). 

Ascostromata erumpent through the host surface in linear 

rows parallel to the host fibers, 500–750 μm long and 140– 

200 μm wide, with three to ten ascomata arranged in a line 

(Fig. 59a). Ascomata 115–160 μm diam., semi-immersed in 

substrate to erumpent, globose, subglobose, black, coriaceous 

(Fig. 59b). Cells of ascostromata heavily pigmented and 

thick-walled, cells of peridium composed of large lightly 

pigmented cells of textura angularis, cells 5–15 μm diam.,


Fig. 58 Melanomma pulvis-pyrius (a–b, d–e, h–j from UPS, holotype; c, g, k, l from epitype). a Ascomata gregarious on the host surface. b 

Vertical section of an ascoma. c–f Asci with pedicels. g Dehiscent ascus. h–l Ascospores. Scale bars: a=0.5 mm, b=200 μm, c–l=10 μm


cell wall



None. 


Its small-sized ascomata, broadly cylindrical to slightly 

obclavate asci with a short, thick, knob-like pedicel, as well 

as its monocotyledonous host preference point Metameris 

to the Phaeosphaeriaceae. But DNA comparisons are 

needed for confirmation. 

Mixtura O.E. Erikss. & J.Z. Yue, Mycotaxon 38: 203 

(1990). (Phaeosphaeriaceae) 


Habitat terrestrial, parasitic. Ascomata small-sized, scattered 

or clustered on the leaf spots, immersed, erumpent, 

minutely papillate, ostiolate. Papilla slightly raised. Peridium 

thin, comprising one cell type of lightly pigmented 

thin-walled cells of textura angularis. Hamathecium of 

dense, filliform, septate, cellular pseudoparaphyses, 4– 

6.3 μm broad, embedded in mucilage. Asci bitunicate, 

ovoid, with a very short stumpy pedicel. Ascospores fusoid 

to narrowly fusoid with broadly to narrowly rounded ends, 

curved, dark brown, multi-septate, distoseptate, with a germ 

pore at the lower end. 


Literature: Eriksson and Yue 1990. 

Type species 

Mixtura saginata (Syd.) O.E. Erikss. & J.Z. Yue, Mycotaxon 

38: 203 (1990). (Fig. 60) 

≡ Leptosphaeria saginata Syd., Annls mycol. 37: 376 (1939). 

Producing elongated yellow spots with brownish margins, 

leaf spots up to 45×3–5 mm, opposite side visible as a 

brownish spots (Fig. 60a). Ascomata 170–200 μm high×210– 

280 μm diam., scattered on the lower side of the leaf, 

immersed, erumpent, breaking through the epidermis, minutely 

papillate. Papilla central, slightly raised, ostiolate, 

ostiole surrounded by a white margin (Fig. 60b). Peridium 

22–34 μm wide, thicker at the apex, thinner at the base, 

comprising one cell type of lightly pigmented thin-walled 

cells of textura angularis, cells up to 6×8 μm diam., cell wall 

0.5–1.2 μm thick, apex cells smaller and walls thicker 

(Fig. 60c). Hamathecium of dense, filliform, septate, cellular 

pseudoparaphyses, 4–6.3 μm broad, embedded in mucilage. 

Asci 80–128×41–53(−69) μm (x ¼ 100:9 52:8mm, n =10), 

8-spored, bitunicate, fissitunicate dehiscence not observed, 

sac-like, with a very short stumpy pedicel and a small ocular 

chamber (Fig. 60d). Ascospores 86–94(−106)×20.5–23.5 μm 

(x ¼ 92:7 21:7mm, n=10), fasciculate, fusoid to narrowly 

fusoid, slightly curved, dark brown, 7-septate, distoseptate, 

with or without constriction at the primary septum, smoothwalled, 

with a germ pore at the lower end (Fig. 60e and f). 


Material examined: ECUADOR, Tungurahua, Hacienda 

San Antonio pr. Baños, Province, on the leaves of Chusqueae 

serrulatae Pilger., 9 Jan. 1938, H. Sydow. (S reg. nr F8934 

type, F8935isolectotype, asLeptosphaeria saginata). 

Notes 

Morphology 

Mixtura was formally established by Eriksson and 

Yue (1990) as a monotypic genus represented by M. 

saginata basedonitsimmersedandthin-walledascomata, 

sparse, broad pseudoparaphyses, sac-like asci with a short 

pedicel and thick apex. Mixtura has a “mixture” of 

characters found in other pleosporalean genera. The 

peridium structure is comparable with Phaeosphaeria, 

the ascospores with Trematosphaeria and asci with 

Wettsteinina (Eriksson and Yue 1990).Accordingtothe 

structure of ascomata and hamathecium, Mixtura was 

provisionally assigned to Phaeosphaeriaceae (Eriksson 

and Yue 1990). 


None. 


Morphologically, the sparse broad pseudoparaphyses and 

sac-like asci with a thick apical structure in Mixtura seem 

more comparable with the generic type of Teratosphaeria 

(T. fibrillose Syd. & P. Syd., Teratosphaeriaceae, Capnodiales, 

Dothideomycetidae) than that of Phaeosphaeria (P. 

oryzae). The heavily pigmented, multi-septate ascospores 

and the persistent pseudoparaphyses of Mixtura however, 

differ from those of Teratosphaeria. Thus, here we assign 

Mixtura under Teratosphaeriaceae as a distinct genus until 

phylogenetic work is carried out. 

Montagnula Berl., Icon. fung. (Abellini) 2: 68 (1896). 

(Montagnulaceae) 



immersed to erumpent, gregarious or grouped, 

globose to subglobose, black. Hamathecium of dense, 

narrowly cellular, septate pseudoparaphyses. Asci bitunicate, 

fissitunicate, usually cylindro-clavate to clavate with a 

long pedicel. Ascospores oblong to narrowly oblong, 

straight or somewhat curved, reddish brown to dark 

yellowish brown, muriform or phragmosporous. 

Anamorphs reported for genus: Aschersonia (Hyde et al. 2011).


Fig. 60 Mixtura saginata 

(from S reg. nr F8934, type). a, 

b Leaf spots in leaves of Chusquea 

serrulatae. Note the erumpent 

ascomata surrounded by 

white material in (b). c Section 

of an ascoma. Note the peridium 

structure which comprises cells 

of textura angularis. The arrangement 

of the asci and pseudoparaphyses 

can also be seen. 

d Immature asci in pseudoparaphyses. 

Note the stumpy pedicel 

and thickened apex with 

flattened ocular chamber. e, f 

Mature ascospores. Note the 

hyaline ends and distosepta. 

Scale bars: a=10 mm, b, c= 

100 μm, d=50 μm, e–f=20 μm 

Literature: Aptroot 1995; Barr2001; Berlese1896; Clements 

and Shear 1931; Crivelli1983; Leuchtmann 1984; Ramaley 

and Barr 1995; Schoch et al. 2006; Wehmeyer1957, 1961; 


Type species 

Montagnula infernalis (Niessl) Berl., Icon. fung. (Abellini). 

2: 68 (1896). (Fig. 61) 

≡ Leptosphaeria infernalis Niessl, Inst. Coimbra 31: 13 

(1883). 

Ascomata 220–280 μm high×250–310 μm diam., immersed 

to erumpent, gregarious or clustered, globose to 

subglobose, sometimes triangular in dried material, short 

ostiole always filled with hyaline closely adhering cells, black 

(Fig. 61a and b). Peridium 40–55 μm thick at sides, up to 

80 μm thick near the apex, 3-layered, outer layer composed of 

heavily pigmented thick-walled small cells of textura angularis, 

cells3–8 μm diam., wall 1.5–3 μm thick, apex thicker 

with smaller cells and thicker cell wall, thinner near the base; 

mid layer less pigmented, cells 4–13 μm diam.; innermost 

layer of narrow compressed rows of cells, merging with


pseudoparaphyses (Fig. 61c). Hamathecium of dense, narrow 

cellular pseudoparaphyses, 2–4.5 μm broad, septate (Fig. 61f). 

Asci 153–170(−200)×17.5–21.5 μm (including pedicel), 

bitunicate, fissitunicate, cylindro-clavate to clavate, pedicel 

28–60(−85) μm long, 8-spored, biseriate, with an ocular 

chamber best seen in immature ascus (to 3 μm wide×3 μm 

high) (Fig. 61d and e). Ascospores 24–29×9–11 μm, oblong to 

narrowly oblong, straight or somewhat curved, reddish brown 

to dark yellowish brown, verruculose, with five transverse 

septa and one vertical septum in each middle cells, constricted 

at the primary and secondary primary septa (Fig. 61g). 


Material examined: PORTUGAL, Coimbra Lusitania, 

on leaves of Fourcroya longava pr., Feb., 1881, leg. Moller. 

(M 1183, holotype). 

Notes 

Morphology 

Montagnula was introduced to accommodate two Pleospora 

species, i.e. P. infernalis (Niessl) Wehm. and P. 

gigantea Mont. by Berlese (1896), based on the presence of 

hyphal stromatic tissues over the ascomata and asci with 

relatively long pedicels (Barr 2001). Montagnula infernalis 

was selected as the lectotype species (Clements and Shear 

1931). Subsequently, Wehmeyer (1957, 1961) treated Montagnula 

as a subgenus of Pleospora. Crivelli(1983) accepted 

Montagnula as a separate genus, and divided it into two 

subgenera, i.e. Montagnula and Rubiginospora. Montagnula 

was characterized by having dark brown ascospores and 

exclusively occurring on Agavaceae, while Rubiginospora 

has reddish brown ascospores and occurs on Poaceae. This 

proposal was not accepted by many workers (Barr 2001). 

Subsequently, more species with various ascospores (such as 

phragmosporous species by Leuchtmann (1984) and didymosporous 

species by Aptroot (1995) were added in this 

genus), which has obviously become heterogenic. Barr 

(2001) assigned species of Montagnula into different genera, 

i.e. Kalmusia and Didymosphaerella, respectively and introduced 

Montagnulaceae to accommodate all of these genera. 


Montagnula opulenta forms a robust phylogenetic clade 

with species of Bimuria, Curreya, Didymocrea, Letendraea, 

Paraphaeosphaeria, Phaeodothis and Karstenula, 

which might represent a familial group (Schoch et al. 2006; 

Zhang et al. 2009a). A more convincing conclusion can 

only be obtained following sequence data from more 

verified fungi being added to the phylogenetic tree. 


One striking character of Montagnula infernalis is the very 

long ascal pedicel once it is released from the ascomata. 

However, this character appears to have evolved more than 

once and can be found in Kirschsteiniothelia elaterascus 

Shearer which clusters with Helicascus (Shearer et al. 2009). 

The same ascus character is also found in Xenolophium and 

Ostropella in the Platystomaceae (Mugambi and Huhndorf 

2009b). Montagnula opulenta is a didymosporous species, 

but phylogenetically closely related to those dictyosporous 

(Karstenula rhodostoma) and phragmosporous (Paraphaeosphaeria 

michotii) members of Montagnulaceae (Zhang et al. 

2009a). This might indicate that compared to other morphological 

characters, ascospore type is not a valid character at 

family level classification. 

Moristroma A.I. Romero & Samuels, Sydowia 43: 246 




scattered, or in small groups, superficial, cushion-like, circular 

in outline, wall black, roughened, containing numerous 

locules. Peridium thin, 1-layered. Hamathecium of dense, long 

filliform pseudoparaphyses, 2–3 μm broad, septate, branching. 

Asci polysporous, with a short, laterally displaced, sometimes 

papillate knob-shaped pedicel, apex very thick walled, 

bitunicate, fissitunicate, obclavate, ocular chamber not observed. 

Polyspores oblong to cylindrical, hyaline, non-septate. 


Literature: Eriksson 2006; Romero and Samuels 1991. 

Type species 

Moristroma polysporum A.I. Romero & Samuels, Sydowia 

43: 246 (1991). (Fig. 62) 


scattered, or in small groups of 2–3, superficial, with basal 

wall remaining immersed in host tissue, cushion-like, 

circular in outline, wall black, roughened, containing 

numerous locules, each locule 120–240 μm diam., ostiolate 

(Fig. 62a and b). Peridium 14–30 μm thick, 1-layered, 

composed of small heavily pigmented thick-walled cells of 

textura angularis, cells2–4 μm diam., cell wall 1.5–3 μm 

thick, peridium between the locules hyaline (Fig. 62b and c). 

Hamathecium of dense, long filliform pseudoparaphyses, 2– 

3 μm broad, septate, branching. Asci 44–60×12–14 μm 

(x ¼ 54:3 13mm, n=10), polysporous, with a short, papillate 

knob-shaped pedicel, apex very thick-walled, bitunicate, 

fissitunicate, obclavate, ocular chamber not observed 

(Fig. 62d and e). Polyspores 3–4(−5)×0.6–1.2 μm, oblong 

to cylindrical, hyaline, non-septate, smooth (Fig. 62f). 


Material examined: ARGENTINA, Buenos Aires, 

Ramallo, on Eucalyptus viminalis Labill., May 1982, Romero


Fig. 61 Montagnula infernalis 

(from M 1183, holotype). a 

Appearance of ascomata immersed 

in host tissue. b Section 

of an immersed ascoma. Note 

the hyaline closely adhering 

cells in the ostiole region. c 

Section of the peridium comprising 

a few layers of cells. d 

An immature ascus with a long 

pedicel. e, g Mature muriform 

ascospores in asci. f Cellular 

pseudoparaphyses. Scale bars: 

a=0.5 mm, b, c=100 μm, 

d–g=20 μm 

27/4-13 (BAFC 32036, holotype); Nov. 1982, on decorticated 

wood, Romero 35/4-13 (BAFC 32037, paratype). 

Notes 

Morphology 

Moristroma was formally established by Romero and 

Samuels (1991) based on its “cushion-shaped ascomata 

containing lots of locules with numerous asci inside, asci 

obclavate, polysporous, with a knob-shaped pedicel”. The 

bitunicate asci and numerous cellular pseudoparaphyses 

undoubtedly point it to Pleosporales, while the familial 

placement of Moristroma is uncertain, and it was 

temporarily assigned to Dacampiaceae by Romero and 

Samuels (1991), but no 3-layered peridium is found. 

Eriksson (2006) assignedittoTeichosporaceae. 


None. 


The familial status of Moristroma cannot be determined yet.


Fig. 62 Moristroma polysporum 

(from BAFC 32036, holotype). 

a Two multiculate 

ascostroma on the host surface. 

b Section of an ascostroma. 

Note the multilocula. c Section 

of the peridium. Note the thick 

walled cells. d, e Broadly cylindrical 

to fusoid asci containing 

numerous part spores. f Released 

part spores. Scale bars: 

a=0.5 mm, b=200 μm, c= 

50 μm, d–f=10 μm 

Morosphaeria Suetrong, Sakay., E.B.G. Jones & C.L. 

Schoch, Stud. Mycol. 64: 161 (2009). (Morosphaeriaceae) 


Habitat marine, saprobic. Ascomata large, solitary or 

gregarious, immersed to erumpent, subglobose or depressed 

with a flatted base, ostiolate, papillate, brown to black, 

coriaceous. Peridium thick. Hamathecium of dense, long 

cellular pseudoparaphyses, septate. Asci 8-spored, bitunicate, 

cylindrical, with short pedicels. Ascospores uniseriate 

to partially overlapping, ellipsoidal, hyaline, 1-3-septate, 

constricted at the septa, central cells larger, apical cells if 

present small and elongated, surrounded with mucilaginous 

sheath. 


Literature: Hyde and Borse 1986; Hyde 1991a, b; 

Suetrong et al. 2009; Zhang et al. 2009a. 

Type species 

Morosphaeria velataspora (K.D. Hyde & Borse) Suetrong, 

Sakay., E.B.G. Jones & C.L. Schoch, Stud. Mycol. 64: 161 

(2009). (Fig. 63) 

≡ Massarina velataspora K.D. Hyde & Borse, Mycotaxon 

27: 163 (1986). 

Ascomata 0.7–1.2 mm diam., solitary or gregarious, 

immersed to erumpent, subglobose or depressed, with a 

flattened base not easily removed from the substrate, ostiolate, 

epapillate or papillate, brown to black, coriaceous (Fig. 63a). 

Peridium thick, the upper part of the peridium composed of 

brown thick-walled cells of textura angularis, cells are 

smaller and wall thicker near the apex, at the rim is 

composed of vertical, parallel, brown, elongate cells, 

wedge-shape in section (Fig. 63a). Hamathecium of dense, 

long cellular pseudoparaphyses, 1.1–1.7 μm broad, septate. 

Asci 220–320×23–34 μm (x ¼ 251 28:2mm, n=10), 8-


spored, bitunicate, cylindrical, with short pedicels (Fig. 63b). 


uniseriate to partially overlapping, ellipsoidal, hyaline, 1-3- 

septate, constricted at the septa, central cells larger, apical 

cells if present small and elongated, surrounded with 

mucilaginous sheath, 5–22 μm wide (Fig. 63c, d and e). 


Material examined: Jan. 1984, Herb. IMI 297770, 

slides 1–10 (holotype) and dried wood (isotype). 

Notes 

Morphology 

Two Massarina sensu lato species described from the 

marine environment, viz. M. ramunculicola (Sacc.) O.E. 

Erikss. & J.Z. Yue and M. velataspora K.D. Hyde & Borse, 

form a robust clade, and a new genus Morosphaeria was 

established for them (Suetrong et al. 2009). Together with 

two Helicascus species, they belong to Morosphaeriaceae 

(another marine family) (Suetrong et al. 2009). Morphologically, 

Morosphaeria is characterized by solitary to 

gregarious, subglobose to lenticular, immersed to superficial 

ascomata which are ostiolate and papillate, numerous, 

filliform pseudoparaphyses, 8-spored, clavate to cylindrical, 

bitunicate, fissitunicate asci, and hyaline, 1-3-septate, 

fusoid to ellipsoidal ascospores which are surrounded with 

mucilaginous sheath. 


Species of Morosphaeria form a sister group with 

Helicascus and both of these genera were assigned to a 

new family, i.e. Morosphaeriaceae (Suetrong et al. 2009). 

In this study, a strain of Asteromassaria pulchra, occuring 

on dead twigs of Prunus spinosa, is basal to other species 

of Morosphaeriaceae, and gets well support. Thus here we 

tentatively assign Asteromassaria in Morosphaeriaceae. 

Fig. 63 Morosphaeria velataspora (from IMI 297770, type). a Section of an ascoma. b Cylindrical asci embedded in pseudoparaphyses. c–e 

Hyaline, 1-3-septate, ascospores with mucilaginous sheath. Scale bars: a=100 μm, b=50 μm, c–e=20 μm



The only morphological difference between M. velataspora 

and M. ramunculicola are their morphology of 

ascomata and size of ascospores (Hyde 1991b). But M. 

velataspora was reported staining the woody substrate (or 

agar in culture) purple (Hyde and Borse 1986; Hyde 

1991b). Although this character could not be verified in 

the strain used by Suetrong et al. (2009), purple staining has 

been reported to have phylogenetic significance at familial 

rank in freshwater fungi (Zhang et al. 2009a). 

Murispora Yin. Zhang, C.L. Schoch, J. Fourn., Crous & K. 

D. Hyde, Stud. Mycol. 64: 95 (2009b). (Amniculicolaceae) 


Habitat freshwater, saprobic. Ascomata medium-sized, 

scattered to gregarious, immersed, lenticular, apex slightly 

protruding, opening through a small rounded pore, 

substrate stained purple. Peridium thin, composed of a 

few layers cells of textura angularis, thicker at the apex 

with pseudoparenchymatous cells. Hamathecium of narrowly 

cellular pseudoparaphyses, embedded in mucilage. 

Asci 8-spored, bitunicate, fissitunicate, biseriate, cylindroclavate 

with short pedicels. Ascospores curved- fusoid 

with narrowly rounded ends, golden yellow turning brown 

when senescent, multi-septate, constricted at the septa, 

with one, rarely two longitudinal septa in all cells except 

end cells, smooth or finely verruculose, surrounded by a 

wide mucilaginous sheath. 

Anamorphs reported for genus: Phoma (Webster 1957). 

Literature: Zhang et al. 2009a, b. 

Type species 

Murispora rubicunda (Niessl) Yin. Zhang, J. Fourn. & K. 

D. Hyde, Stud. Mycol. 64: 96 (2009a). (Fig. 64) 

≡ Pleospora rubicunda Niessl, Notiz. Pyr.: 31 (1876). 


scattered to gregarious, immersed, lenticular, apex 

laterally flattened, black, slightly protruding, opening 

through a small rounded pore, substrate stained purple 

(Fig. 64a). Peridium 15–18 μm thick at sides, composed 

of 3–4 layers cells of textura angularis, upto28–30 μm 

thick at the apex with very thick-walled cells, pseudoparenchymatous, 

nearly absent at the base (Fig. 64b). 

Hamathecium of narrowly cellular pseudoparaphyses, 

1–1.7 μm broad, embedded in mucilage. Asci 124– 

142×19–21 μm, 8-spored, bitunicate, fissitunicate, 

biseriate, cylindro-clavate with a small ocular chamber, 

with short pedicels (Fig. 64c). Ascospores 30–38×10– 

12 μm, curved-fusoid with narrowly rounded ends, 

golden yellow turning brown when senescent, 7–9 transversally 

septate, constricted at the septa, with one, rarely 

two longitudinal septa in all cells except end cells which 

are often slightly paler, all cells filled with a large 

refractive guttule, smooth to finely verruculose, surrounded 

by a wide mucilaginous sheath (Fig. 64d). 

Anamorph: Phoma sp. (Webster 1957). 

Material examined: FRANCE, Haute Garonne, 

Avignonet, Lac de Rosel, 16 Jan. 2007, on submerged 

dead herbaceous stem (Dipsacus?), leg. Michel Delpont, 

det. Jacques Fournier (IFRD 2017). 

Notes 

Morphology 

Murispora was introduced based on Pleospora rubicunda 

which is characterized by immersed, erumpent or 

nearly superficial, globose to subglobose, elongated weakly 

papillate ascomata which stain the woody substrate purple, 

trabeculate pseudoparaphyses, 8-spored, bitunicate, fissitunicate, 

oblong to clavate asci, fusoid, pale or reddish 

brown, muriform ascospores (Zhang et al. 2009a). A 

phylogenetic study indicated that Murispora forms a robust 

clade with species of Amniculicola, and Amniculicolaceae 

was introduced to accommodate them (Zhang et al. 2009a). 


Murispora rubicunda forms a robust clade with species of 

Amniculicola and Neophaeosphaeria (Zhang et al. 2009a). 


As has mentioned by Eriksson (1981, P. 135), the purplestaining 

species of Pleospora, treated by Webster (1957), 

should not belong to the Pleosporaceae. Both Pleospora 

straminis and P. rubelloides should be closely related to 

Murispora. 

Neomassariosphaeria Yin. Zhang, J. Fourn. & K.D. Hyde, 

Stud. Mycol. 64: 96 (2009a). (Amniculicolaceae) 


Habitat freshwater, saprobic. Ascomata medium-sized, 

scattered or in small groups, immersed, with a slightly 

protruding elongated papilla, ostiolate, lenticular, stain the 

substrate purple. Peridium thin. Hamathecium of dense, 

long cellular pseudoparaphyses, septate. Asci 8-spored, 

bitunicate, fissitunicate, cylindro-clavate, with short furcate 

pedicels. Ascospores 2-3-seriate, narrowly fusoid, somewhat 

curved, reddish brown, multi-septate, slightly constricted 

at the primary septum. 


Literature: Leuchtmann 1984; Zhang et al. 2009a, b.


Fig. 64 Murispora rubicunda 

(from IFRD 2017). a Habitat 

section of the immersed ascomata. 

b Section of an ascoma. 

Note the thin peridium and cells 

of textura angularis. c Mature 

and immature asci. d Muriform 

ascospores. Scale bars: a, b= 

100 μm, c, d=20 μm 

Type species 

Neomassariosphaeria typhicola (P. Karst.) Yin. Zhang, J. 

Fourn. & K.D. Hyde, Stud. Mycol. 64: 96 (2009a). (Fig. 65) 

≡ Leptosphaeria typhicola P. Karst., Bidr. Känn. Finl. 

Nat. Folk 23: 100 (1873). 


scattered or in small groups, immersed, lenticular, with a 

slightly protruding elongated papilla, ostiolate, stain the 

substrate purple (Fig. 65a). Peridium 15–30 μm thick. 


1.5–2.5 μm thick, septate. Asci 110–160×13–15 μm, 8- 

spored, bitunicate, fissitunicate, cylindro-clavate, with short 

furcate pedicels (Fig. 65b, c and d). Ascospores 30–48×7– 

11 μm, 2-3-seriate, narrowly fusoid, somewhat curved, 

reddish brown, 7-septate, slightly constricted at the primary 

septum, verruculose (Fig. 65c and d). 


Material examined: DENMARK, Sjaeland, Frederikskilde, 

Suserup Skove, Tystrup Lake, 25 May 2007, on 

submerged culm of Phragmites, leg. & det. Jacques Fournier 

(IFRD 2018). 

Notes 

Morphology 

Neomassariosphaeria is most comparable with Murispora, 

and is distinguished from Murispora by its phragmosporous 

ascospores. Both genera were assigned to 

Amniculicolaceae (Zhang et al. 2009a). 


Both Neomassariosphaeria grandispora and N. typhicola 

clustered with species of Murispora and Amniculicola in 

Amniculicolaceae (Zhang et al. 2009a,c). 


Similar with those purple-staining species of 

Pleospora assigned to Murispora, the purple-staining 

species of Phaeosphaeria mentioned by Crivelli 

(1983) and Leuchtmann (1984) might be assigned to 

Neomassariosphaeria. 

Neophaeosphaeria M.P.S. Câmara, M.E. Palm & A.W. 

Ramaley, Mycol. Res. 107: 519 (2003). (Leptosphaeriaceae)


Fig. 65 Neomassariosphaeria 

typhicola (from IFRD 2018). a 

Immersed ascomata gregarious 

in the host substrate. b–d 

Cylindro-clavate asci embedded 

in pseudoparaphyses. Note the 

phragmosporous ascospores. 

Scale bars: a=200 μm, 

b–d=20 μm 


Habitat terrestrial, parasitic or saprobic. Ascomata small, 

forming in leaf spots, scattered or clustered, immersed, 

depressed globose, under clypeus, coriaceous. Peridium thin. 

Hamathecium of dense, cellular pseudoparaphyses, septate, 

embedded in mucilage. Asci 8-spored, bitunicate, fissitunicate 

dehiscence not observed, broadly cylindrical to oblong, 

with a short furcate pedicel. Ascospores obliquely uniseriate 

and partially overlapping, oblong, pale brown, 1-3-septate. 

Anamorphs reported for genus: Coniothyrium-like 

(Câmara et al. 2003). 

Literature: Câmara et al. 2001, 2003; Checa et al. 2002; 

Ellis and Everhart 1892.


Type species 

Neophaeosphaeria filamentosa (Ellis & Everh.) M.P.S. 

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

519 (2003). (Fig. 66) 

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

64 (1888). 

Ascomata 115–157 μm high×115–186 μm diam., forming 

in leaf spots, scattered or clustered in circular areas, immersed, 

depressed globose, with a small ostiolar pore slightly 

penetrating above the surface, under clypeus, coriaceous, 

papilla not conspicuous (Fig. 66a). Peridium 18–30 μm thick, 

composed of large pigmented thin-walled cells of textura 

angularis, cells up to 10 μm diam. (Fig. 66c). Hamathecium 

of dense, cellular pseudoparaphyses 1.5–2.5 μm broad, 

septate, embedded in mucilage (Fig. 66b). Asci 70–105×8– 


fissitunicate dehiscence not observed, broadly cylindrical to 

oblong, with a short, broad, furcate pedicel, 6–13 μm long, 

with a small ocular chamber, best seen in immature asci, up to 

1.5 μm wide×1 μm high (Fig. 66d, e and f). Ascospores 12– 

15×4–5 μm (x ¼ 13:8 5mm, n=10), obliquely uniseriate 

and partially overlapping, oblong, yellowish brown, (1-2-)3- 

septate, constricted at the primary septum, the upper second 

cell often broader than others, verruculose, containing four 

refractive globules (Fig. 66g). 

Anamorph: Ellis and Everhart (1892) noted that the 

“spermogonial stage is a Coniothyrium (C. concentricum) 

with small (4 μm), globose, brown sporidia.” 


dead parts in living leaves of Yucca filamentosa L., Jul. 

1888, Ellis & Everhart (NY, holotype). 

Notes 

Morphology 

Neophaeosphaeria was formally established by 

Câmara et al. (2003) by segregating Paraphaeosphaeria 

species with 3-4-septate ascospores and anamorphs of 

ovoid to ellipsoid, non-septate, brown, verrucose to 

punctuate conidia forming from percurrently proliferating 

conidiogenous cells. Neophaeosphaeria filamentosa was 

selected as the generic type. Currently, four species are 

included under Neophaeosphaeria, i.e. N. barrii, N. 

conglomerate (M.E. Barr) M.P.S. Câmara, M.E. Palm & 

A.W. Ramaley, N. filamentosa and N. quadriseptata (M. 

E. Barr) M.P.S. Câmara, M.E. Palm & A.W. Ramaley 

(Câmara et al. 2003). At present all species in Neophaeosphaeria 

occur on Yucca (Agavaceae). 


The four Neophaeosphaeria species form a monophyletic 

clade based on both ITS and SSU rDNA 

sequences (Câmara et al. 2001; Checa et al. 2002), and 

they fall in the group comprising members of Phaeosphaeriaceae 

and Leptosphaeriaceae (Câmara et al. 

2003). Neophaeosphaeria filamentosa, the generic type 

of Neophaeosphaeria, nestedinLeptosphaeriaceae with 

low to moderate bootstrap values (Schoch et al. 2009; 

Zhang et al. 2009a). 


The familial status of Neophaeosphaeria under Leptosphaeriaceae 

is confirmed, although this family remains 

poorly supported in phylogenetic studies. 

Nodulosphaeria Rabenh., Klotzschii Herb. Viv. Mycol., 

Edn 2: no. 725 (in sched.) (1858). (Phaeosphaeriaceae) 


Habitat terrestrial, saprobic or hemibiotrophic. Ascomata 

small, immersed to erumpent, globose or subglobose, 

black, papillate, ostiolate. Papilla with numerous setae in 

the pore-like ostiole. Peridium thin, composed of thickor 

thin-walled large cells. Hamathecium of cellular 

pseudoparaphyses, septate and branching. Asci 8- 

spored, bitunicate, fissitunicate, clavate to cylindroclavate, 

with a very short, furcate pedicel and a small 

ocular chamber. Ascospores filamentous, hyaline or pale 

brown, multi-septate, one of the upper cells swollen. 


Literature: Barr 1992a; Holm 1957, 1961; Shoemaker 

1984b; Shoemaker and Babcock 1987. 

Type species 

Nodulosphaeria hirta Rabenh., Klotzschii Herb. Viv. 

Mycol., Edn 2: no. 725 (in sched.) (1858). (Fig. 67) 


scattered, or in small groups, immersed to erumpent, 

globose or subglobose, black, papillate, ostiolate. Papilla 

50–80 μm high, numerous setae occur in the pore-like 

ostiole (Fig. 67a and b). Peridium 15–30 μm wide at the 

sides, thinner at the base, coriaceous, comprising two types 

of cells, outer cells of 1–2 layers of heavily pigmented cells 

of textura angularis, cells 6–8 μm diam., cell wall 1.5– 

3 μm thick, inner of compressed cells, 5×13–3×8 μm 

diam., wall 2–3 μm thick (Fig. 67c). Hamathecium of long 

cellular pseudoparaphyses 2–3 μm broad, septate and 

branching, mucilage not observed. Asci 100–123×12.5–15 

(−17.5) μm (x ¼ 110:8 14:3mm, n=10), 8-spored, bitunicate, 

fissitunicate, clavate to cylindro-clavate, with a very 

short, furcate pedicel, with a small ocular chamber (to 2 μm 

wide×1 μm high) (Fig. 67d). Ascospores 48–63×5–6.5 μm 

(x ¼ 55:3 5:6mm, n=10), 4-seriate, filamentous, pale


Fig. 66 Neophaeosphaeria filamentosa (from NY, holotype). a 

Ascomata as a circular cluster on the host surface. b Hamathecium of 

wide psuedoparaphyses. c Section of peridium comprising cells of textura 

angularis. d–f Cylindrical asci with thickened apex. Note the short furcate 

pedicel. g Pale brown, 3-septate ascospores. Note the verruculose 

ornamentation. Scale bars: a=200 μm, b, c=20 μm, d–g=10 μm


brown, 8-septate, the 4th upper cell broader than the others, 

smooth-walled, without sheath (Fig. 67e and f). 


Material examined: GERMANY, Dresdae, in herbarum 

caulibus emortuis perrara, exeunte majo, 1858 (BR 

101945–95, holotype, asNodulosphaeria hirta). 

Notes 

Morphology 

The name Nodulosphaeria was first used by Rabenhorst 

(1858) but was considered as a synonym of Leptosphaeria 

for many years (Clements and Shear 1931). The name was 

reinstated by Holm (1957) and was represented by N. hirta, 

which was concurrently treated as a synonym of N. derasa 

(Berk. & Broome) L. Holm. The most outstanding morphological 

characters of Nodulosphaeria were considered to be 

apex of ascomata often covered with setae, ascospore with 

three or more transverse septa with a supramedian enlarged 

cell or elongated to a scolecospore, mostly with terminal 

appendages (Barr 1992a; Holm 1961; Shoemaker 1984b). 

The ascomata are usually immersed and the peridium 

comprises a few layers of brown, relatively thin-walled cells 

of textura angularis and textura prismatica similar to those 

of Phaeosphaeria. Thus, Nodulosphaeria is likely to be a 

member of Phaeosphaeriaceae. However,thisneedstobe 

confirmed by molecular analysis. The boundary between 

Nodulosphaeria and Ophiobolus is not clear-cut, and the 

circumscriptions of them usually depend on the viewpoint of 

different mycologists. For instance, Shoemaker (1976) has 

assigned some Nodulosphaeria species such as N. erythrospora, 

N. fruticum, N. mathieui and N. megalosporus to 

Ophiobolus. Subsequently, more species were added to 

Nodulosphaeria (Barr 1992a; Shoemaker 1984b; Shoemaker 

and Babcock 1987). Currently, more than 60 names are 

included in Nodulosphaeria (http://www.mycobank.org/, 06/ 

2010). 


None. 


All species included in Nodulosphaeria have an inflated 

ascospore cell as mentioned above. However, it is likely 

that this character would have evolved more than once as it 

is probably an adaption for ascospore ejection from the 

ascus (Shoemaker 1976). It occurs in Ophiobolus species 

and the ascomata of these species are quite dissimilar to 

Nodulosphaeria species and their exclusion from Nodulosphaeria 

seems warranted. When considering whether a 

species belongs in Nodulosphaeria, one must also consider 

the ascomata and peridium structure until DNA sequences 

are available. 

Ohleria Fuckel, Fungi rhenani exsic.: no. 2173 (1868). 



Habitat terrestrial, saprobic. Ascomata small to medium size, 

solitary, scattered, or in small groups, erumpent to nearly 

superficial, papillate, ostiolate. Peridium thin, thicker at the 

apex, 1-layered. Hamathecium of dense, long trabeculate 

pseudoparaphyses. Asci 8-spored, bitunicate, fissitunicate, 

cylindrical, with a short pedicel. Ascospore brown to reddish 

brown, broadly to narrowly fusoid, 3-septate, easily separating 

into two parts at the primary septum. 

Anamorphs reported for genus: Monodictys (Samuels 1980). 

Literature: Barr 1990b; Clements and Shear 1931; Patel et 

al. 1997; Samuels 1980. 

Type species 

Ohleria modesta Fuckel, Fungi rhenani exsic. (1868) (Fig.68) 


solitary, scattered, or in small groups of 2–3, erumpent to 

nearly superficial, coriaceous, with basal wall remaining 

immersed in host tissue, broadly or narrowly conical, with a 

flattened base not easily removed from the substrate, black; 

apex with a conical protruding papilla and an often porelike 

ostiole (Fig. 68a). Peridium 22–53 μm thick laterally, 

thicker at the apex, 1-layered, composed of heavily pigmented 

thick-walled cells of textura angularis, cells to 7 μm diam., 

cell wall 1.5–3 μm thick, apex cells smaller and walls 

thicker, base cells walls thinner (Fig. 68b). Hamathecium of 

dense, long trabeculate pseudoparaphyses 1–2 μm broad, 

septate, branching and anastomosing (Fig. 68c). Asci 90– 

130×(5.5-)7–10 μm (x ¼ 107:3 8mm, n=10), 8-spored, 

with a short pedicel up to 20 μm long, bitunicate, 

fissitunicate, cylindrical, with a small ocular chamber (to 

1.5 μm wide×1.5 μm high) (Fig. 68c, d and e). Ascospore 

15–22×4–5 μm (x ¼ 20 4:4mm, n=10), biseriate near the 

top and uniseriate at the base, broadly fusoid to fusoid with 

broadly to narrowly rounded ends, brown to reddish brown, 

3-septum, deeply constricted at the median septum and 

breaking into two conical partspores, no constriction at the 

secondary septum, smooth (Fig. 68d and e). 


Material examined: GERMANY, on decorticated, 

decaying roots of Fagus sylvatica, very rare, collected in 

autumn (G: F. rh. 2173, isotype). 

Notes 

Morphology 

Ohleria is characterized by its subglobose to conic 

ascomata, produced on decorticated woody substrates, as


Fig. 67 Nodulosphaeria hirta (from BR 101945–95, holotype). a 

Appearance of ascomata on the host surface. b Vertical section of an 

ascoma. Note the setae at the apex and in the ostiole. c Section of a 

partial peridium. Note the outer layer cells of textura angularis and 

inner layer compressed cells. d Squash mount showing asci in 

pseudoparaphyses. e, f. The light brown filiform ascospores. Scale 

bars: a=0.5 mm, b=100 μm, c=50 μm, d=20 μm, e, f=10 μm 

well as its brown and phragmosporous ascospores which 

break into two parts at the median septum (Samuels 1980). 

Some species of Ohleria are widespread. For instance, O. 

brasiliensis is reported from New Zealand, Brazil as well as 

United States (Samuels 1980). Ohleria has been considered 

closely related to Sporormia and Preussia based on the 

ascosporic characters, and several species of Ohleria, suchas 

O. aemulans Rehm, O. haloxyli Kravtzev, O. silicata 

Kravtzev and O. kravtzevii Schwarzman, have been transferred 

to these genera. Clements and Shear (1931) treated 

Ohleria as a synonym of Ohleriella, despite the fact that 

Ohleriella is a coprophilous fungus. When the ascomata and 

habitats are considered, Ohleria seems closely related to 

Melanomma and Trematosphaeria (Samuels 1980). 


None. 


To some degree, habitats show phylogenetic significance 

(Zhang et al. 2009a). Thus, Ohleria seems less likely related 

to Sporormia and Preussia. But its relationship with 

Melanomma is uncertain, because of their differences in 

hamathecium and ascospores.


Fig. 68 Ohleria modesta (from g: f. rh. 2173, isotype). a Ascomata scattering on host surface. b Section of a partial peridium. c Asci embedded 

in pseudoparaphyses. d, e Cylindrical asci with short pedicels. Scale bars: a=1 mm, b, c=50 μm, d, e=20 μm 

Ohleriella Earle, Bull N Y Bot Gard 2: 349 (1902). 

(Delitschiaceae) 



immersed, erumpent to nearly superficial, scattered or in 

small groups, usually with a wide papilla, ostiolate, 

coriaceous. Peridium composed of small pigmented cells 

of textura angularis. Asci 8-spored or fewer, cylindroclavate, 

with a furcate pedicel. Hamathecium of trabeculate 

pseudoparaphyses. Ascospores brown to dark brown, 

cylindrical to nearly clavate with broadly to narrowly round 

ends, multi-septate, easily broken into partspores, smooth, 

with elongated germ slit in each cell. 


Literature: Ahmed and Cain 1972; von Arx and Müller 

1975; Barr 1990a; Clements and Shear 1931. 

Type species 

Ohleriella neomexicana Earle, Bull N Y Bot Gard 2: 349 

(1902). (Fig. 69) 


solitary, scattered, or in small groups, immersed to 

erumpent, to nearly superficial, with basal wall remaining


immersed in host tissue, coriaceous, globose or subglobose, 

usually a somewhat thick, short papilla, up to 

100 μm high, with a pore-like ostiole (Fig. 69a). Peridium 

27–35 μm thick laterally, up to 55 μm thick at the apex, 1- 

layered, composed of small pigmented cells of textura 

angularis, cells up to 5×8 μm diam., cell wall 1.5–2 μm 

thick, apex cells smaller and walls thicker (Fig. 69b). 

Hamathecium of dense, long trabeculate pseudoparaphyses, 

1–1.5 μm broad, anastomosing and branching between 

and above the asci. Asci 150–208×17.5–25 μm 


cylindrical to cylindro-clavate, with a narrowed, furcate, 

thin pedicel, 15–55 μm long, 2–3.5 μm broad, with a large 

truncate ocular chamber best seen in immature asci (to 4 μm 

wide×3 μm high) (Fig. 69e, f and g). Ascospores 55–72.5× 

10–12 μm (x ¼ 63 10:4mm, n=10), 3–4 seriate to uniseriate 

near the base, cylindrical to clavate, with broadly to 

narrowly rounded ends, brown, 6–7 transversesepta,easily 

separating into partspores, with germ slits, central partspores 

of the ascospores shorter than broad, rectangular in vertical 

section, round in transverse section, 7–8×6–10 μm diam., 

apical cells usually longer than broad, 11–17.5×6–7 μm 

diam. (Fig. 69c and d). 


Material examined: USA, Albuquerque, Bernalillo Co., 

New Mexico, dry gravelly hill, on wood, 29 Nov. 1901, T. 

S.A. Cockerell (NY, holotype). 

Notes 

Morphology 

Ohleriella was formally established by Earle (1902) based 

on its “medium to large ascomata with a wide papilla, 

relatively wide peridium, cylindro-clavate asci, brown to 

deep brown multi-septate ascospore, with elongated germ slit 

on each cell”, and was monotypified by O. neomexicana 

(Barr 1990a). Ohleriella subsequently has been treated as a 

synonym of Ohleria, Sporormiella or Preussia (Ahmed and 

Cain 1972; vonArxandMüller1975; Clements and Shear 

1931). Spororminula tenerifae, the generic type of Spororminula, 

was assigned to Ohleriella, thus Spororminula was 

treated as a synonym of Ohleriella (Barr 1990a). Two new 

species were introduced by Barr (1990a) from North America. 

Currently, three species are included in this genus, i.e. O. 

herculean (Ellis & Everh.) M.E. Barr, O. neomexicana and O. 

nudilignae M.E. Barr & Malloch (http://www.index 

fungorum.org; http://www.mycobank.org, 01/03/2009). 

The generic type, O. neomexicana, is morphologically 

similar to the coprophilous genus Sporormiella, but is 

saprobic on grass stems. 


None. 


Although we maintain Ohleriella as a separate genus 

here, its saprobic habitat on grasses and similarity to the 

coprophilous Sporormiella may indicate a close evolutionary 

relationship, with the grass saprobic possibly 

being an early relative of the coprophilous Sporormiella. 

Alternatively, the species/genera may simply occupy 

different ecological niches (i.e. dead grass vs dead grass 

in dung). Molecular studies are needed to resolve this 

issue. 

Ophiobolus Reiss, Hedwigia 1:27 (1854). 




medium-sized, solitary, scattered, or in groups, globose or 

pyriform, coriaceous, black, papillate, ostiolate, periphysate. 

Peridium thin, thicker near the apex, thinner at the base. 

Hamathecium of long cellular pseudoparaphyses, septate, 

anastomosing or branching not observed. Asci 8-spored, 

bitunicate, fissitunicate dehiscence not observed, cylindrical, 

with a short, furcate pedicel. Ascospores filamentous, 

narrower toward the lower end, pale brown, multiseptate, 

separating into two partspores from the middle 

septum, from the breaking point, the second cell of each 

partspore enlarged. 

Anamorphs reported for genus: Coniothyrium-like, 

Rhabdospora, Phoma-like and Scolecosporiella (Hyde et 

al. 2011; Shoemaker 1976; Sivanesan 1984). 

Literature: Holm 1948, 1957; Müller 1952; Reiss 1854; 

Shoemaker 1976; Sivanesan 1984. 

Type species 

Ophiobolus disseminans Reiss, Hedwigia 1:27 (1854) 

(Fig. 70). 


scattered, or in groups often arranged in a row, immersed with 

a protruding papilla, globose, pyriform, coriaceous, black, 

periphysate. Papilla 40–90 μm high, with a pore-like ostiole 

(Fig. 70a and b). Peridium 40–55 μm wide at the sides, up to 

70 μm thick at the apex, thinner at the base, comprising two 

cell types, outer layer composed of small heavily pigmented 

thick-walled cells of textura angularis, cells 2–5 μm diam., 

cell wall 2–3 μm thick, apex cells smaller and walls thicker, 

inner layer composed of lightly pigmented or hyaline thinwalled 

cells of textura angularis, 5–7 μm diam., wall 1.5– 

2 μm thick, merging with pseudoparaphyses (Fig. 70c). 

Hamathecium of long cellular pseudoparaphyses, 2–3 μm 

broad, septate, anastomosing or branching not observed 

(Fig. 70e). Asci 150–195×8–12.5 μm (x ¼ 169:5 10:7mm,


Fig. 69 Ohleriella neomexicana (NY, holotype). a Ascoma scattering 

on the host surface. b Section of a partial peridium. Note the small 

cells of textura angularis. c Ascospore in ascus. d Ascospore breaking 

into part spores. Note the sigmoid germ slit. e Dehiscent ascus. f, g 

Asci with short pedicels. Scale bars: a=100 μm, b=50 μm, c–g= 

10 μm


n=10), 8-spored, bitunicate, fissitunicate dehiscence not 

observed, cylindrical but narrowing towards the base, with a 

short, furcate pedicel which is 10–25 μm long, ocular 

chamber not observed (Fig. 70d and e). Ascospores 110– 

160×2.5–4 μm (x ¼ 135:3 3mm, n=10), filamentous, 

narrower toward the lower end, pale brown, 22–30- 

septate, separating into two partspores from the middle 

septum, from the breaking point the second cell of each 

partspore enlarged. 


Material examined: GERMANY, near Kassel, on dead 

stem of Cirsium arvense (L.) Scop., Spring 1853 (BPI- 

629021, type). 

Notes 

Morphology 

Ophiobolus was established by Reiss (1854) as a 

monotypic genus represented by O. disseminans based on 

its “Perithecia discreta, ostiolis prominentibus: sporae ascis 

inclusae, binatae, filliformes, multiseptatae”. 

A broad generic concept was adopted for the genus by 

Holm (1948) and Müller (1952). Shoemaker (1976)surveyed 

Canadian species of Ophiobolus using the broad concept of 

Holm (1948) and Müller (1952). A narrower generic concept 

was used by Holm (1957), which only included species with 

ascospores separating into two halves. Holm (1957) assigned 

species with enlarged ascospore cells to Nodulosphaeria,and 

those with long spirally coiled ascospores to Leptospora 

(Shoemaker 1976). This left only three species accepted 

under Ophiobolus (Holm 1957), although this concept has 

rarely been followed with new species recently being 

described (Raja and Shearer 2008). 

Walker (1980) provided a detailed description from the 

type material and dealt with many species of scolecospored 

fungi that had been placed in Ophiobolus by Saccardo 

(1883). Thus, currently several Ophiobolus sensu lato 

species are separated into Acanthophiobolus, Entodesmium, 

Leptosphaeria and Leptospora. Ophiobolus sensu lato 

contains about 300 species names (Sivanesan 1984; http:// 

www.mycobank.org/, 04/02/2009). 


Ophiobolus fulgidus (Cooke & Peck) Sacc. (as Leptosphaeria 

fulgida (Cooke & Peck) M. E. Barr in Dong et al. 

1998) lacks support in the clade of Leptosphaeriaceae 

(Dong et al. 1998). We expect it may closely related to 

Phaeosphaeriaceae. 


We agree from morphological data that Ophiobolus 

should comprise species that have filamentous spores that 

break easily into two halves at the central septum, with the 

second cell on either side being swollen (Walker 1980) and 

that the genus presently comprises three species (i.e. O. 

anthrisci (L. Holm) L. Holm, O. ophioboloides (Sacc.) L. 

Holm and O. acuminatus). All other Ophiobolus species 

need to be re-examined and should be placed in other 

genera such as Nodulosphaeria and Leptospora. The genus 

is in need of revision and molecular phylogenetic study. 

Ophiosphaerella Speg., Anal. Mus. nac. Hist. nat. B. Aires 

19: 401–402 (1909). (Phaeosphaeriaceae) 



small- to medium-sized, solitary or scattered, immersed, 

globose or subglobose, papillate, ostiolate. Peridium thin. 

Hamathecium of dense, filliform, septate pseudoparaphyses. 

Asci bitunicate, fissitunicate dehiscence not observed, cylindrical 

often narrower near the base, with a short furcate 

pedicel. Ascospores filamentous, pale brown, multi-septate. 

Anamorphs reported for genus: Scolecosporiella (Farr et 

al. 1989). 

Literature: von Arx and Müller 1975; Schoch et al. 2006, 

2009; Spegazzini 1909; Walker 1980; Wetzel et al. 1999; 


Type species 

Ophiosphaerella graminicola Speg., Anal. Mus. nac. Hist. 

nat. B. Aires 19: 401 (1909). (Fig. 71) 


or scattered, immersed with a short papilla protruding out of 

the substrate, globose or subglobose, often laterally flattened, 

dark brown to black, papillate, papilla ca. 100μm high, 140– 

180 μm broad, disk-like in appearance from above, periphysate 

(Fig. 71a and b). Peridium 11–25 μm wide, thicker near 

the apex, comprising two cell types of small cells, outer wall 

composed 6–10 layers of lightly brown flattened cells of 

textura angularis, inner layer composed of paler and thinwalled 

cells, both layers thicker near the apex (Fig. 71b). 

Hamathecium of dense, long pseudoparaphyses 0.8–1.5 μm 

broad near the apex, septate, 2–3 μm broad between the asci. 

Asci 105–135×5.5–10 μm (x ¼ 118:5 7mm, n=10), 8- 

spored, bitunicate, cylindrical and narrower near the 

base, with a short, furcate pedicel, up to 30 μm long, 

small inconspicuous ocular chamber (to 1.5 μm wide× 

1 μm high) (Fig. 71c, d, e and f). Ascospores 100–125× 

1.8–2.2 μm (x ¼ 118 2mm, n=10), filamentous, pale 

brown, 12–20 septa, smooth-walled. 


Material examined: ARGENTINA, Tucumán, on leaf 

sheath of Leptochloa virgata (L.) P. Beauv., 14 Apr. 1906, 

C. Spegazzini (LPS 858, holotype).


Fig. 70 Ophiobolus 

disseminans (from BPI-629021, 

type). a Immersed ascomata 

scattered on the host surface. 

Note the erumpent papilla. b 

Section of an ascoma. c. Section 

of a partial peridium. Note the 

thick-walled outer layer and 

thin-walled inner layer (orange 

colour due to DIC). d Ascus 

with a short furcate pedicel. e 

Squash mount showing asci in 

pseudoparaphyses. Scale bars: 

a=0.5 mm, b=100 μm, 

c=50 μm, d, e=20 μm 

Notes 

Morphology 

Ophiosphaerella was introduced by Spegazzini (1909) 

who described and illustrated a single new species, O. 

graminicola, and thus the genus was validly published 

(Walker 1980, p. 70). After checking the type specimen, 

Petrak and Sydow (1936) transferred the generic type to 

Ophiobolus graminicolus (Speg.) Petrak & Syd, and 

assigned Ophiosphaerella as a synonym of Ophiobolus. 

This was followed by von Arx and Müller (1975). 

Ophiosphaerella differs from Phaeosphaeria by its scolecospores 

without swollen cells or appendages, and from 

Ophiobolus by its ascospores without swollen cells or 

separating into partspores, thus was kept as a separating 

genus (Eriksson 1967a; Walker 1980). 


Ophiosphaerella forms a monophyletic group as a sister 

group of Phaeosphaeria located in Phaeosphaeriaceae (Schoch 

et al. 2006, 2009; Wetzel et al. 1999; Zhang et al. 2009a). 


Numerous Ophiobolus species are likely to belong in 

Ophiosphaerella. The two genera are distinguished as 

Ophiobolus sensu Shoemaker (1976) has swollen central 

cells or breaking into partspores or with long spirally 

coiled ascospores, and Ophiosphaerella (sensu Walker


1980) has scolecospores without swollen central cells or 

breaking into partspores.The recent introduction of Ophiobolus 

shoemakeri Raja & Shearer (Raja and Shearer 2008) 

is probably incorrect since the ascospores do not split up 

into partspores and there is no swelling above septum 

either. In particular, its freshwater habitat also distinguishes 

it from other species of Ophiobolus. Like 

Ophiobolus, Ophiosphaerella is in need of phylogenetic 

analysis but appears to be closely related to Phaeosphaeriaceae 

(Schoch et al. 2006). 

Ostropella (Sacc.) Höhn., Annls mycol. 16: 144 (1918). 


≡ Ostropa subgen. Ostropella Sacc., Syll. fung. (Abellini) 

2: 805 (1883). 


Habitat terrestrial, saprobic. Ascomata large, erumpent 

to superficial, solitary or gregarious, globose to subglobose, 

with broad and compressed papilla and slit-like 

ostiole. Peridium carbonaceous. Hamathecium of dense, 

long trabeculate pseudoparaphyses, anastomosing and 

branching, rarely septate, embedded in mucilage. Asci 

clavate with very long and thin and furcate pedicels. 

Ascospores pale brown, ellipsoid to fusoid, 1-septate, 

constricted. 



Yue 1987; Huhndorf 1993; Müller and von Arx 1962; 

Müller and Dennis 1965; Saccardo 1883. 

Fig. 71 Ophiosphaerella 

graminicola (from LPS 858, 

holotype). a Ascomata on the 

host surface. Note the protruding 

disk-like papilla. b Section 

of an ascoma. c Asci in 

pseudoparaphyses with short 

pedicels. d–f Cylindrical asci 

with short pedicels. Scale bars: 

a=0.5 mm, 

b=100 μm, c–f =10 μm


Type species 

Ostropella albocincta (Berk. & M.A. Curtis) Höhn., Annls 

mycol. 16: 144 (1918). (Fig. 72) 

≡ Ostropa albocincta Berk & M.A. Curtis, in Berkeley, 

J. Linn. Soc., Bot. 10: 372 (1868). 


scattered to gregarious, erumpent to superficial, globose to 

subglobose, roughened, often covered with white crustose 

covering, with subiculum, with a broad compressed papilla 

and long and slit-like ostiole (Fig. 72a). Peridium 100– 

250 μm thick, not of uniform thickness throughout entire wall 

area, composed of two cell types, one is of lightly pigmented 

thin-walled cells of textura prismatica, cells up to 17×3 μm 

diam., cell wall


Ascostromata black, immersed, penetrating into the 

substrate with dark brown hyphae. Ascomata up to 

680 μm high×540 μm diam., solitary, immersed or 

erumpent, subglobose to pyriform, subiculate or nonsubiculate, 

papillate or epapillate, ostiolate, periphysate, carbonaceous 

(Fig. 73a). Peridium thick. Hamathecium of long 

trabeculate pseudoparaphyses, 1–1.5 μm broad. Asci 90– 

130×12–17 μm (x ¼ 116 15mm, n=10), bitunicate, fissitunicate, 

cylindrical, 8-spored, uniseriate, with a short 

furcate pedicel, without apical apparatus (Fig. 73b, c and 

d). Ascospores 17.5–25×10–12.5 μm (x ¼ 21 11mm, n= 

10), ellipsoid to broadly fusoid with broadly rounded ends, 

1-septate, constricted at the septum, hyaline, smooth-walled, 

surrounded by a gelatinous sheath that contracts to form a 

lateral, lentiform, viscous appendage over the septum, 7.5– 

12.5 μm diam., 1–3 μm thick (Fig. 73e, f, g and h). 


Material examined: USA, Florida, Charlotte Harbor in 

Punta Garda, 10 Jan. 1964, leg., det. J. J. Kohlmeyer (Herb. 

J. Kohlmeyer No. 1720). 

Notes 

Morphology 

Paraliomyces was introduced to accommodate the 

marine fungus P. lentifer, which is characterized by 

immersed ascomata produced within the ascostroma, 

trabeculate pseudoparaphyses, cylindrical, 8-spored asci, 

ellipsoidal, hyaline, 1-septate ascospores surrounded by a 

gelatinous sheath, which forms a lentiform, viscous 

appendage over the septum (Kohlmeyer 1959). 


Based on analysis of SSU sequences, Paraliomyces 

lentifer nested within Pleosporales, but its familial status 

was left undetermined (Tam et al. 2003). 


None. 

Phaeosphaeria I. Miyake, Bot. Mag., Tokyo 23: 93 (1909). 




small, solitary, scattered, or in small groups, immersed, 

globose, subglobose, wall black. Apex with a pore-like 

ostiole. Peridium thin. Hamathecium of dense, filliform, 

septate pseudoparaphyses. Asci 8-spored, bitunicate, fissitunicate, 

broadly cylindrical to narrowly fusoid, with a 

short pedicel. Ascospores fusoid to narrowly fusoid, pale 

brown to brown, 3-septate. 

Fig. 72 Ostropella albocincta (K(M): 143941, syntype). a Ascomata 

gregarious on host surface. b Section of the partial peridium. Note the 

peridium comprising two cell types and the whitening tissue 

(arrowed). c Pseudoparaphyses. d, e Asci with long pedicel. f– 

h Ascospores, which are strongly constricted at the central septum. 

Scale bars: a=1 mm, b=100 μm, d, e, h=20 μm, c, g, f=10 μm 

Anamorphs reported for genus: Amarenographium, 

Hendersonia-like, Phaeoseptoria, Scolecosporiella and 

Stagonospora (Hyde et al. 2011; Leuchtmann 1984; 

Shoemaker and Babcock 1989b). 

Literature: von Arx and Müller 1975; Câmara et al. 

2002; Eriksson 1967a, 1981; Holm 1957; Khashnobish and 

Shearer 1996; Leuchtmann 1984; Miyake 1909; Shoemaker 

and Babcock 1989b. 

Type species 

Phaeosphaeria oryzae I. Miyake, Bot. Mag., Tokyo 23: 

136 (1909). (Fig. 74) 


solitary, scattered, or in small groups, immersed, globose, 

subglobose, wall black, forming black spots on the leaves 

of hosts (Fig. 74a). Apex with a pore-like ostiole. Peridium 

4–8 μm wide at the sides, composed of heavily pigmented 

thin-walled cells of textura angularis, cells 2–2.5×3–5 μm 

diam., cell wall less than 1 μm thick (Fig. 74b). Hamathecium 

of dense, long cellular pseudoparaphyses 2–2.5 μm 

broad, embedded in mucilage, rarely branched, septate. Asci 

53–80(−90)×7–10 μm (x ¼ 65:3 8:3mm, n=10), 8- 

spored, bitunicate, fissitunicate, broadly cylindrical to 

narrowly fusoid, with a short pedicel which is ca. 8μm 

long, with a small ocular chamber and an inconspicuous 

apical apparatus (to 2 μm wide×1 μm high) (Fig. 74c, d 

and e). Ascospores 17– 22(− 28)×4– 5 μ m 

(x ¼ 20:5 4:6mm, n=10), obliquely uniseriate, partially 

overlapping or biseriate, narrowly fusoid with rounded ends, 

pale brown, 3-septate, slightly constricted at primary septum, 

granulate (Fig. 74f and g). 


Material examined: JAPAN, Suruya, Shizuoka, on the 

leaves of Oryza sativa, Sept. 1907 (S nr F9572, F9573, 

lectotype). 

Notes 

Morphology 

Phaeosphaeria was introduced by Miyake (1909), but 

was regarded as a synonym of Leptosphaeria for a long 

time. Holm (1957), however, reinstated Phaeosphaeria, 

assigning some Leptosphaeria sensu lato species with 

relatively small ascomata and which occurred on monocotyledons 

to Phaeosphaeria. Although this division based 

b

Fungal Diversity


Fig. 73 Paraliomyces lentifer (from Herb. J. Kohlmeyer No. 1720). a 

Section of an immersed ascoma. b Eight-spored cylindrical asci 

embedded in pseudoparaphyses. c, d Cylindrical asci with short 

on host range is considered unnatural by some workers 

(Dennis 1978; Sivanesan 1984), it has been widely 

accepted (von Arx and Müller 1975; Eriksson 1967a; 

Hedjaroude 1969; Shoemaker and Babcock 1989b). 

Eriksson (1981) further revised the generic concept of 

pedicels. e–h One-septate hyaline ascospores. Scale bars: a=100 μm, 

b–d=20 μm, e–h=10 μm 

Phaeosphaeria by including dictyosporous taxa as well as 

some perisporium taxa. Phaeosphaeria was further divided 

into six subgenera, i.e. Ovispora, Fusispora, Phaeosphaeria, 

Spathispora, Vagispora and Sicispora, based on 

differences in ascospore shape and the number of septa


(Shoemaker and Babcock 1989b). Phaeosphaeria species 

are usually associated or parasitic on annual monocots, 

such as Cyperaceae, Juncaceae or Poaceae but have also 

been recorded as saprobes and on dicotyledons (e.g. P. 

viridella and P. vagans). 


The separation of Phaeosphaeria from Leptosphaeria 

sensu stricto was supported by phylogenetic studies based 

on ITS sequences. The peridium structure, pseudoparenchymatous 

cells in Phaeosphaeria versus scleroplectenchymatous 

cells in Leptosphaeria had phylogenetic 

significance in the distinction between these two genera, 

while the subgenus division was not supported by the 

phylogenetic results (Câmara et al. 2002; Morales et al. 

1995). The familial status of both Phaeosphaeriaceae and 

Leptosphaeriaceae was verified by multigene phylogenetic 

analysis (Schoch et al. 2009; Zhang et al. 2009a). 


Phaeosphaeria was originally thought to be a synonym of 

Leptosphaeria (Müller 1950; Munk1957), however, molecular 

analysis has shown these two genera differ with 

Phaeosphaeria having pseudoparenchymatous peridium, Stagonospora-like 

anamorph and mostly monocotyledonous 

hosts and Leptosphaeria having scleroplectenchymatous 

peridium, Phoma-like anamorph and mostly dicotyledonous 

hosts (Câmara et al. 2002;Schochetal.2009; Shoemaker and 

Babcock 1989b; Zhang et al. 2009a). It is now recognized 

that Phaeosphaeria is the type genus of Phaeosphaeriaceae 

and related genera include Entodesmium and 

Setomelanomma and probably Ophiosphaerella (Schoch 

et al. 2009; Zhangetal.2009a). Paraphaeosphaeria was 

introduced as an off-shoot of Phaeosphaeria and differs 

in ascospore shape and septation as well as anamorphic 

stages (Eriksson 1967a, b). Similarly, Nodulosphaeria 

was recently reinstated and differs from Phaeosphaeria 

because of setae over the apex as well as its ascospores 

with swelling supramedian cells and terminal appendages 

(Holm 1957, 1961). While the newly reinstated Phaeosphaeria 

was confined to monocotyledons and particularly 

grasses, there are now many species that have been 

described from dicotyledons (Farr et al. 1989). Whether 

these taxa form a monophyletic group needs to be 

investigated with fresh collections and molecular data. 

Phaeosphaeriopsis M.P.S. Câmara, M.E. Palm & A.W. 

Ramaley, Mycol. Res. 107: 519 (2003). (Phaeosphaeriaceae) 


Habitat terrestrial, saprobic or hemibiotrophic? Ascomata 

small, scattered or in small groups, immersed, 

globose, subglobose. Peridium thin, comprising one cell 

type of textura angularis. Hamathecium of dense, wide 

cellular pseudoparaphyses. Asci 8-spored, bitunicate, 

cylindrical to broadly fusoid, with a short pedicel and a 

small ocular chamber. Ascospores obliquely uniseriate 

and partially overlapping to biseriate even triseriate, 

cylindrical, pale brown, multi-septate, primary septum 

submedian, with or without constriction, verrucose or 

baculate. 

Anamorphs reported for genus: Coniothyrium-like, 

Phaeostagonospora (Câmara et al. 2003). 

Literature: Câmara et al. 2003. 

Type species 

Phaeosphaeriopsis glaucopunctata (Grev.) M.P.S. Câmara, 

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

(Fig. 75) 

≡ Cryptosphaeria glaucopunctata Grev. Fl. Edin.: 362 

(1824). 


scattered, or in small groups, immersed, globose, subglobose 

(Fig. 75a). Peridium 10–25 μm wide, comprising one 

type of cells, composed of thick-walled cells of textura 

angularis, cells 4–9 μm diam., cell wall 2–3 μm thick, 

almost equal in thickness. Hamathecium of dense, wide 

cellular pseudoparaphyses, 3–5 μm broad. Asci (50-)60– 

110×10–15 μm (x ¼ 82:3 12mm, n =10), 8-spored, 

bitunicate, fissitunicate dehiscence not observe, cylindrical 

to broadly fusoid, with a short pedicel, with a small 

ocular chamber (to 0.8 μm wide×1 μm high) (Fig. 75b). 

Ascospores 18–28×5–7.5 μm (x ¼ 23:5 6:2mm, n=10), 

obliquely uniseriate and partially overlapping to biseriate 

even triseriate, cylindrical, pale brown, 4(−5)- 

septate, without constriction or slightly constricted at 

the basal septum, the forth cell from the apex usually 

slightly inflated, the basal cell often longer, baculate 

(Fig. 75c, d, e and f). 


Material examined: UK, Epping, Sept. 1863 (E, M.C. 

Cooke 166, barcode: E00074286). 

Notes 

Morphology 

Phaeosphaeriopsis was introduced to accommodate 

some species of Paraphaeosphaeria based on both morphological 

characters and results of SSU rDNA sequence 

analyses (Câmara et al. 2003). Most of the Phaeosphaeriopsis 

species occur on the Agavaceae, although P. 

glaucopunctata occurs on Liliaceae (Ruscus). Phaeosphaeriopsis 

is characterized by having uni- or multioculate 

stromata and 4- or 5-septate ascospores. Although the


Fig. 74 Phaeosphaeria oryzae 

(from S nr F9572, F9573, lectotype). 

a Appearance of ascomata 

on the host surface. b 

Section of an ascoma. c Squash 

mount showing asci in pseudoparaphyses. 

Note that asci with 

short pedicels. d, e Asci with 

short pedicels. F, G. Light 

brown 3-septate ascospores. 

Scale bars: a=100 μm, b–g= 

10 μm 

morphological characters of Phaeosphaeriopsis species is 

more diverse than those of Paraphaeosphaeria sensu 

stricto or Neophaeosphaeria, the ITS sequences are more 

similar to each other than those of the other two genera 

(Câmara et al. 2003). Currently, Phaeosphaeriopsis comprises 

seven species, namely P. agavensis (A.W. Ramaley, 

M.E. Palm & M.E. Barr) M.P.S. Câmara, M.E. Palm & A. 

W. Ramaley, P. amblyospora A.W. Ramaley, P. glaucopunctata, 

P. musae Arzanlou & Crous, P. nolinae (A.W. 

Ramaley) M.P.S. Câmara, M.E. Palm & A.W. Ramaley, P. 

obtusispora (Speg.) M.P.S. Câmara, M.E. Palm & A.W. 

Ramaley and P. phacidiomorpha (Ces.) D.F. Farr & M.E. 

Palm (http://www.mycobank.org/, 06/2010). 


The generic type of Phaeosphaeriopsis, P. glaucopunctata, 

located in Phaeosphaeriaceae based on SSU rDNA 

sequences (Câmara et al. 2003). Phaeosphaeriopsis musae 

is also shown to belong to Phaeosphaeriaceae in recent 

phylogenetic studies (Schoch et al. 2009; Plate 1). 


None. 

Platysporoides (Wehm.) Shoemaker & C.E. Babc., Can. J. 

Bot. 70: 1648 (1992). (Pleosporaceae) 

≡ Pleospora subgenus Platysporoides Wehmeyer, A 

World Monograph of the genus Pleospora and its Segregates, 

p. 236. 1961. 


Habitat terrestrial, saprobic? Ascomata small, scattered, 

immersed, semi-immersed to nearly superficial, globose, 

subglobose, black, smooth; apex with a protruding 

papilla and pore-like ostiole, without periphyses. Perid-


ium thin, composed of a few layers of textura 

angularis. Hamathecium of numerous, cellular pseudoparaphyses, 

anastomosing, septate. Asci bitunicate, fissitunicate, 

cylindrical to cylindro-clavate, with a short, 

furcate pedicel. Ascospores broadly ellipsoid, reddish 

brown, muriform. 


Literature: Shoemaker and Babcock 1992; Wehmeyer 

1961. 

Type species 

Platysporoides chartarum (Fuckel) Shoemaker & C.E. 

Babc., Can. J. Bot. 70: 1650 (1992) (Fig. 76) 

≡ Pleospora chartarum Fuckel, Jb. nassau. Ver. Naturk. 

23–24: 133–134 (1870). 


scattered, immersed, semi-immersed to rarely superficial, 

globose, subglobose, black, smooth; apex with a protruding 

papilla, 50–85 μm long, 60–85 μm broad, ostiolate 

(Fig. 76a and b). Peridium 8–22 μm wide, composed of 

2–4 layers of brown cells of textura angularis, cells5– 

9 μm diam., cell wall 1–2.5 μm thick, without periphyses. 


2–3 μm broad, anastomosing, septate (Fig. 76c). Asci 

110–140×12.5–16.5 μm (x ¼ 121:5 14:7mm, n=10), 

(6-)8-spored, bitunicate, fissitunicate, cylindrical to 

cylindro-clavate, with a short, furcate pedicel, 8–17 μm 

long, ocular chamber not observed (Fig. 76c, d and e). 

Ascospores 20–26×8–11 μm (x ¼ 23:7 9mm, n=10), 

obliquely uniseriate and partially overlapping, flattened, 

broadly ellipsoid in front view, reddish brown, 3 

Fig. 75 Phaeosphaeriopsis 

glauco-punctata (from M.C. 

Cooke 166). a Ascomata immersed 

in the substrate. b Eightspored 

cylindrical asci. c–f. Pale 

brown baculate ascospores 

which are released from asci. 

Scale bars: a=200 μm, 

b=20 μm, c, d–f=10 μm


transverse septa, 1 longitudinal septum in each central 

cell, 1 oblique septum in each end cell, constricted at all 

septa, granulate, with a sheath 2–3 μm wide (as reported 

in Shoemaker and Babcock 1992) (Fig.76f, g and h). 


Material examined: GERMANY, Budenheim, Leopold 

Fuckel, Nassau’s Flora, on old paper (G NASSAU: 210558 

(a), as Sphaeria chartarum Wallr., type). 

Notes 

Morphology 

Platysporoides was introduced as a subgenus of Pleospora 

by Wehmeyer (1961) and was typified by Pleospora 

chartarum. Shoemaker and Babcock (1992) raised Platysporoides 

to generic rank and placed it in the Pleosporaceae 

based on its “applanodictyospore” and “terete pored 

beak of the ascomata”. Currently, eleven species are 

included in this genus (Shoemaker and Babcock 1992). 

Another comparable pleosporalean family is Diademaceae, 

which is distinguished from Platysporoides by its ascoma 

opening as “an intraepidermal discoid lid” (Shoemaker and 

Babcock 1992). 


None. 


Aigialus grandis is another pleosporalean fungus 

with flattened and muriform ascospores as well as 

papilla and ostioles, which belongs to Aigialaceae, a 

phylogenetically well supported marine family 

(Suetrong et al. 2009). Thus, it is highly likely that 

flattened and muriform ascospores are of little phylogenetic 

significance. 

Pleomassaria Speg., Anal. Soc. cient. argent. 9: 192 

(1880). (Pleomassariaceae) 



solitary, scattered, or in small groups, immersed, erumpent 

by a minute slit or a small conical swelling in the bark, 

flattened, papillate, ostiolate. Hamathecium of dense, 

cellular pseudoparaphyses, embedded in mucilage. Asci 

bitunicate, fissitunicate, broadly cylindrical to broadly 

cylindro-clavate, with a short, thick pedicel. Ascospores 

muriform, brown, constricted at the septa. 

Anamorphs reported for genus: Prosthemium and Shearia 

(Barr 1982b; Sivanesan 1984). 

Literature: Barr 1982b, 1990b, 1993a; Clements and Shear 

1931; Eriksson 2006; Lumbsch and Huhndorf 2007; 

Shoemaker and LeClair 1975; Sivanesan 1984; Tanaka 

et al. 2005. 

Type species 

Pleomassaria siparia (Berk. & Broome) Sacc., Syll. fung. 

2: 239 (1883) (Fig. 77) 

≡ Sphaeria siparia Berk. & Broome, Ann. Mag. nat. 

Hist., Ser. 2 9: 321 (1852). 


solitary, scattered, or in small groups, immersed, erumpent 

by a minute slit or a small conical swelling in the bark, 

depressed globose, papillalte, ostiolate (Fig. 77a). Peridium 

45–60 μm wide, thicker at the apex, thinner at the 

base, 1-layered, composed of small pigmented thickwalled 

compressed cells, cells ca. 15×3μm diam., cell 

wall 2–3.5 μm thick, apex cells larger, base composed of 

small pigmented thick-walled cells of textura angularis, 

ca. 5μm diam. (Fig. 77b). Hamathecium of dense, cellular 

pseudoparaphyses, 1–2 μm broad, embedded in mucilage, 

anastomosing or branching not observed. Asci 180–250× 

28–42 μm (x¼206:3 36:8mm, n=10), 8-spored, bitunicate, 

fissitunicate, broadly cylindrical to broadly cylindroclavate, 

with a short, thick pedicel, 15–45 μm long, with 

inconspicuous ocular chamber (Fig. 77c and d). Ascospores 

45–58×12.5–17.5 μm (x ¼ 50:5 14:8mm, n=10), 

biseriate, narrowly oblong with broadly to narrowly 

rounded ends, brown, muriform with 5–8 transverse septa 

and 1–2 vertical septa in some cells, smooth to verrucose, 

constricted at the septa, surrounded by a mucilaginous 

sheath (Fig. 77e, f and g). 

Anamorph: Prosthemium betulinum Kunze (Sivanesan 

1984). 

Conidia to 120 μm diam., with 3–5 arms, each arm 3–5- 

septate, 40–55×13–16 μm, connected to a central cell 

(Fig. 77h, i and j). 

Material examined: UK, Wiltshire, Spye Park, on 

branch of Betulina with Hendersonia polycystis Berk., et 

Br. leg. C.E. Broome, 1850? (BR, type). 

Notes 

Morphology 

Pleomassaria as characterized by Barr (1982b) has 

medium- to large-sized, immersed ascomata, cellular 

pseudoparaphyses, clavate to oblong asci and large, 

muriform ascospores (Barr 1982b; Sivanesan 1984). The 

muriform and somewhat asymmetrical ascospores with a 

submedian primary septum distinguish Pleomassaria from 

Asteromassaria in the family Pleomassariaceae, while in 

Splanchnonema ascospores have distinct bipolar asymmetry. 

Barr (1982b) included five North American species in 

the genus, while Kirk et al. (2008) listedfourspecies.


Fig. 76 Platysporoides chartarum 

(from G NASSAU: 

210558, type). a, b Ascomata 

scattered among fibers. Note the 

central ostioles. c Asci in numerous 

cellular pseudoparaphyses. 

d, e Cylindro-clavate asci 

with short pedicels. f–h. Muriform 


a, b=200 μm, c–e=20 μm, 

f–h=10 μm 

Barr (1993a) treated Pleomassaria as a synonym of 

Splanchnonema based on a morphological cladistic analysis, 

but this proposal was not followed by later workers 

(Eriksson 2006; Lumbsch and Huhndorf 2007; Tanaka et 

al. 2005). 


Pleomassaria siparia forms a robust phylogenetic clade 

with Melanomma pulvis-pyrius (generic type) (Schoch et al. 

2009; Zhang et al. 2009a), which might represent a 

phylogenetic family (or suborder?).



The genera Asteromassaria, Pleomassaria and Splanchnonema 

of Pleomassariaceae areconsideredtobeclosely 

related and difficult to separate (Barr 1982b; Crivelli 1983). 

They all have ascomata which are immersed in bark and are 

visible as slightly raised pustules with small ostioles, but 

may eventually become erumpent (e.g. Asteromassaria 

macrospora). Pseudoparaphyses are cellular, asci are bitunicate, 

while ascospores vary from 1-septate and pale brown 

(e.g. Asteromassaria macrospora) tomuriform(e.g.Pleomassaria 

siparia) and may be symmetrical (e.g. Asteromassaria 

macrospora) or highly asymmetrical (e.g. 

Splanchnonema pustulatum). The peridium ranges from 

thick-walled textura angularis (e.g. Asteromassaria macrospora) 

to thin-walled compressed cells (e.g. Splanchnonema 

pustulatum) and medium textura prismatica (e.g. Pleomassaria 

siparia). Anamorphs also vary distinctly, Prosthemium 

in Pleomassaria siparia, Scolicosporium in Asteromassaria 

macrospora but no anamorphic stage reported for Splanchnonema 

pustulatum. Furthermore, Asteromassaria pulchra 

clusters in Morosphaeriaceae in this study, thus here we 

tentatively assign Asteromassaria in Morosphaeriaceae 

(Plate 1). There seems to be considerable confusion in this 

family, especially when Pleomassaria siparia forms a robust 

phylogenetic clade with Melanomma pulvis-pyrius (Melannomataceae). 

Thus in this study, Pleomassariaceae is 

restated as a separate family from Melannomataceae. 

Therefore, fresh collections of the types of these genera are 

needed for molecular analysis and to establish which 

characters are important for classification. 

Pleophragmia Fuckel, Jb. nassau. Ver. Naturk. 23–24: 243 

(1870). (Sporormiaceae) 


Habitat terrestrial, saprobic (coprophilous). Ascomata smallto 

medium-sized, gregarious, immersed to erumpent, globose 

to subglobose, black, coriaceous; apex with a short papilla, 

or sometimes forming an ostiolar pore. Peridium thin, 

composed of several layers of thin-walled cells of textura 

angularis. Hamathecium of dense, delicate pseudoparaphyses. 

Asci 8-spored, bitunicate, fissitunicate, clavate to 

cylindro-clavate, with a relatively long pedicel and an ocular 

chamber. Ascospores muriform, narrow oblong to cylindrical 

with rounded ends, dark brown, constricted at each septum. 


Literature: von Arx and Müller 1975; Cain 1934. 

Type species 

Pleophragmia leporum Fuckel, Jb. nassau. Ver. Naturk. 

23–24 (1870) [1869–70]. (Fig. 78) 


gregarious, immersed to slightly erumpent, globose to 

subglobose, black; apex with a short papilla, sometimes 

forming a ostiolar pore (Fig. 78a). Peridium 25–35 μm 

thick at the sides, composed of one cell type of lightly 

pigmented thin-walled cells of textura angularis, cells 6– 

10 μm diam., cell wall 1.5–2 μm thick (Fig. 78b). 

Hamathecium of numerous, long pseudoparaphyses, 1– 

2 μm broad, anastomosing not observed. Asci 160–250× 

22.5–27.5 μm (x ¼ 203:6 25mm, n=10), 8-spored, bitunicate, 

fissitunicate, clavate to cylindro-clavate, with a 20– 

50 μm long pedicel and an ocular chamber (to 5 μm wide× 

2 μm high) (Fig. 78e and f). Ascospores 42–50×8–10 μm 

(x ¼ 46 10mm, n=10), biseriate to uniseriate and partially 

overlapping, narrowly oblong to cylindrical with rounded 

ends, dark brown, often slightly curved, with 9 transverse 

septa with two crossing longitudinal septa in the centre, 

constricted at each septum, smooth-walled (Fig. 78c, d, g 

and h). 


Material examined: GERMANY, between Königstein 

and Glashütten, on the same dung with Delitschia minuta. 

s.d. (G, Fungi rhenani n2272, type). 

Notes 

Morphology 

Pleophragmia was formally established by Fuckel 

(1870) and monotypified by Pleophragmia leporum. The 

most comparable genus to Pleophragmia is Sporormia, as 

ascospores of both have no germ slits and the inner layer of 

wall is considerably thinner than the outer layer (Barr 

1990a, b). But the muriform ascospores of Pleophragmia 

can be readily distinguished from the phragmosporous 

ascospores of Sporormia. Currently, only four species are 

accommodated under this genus (http://www.mycobank. 

org, 28-02-2009). 


None. 


The presence of both transverse and crossing longitudinal 

septa is the most striking character of Pleophragmia, although 

the phylogenetic significance of this character is unclear. 

Pleoseptum A.W. Ramaley & M.E. Barr, Mycotaxon 54: 76 

(1995). (Phaeosphaeriaceae) 


Habitat terrestrial, saprobic? Ascomata medium-sized, 

scattered, or in small groups, immersed, globose to conoid,


Fig. 77 1 Pleomassaria siparia 

(from BR, type). a Ascomata on 

the host surface (after removing 

the cortices). b Section of a 

partial peridium. c, d Asci with 

short pedicels. e–g Ascospores 

with thin sheath. Scale bars: 

a=0.5 mm, b–d=50 μm, e–g= 

20 μm. 2 Prosthemium betulinum 

(from BR, type). h–j Conidia 

with arms. Scale bars: 

h–j=20 μm 

black, papillate, ostiolate. Peridium 1-layered. Hamathecium 

of dense, long cellular pseudoparaphyses, septate, 

branching. Asci 8-spored, bitunicate, fissitunicate, cylindrical 

to cylindro-clavate, with furcate pedicel. Ascospores 

obliquely uniseriate and partially overlapping, muriform, 

ellipsoid, ovoid to fusoid, yellowish to dark brown. 

Anamorphs reported for genus: Camarosporium (Ramaley 

and Barr 1995).


Fig. 77 (continued) 

Literature: Ramaley and Barr 1995. 

Type species 

Pleoseptum yuccaesedum A.W. Ramaley & M.E. Barr, 

Mycotaxon 54: 76 (1995). (Fig. 79) 

Ascomata 300–500 μm diam., scattered, or in small 

groups of 2–3, immersed with a flattened top, globose to 

conoid, black, papillate, ostiolate (Fig. 79a). Papilla 

small, slightly protruding from the host surface. Peridium 

30–50 μm thick at sides, up to 100 μm thick at the apex, 

1-layered, composed of 5–8 layers of heavily pigmented 

purplish-brown cells of textura angularis, cells 5–12 μm 

diam., cell wall 1–2 μm thick, apex cells smaller and 

walls thicker (Fig. 79c). Hamathecium of dense, long 

cellular pseudoparaphyses 1–2 μm broad, septate, 

branching (Fig. 79b). Asci 125–170(−195)×15–22 μm 


cylindrical to cylindro-clavate, with a short, 

narrowed, furcate pedicel which is 10–20 μm long, with 

an ocular chamber best seen in immature asci (to 5 μm 

broad×3 μm high) (Fig. 79d and e). Ascospores 22–30× 

11–14 μm (x ¼ 27:1 12:6mm, n=10) obliquely uniseriate 

and partially overlapping, ellipsoid, ovoid to fusoid, 

yellowish to yellowish brown, becoming reddish brown 

to dark brown, muriform, with 3-(4) transverse septa, 

constricted at the primary septum, part above central 

septum wider, vertical septa exist in each cell, ornamentation 

of foveolae in linear rows (Fig. 79f and g). 

Anamorph: Camarosporium yuccaesedum Fairm. 

(Ramaley and Barr 1995). 

Conidiomata 200–450 μm diam., pycnidial, immersed, 

scattered, subglobose to conoid, ostiolate. Macroconidiogenous 

cells determinate or indeterminate, enteroblastic, 

hyaline, smooth. Macroconidia holoblastic, 20–36×10– 

15 μm diam., ellipsoid to narrowly ovoid, muriform, 

yellowish brown, 3–7 transverse septa, constricted at the 

septa. Microconidiogenous cells produced near or in the 

ostiole, hyaline, smooth. Microconidia 5–10×5–7 μm 

diam., globose to ovoid, aseptate, hyaline, smooth. 

Material examined: USA, Colorado, Montezuma County, 

hillside near entrance to Mesa Verde National Park, on dead 

leaves of Yucca baccata, 11 Oct. 1992, Ramaley Annette 

(9237A) (BPI 802381, holotype). 

Notes 

Morphology 

Pleoseptum is a monotypic genus established by 

Ramaley and Barr (1995) and represented by P. yuccaesedum 

basedonits“immersed ascomata, thick peridium, 

muriform ascospores, anamorphic stage and the linoeate 

ornamentation of the ascospores and conidia”. Theshape 

of ascomata of Pleoseptum is comparable with that of 

Chaetoplea, but the peridium structure easily distinguishes 

them. Some species of Curreya, Leptosphaeria and 

Heptameria are comparable with Pleoseptum, but their 

anamorphic stages differ. 

Pleoseptum yuccaesedum and its Camarosporium yuccaesedum 

anamorph both formed in the leaves of Yucca 

baccata and the ascomata and conidiomata were indistinguishable. 

Camarosporium is the anamorph of diverse 

teleomorph genera included in Botryosphaeriales and 

Cucurbitariaceae (Kirk et al. 2008). The genus is in need 

of revision (Sutton 1980) and is no doubt polyphyletic. 


None. 


The placement of Pleoseptum under Phaeosphaeriaceae 

is still tentative. 

Pleospora Rabenh. ex Ces. & De Not., Comm. Soc. crittog. 

Ital. 1: 217 (1863). (Pleosporaceae) 


Habitat terrestrial, saprobic or parasitic. Ascomata smallto 

medium-sized, immersed, erumpent to superficial,


Fig. 78 Pleophragmia leporum (from G. Fungi rhenani n2272, 

type). a Appearance of ascomata on the substrate surface. Note the 

ostiolar pore. b Section of a partial peridium. c, h Apical part of an 

ascus. Note the apical apparatus in (c). d Released ascospores. e, f 

Clavate Asci with pedicels. g Part of a broken ascospore. Note the 

crossing septa. Scale bars: a=0.5 mm, B=50 μm, c–f=20 μm, g, 

h=10 μm


Fig. 79 Pleoseptum yuccaesedum (from BPI 802381, holotype). a 

Appearance of ascomata scattered on the host surface. Only the 

upper region is visible. b Squash mount of asci in pseudoparaphyses. 

c Section of an ascoma. Note the peridium comprising cells of 

textura angularis. d, e Asci with short furcate pedicels. f, g 

Muriform dark-brown ascospores. Scale bars: a=0.5 mm, b= 

40 μm, c=100 μm, d, e=20 μm, f, g=10 μm


papillate, ostiolate. Peridium thin. Hamathecium of dense, 

cellular pseudoparaphyses. Asci 8-spored, bitunicate, 

fissitunicate, cylindrical to clavate, with furcate pedicel 

and small inconspicuous ocular chamber. Ascospores 

muriform, brown or pale brown, with or without 

sheath. 

Anamorphs reported for genus: Stemphylium (Simmons 

1985). 

Literature: Barr 1981; Frisullo and Braun 1996; Kodsueb 

et al. 2006a; Luttrell1951; Wehmeyer1946, 1961, 1975; 


Type species 

Pleospora herbarum (Pers.) Rabenh., Klotzschii Herb. Viv. 

Mycol. 2: no. 547 (1854). (Fig. 80) 

≡ Sphaeria herbarum Pers., Syn. meth. fung. (Göttingen) 

1: 78 (1801). 


scattered, or in small groups of 2–3, immersed, semiimmersed 

to erumpent, broadly to narrowly oblong and 

flattened, with flattened base not easily removed from the 

substrate, wall black, papillate, ostiolate (Fig. 80a and b). 

Peridium 30–50 μm thick on sides, thinner at the base, 

coriaceous, 2-layered, outer layer composed of one or two 

layers of heavily pigmented thick-walled cells of textura 

angularis, cells 5–10 μm diam., cell wall 2–4 μm thick, 

apex cells smaller and walls thicker, inner layer composed 

of hyaline thin-walled cells of textura angularis, 8–12 μm 

diam., wall hyaline, 0.5–1.5 μm thick (Fig. 80c). Hamathecium 

of dense, cellular pseudoparaphyses, 2–3 μm 

broad, filling the gaps between the asci. Asci 100–210× 

27.5–30 μm (x ¼ 142:2 28:3mm, n=10), 8-spored, bitunicate, 

fissitunicate, broadly cylindrical to clavate, with a 

short, thick, furcate pedicel, 8-12(−20) μm long, with 

small inconspicuous ocular chamber (ca. 3 μm wide× 

1 μm high) (Fig. 80d and e). Ascospores 28–38×12.5– 

15 μm (x ¼ 33 14:5mm, n=10), ellipsoidal, straight or 

sometimes curved, with broadly rounded ends and upper 

hemispore slightly shorter and broader; spores usually 

divided by 3 A-transsepta, all 4 segments by longisepta 

andthenbyonestratumofB-transsepta(maturesporesas 

a rule with 7 transsepta, 3A+4B), yellowish brown, 

smooth; each hemispore with thick gelatinous sheath, the 

lower one with umbilicus (sheaths fused in mature spores) 

(Fig. 80f, g, h, i, j and k). 

Anamorph: Stemphyllium herbarum E. Simmons 

(Simmons 1985). 

Material examined: GERMANY, on stalks of Melilotusalla? 

at the bank of the Elbe in Konigstein, 1882 (E, 

Krieger 683); as Sphaeria herbarum Persoon Syn. fung. p. 

78 (E, 81); as Sphaeria herbarum Fries, Scleromyceti 

Sueciae 38 (E, lectotype). 

Notes 

Morphology 

Pleospora was originally assigned within Sphaeriales. 

Subsequently, it was assigned within Pseudosphaeriales and 

Pleosporales (Wehmeyer 1961). Pleospora is a large group, 

which is widely distributed and associated with a wide range 

of species of monocotyledons as well as dicotyledons 

(Wehmeyer 1975). All species of Pleospora have muriform 

ascospores (Wehmeyer 1961, 1975). Pleospora has downward 

growing pseudoparaphyses within the ascomata of 

“Pleospora-type” development (Luttrell Univ. Mo. Stud. 

1951), which subsequently served as a diagnostic character. 

However, only a limited number of species had detailed 

studies on this character (Wehmeyer 1961). The heterogeneous 

nature of Pleospora has been noted, and several 

subgenera have been erected, such as Scleroplea to include 

all “sclerotioid” species of Pleospora, Teichosporoides to 

accommodate species of Pleospora with immersed ascomata, 

Pleosphaeria for those having superficial and setose ascomata 

(Wehmeyer 1961). Similarly, Cucurbitaria, Fenestella and 

Montagnula are also separated as a section from Pleospora. 

Most of these subgenera are currently at genus rank. 


The polyphyletic nature of Pleospora is clear (Kodsueb et 

al. 2006a), and those that stain the woody substrate purple 

should be assigned to Amniculicolaceae (Zhang et al. 2009a). 


As some Pleospora species have a wide range of host 

spectrum, especially on both monocotyledons and dicotyledons, 

it is highly possible they are cryptic species. 

Preussia Fuckel, Hedwigia 6: 175 (1867) [1869–70]. 

(Sporormiaceae) 


Habitat terrestrial, saprobic (on decaying fibers or coprophilous). 

Ascomata small- to medium-sized, cleistothecial or 

perithecial, solitary or scattered on substrate surface, 

globose, membraneous, black. Peridium thin, composed 

of thick-walled, poly-angular cells from the surface view. 

Pseudoparaphyses not observed. Asci (4-) 8-spored, bitunicate, 

clavate to broadly clavate, with a long and thin and 

furcate pedicel. Ascospores 3–6 seriate to uniseriate near 

the base, cylindrical with rounded ends, brown, septate, 

easily breaking into partspores, with germ slits in each cell. 

Anamorphs reported for genus: Phoma (von Arx 1973; 

Cain 1961; Malloch and Cain 1972). 

Literature: Ahmed and Cain 1972; Arenaletal.2005; von 

Arx 1973; von Arx and van der Aa 1987; Auerswald1866;


Fig. 80 Pleospora herbarium (from E, Krieger 683). a Immersed 

ascomata scattering on host surface. b Ascomata in small groups. Note: 

the surface layer of the host is removed. c Section of an ascoma. Note 

the peridium cells of textura angularis. d, e. Asci with short pedicels f–j 

Ascospores. Scale bars: a, b=0.5 mm, c=100 μm, d, e=30 μm, f–j= 

20 μm 

Barr 1987b, 1990a;Boylan1970; Cain 1961; Eriksson 1992; 

Fuckel 1866; Guarroetal.1981, 1997a, b; KhanandCain 

1979a, b; Kruys and Wedin 2009; Lodha 1971; Lorenzo 

1994; Luck-Allen and Cain 1975; Maciejowska and


Williams 1963; Malloch and Cain 1972; Munk 1957; 

Narendra and Rao 1976; Rai and Tewari 1963; Sultana 

and Malik 1980. 

Type species 

Preussia funiculata (Preuss) Fuckel, Jb. nassau. Ver. 

Naturk. 23–24: 91 (1870) [1869–70]. (Fig. 81) 

≡ Perisporium funiculatum Preuss, Fung. Hoyersw.: no. 

145 (1851). 

Ascomata 240–500 μm diam., cleistothecial, solitary, 

scattered on substrate, superficial, globose, membraneous, 

black (Fig. 81a). Peridium thin, composed of thick-walled, 

poly-angular cells in front view (Fig. 81b). Pseudoparaphyses 

not observed. Asci 42–65×20–25 μm (x ¼ 55:8 21:8mm, n= 

10), (4-)8-spored, bitunicate, broadly clavate, with a long and 

thin and furcate pedicel, up to 115 μm long, ocular chamber 

not observed (Fig. 81c and d). Ascospores 30–40×6.3– 

7.5 μm (x ¼ 35:6 6:9mm, n=10), 3–6 seriate to uniseriate 

near the base, cylindrical with rounded ends, brown, with 

3 transverse septa, easily breaking into partspores, central 

cells round in transverse section but rectangular in 

vertical section, with a germ slit in each cell, 6.5–8.5× 

4–7.5 μm broad, apical cells 8.8–10×5–7 μm broad, 

sheath not observed. 


Material examined: USA, Ontario, York Co., Nashville, 

on old jute sack on ground, 1 Jul. 1960, leg. & det. R.F. 

Cain (in part Preussia typharum) (TRTC 46985). 

Notes 

Morphology 

Preussia was introduced by Fuckel (1866) to accommodate 

species having cleistothecioid ascomata, bitunicate 

asci, multi-septate ascospores with a germ slit in 

each cell and with a gelatinous sheath, and occurring in 

soil or plant debris. Preussia, Sporormia and Sporormiella 

are regarded as closely related genera, which share 

numerous morphological characters. Sporormia can be 

distinguished from Preussia by its perithecioid ascomata 

and cylindrical asci. The only distinguishing morphological 

character for Preussia from Sporormiella are the 

cleistothecioid ascomata in Preussia (Barr 2000; Cain 

1961), but this has been shown to have little phylogenetic 

significance (von Arx 1973; Zhang et al. 2009a). 

Substrate preference has been used to distinguish species 

of Sporormiella and Preussia, with Sporormiella being 

restricted to a coprophilous habitat, while Preussia grows 

in plant debris, wood or soil (von Arx and van der Aa 

1987). This proposal was rejected, as P. intermedia 

(Clum) Cain can be isolated from either soil or dung 

(Guarro et al. 1997b). In a review of Preussia, Cain 

(1961) accepted 12 species, and some of them are coprophilous. 

Subsequently, numerous additional new species have 

been published (Arenal et al. 2005; Barr 1987b, 1990a; 

Boylan 1970; Eriksson1992; Guarro et al. 1981, 1997a, b; 

Khan and Cain 1979a; Lodha1971; Lorenzo 1994; Luck- 

Allen and Cain 1975; Maciejowska and Williams 1963; 

Malloch and Cain 1972; Narendra and Rao 1976; Rai and 

Tewari 1963; Sultana and Malik 1980). Currently, 84 species 

are listed under Preussia (http://www.mycobank.org/ 

mycotaxo.aspx, 10/2010) and Kirk et al. (2008) estimates 

there are 51 species. 


In phylogenetic analysis based on ITS, nLSU, mtSSU 

and β-tubulin gene fragments, Preussia, Sporormiella and 

Spororminula clustered together. Thus, Sporormiella together 

with Spororminula are treated as synonyms of 

Preussia (Kruys and Wedin 2009). 


Preussia sensu lato (including Sporormiella and Spororminula) 

based on both morphology and molecular data 

should be accepted pending further research. 

Quintaria Kohlm. & Volkm.-Kohlm., Bot. Mar. 34: 34 


Habitat marine, saprobic. Ascomata medium-sized, scattered 

or loosely gregarious, immersed, mostly subglobose, 

rarely globose, with a protruding papilla, ostiolate. Peridium 

thin, 2-layered, coriaceous, thicker near the apex. 

Hamathecium of dense, filamentous, trabeculate pseudoparaphyses, 

branching and anastomosing between and 

above asci. Asci 8-spored, bitunicate, fissitunicate, 

cylindro-clavate, with a short furcate pedicel. Ascospores 

biseriate, broadly fusoid to fusoid, hyaline, mostly 5- 

septate, rarely up to 7-septate. 


Literature: Hyde and Goh 1999; Kohlmeyer and 

Volkmann-Kohlmeyer 1991; Suetrong et al. 2009; Zhang 

et al. 2008b. 

Type species 

Quintaria lignatilis (Kohlm.) Kohlm. & Volkm.-Kohlm., 

Bot. Mar. 34: 35 (1991). (Fig. 82) 

≡ Trematosphaeria lignatilis Kohlm., Marine Ecology, 

[Pubblicazioni della Stazione Zoologica Napoli I] 5(4): 365 

(1984). 

Ascomata 240–500 μm diam., scattered or loosely 

gregarious, immersed, globose to subglobose, coriaceous, 

ostiolate, ostiole is encrusted with thick-walled black cells, 

papilla up to 400 μm long (Fig. 82a). Peridium thin, 20– 

30 μm wide, thinner at the base, thicker near the apex, up to


Fig. 81 Preussia funiculata 

(from TRTC 46985). a Superficial 

cleistothecoid ascomata. b 

Part of peridium from front 

view. c Squash mounts showing 

a large number of asci. d A 

clavate ascus with a long and 

thin pedicel. Scale bars: a= 

0.5 mm, b=20 μm, c, d=50 μm 

300 μm, 2-layered, outer layer composed of hyphoid cells, 

inner layer composed of compressed cells of textura 

angularis (Fig. 82b). Hamathecium of dense, filamentous, 

trabeculate pseudoparaphyses, 0.8–1.5 μm broad, branching 

and anastomosing between and above asci (Fig. 82e). 

Asci 175–250×25–35 μm (x ¼ 220 28mm, n=10), 8- 

spored, bitunicate, fissitunicate, cylindro-clavate, with a 

short, furcate pedicel, to 20 μm long (Fig. 82c and e). 

Ascospores 55–73×12.5–15 μm (x ¼ 63:3 13:1mm, n= 

10), biseriate, broadly fusoid to fusoid, usually slightly 

curved, smooth, hyaline, mostly 5-septate, rarely up to 7- 

septate, smooth-walled, lacking a sheath. 


Material examined: BELIZE, Twin Cays, on attached 

dead tip of prop root of Rhizophora mangle, with shipworms, 

3 Apr. 1983, leg. & det. J.K. Kohlmeyer (J. 

Kohlmeyer No. 4365a, holotype). 

Notes 

Morphology 

Quintaria was introduced to accommodate the marine 

fungus, Trematosphaeria lignatilis, based on its immersed 

ascomata with rounded bases, black incrustations surrounding 

the sides of the ostiolar canal as well as its hyaline 

ascospores (Kohlmeyer and Volkmann-Kohlmeyer 1991). 

Subsequently, three more species were introduced to this 

genus, viz. Q. aquatica K.D. Hyde & Goh, Q. microsporum 

Yin. Zhang, K.D. Hyde & J. Fourn. and Q. submerse K.D. 

Hyde & Goh, which are all from freshwater (Hyde and Goh 

1999; Zhang et al. 2008b). 


Multigene phylogenetic study indicated that Quintaria 

lignatilis forms a separate sister clade to other families of 

Pleosporales, which may represent a new familial linage 

(Suetrong et al. 2009). This was supported by phylogenetic 

studies which place the freshwater Q. submersa separate from 

Q. lignatilis (Schoch et al. 2009;Suetrongetal.2009; Plate 1). 


The freshwater members of Quintaria should likely be 

excluded from this genus, and only the generic type, Q. 

lignatilis retained, but this needs confirmation. 

Roussoëlla Sacc., in Saccardo & Paoletti, Atti Inst. Veneto 

Sci. lett., ed Arti, Sér. 3 6: 410 (1888). (Arthopyreniaceae 

(or Massariaceae))



Habitat terrestrial, saprobic. Ascomata medium-sized, clustered, 

immersed in host tissue, forming under darkened, 

slightly raised, somewhat liner or dome-shaped stroma on the 

host, with a flush intra-epidermal papilla; immersed under 

clypeus, papillate, ostiolate. Peridium thin, comprising 

several layers of compressed cells. Hamathecium of dense, 

long trabeculate pseudoparaphyses, embedded in mucilage, 

hyaline, anastomosing and septate. Asci 8-spored, bitunicate, 

cylindrical, with furcate pedicel, and a conspicuous ocular 

chamber. Ascospores uniseriate to partially overlapping, fusoid 

or ellipsoidal, brown, 1-septate, constricted at the septum. 

Anamorphs reported for genus: Cytoplea (Hyde et al. 

1996a). 

Literature: Hyde et al. 1996a; Hyde 1997; Ju et al. 1996; 

Tanaka et al. 2009. 

Type species 

Roussoëlla nitidula Sacc. & Paol., Atti Ist. Veneto Sci., Ser. 

6, 6:410. (1888). (Fig. 83) 

Ascomata 160–200 μm high×400–500 μm diam., clustered, 

immersed in host tissue, forming under darkened, 

slightly raised, somewhat liner or dome-shaped stroma on the 

host, with a flush intra-epidermal papilla; in vertical section 

subglobose with a flattened base, immersed under clypeus, 

subglobose with a flattened base, papillate, ostiolate (Fig. 83a). 

Peridium up to 20 μm thick, comprising several layers of 

compressed cells. Hamathecium of dense, long trabeculate 

pseudoparaphyses, 1–1.5 μm broad, embedded in mucilage, 

anastomosing and septate. Asci 123–220×7–11 μm,8-spored, 

bitunicate, cylindrical, with furcate pedicels, and a conspicuous 

ocular chamber (Fig. 83b, c and d). Ascospores 17.5– 

22×5.5–8 μm, uniseriate to partially overlapping, fusoid or 

ellipsoidal, brown, 1-septate, constricted at the septum, 

ornamented with longitudinal wall striations and surrounded 

by a wide mucilaginous sheath (Fig. 83e, f, g and h). 

Anamorph: Cytoplea hysterioides K.D. Hyde (Hyde et 

al. 1996a). 

Material examined: MALAYSIA, Malacca, on culms of 

Bambusa Bar & Grill, 1885, B. Scortechini 15 (PAD, 

Roussoëlla nitidula Sacc. Paol. 2484, holotype, on a loose 

label Roussoëlla nitidula S. & P. Est Phyllachora phaeodidym./15 

prob. original material from Malacca Peninsula). 

Notes 

Morphology 

Roussoëlla was introduced by Saccardo for the single 

species R. nitidula Sacc. & Paol. (Saccardo and Paoletti 

1888). It was redescribed by Hyde et al. (1996a) and the 

anamorph of Roussoëlla hysterioides (Ces.) Höhn., Cytoplea 

hysterioides K.D. Hyde was determined and described. 

Roussoëlla was then reviewed by Hyde (1997) and a 

modified key for Roussoëlla species was provided based on 

the one proposed by Ju et al. (1996). Roussoëlla is 

characterized as having immersed ascomata containing 

long cylindrical asci and brown 1-septate ornamented 

ascospores. In this study, we have checked the type species 

and it matches Hyde et al. (1996a). The asci are bitunicate, 

but we could not observe the fissitunicate dehiscence. 


Species of Roussoëlla, Roussoellopsis as well as Arthopyrenia 

salicis form a robust phylogenetic clade, which form 

a sister group with pleosporalean families, but the generic 

type of Roussoëlla (R. nitidula) was not included in the 

phylogenetic study (Tanaka et al. 2009). 


The bambusicolous habitat of Roussoëlla is a striking 

character at generic rank classification but its relationship to 

the lichenized Arthopyrenia is unexpected and will require 

more analysis. 

Saccharicola D. Hawksw. & O.E. Erikss., in Eriksson & 

Hawksworth, Mycologia 95: 431 (2003). (Massarinaceae) 


Habitat terrestrial, parasitic. Ascomata medium-sized, solitary, 

scattered, immersed, globose to subglobose, carbonaceous, 

papillate, ostiolate. Peridium relatively thin, composed of one 

cell type of pale brown to hyaline pseudoparenchymatous cells. 

Hamathecium of trabeculate pseudoparaphyses. Asci bitunicate, 

8-spored, cylindro-clavate to clavate. Ascospores biseriate and 

sometimes laterally uniseriate, fusoid with narrowly rounded 

ends, septate, constricted at the septa, the upper second cell 

becoming pigmented when mature, smooth or verruculose. 

Anamorphs reported for genus: Stagonospora (Eriksson 

and Hawksworth 2003; Kaiser et al. 1979; Leuchtmann 

1984). 

Literature: Eriksson and Hawksworth 2003. 

Type species 

Saccharicola bicolor (D.Hawksw.,W.J.Kaiser&Ndimande) 

D. Hawksw. & O.E. Erikss., Mycologia 95: 431 (2003). 

(Fig. 84) 

≡ Leptosphaeria bicolor D. Hawksw., W.J. Kaiser & 

Ndimande, Mycologia 71: 483 (1979). 


solitary, scattered, immersed, globose to subglobose, wall 

black, carbonaceous, with a protruding papilla, with a 

central ostiole (Fig. 84a). Peridium 15–20 μm thick

Fig. 82 Quintaria lignitalis 

(from J. Kohlmeyer No. 4365a, 

holotype). a Ascomata immersed 

in substrate. b Section of 

an ascoma. Note the thin peridium 

and elongated papilla. c, e 

Asci embedded in pseudoparaphyses. 

d Five septate fusoid 

hyaline ascospores. Scale bars: 

a=0.5 mm, b=200 μm, c, e= 

50 μm, d =20 μm 

Fungal Diversity


composed of one cell type of pale brown to hyaline 

pseudoparenchymatous cells, becoming thicker near the apex 

(Fig. 84a). Hamathecium of 1–2 μm broad, filliform, hyaline, 

septate pseudoparaphyses, branching and anastomosing in 

mucilage. Asci (90-)125–150×(20-)25–30 μm, 8-spored, with 

a short pedicel, bitunicate, cylindro-clavate to clavate, with a 

small ocular chamber at the apex (Fig. 84c). Ascospores 29– 

42×8–11 μm, biseriate and sometimes laterally uniseriate, 

fusoid with narrowly rounded ends, (2-)3-septate, deeply 

constricted at the septa, the upper second cell subhyaline to 

pale brown when young and becoming dark brown to almost 

black at maturity, smooth or verruculose (Fig. 84d). (data from 

the original description by Kaiser et al. (1979) because of the 

bad condition of the type material). 

Fig. 83 Roussoëlla nitidula 

(from PAD Paol. 2484, holotype). 

a Appearance of the 

stroma on host surface. b Asci 

and pseudoparaphyses. c, d 

Long cylindrical furcate asci. 

e–h. Ascospores. Note the striate 

ornamentation. Scale bars: 

a=0.5 mm, b–d=20 μm, 

e–h=10 μm


Anamorph: Pycnidia typical of Stagonospora (Sphaeropsidales), 

“scattered,arisingsinglybothonthehostandinpure 

culture, in culture generally surrounded by an envelope of 

mycelial hyphae, numerous, immersed on the host, but nearly 

superficial in culture, subglobose to slightly applanate, black, 

150–250 μm diam., with a central slightly papillate ostiole, 

lacking a distinct neck; walls mainly 15–20 μm thick, 

composed of three to six layers of pseudoparenchymatous 

cells, the outermost layers dark brown and inner pale brown to 

hyaline cells somewhat compressed radially, very variable in 

size, cells of the outer layers mainly 7–12 μm long×4–6 μm 

wide in vertically section and 10–12 μm diam. in surface 

view, wall not or only slightly thicked near the ostiole. 

Conidiogenous cells lining the inner surface of the pycnidial 

cavity, holoblastic, minute and difficult to distinguish from the 

pseudoparenchymatous cells with which they are mixed, 

mammiform with a flattened apex, hyaline, smooth walled, 

about 4–6 μm tall and 4–6 μm wide. Conidia copiously 

produced, ellipsoid, with somewhat truncated ends, hyaline, 

smooth walled, (2-)3 septate, not or slightly constricted at the 

septa, often guttulate, rather thin walled, (21-)24–28(−34) 

μm×7–8.5(−11.5) μm” (from Kaiser et al. 1979). 

Material examined: KENYA, near Nairobi, on leaves of 

Saccharum officinarum L.; 24 Aug. 1977; leg. W.J. Kaiser 

(IMI 215888, holotype). 

Notes 

Morphology 

Saccharicola was separated from Leptosphaeria as a new 

genus based on its Stagonospora anamorph and its biotrophic 

habitat in leaves of sugar cane, and two species were 

included, i.e. Saccharicola bicolor and S. taiwanensis (J.M. 

Yen & C.C. Chi) O.E. Erikss. & D. Hawksw. (Eriksson and 

Hawksworth 2003). Saccharicola is characterized by its 

parasitic habitat on monocots, small ascomata, bitunicate 

asci, presence of pseudoparaphyses as well as its 3-septate 

ascospores (Eriksson and Hawksworth 2003). 


Based on the limited phylogenetic analysis of SSU 

sequences, Saccharicola is considered to be closely related 

to Massarina eburnea, the generic type of Massarina 

(Eriksson and Hawksworth 2003). Thus, Saccharicola was 

assigned to Massarinaceae, which includes Keissleriella, 

Massarina and Saccharicola (Eriksson and Hawksworth 

2003). 


Based on the parasitic habitat on monocots and its small 

ascomata and Stagonospora (or Cercospora? for S. taiwanensis, 

see Eriksson and Hawksworth 2003; Shoemaker 

and Babcock 1989b) anamorph, Saccharicola seems more 

similar to Pleosporineae. Further molecular study is needed 

for confirmation. 

Salsuginea K.D. Hyde, Bot. Mar. 34: 315 (1991). (Pleosporales, 



Habitat marine, saprobic. Ascomata large, solitary, fusoid, 

conical or subglobose, with or without a flattened base, 

immersed under a darkened clypeus, papillate, ostiolate. 

Peridium thin, composed of round cells (in cross section) at 

sides, fusing at the top with the clypeus, thin at the base. 


anastomosing, embedded in mucilage. Asci 8-spored, bitunicate, 

fissitunicate, clavate to cylindro-clavate, pedunculate, 

with a large ocular chamber and conspicuous apical ring. 

Ascospores uniseriate, obovoid, brown to black, with hyaline 

apical germ pores, 1-septate, constricted at the septum, dark 

brown with paler apical cells, lacking sheath, smooth. 


Literature: Hyde 1991a; Suetrong et al. 2009. 

Type species 

Salsuginea ramicola K.D. Hyde, Bot. Mar. 34: 316 (1991). 

(Fig. 85) 


solitary, fusoid, conical or subglobose, with or without a 

flattened base, immersed under a darkened clypeus, papillate, 

ostiolate, ostiole rounded (Fig. 85a). Peridium up to 39 μm 

thick, composed of round cells (in cross section) at sides, 

fusing at the top with the clypeus, thin at the base (Fig. 85b). 


1–2 μm broad, anastomosing, embedded in mucilage. Asci 

440–512×29–34 μm, 8-spored, bitunicate, fissitunicate, 

clavate to cylindro-clavate, pedunculate, with a large ocular 

chamber and conspicuous apical ring (Fig. 85c and e). 

Ascospores 59–72×24–30 μm, uniseriate, obovoid, brown to 

black, with hyaline apical germ pores, 1-septate, constricted 

at the septum, dark brown with paler apical cells, lacking 

sheath, smooth (Fig. 85d and f). 


Material examined: THAILAND, Ranong mangrove, 

Aegiceras corniculatum (L.) Blanco., Oct. 1988, leg. & det. 

K.D. Hyde (BRIP 17102, holotype). 

Notes 

Morphology 

Salsuginea was introduced to accommodate the mangrove 

fungus, S. ramicola, which is characterized by large, 

immersed, ostiolate and papillate ascomata under a clypeus,


Fig. 84 Saccharicola bicolor 

(from IMI 215888, holotype). a 

Section of an ascomata immersed 

in the host tissue. b 

Section of a partial pycnidia. 

Note the phragmosporous conidia. 

c Clavate ascus with ocular 

chamber and short pedicel. d 

Ascospores. Note the pigmented 

central cell(s). Scale bars: a, b= 

50 μm, c=20 μm, d=10 μm 

dense, trabeculate pseudoparaphyses embedded in gel matrix, 

fissitunicate, 8-spored, cylindrical asci with short pedicel and 

conspicuous apical apparatus, 1-septate, dark brown ascospores 

with paler apical cells (Hyde 1991a). Salsuginea is 

considered closely related to Helicascus and Caryospora, and 

they are all proposed to Melanommataceae (Hyde 1991a). 


Based on a multigene phylogenetic analysis, Salsuginea 

ramicola nested in a paraphyletic clade within Pleosporales; 

its familial status is undetermined (Suetrong et al. 2009). 


It has been shown that trabeculate pseudoparaphyses has 

no phylogenetic significance at familial rank, so a well 

resolved phylogeny based on DNA comparisons will be 

necessary to categorize this genus. 

Semidelitschia Cain & Luck-Allen, Mycologia 61: 581 

(1969). (Delitschiaceae) 



immersed to slightly erumpent, scattered, coriaceous, 

papillate, ostiolate. Hamathecium of non-typical trabeculate 

pseudoparaphyses, thin, septate, rarely branching. 

Asci cylindrical, pedicellate, each with a conspicuous 

large apical ring. Ascospores non-septate, dark brown to 

nearly black, each with an elongated germ slit. 


Literature: Barr 2000; Cain and Luck-Allen 1969. 

Type species 

Semidelitschia agasmatica Cain & Luck-Allen, Mycologia 

61: 581 (1969). (Fig. 86) 

Ascomata 550–900 μm diam., solitary, immersed to 

erumpent, globose to subglobose, black, semicoriaceous, 

smooth-walled, with a protruding papilla and a conspicuous 

ostiole (Fig. 86a). Peridium thin, comprising multi-angular


cells from front view. Hamathecium of non-typical trabeculate 

pseudoparaphyses, 1–2 μm broad, septate, rarely 

branching, anastomosing not observed. Asci 410–505×(38-) 

43–50 μm (x ¼ 470:6 46:4mm, n=10), 8-spored, bitunicate, 

fissitunicate dehiscence not observed, cylindrical, with 

a thick pedicel which is up to 90 μm long, and with a large 

and conspicuous dome-shaped ocular chamber surrounded by 

apical ring (to 18 μm wide×4 μm high) (Fig. 86b and e). 


uniseriate to obliquely uniseriate and partially overlapping, 

broad fusoid to subglobose, hyaline when 

young, then becoming yellowish brown, reddish brown 

Fig. 85 Salsuginea ramicola 


a Habitat section of an ascoma. 

b Section of the partial peridium. 

c Clavate mature and immature 

asci. d Ascospores 

within ascus. e Apical part of 

immature asci. f Ascospores 

with an apical chamber at each 

end. Scale bars: a=0.5 mm, 

b–e=50 μm, f=10μm


and nearly black and opaque when mature, non-septate, 

smooth-walled, with a full length germ slit, surrounded 

by a broad gelatinous sheath (Fig. 86c and d). 


Material examined: CANADA, Alberta, North of 

Beaver Mines, on sheep dung, 28 Jul. 1962, E.R. Luck- 

Allen, (TRTC 41607, paratype); USA, Montana: Gallatin 

County, 60 min S of Bozeman, on sheep dung, 2 Sept. 

1957, Cain (TRTC 42032, paratype); Stillwater County 

Columbus, on cow dung, 3 Sept. 1957, Cain (TRTC 42031, 

paratype); South Dakota, Meade Co.: South of Wall, on 

cow dung, 3 Sept. 1962, Cain (TRTC 40697, holotype). 

Notes 

Morphology 

Semidelitschia was formally establishedbyCainandLuck- 

Allen (1969) and was assigned to Sporormiaceae. Although it 

is similar to Delitschia, it differs as the ascospores are 1- 

celled, as opposed to 2-celled. Subsequently, Semidelitschia 

was transferred to Delitschiaceae together with Delitschia 

(Barr 2000). Currently, three species are listed under this 

genus, i.e. S. agasmatica Cain & Luck-Allen, S. nanostellata 

A.E. Bell & Mahoney and S. tetraspora J.H. Mirza & S.M. 

Khan (Index Fungorum) although the number of species in 

the genus are given as only two in Kirk et al. (2008). 


None. 


This is a clearly defined genus that differs from 

Delitschia in having 1-celled ascospores. Cultures of S. 

agasmatica are needed for sequencing and for establishing 

the placement and uniqueness of the genus. 

Setomelanomma M. Morelet, Bull. Soc. Sci. nat. Arch. 

Toulon et du Var 227:15 (1980). (Phaeosphaeriaceae) 


Habitat terrestrial, hemibiotrophic or biotrophic. Ascomata 

small, solitary, scattered, immersed, erumpent to superficial, 

globose to subglobose, black; with or without a small 

papilla, apex covered with setae and a periphysate ostiole. 

Peridium thin, 1-layered, composed of several layers of 

cells of textura angularis. Hamathecium of dense, 1–2 μm 

broad pseudoparaphyses, septate, anastomosing. Asci 8- 

spored, bitunicate, broadly cylindrical. Ascospores fusoid to 

broadly clavate, pale brown to brown, 3-septate. 


Literature: Leonard and Suggs 1974; Morelet 1980; 

Rossman et al. 2002; Schochetal.2009; Zhang et al. 2009a. 

Type species 

Setomelanomma holmii M. Morelet, Bulletin de la Société 

des Sciences naturelles et d’Archéologie de Toulon et du 

Var 36 (no. 227): 15 (1980). (Fig. 87) 

(Some information in the following description is from 

Rossman et al. (2002)) 

Ascomata 80–250 μm diam., solitary, scattered, immersed, 

erumpent to superficial, globose to subglobose, 

black, with setae; with or without a small papilla, apex 

covered with setae and a periphysate ostiole. Peridium 15– 

25 μm thick, 1-layered, composed of several layers of cells 

of textura angularis, cell wall thinner and more lightly 

pigmented towards centrum, cell wall thicker near the apex. 

Hamathecium of dense, 1–2 μm broad pseudoparaphyses, 

thicker near the base, septate, anastomosing (Fig. 87a and 

d). Asci 70–100×11–14 μm, 8-spored, bitunicate, broadly 

cylindrical with a short, thick, furcate pedicel, with a small 

ocular chamber (Fig. 87a, b and c). Ascospores 16–21×5– 

6.5 μm, obliquely uniseriate and partially overlapping to 

biseriate, fusoid to broadly clavate with broadly to narrowly 

rounded ends, pale brown to brown, 3-septate, slightly 

constricted at the median septum, smooth (Fig. 87e). 


Material examined: FRANCE, Leuglay, on dying twigs 

of Picea pungens. 8 May 1987, leg. M. Morelet (UPS F- 

117969 (slide), isotype). 

Notes 

Morphology 

Setomelanomma was formally established by Morelet 

(1980) as a monotypic genus represented by S. holmii, 

which was collected in France. The description, however, is 

not detailed and lacks illustrations. Rossman et al. (2002) 

collected this species in North America and detailed studies 

were conducted including both morphology and phylogeny. 

The bitunicate, broadly cylindrical asci, cellular pseudoparaphyses 

as well as the pale brown, septate ascospores with 

a median primary septum point Setomelanomma to 

Phaeosphaeriaceae as defined by Barr (1992a) and 

Eriksson et al. (2002) (Rossman et al. 2002). However, its 

setose ascomata, brown and 3-septate ascospores together 

with its residence in conifers distinguish it from all other 

genera under Phaeosphaeriaceae (Rossman et al. 2002). 

Setomelanomma is mostly comparable with Kalmusia and 

Phaeosphaeria. Setomelanomma can be distinguished from 

Kalmusia by its erumpent to superficial ascomata with 

almost no papilla, and Phaeosphaeria differs from Setomelanomma 

by its host spectrum and reported anamorphic 

stages (Rossman et al. 2002). Currently, five species are 

included in Setomelanomma, namely S. holmii, S. monoceras, 

S. prolata K.J. Leonard & Suggs, S. rostrata (K.J. Leonard) K.


Fig. 86 Semidelitschia agasmatica 

(from TRTC 40697, holotype). 

a Immersed ascomata 

scattered on the surface of the 

substrate. b Squash of ascoma. 

Note the numerous released asci. 

c Apical ring of cylindrical 

asci. d One-celled ascospores. 

Note the germ slits (see arrow). 

e Cylindrical ascus. Note the 

tapering pedicel. Scale bars: 

a=0.5 mm, b–e=100 μm 

J. Leonard & Suggs and S. turcica (Luttr.) K.J. Leonard & 

Suggs (http://www.mycobank.org/, 06/2010). 


Setomelanomma forms a well supported phylogenetic 

clade with other members of Phaeosphaeriaceae (Schoch 

et al. 2009; Zhang et al. 2009a). 


None. 

Shiraia Henn., Bot. Jb. 28: 274 (1900). (Pleosporales, 



Habitat terrestrial, parasitic. Ascostroma warty-like or 

tuber-like. Ascomata medium to large, subglobose, gregarious 

on the surface layer of ascostroma, immersed, 

ostiolate, with a small black opening seen on the surface 

of the ascostroma, ostiole rounded. Hamathecium of


dense, long trabeculate pseudoparaphyses, anastomosing 

and branching between the asci. Asci bitunicate, fissitunicate, 

cylindrical to cylindro-clavate, with a short furcate 

pedicel, with a big and truncate ocular chamber. Ascospores 

obliquely uniseriate and partially overlapping, 

narrowly fusoid to fusoid or broadly fusoid with tapering 

or narrowly rounded ends, hyaline to pale brown or 

brown, muriform. 

Anamorphs reported for genus: coelomycetous with 

muriform conidia (see Liu 2009). 

Literature: Cheng et al. 2004; Hino 1961; Kishi et al. 

1991; Liu 2009; Morakotkarn et al. 2008. 

Type species 

Shiraia bambusicola Henn., Bot. Jb. 28: 274 (1900). 

(Fig. 88) 

Ascostroma 1–1.5 cm high×1–2.5 cm diam., subglobose, 

oblong to irregular, slightly pink with cracking 

surface. Ascomata 350–800 μm high×300–700 μm diam., 

subglobose, gregarious on the surface layer of ascostroma, 

immersed, ostiolate, with a small black opening seen on 

the surface of the ascostroma, ostiole rounded, the inner 

tissue of ascostroma carnation red (Fig. 88a and b). 


0.8–1.5 μm broad, anastomosing and branching 

between the asci. Asci 300– 425×20– 35 μm 


cylindrical to cylindro-clavate, with a short furcate 

pedicel, up to 50 μm long, with a big and truncate ocular 

chamber (Fig. 88c and d). Ascospores 62.5–80×17.5– 

22.5 μm (x ¼ 72:3 19:3mm, n=10), obliquely uniseriate 

and partially overlapping, narrowly fusoid to fusoid with 

tapering or narrowly rounded ends, hyaline turning pale 

brown when mature, muriform, with 9–13 transversal 

septa, 1–3 longitudinal septa in central cells, slightly 

constricted at the septa, usually with a gelatinous cap at 

each end (Fig. 88e, f and g). 

Anamorph: coelomycetous with muriform conidia (see 

Liu 2009). 

Material examined: CHINA, Zhejiang, Hangzhou, 

Panan, on bamboom, 15 Jun. 2009, leg. Liu Yongxiang 

(IFRD 2040). 

Notes 

Morphology 

Shiraia is reported as a parasite on branches of several 

genera of bamboo distributed mainly in southern regions of 

China and Japan (Hino 1961; Kishi et al. 1991; Liu 2009). 

Shiraia is characterized by its bambusicolous habitat, large 

ascostroma and muriform ascospores. Asci comprise 6 

ascospores in this study and some previous studies (Hino 

1961; Liu 2009). Shiraia bambusicola is well studied 

because of its medical effect in anticancer treatment (Kishi 

et al. 1991). 


Based on the SSU and ITS rDNA sequences analysis, its 

pleosporalean status was verified, and Shiraia was suggested 

to be closely related to Leptosphaeriaceae and/or 

Phaeosphaeriaceae (Pleosporineae) (Cheng et al. 2004). 

Based on the molecular phylogenetic analysis, another 

Shiraia-like fungus was reported which produced distinctive 

prawn-shaped conidioma-like structures (Morakotkarn 

et al. 2008), and differed from conidiomata in the anamorph 

of S. bambusicola described by Liu (2009). 

Concluding remarks A relatively broad species concept of 

Shiraia bambusicola is currently used, which could 

comprise several species. 

Sinodidymella J.Z. Yue & O.E. Erikss., Mycotaxon 24: 295 

(1985). (Teichosporaceae) 


Habitat terrestrial, saprobic? Ascomata medium to large, 

scattered, or in small groups, immersed, erumpent, to 

superficial, globose, subglobose, coriaceous, apex flattened, 

with radial ridges arranged around the central 

region. Peridium thick, 2-layered. Hamathecium of 

dense, broadly trabeculate pseudoparaphyses, anastomosing 

and branching between the asci. Asci 8-spored, with a 

short, furcate pedicel, bitunicate, cylindrical. Ascospores 

broadly ellipsoid, hyaline, becoming pale brown when 

mature, 1-septate, constricted at the median septum. 


Literature: Yue and Eriksson 1985. 

Type species 

Sinodidymella verrucosa (Petr.) J.Z. Yue & O.E. Erikss., 

Mycotaxon 24: 295 (1985). (Fig. 89) 

≡ Amphididymella verrucosa Petr., Meddn Göteb. Bot. 

17: 129 (1947). 


or in small groups, immersed, becoming erumpent, to 

nearly superficial, globose, subglobose, coriaceous, apex 

flattened, with 3–6 radial ridges arranged around the central 

region, with a flattened base not easily removed from the 

substrate, wall black, roughened (Fig. 89a and b). Peridium 

100–150 μm thick, thinner at the base, 2-layered, outer layer 

thin, up to 40 μm thick, composed of small heavily pigmented 

thick-walled cells of textura globulosa, cells up to 5 μm 

diam., cell wall 3–6 μm thick, inner layer thick, composed of


hyaline small cells of textura epidermoidea, 2–4 μm diam., 

cell wall 1–3 μm thick, interspersed with interwoven 

mycelium in places (Fig. 89b and c). Hamathecium of dense, 

broadly trabeculate pseudoparaphyses 1–2 μm broad, anastomosing 

between and above the asci (Fig. 89d). Asci 140–190 

(−205)×12.5–15(−17.5) μm (x ¼ 164 14:3mm, n=10), 8- 

spored, bitunicate, cylindrical, with a short, furcate pedicel, 

20–45 μm long, and an inconspicuous ocular chamber (to 

2 μm wide×1 μm high) (Fig. 89d and e). Ascospores 20–25× 

10–12 μm (x ¼ 22:1 10:3mm, n=10), obliquely uniseriate 

and partially overlapping, broadly ellipsoid with rounded 

ends, hyaline, becoming pale brown when mature, 1-septate, 

constricted at the median septum, smooth (Fig. 89f). 


Material examined: CHINA, Kansu Prov., between 

Scharakuto and Kweite, on rotten stems of Salsola gemmascens 

Pall., 25 Jul. 1935, G. Fenzel 2400 (W 16366, type). 

Notes 

Morphology 

Sinodidymella was formally established by Yue and 

Eriksson (1985) as they noticed that Amphididymella 

verrucosa Petr. was not congeneric with the generic type, 

A. adeana Petr., which is a pyrenolichen. Thus a new 

monotypic genus, Sinodidymella was introduced to accommodate 

it. The most outstanding morphological character of 

Sinodidymella is its radial ridges, which are somewhat 

comparable with that of Lophiostoma rugulosum Yin. 

Zhang, J. Fourn. & K.D. Hyde, although their pseudoparaphyses 

are dissimilar. Lophiostoma rugulosum has “tightly 

aggregated cellular pseudoparaphyses” and “apically ending 

into bunches of clavate cells” (Zhang et al. 2009b). 


None. 


The radial ridges have little phylogenetic significance in 

genus level classification (Zhang et al. 2009b), but the 

broadly trabeculate pseudoparaphyses of Sinodidymella 

may fit Melanommataceae. 

Splanchnonema Corda, in Sturm, Deutschl. Fl., 3 Abt. 

(Pilze Deutschl.) 2(9), Tome 3: 115 (1829). (?Pleomassariaceae) 



solitary or scattered, immersed in cortex with a pseudostromal 

covering, with a small ostiole appearing on the host 

surface, flattened subglobose. Peridium thin. Hamathecium 

Fig. 87 Setomelanomma holmii (from UPS F-117969 (slide), isotype). b 

a, b Asci with short pedicels in pseudoparaphyses. c Partial view of 

ascus. d Branching and septate pseudoparaphyses. a Three-septate 

lightly pigmented ascospores in ascus. Scale bars: a–e=10 μm 

of dense, cellular pseudoparaphyses, embedded in mucilage, 

anastomosing and branching. Asci bitunicate, fissitunicate, 

clavate to broadly cylindrical, with a short, 

narrowed, furcate pedicel. Ascospores clavate with a 

rounded apex and acute base, reddish brown, constricted 

at the septa. 

Anamorphs reported for genus: Myxocyclus, Steganosporium 

(Barr 1982b). 

Literature: Barr 1982b, 1993a; Boise 1985; Corda 1829; 

Eriksson 1981; Ramaley and Barr 1995; Shoemaker and 

LeClair 1975; Sivanesan 1984; Tanaka et al. 2005. 

Type species 

Splanchnonema pustulatum Corda, in Sturm, Deutschl. Fl., 

3 Abt. (Pilze Deutschl.) 2(9), Tome 3: 115 (1829). (Fig. 90) 


solitary or scattered, immersed in cortex with a pseudostromal 

covering, with a small ostiole appearing on the host surface, 

flattened subglobose (Fig. 90a). Peridium 15–25 μm thick, 

composed of small lightly pigmented thin-walled compressed 

cells (Fig. 90b). Hamathecium of dense, long cellular 

pseudoparaphyses 2–3 μm broad, embedded in mucilage, 

anastomosing and branching. Asci 200–250×30– 


fissitunicate, clavate to broadly cylindrical, with a short, 

narrowed, furcate pedicel up to 35 μm long, without 

conspicuous ocular chamber (Fig. 90c and d). Ascospores 

45–53×20–24 μm (x ¼ 48:5 22:3mm, n=10), obliquely 

uniseriate and partially overlapping to biseriate, clavate 

with a rounded apex and acute base, reddish brown, 2- 

septate, apical cell largest, broader than the lower cells, 

basal cell smallest, constricted at the septa, smoothwalled, 

surrounded by a regular hyaline gelatinous 

sheath, 3–6 μm thick (Fig. 90e and f). 


Material examined: UK, Avon, nr Bath, Batheaston, on 

branch of Ulmus, C.E. Broome (L, No. 910.251-352, No. 

910.251-371). 

Notes 

Morphology 

A confusing outline of the history of Splanchnonema 

was provided by Shoemaker and LeClair (1975), which at 

the time was a valid, but little used name. Eriksson (1981) 

and Sivanesan (1984) stated (without comment) that the 

lectotype of Splanchnonema is S. pupula (Fr.) O. Kuntze. 

However, S. pustulatum is listed as the generic type in the

Fungal Diversity


Fig. 88 Shiraia bambusium 

(from IFRD 2040). a Ascostroma 

form a nubby structures 

on the twigs of host. b Vertical 

section of an ascostroma. Note 

the reddish staining of the inner 

tissue. c, d Cylindrical asci with 

a short pedicel. e–g Muriform 

fusoid hyaline ascospores. Scale 

bars: a=1 cm, b=1 mm, c, d= 

50 μm, e–g=20 μm 

online databases MycoBank and Index Fungorum. We 

assume Eriksson (1981) gained his data from Shoemaker 

and LeClair (1973), who considered S. pustulatum to be a 

synonym of S. pupula. Since we were unable to locate 

material of Corda or Fries we used a later collection of C.E. 

Broome. 

Splanchnonema can be distinguished from the morphologically 

comparable genera, i.e. Pleomassaria or 

Splanchospora by its depressed ascomata, and obovoid 

and asymmetrical ascospores (Barr 1982b). Currently, 

about 40 species are included in this genus. Barr 

(1993a) provided a key to 27 North American species, 

however, the inclusion of species with a range of 

ascospore types and immersed to superficial ascomata 

suggests the genus to be polyphyletic. Tanaka et al. 

(2005) suspected that the genus might include species of 

Pleomassaria, thus this genus needs further study. 


Splanchnonema platani (= Massaria platani) is poorly 

supported to be related to Lentitheciaceae (Schoch et al. 

2009). 

Fig. 89 Sinodidymella verrucosa (from W 16366, type). a Ascomata b 

on the host surface. Note the radial ridges around the pseudostiolar 

region. b Section of an ascoma. c Section of peridium. Note the 

hyaline small cells and interwoven hyphae. d Cylindrical asci in 

pseudoparaphyses. e Eight-spored ascus with short pedicel. f Hyaline, 

1-septate ascospores which turn pale brown when mature. Scale bars: 

a=1 mm, b=100 μm, c=50 μm, d–f=20 μm

Fungal Diversity



Splanchnonema pustulatum has unique ascospores formed 

in immersed ascomata with thin walls, indicating that Splanchnonema 

sensu stricto shouldbeconfinedtoafewsimilar 

species. The type needs recollecting, sequencing and epitypifying 

in order to establish the phylogenetic relationships of 

this genus and to study what may be important defining 

characters. Also see entry under Pleomassaria. 

Sporormia De Not., Micromyc. Ital. Novi 5: 10 (1845). 



Habitat terrestrial, saprobic (coprophilous). Ascomata small, 

solitary, scattered, immersed to erumpent, globose, subglobose, 

wall black; apex without obvious papilla, ostiolate. 

Peridium thin. Hamathecium of rare, broad, septate pseudoparaphyses. 

Asci 8-spored, bitunicate, fissitunicate dehiscence 

not observed, short cylindrical, with a short, narrowed, 

furcate pedicel. Ascospores fasciculate, broadly filliform, 

reddish brown, multi-septate, easily separating into partspores, 

without visible germ-slits or pores. 

Fig. 90 Splanchnonema pustulatum (from L, No. 910.251–352, No. 

910.251–371). a Appearnce of ascomata on the host surface beneath a 

slightly raised area with minute ostiolar opening. b Section of the 

partial peridium. Note the compressed cells. c Dehiscent ascus. d 

Cluster of three asci joined in hymenium and pseudoparaphyses. e, f 

Asymmetric ascospores. Note the conspicuous sheath. Scale bars: a= 

1 mm, b–d=50 μm, e, f=20 μm



Literature: Ahmed and Asad 1968; Ahmed and Cain 1972; 

Kirschstein 1944; de Notaris 1849. 

Type species 

Sporormia fimetaria De Not., Micromyc. Ital. Novi 5: 10 

(1845). (Fig. 91) 

Ascomata 100–150 μm diam., solitary, scattered, 

immersed to erumpent, globose, subglobose, wall 

black; apex without obvious papilla, ostiolate 

(Fig. 91a). Peridium thin (other characters unknown). 

Hamathecium of rare, 2–3 μm wide, septate pseudoparaphyses. 

Asci 70–100×13–18 μm (x ¼ 86:4 14:9mm, 

n=10), 8-spored, bitunicate, fissitunicate dehiscence not 

observed, shortly cylindrical, with a short, narrowed, 

furcate pedicel up to 20 μm long, no apical apparatus 

could be observed (Fig. 91b, c and d). Ascospores 50– 

58×4–5 μm (x ¼ 54:7 4:8mm, n =10), fasciculate, 

broadly filliform, reddish brown, with 16 cells, easily 

separating into partspores, central cells of the ascospores 

shorter than broad, rectangular in vertical section, round 

in transverse section, 4–5×2.5–3.5 μm, without visible 

germ-slits or pores, apical cells usually longer than 

broad, 5–6.5 μm long, also without apertures (sheath is 

reported (Ahmed and Cain 1972), but not observed in 

this study) (Fig. 91e). 


Material examined: 1832, (RO, type, as Hormospora 

fimetaria De Not.). 

Notes 

Morphology 

Sporormia was formally established by de Notaris (1849), 

and only one species was described, i.e. S. fimetaria, which 

subsequently was selected as the generic type. Sporormia 

sensu stricto was accepted by several workers, and only 

includes members with a fasciculate ascospore arrangement, 

parallel to the ascus, and the part cells of the ascospores 

lacking germ-slits (Ahmed and Asad 1968; Ahmed and Cain 

1972; Kirschstein 1944). Species whose ascospores are not 

fasciculate and have partspores with germ-slits were assigned 

to Sporormiopsis by Kirschstein (1944) andtoSporormiella 

by Ahmed and Cain (1972). 


The generic status of Sporormia in Pleosporales was 

verified based on a phylogenetic analysis of ITS-nLSU 

rDNA, mtSSU rDNA and ß-tubulin sequences (Kruys and 

Wedin 2009). Sporormia clustered together with species of 

Westerdykella (including Eremodothis and Pycnidiophora), 

but lacks clear statistical support. Thus, the relationship of 

Sporormia with other genera of Sporormiaceae is unclear 

and not resolved yet. 


Several coprophilous taxa (e.g. Chaetopreussia and 

Pleophragmia as well as Sporormiella nigropurpurea) in 

the Pleosporales were not included in the study by Kruys 

and Wedin (2009). Strains of these genera need to be 

collected and analyzed and their relationship with Sporormia 

established. 

Trematosphaeria Fuckel, Jb. nassau. Ver. Naturk. 23–24: 

161 (1870). (Trematosphaeriaceae) 


Habitat terrestrial or freshwater, saprobic. Ascomata 

subglobose, unilocular, erumpent to superficial, with 

papillate ostiole. Peridium thin, comprising several cell 

types. Hamathecium of dense, delicate, filliform, septate 

pseudoparaphyses. Asci bitunicate, fissitunicate, 

cylindro-clavate, normally 8-spored. Ascospores 

ellipsoid-fusoid to biconic, septate, smooth to finely 

verruculose, brown. 

Anamorphs reported for genus: hyphopodia-like (Zhang 

et al. 2008a). 

Literature: von Arx and Müller 1975; Barr 1979a; 

Boise 1985; Clements and Shear 1931; Zhang et al. 

2008a. 

Type species 

Trematosphaeria pertusa (Pers.) Fuckel, Jb. nassau. Ver. 

Naturk. 23–24: 161 (1870). (Fig. 92) 

≡ Sphaeria pertusa Pers., Syn. meth. fung. (Göttingen) 

1: 83 (1801). 


solitary, scattered, or in groups, initially immersed, becoming 

erumpent, to semi-immersed, subglobose, black; apex 

with a short ostiole usually slightly conical and widely 

porate, to 100 μm high (Fig. 92a and b). Peridium 48– 

55 μm wide laterally, to 80 μm at the apex, thinner at the 

base, 30–40 μm thick, coriaceous, 3-layered, comprising 

several cell types, one is of small heavily pigmented thickwalled 

cells of textura angularis, cells 4–8 μm diam., cell 

wall 1.5–3 μm thick in places with columns of textura 

prismatica orientated perpendicular to the ascomatal surface, 

apex cells smaller and walls thicker, forming thickwalled 

cells of textura pseudoparenchymata, and larger, 

paler cells of mixture of textura epidermoidea and textura 

angularis at the base (Fig. 92b, c and h). Hamathecium of 

dense, filamentous, 1.5–2.5 μm broad, septate pseudoparaphyses, 

embedded in mucilage, branching and anastomos-


ing between and above the asci (Fig. 92d, e and f). Asci 

100–145×15–17 μm (x ¼ 118 15:5mm, n=10), 8-spored, 

bitunicate, fissitunicate, cylindro-clavate, with a short, 

thick, furcate pedicel which is 12–30 μm long, with a 

truncate ocular chamber (Fig. 92d, e, f, g and i). Ascospores 

27.5–32.5×7.5–8.5 μm (x ¼ 29:5 8mm, n=10), biseriate 

to uniseriate near the base, fusoid with broadly to narrowly 

rounded ends, dark brown, 1–3-septate, secondary septum 

forming late or often absent, constricted at the median 

septum, the upper cell often shorter and broader than the 

lower one, smooth to finely verruculose, containing 

refractive globules (Fig. 92j and k). 

Anamorph: Only hyphopodia-like structures (or conidia?) 

observed (Zhang et al. 2008a). 

Colonies (of epitype) reaching 5 cm diam. after 20 days 

growth on MEA at 25°C, raised, woolly, deep grey, with 

irregular to rhizoidal margin, reverse darkened. Hyphopodialike 

structures (or conidia?) produced after 6 months, hyaline 

to pale brown, lobed, 4–4.5(−5) μm long and 3–3.5 μm diam. 

Material examined: EUROPE, Upsala, on decaying 

wood, designated by Boise (1985), (L-Pers 910269–172, 

as Sphaeria pertusa Pers., neotype); FRANCE, Deux 

Sèvres, Sansais, Le Vanneau, Les Grandes Mottines, 

swamp, on bark of a dead stump of Fraxinus excelsior, 

25 Apr. 2004, J. Fournier (IFRD 2002, epitype); Haute 

Garonne, Avignonet, Canal du Midi, on submerged wood 

of Platanus in a canal, 23 Nov. 2006, Michel Delpont, det. 

J. Fournier (IFRD2003). 

Notes 

Morphology 

Trematosphaeria was formally established in ‘Rhenish 

fungi’ by Fuckel (1870) based on the broadly pertuse 

ascomata, and Fries (1823) assigned it under Ascomycetes, 

Pyrenomycetes, Lophiostomataceae. Subsequently, Winter 

(1885) placed Trematosphaeria in Amphisphaeriaceae. 

Berlese (1890), however, treated Trematosphaeria as a 

synonym of Melanomma (Melanommataceae). After establishment 

of Loculoascomycetes (Luttrell 1955), Trematosphaeria 

was assigned to Pleosporaceae (Loculoascomycetes, 

Pleosporales) (Holm1957), and this was followed by von 

Arx and Müller (1975). Trematosphaeria was assigned to 

Melanommataceae by Barr (1979a), and this has been 

widely followed (Eriksson 2006; Kirketal.2001; Lumbsch 

and Huhndorf 2007). 

Trematosphaeria pertusa, the lectotype species of Trematosphaeria 

(Clements and Shear 1931), is characterized by 

having semi-immersed to erumpent ascomata, filamentous 

pseudoparaphyses, cylindro-clavate asci, fusoid, 1-septate 

reddish brown to dark brown ascospores (Zhang et al. 

2008a). All of these characters are quite different from those 

of Melanomma, the familial type of Melanommataceae. 

Fig. 91 Sporormia fimetaria (from RO, type). a Appearance of 

ascomata on the host surface. Note the scattered distribution. b–d 

Broad cylindrical asci with a short and thick pedicel. e Released 

filiform ascospores which may break up into part spores. Scale bars: 

a=0.5 mm, b–d=20 μm, e=10 μm 


Trematosphaeria pertusa forms a robust phylogenetic 

clade with Falciformispora lignatilis and Halomassarina 

thalassiae, and they are all assigned to Trematosphaeriaceae 

(Suetrong et al. 2009; Zhang et al. 2009a; Plate 1). 


Trematosphaeria pertusa is a terrestrial species which 

can also survive in a freshwater environment. However, 

both Falciformispora lignatilis and Halomassarina thalassiae 

are marine fungi. Their habitat difference may indicate 

their distant relationship, at least above genus level. 

Verruculina Kohlm. & Volkm.-Kohlm., Mycol. Res. 94: 

689 (1990). (Testudinaceae) 


Habitat marine, saprobic. Ascomata medium-sized, solitary 

under clypeate, immersed to semi-immersed, subglobose to 

depressed ellipsoidal, papillate, ostiolate, periphysate, 

black, carbonaceous. Peridium thin, comprising a few 

layers of cells of textura angularis. Hamathecium of long 

cellular pseudoparaphyses, embedded in mucilage, hyaline, 

septate and sparsely branching. Asci 8-spored, bitunicate, 

fissitunicate, cylindrical, with short pedicels, ocular chamber 

not observed. Ascospores biseriate, ovoid or ellipsoidal, 

dark brown, 1-septate, constricted at the septum, verrucose 

or verruculose, with or without germ pore. 


Literature: Kohlmeyer and Volkmann-Kohlmeyer 1990; 

Suetrong et al. 2009. 

Type species 

Verruculina enalia (Kohlm.) Kohlm. & Volkm.-Kohlm., 

Mycol. Res. 94: 689 (1990). (Fig. 93) 

≡ Didymosphaeria enalia Kohlm., Ber. dt. bot. Ges. 79: 

28 (1966). 


under clypeate, immersed to semi-immersed, subglobose to 

depressed ellipsoidal, ostiolate, papillate, periphysate, black, 

carbonaceous. Peridium thin, comprising a few layers of cells 

of textura angularis. Hamathecium of long cellular pseudoparaphyses, 

1.5–2 μm broad, embedded in mucilage, hyaline, 

septate and sparsely branching. Asci 177–135×12.5–15.5 μm, 

8-spored, bitunicate, fissitunicate, cylindrical, with short 

furcate pedicels, ocular chamber not observed (Fig. 93a). 

b

Fungal Diversity


Ascospores 16.5–23×7.5–10 μm, biseriate, ovoid or ellipsoidal, 

dark brown, 1-septate, constricted at the septum, verrucose 

or verruculose, with or without germ pore (Fig. 93b). 


Material examined: SEYCHELLES, Victoria, on submerged 

branch of Rhizophora mangle L., Mar. 2004, K.D. 

Hyde (KDH 2137, slide). 

Notes 

Morphology 

Verruculina was introduced to accommodate an obligate 

marine species, i.e. Verruculina enalia (Kohlmeyer 

and Volkmann-Kohlmeyer 1990). Verruculina is characterized 

by immersed, clypeate, carbonaceous, ostiolate and 

papillate ascomata. The peridium is composed of cells of 

textura angularis. Pseudoparaphyses are trabeculate and 

embedded in mucilage. Asci are 8-spored, cylindrical with 

short pedicels and ocular chamber, and ascospores are 

ellipsoidal, 1-septate, dark brown, verrucose or verruculose. 

The partly or completely immersed clypeate ascomata 

of V. enalia is comparable with those of 

Didymosphaeria futilis, but it differs from the later by 

the dark peridium, gelatinous matrix around the pseudoparaphyses, 

stipitate asci with an ocular chamber, and the 

verruculose ascospores (Kohlmeyer and Volkmann- 

Kohlmeyer 1990). 


Based on multigene phylogenetic analysis, Verruculina 

enalia nested within Testudinaceae (Suetrong et al. 2009). 

Thus, its familial placement seems clarified. 


None. 

Westerdykella Stolk, Trans. Br. Mycol. Soc. 38: 422 




small, scattered on the upper layer of the culture medium, 

wall black. Peridium thin, composed of one layer of cells of 

polygonal, dark brown, thick-walled cells. Hamathecium 

not observed. Asci 32-spored, bitunicate nature undetermined, 

fissitunicate dehiscence not observed, subglobose to 

ellipsoid, arranged in the centre of the ascomata, with or 

without a short pedicel. Ascospores globose, brown, 1-celled, 

without germ pore. 

Anamorphs reported for genus: Phoma-like (von Arx 1974). 

Literature: von Arx 1973, 1981; Kruys et al. 2006; Kruys 

and Wedin 2009; Stolk 1955a. 

Fig. 92 Trematosphaeria pertusa (a, d, f–i from epitype, b, c, e, j b 

from neotype). a Ascomata on the host surface. b Section of an ascoma. 

c, h Section of the peridium. c shows the peridium structure at sides, 

and h indicates the basal peridium structure. Note the hyaline and thinwalled 

cells in (h). d Asci amongst pseudoparaphyses. e Ascus with 

pedicle. f, g Dehiscent ascus. i Upper part of the ascus, showing the 

ocular chamber and the mucilage covering the apex. j, k Ascospores. 

Scale bars: a=0.5 mm, b, c=100 μm, d–h=20 μm, i–k=10 μm 

Type species 

Westerdykella ornata Stolk, Trans. Br. Mycol. Soc. 38: 422 

(1955). (Fig. 94) 

Ascomata 100–300 μm diam., cleistothecoid, scattered 

on the upper layer of the culture medium, wall black 

(Fig. 94a). Peridium composed of one layer of cells of 

polygonal in front view, dark brown, thick-walled cells, 

ca. 5μm diam. Hamathecium not observed. Asci 25–32× 

16–22 μm, 32-spored, bitunicate nature undetermined, 

fissitunicate dehiscence not observed, subglobose to 

ellipsoid, arranged in the centre of the ascomata, with 

a short furcate pedicel best seen in immature asci 

(Fig. 94b, c, d and f). Ascospores 6.2–7×6–6.8 μm, 

globose, brown, 1-celled, ornamented with irregular spiral 

bands, which occur in four to five coils, without germ 

pore (Fig. 94e). 


On MEA colonies spreading, but somewhat erumpent, 

with moderate aerial mycelium and even, lobate margins; 

surface dirty white with luteous to orange patches; reverse 

orange to sienna. On PDA similar but with sparse aerial 

mycelium; surface with patches of orange to luteous and 

dirty white; reverse luteous with cream margins. On OA 

flat, spreading with sparse aerial mycelium; surface with 

luteous and dirty white patches and transparent margins; 

sporulating on OA, visible as black masses of aggregated 

ascomata; colonies reaching 4 cm diam. on all media (based 

on CBS 379.55). 

Material examined: MOZAMBIQUE, Inhaca, leg. H.J. 

Swart, mangrove mud (CBS 379.55, holotype). 

Notes 

Morphology 

Westerdykella was introduced to accommodate a 

coprophilous fungus, which is characterized by cleistothecioid 

and membraneous ascomata (Stolk 1955a). Asci 

are subglobose to ellipsoid, stalked, many-spored and 

evanescent. Ascospores are globose to subglobose, 

brown, ornamented with spiral bands, without germ pores 

(Stolk 1955a). Westerdykella was assigned under Phaeosporeae 

of the Eurotiaceae (Stolk 1955a), and was 

assigned to Sporormiaceae by von Arx and Müller 

(1975). Based on the spore ornamentation, von Arx and

Fungal Diversity


Fig. 93 Verruculina enalia (from KDH 2137, slide). a Cylindrical asci with short pedicels. b One-septate verruculose ascospores. Scale bars: 

a=20 μm, b=10 μm 

van der Aa (1987) andBarr(2000) acceptedWesterdykella as 

a separate genus, but this is not supported by molecular 

phylogenetic analysis (Kruys and Wedin 2009). 


Phylogenetic reconstructions indicated that both Pycnidiophora 

and Eremodothis should be treated as synonyms 

of Westerdykella (Kruys and Wedin 2009). 


Westerdykella is another example where ascospore ornamentation 

can be phylogenetically uninformative. Westerdykella 

is proved a good genus of Sporormiaceae (Kruys and Wedin 

2009). 

Wettsteinina Höhn., Sber. Akad. Wiss. Wien, Math.-naturw. 

Kl., Abt. I 116: 126 (1907). (?Lentitheciaceae) 


Habitat terrestrial or freshwater? hemibiotrophic or 

saprobic. Ascomata generally small, scattered, immersed 

with a protruding broad papilla. Peridium very thin, 

composed of few layers of thin-walled large polygonal 

cells in surface view. Hamathecium deliquescing at 

maturity. Asci bitunicate, fissitunicate, subglobose to 

obpyriform, without a pedicel, with small truncate ocular 

chamber. Ascospores hyaline and turning pale brown 

when mature, septate, upper second cell enlarged, slightly 

constricted at the second septum, smooth, surrounded by a 

hyaline gelatinous sheath. 

Anamorph reported for genus: Stagonospora (Farr et al. 

1989). 

Literature: Barr 1972; Müller 1950; Shoemaker and 

Babcock 1987, 1989b. 

Type species 

Wettsteinina gigaspora Höhn., Sber. Akad. Wiss. Wien, 

Math.-naturw. Kl., Abt. 1 116: 126 (1907). (Fig. 95) 

Ascomata 150–250 μm diam., scattered, immersed 

with protruding broad papillae, 50–90 μm diam. 

Peridium thin, composed of few layers of thin-walled 

large polygonal cells in surface view, 6–15 μm diam. 

(Fig. 95a). Hamathecium deliquescing at maturity. Asci 

140–200×75–120 μm, 8-spored, bitunicate, fissitunicate, 

subglobose to obpyriform, lacking a pedicel, with a small 

truncate ocular chamber (to 8 μm wide×5 μm high) 

(Fig. 95b and c). Ascospores 90–110×25–30 μm, 2–4- 

seriate, hyaline and turning pale brown when mature, 

broadly clavate, 4-septate, primary septum distinct and 

constricted forming 1/3 rd from the apex of the ascospore, 

complete, secondary septa less distinct and slightly 

constricted, incomplete, with one forming above and two 

forming below the primary septum, largest cell the second 

cell from apex, smooth, surrounded by a hyaline gelatinous 

sheath 5–8 μm thick (Fig. 95d and e). 


Material examined: SLOVENIA, Postojna, on Genista 

sagittalis leg. Stapf. det. H. Rehm. (S, holotype of 

Massarina gigantospora).


Fig. 94 Westerdykella ornata (from CBS 379.55, holotype). a Appearance of the ascomata on culture substrate surface. b–f Mature and 

immature asci as well as the released ascospores. Note the spiral bands around the ascospores. Scale bars: a=1 mm, b–f=10 μm 

Notes 

Morphology 

Confusion exists in the generic type of Wettsteinina. 

Höhnel (1907) described W. gigaspora when introducing 

Wettsteinina, and listed it as the first species of Wettsteinina. 

ClementsandShear(1931) accepted W. gigaspora as 

the generic type of Wettsteinina, which is followed by 

Shoemaker and Babcock (1987). But according to http:// 

www.indexfungorum.org (June 2011), W. gigantospora is 

the generic type of Wettsteinina. Both W. gigantospora 

and W. gigaspora were treated as the synonyms of W. 

mirabilis (Niessl) Höhn. http://www.indexfungorum.org 

(June, 2011, Synonymy Contributor: CBS (2010)). We


tentatively described the generic type of W. gigantospora 

as a representing of the type of W. gigaspora here. 

New family names, i.e. Pseudosphaeriaceae and Wettsteininaceae 

(as Wettsteiniaceae) and a new order, Pseudosphaeriales 

had been introduced to accommodate Wettsteinina and 

its synonym Pseudosphaeria (Höhnel 1907; Locquin 1972). 

After a systematic study, Wettsteinina was included in 

Pleosporaceae basedonits“Pleospora-type” centrum, and 

Pseudosphaeriaceae and Wettsteininaceae are treated as 

synonyms of Pleosporaceae (Shoemaker and Babcock 1987). 


Wettsteinina macrotheca (Rostr.) E. Müll., W. pachyasca 

(Niessl) Petr. and W. dryadis (Rostr.) Petr. were reported to be 

closely related to Pleomassaria siparia (Melanommataceae) 

(Kodsueb et al. 2006a), and W. lacustris (Fuckel) Shoemaker 

& C.E. Babc. nested within Lentitheciaceae (Schoch et al. 

2009). The generic type has not been sequenced. 


The most striking character for Wettsteinina is its 

asymmetrical ascospores, thick-walled obpyriform asci 

and lack of pseudoparaphyses at maturity. These 

characters are comparable with genera in the Capnodiales 

and Venturiales. The phylogenetic significance of 

these characters are not fully understood, while the 

hemibiotrophic or saprobic life style may indicate its 

polyphyletic nature (Shoemaker and Babcock 1987). 

Strains from the genus, in particular the generic type 

require DNA sequence data so that the phylogenetic 

placement can be investigated. 

Wilmia Dianese, Inácio & Dorn. -Silva, Mycologia 93: 

1014 (2001). (Phaeosphaeriaceae) 


Habitat terrestrial, hemibiotrophic or biotrophic. Ascomata 

small, scattered, immersed, globose to subglobose, papillate. 

Peridium thin, composed of a few layers of brown, thickwalled 

cells of textura angularis to prismatica. Hamathecium 

comprising filliform, septate, rarely branching, evanescent, 

cellular pseudoparaphyses embedded in mucilage. Asci 

bitunicate, fissitunicate, cylindrical to clavate, with a short, 

furcate pedicel and ocular chamber. Ascospores fusoid, pale 

brown, 1-septate. 

Anamorphs reported for genus: see below. 

Literature: Dianese et al. 2001. 

Type species 

Wilmia brasiliensis Dianese, Inácio & Dorn.-Silva, Mycologia 

93: 1014 (2001). (Fig. 96) 


immersed, globose to subglobose; apex with a short 

papilla, 40–80 μm long, ostiolate, periphysate, periphyses 

up to 90 μm long (Fig. 96a and b). Peridium 6–15 μm 

wide, 1-layered, composed of 3–7 layers of brown, thickwalled 

cells of textura angularis to prismatica, cells 4– 

9 μm diam., cell wall 2–4 μm thick (Fig. 96a and b). 

Hamathecium of long cellular pseudoparaphyses 2–3 μm 

broad, septate, rarely branching, embedded in mucilage, 

evanescent. Asci 65–95×9.5–14 μm (x ¼ 78:5 11:5mm, 

n=10), 8-spored, bitunicate, fissitunicate, cylindrical to 

clavate, with a short, furcate pedicel and a small ocular 

chamber (Fig. 96c, d and f). Ascospores 22.5–28×5– 

8.5 μm (x ¼ 26:5 6:8mm, n=10), biseriate, fusoid with 

narrowly rounded ends, pale brown, 1-septate, constricted 

at the septum, the upper cell often shorter and broader than 

the lower one, smooth, with or without sheath (Fig. 96d and 

e). 

Anamorph: Conidiomata 170–200 μm high×85–130 μm 

diam., eustromatic, immersed, subglobose to irregular, 

ostiolate, brown. Peridium thin, 1–2 walllayers,6–8 μm 

thick, thicker near the apex. Ostiole 50–63 μm high×30–35 

broad. Conidiogenous cells ampulliform or lageniform, 

phialidic, aseptate. Conidia 13–20×4–7 μm, ellipsoid, 

oblong, ovoid, hyaline (Dianese et al. 2001). 

Material examined: BRAZIL, Distrito Federal, Vargem 

Bonita, Fazenda Agua Limpa, on leaves of Memora 

pedunculata (Vell.) Miers, 18 May 1995, Carlos A. Inácio 

(UB Col. Microl 8438 holotype). 

Notes 

Morphology 

Wilmia was formally established by Dianese et al. (2001) 

as a monotypic genus represented by W. brasiliensis, which 

causes leaf spots on Memora pedunculata. The peridium of 

W. brasiliensis comprises a few layers of brown, thickwalled 

textura angularis to prismatica cells, and it also has 

cellular pseudoparaphyses, clavate asci, 1-septate pale 

brown ascospores (Dianese et al. 2001). 


None. 


The dicotyledonous host habit of Wilmia brasiliensis seems 

in agreement with Leptosphaeriaceae rather than Phaeosphaeriaceae. 

But a verified conclusion can only be reached by 

further molecular phylogenetic study. 

Xenolophium Syd., Bulletin of the Bernice P. Bishop 

Museum, Honolulu, Hawaii 19: 96 (1925). (Pleosporales, 

genera incertae sedis)


Fig. 95 Wettsteinina gigantospora 

(from S, holotype of 

Massarina gigantospora). a 

Ascomata with protruding papilla 

scattered on the host surface. 

b Obpyriform thick-walled 

ascus with small apical apparatus. 

c Fissitunicate ascus. d 

Released hyaline ascospores. 

Note the distinct primary septum 

and less distinct secondary septa. 

e Ascospore with sheath. 

Scale bars: a=0.5 mm, b–d= 

100 μm, e=50 μm 


Habitat terrestrial, saprobic on wood. Ascomata nearly 

superficial, scattered to gregarious, globose, large, with a 

conspicuous compressed papilla and large slit-like ostiole. 

Peridium carbonaceous. Hamathecium of dense, long 

trabeculate pseudoparaphyses, branching and anastomosing 

between and among asci. Asci 8-spored, clavate, with very


long furcate pedicels. Ascospores fusoid to narrowly fusoid, 

light to dark brown, 1-septate, constricted at the septum. 


Literature: Chesters and Bell 1970; Huhndorf 1993; 

Mugambi and Huhndorf 2009b; Müller and von Arx 

1962; Stevens 1925. 

Type species 

Xenolophium leve Syd., Bulletin of the Bernice P. Bishop 

Museum, Honolulu, Hawaii 19: 97 (1925) (Fig. 97) 

Current name: Xenolophium applanatum (Petch) 

Huhndorf, Mycologia 85: 493 (1993). 

≡ Schizostoma applanatum Petch, Ann. Roy. Bot. Gard. 

(Peradeniya) 6: 231 (1916). 

Ascomata 1–1.5 mm diam., scattered to clustered, erumpent 

to superficial, globose with base immersed in host tissue, wall 

black, carbonaceous, roughened with ridges, papillate. Apex 

with a conspicuous hysteriform papilla extending on the sides, 

1–1.4 mm long, 0.4–0.5 mm wide, 0.2–0.3 mm high, smooth, 

ostiole slit-like, nearly as long as papilla length (Fig. 97a). 

Peridium 140–160 μm thick, pseudoparenchymatous, composed 

of two distinct layers: outer crust 16–45 μm thick, 

blackish, of heavily melanized, nearly opaque thick-walled 

angular cells, of uneven thickness forming irregular strands 

extending into the inner layer; inner layer subhyaline, 

composed of thick-walled prismatic to angular cells, with 

columns or patches of darker thick-walled cells extending 

inwardly from the outer layer; papilla wall 200–220 μm thick, 

of heavily melanized angular thick-walled cells (Fig. 97b and 

c). Hamathecium of dense, very long trabeculate pseudoparaphyses 

0.8–1.5 μm broad, embedded in mucilage, anastomosing 

and branching between and above the asci. Asci 104–152× 

9–12 μm (excluding pedicel) (x ¼ 149 10:2mm, n=10), 8- 

spored, bitunicate, fissitunicate dehiscence not observed, 

clavate, with a long, narrowed, furcate pedicel which is 50– 

75 μm long (Fig. 97d). Ascospores 17–26×4–5.5 μm 

(x ¼ 22:5 4:8mm, n=10), upper biseriate and lower uniseriate, 

fusoid, straight to slightly curved, equally 1-septate, constricted 

at the septum, the upper cell slightly wider, with one or rarely 

two additional septa appearing on a small number of senescent 

ascospores, pale brown, median septum darker, constricted, 

smooth, without sheath or appendages (Fig. 97e, f and g). 


Material examined: MARTINIQUE, Morne Rouge, on 

rotten wood, leg C. Lécuru, det Jacques Fournier, 29 Aug. 

2007, IFRD 2038. 

Notes 

Morphology 

Xenolophium was formally established by Sydow (in 

Stevens 1925) to accommodate two species, i.e. X. leve and 

X. verrucosum, ofwhichX. leve is selected as the generic type 

(Huhndorf 1993). Because of its morphological similarity with 

some genera, such as Ostropella and Schizostoma, Xenolophium 

has been treated as a synonym of Ostropella (Müller 

and von Arx 1962)orevenofLophiostoma (Chesters and Bell 

1970). Huhndorf (1993) clarified the circumscription of 

Xenolophium and treated X. leve as a synonym of Schizostoma 

applanata. Xenolophium mainly differs from Ostropella in 

lack of “organized cell composition and triangular pattern of 

melanization” in the peridium (Huhndorf 1993). 


The polyphyletic nature of Xenolophium has been 

demonstrated (Mugambi and Huhndorf 2009b). The generic 

type of Xenolophium (X. leve, current name X. applanatum) 

clustered together with Ostropella albocincta (generic type 

of Ostropella), and both locate in Platystomaceae 

(Mugambi and Huhndorf 2009b). 


The large ascomata with slit-like ostioles, hamathecium of 

numerous and trabeculate pseudoparaphyses, clavate asci with 

long pedicels, and the pale brown, 1-septate ascospores of 

Xenolophium leve are all comparable with those of Ostropella 

albocincta. However, the phylogenetic results do not support 

them being congeneric (Mugambi and Huhndorf 2009b). 

Synonyms 

Javaria Boise, J.R., Acta Amazonica 14(Supl.): 50 (1984). 


Current name: Astrosphaeriella Syd. & P. Syd., Annls 

mycol. 11: 260 (1913). 


Habitat terrestrial, saprobic. Ascomata medium-sized, scattered, 

erumpent to nearly superficial, reflexed pieces of the 

ruptured host tissue usually persisting around the surface of 

the ascomata; ascomata broadly conical, with a flattened 

base not easily removed from the substrate, wall black, 

papillate. Peridium carbonaceous. Hamathecium of trabeculate 

pseudoparaphyses. Asci 8-spored, bitunicate, fissitunicate, 

cylindro-clavate to narrowly fusoid, with a short, 

narrowed, furcate pedicel. Ascospores elongate-fusoid, 

hyaline, 1-septate, constricted at the septum. 


Literature: Barr 1990a; Boise 1984. 

Type species 

Javaria samuelsii Boise, J.R., Acta Amazonica 14(Supl.): 

50 (1984) (Fig. 98)


Fig. 96 Wilmia brasiliensis (from UB Col. Microl 8438, holotype). a 

Section of an ascoma. Note the setae in the ostiole. b Conidioma of 

the coelomycetous anamorphic stage. c, d Clavate asci with short 

furcate pedicels. e, f Released 1-septate pale brown ascospores. Scale 

bars: a, b=100 μm, c, d=20 μm, e, f=10 μm


Fig. 97 Xenolophium applanatum (from IFRD 2038). a Gregarious 

ascomata on the host surface. Note protruding papilla and slit-like 

ostiole. b Vertical section of the papilla and ostiole. c Section of the 

partial peridium. Note the two layers of the peridium. d Eight-spored 

asci in trabeculate pseudoparaphyses. Note the long pedicels. e–g Pale 

brown ascospores. Scale bars: a=2 mm, b=200 μm, c=50 μm, d= 

20 μm, e–g=10 μm


Current name: Astrosphaeriella samuelsii Boise, Acta 

Amazon., Supl. 14(1–2, Suppl.): 50 (1986) [1984]. 

Ascomata 300–380 μm diam., scattered, erumpent 

through the outer layers of the host tissues, to nearly 

superficial, reflexed pieces of the ruptured host tissue 

usually persisting around the surface of the ascomata; 

ascomata broadly conical, with a flattened base not easily 

removed from the substrate, wall black, papillate (Fig. 98a). 

Peridium 50–80 μm thick, carbonaceous and crisp, 1- 

layered. Hamathecium of dense, long trabeculate pseudoparaphyses, 

0.8–1.5 μm broad, embedded in mucilage, 

anastomosing between and above the asci. Asci 140–185× 

17.5–20 μm (x ¼ 158 19:4mm, n=10), 8-spored, bitunicate, 

fissitunicate, cylindro-clavate to narrowly fusoid, with 

a short, narrowed, furcate pedicel up to 20 μm long 

(Fig. 98b and c). Ascospores 48–55(−60)×6–7.5(−10) μm 

(x ¼ 52:2 7:7mm, n=10), biseriate, elongate- fusoid, 

gradually tapering towards the ends, hyaline, surrounded 

with sheath, 2–5 μm thick, 1-septate, constricted at the 

septum (Fig. 98d). 


Material examined: Serra Araca, 60 m, terra firme, 

open forest, deep litter. Dry. 10–13 Mar. 1984, det. Jean R. 

Boise, G.J. Samuels (isotype). 

Notes 

Morphology 

Javaria was introduced by Boise (1984) based on 

seven Amazonian collections on decaying palm petioles; it 

is comparable with Astrosphaeriella in numerous characters. 

But Javaria differs from Astrosphaeriella by its 

hyaline ascospores with sheath, and its apical ring can be 

stained with Congo Red, as well as its small ascomata. 

Barr (1990a) introduced a second species J. shimekii 

which occurs on woody substrate. Some mycologists treat 

Javaria as a synonym of Astrosphaeriella (Hyde and 

Fröhlich 1998). 


None. 


The size of ascomata and pigmentation of ascospores has 

little significance at generic level classification (Zhang et al. 

2009a). Likewise, the staining of endotunica with Congo 

Red has not been shown to have great significance. Thus, 

we accept Javaria as a synonym of Astrosphaeriella. 

Pycnidiophora Clum, Mycologia 47: 899 (1955). 


Current name: Westerdykella Stolk, Trans. Br. Mycol. 

Soc. 38(4): 422 (1955). 



small, cleistothecial, scattered on surface of agar media, 

semi-immersed, globose to subglobose, black. Peridium 

thin, composed of thin-walled, polyangular cells from front 

view. Hamathecium not apparent. Asci numerous, irregularly 

arranged, bitunicate nature undetermined, fissitunicate 

nature undetermined, globose, without pedicel. Ascospores 

gathering in the globose asci, smooth. 

Anamorphs reported for genus: Phoma-like. 

Literature: Cain1961; Clum1955; Stolk 1955b; Thompson 

and Backus 1966. 

Type species 

Pycnidiophora dispersa Clum, Mycologia 47: 900 (1955) 

[1955]. (Fig. 99) 

Current name: Westerdykella dispersa (Clum) Cejp & 

Milko. 

Ascomata 200–290 μm diam., cleistothecial, scattered 

on surface of agar media, semi-immersed, globose to 

subglobose, black (Fig. 99a). Peridium thin, composed of 

thin-walled, poly-angular cells from front view (Fig. 99b). 

Hamathecium not apparent. Asci numerous, 11–14 μm 

diam. (x ¼ 12:3mm, n=10), irregularly arranged, 32-spored 

when mature, bitunicate nature undetermined, fissitunicate 

nature undetermined, globose, without pedicel (Fig. 99b 

and c). Ascospores 4–5.5×2.5–3 μm (x ¼ 4:7 2:8mm, n= 

10), in the globose asci, olivaceous, oblong, 1-celled, 

smooth (Fig. 99d). 

Anamorph: Phoma-like coelomycetes. 

On MEA colonies spreading, flat with sparse aerial 

mycelium, covering the dish after 1 month; surface smokegrey 

with dirty white margins; reverse olivaceous-grey with 

luteous patches. On PDA spreading without aerial mycelium, 

colonies transparent, sporulating profusely with black, globose 

ascomata and pycnidia of a Phoma-like anamorph. On 

OA similar, lacking aerial mycelium, sporulating profusely 

with black, globose ascomata (based on CBS 297.56). 

Material examined: USA, Michigan, East Lansing, 

Science Greenhouse, isolated from damped off Phlox seedling, 

Dec. 1952, F.M. Clum (No. 27) (MSC 133.118, type). 

Notes 

Morphology 

Pycnidiophora was formally established by Clum (1955) 

based on its “imperfect stage of pycnidium”, which was 

subsequently confirmed as the sexual stage (Cain 1961; 

Thompson and Backus 1966). Clum (1955) has described 

and tentatively assigned P. dispersa (Clum) Cain to


Fig. 98 Javaria samuelsii (from isotype). a Ascoma on the host surface. Note reflexed pieces of the ruptured host tissue. b, c Cylindro-clavate 

asci within narrow pseudoparaphyses in gelatinous matrix. d Released ascospore with sheath. Scale bars: a=1 mm, b=50 μm, c, d=20 μm 

Aspergillaceae (= Eurotiaceae), and Stolk (1955b) has 

proposed to assign the morphologically comparable species 

P. multispora Saito & Minoura ex Cain to Eurotiaceae as 

well. Cain (1961), however, suspected that the 32 asco-


spores are actually the disarticulated segments of eight 4- 

celled ascospores, thus assigned it under Preussia (Sporormiaceae). 

After detailed study, Thompson and Backus 

(1966) confirmed that the so-called “eight 4-celled ascospores” 

do not exist in the development of the asci in both 

P. dispersa and P. multisporum. Thus, Pycnidiophora was 

assigned to Eurotiaceae (Eurotiales) (Thompson and 

Backus 1966). 


Phylogenetic study based on the ITS-nLSU rDNA 

sequences indicated that Pycnidiophora dispersa nested 

within clade of Westerdykella (including the generic type, 

W. ornata) (Kruys and Wedin 2009). Morphologically, both 

genera have cleistothecioid ascomata, asci with short or 

without pedicels and ascospores 1-celled and no germ slits. 

Thus, Pycnidiophora is treated as a synonym of Westerdykella 

(Kruys and Wedin 2009). 


Although the pleosporalean status of Pycnidiophora is 

verified, morphological characters such as the cleistothecioid 

ascomata and irregularly arranged asci, which do not 

show typical bitunicate or fissitunicate characters, absence 

of pseudoparaphyses as well as the ascospores separating 

into partspores very early all challenge the traditional 

concept of Pleosporales (Zhang et al. 2009a). Obviously, 

most of these morphological characters overlap with those 

of the Eurotiales. 

Sporormiella Ellis & Everh., N. Amer. Pyren.: 136 (1892). 


Current name: Preussia Fuckel, Hedwigia 6: 175 

(1867) [1869–70]. 



medium-sized, solitary, scattered, or in small groups, 

semi-immersed to nearly superficial, globose, subglobose, 

black, coriaceous, ostiolate, periphysate. Peridium thin, 

composed of small heavily pigmented cells of textura 

angularis, apex cells smaller and walls thicker. Hamathecium 

of dense, septate, cellular pseudoparaphyses, embedded 

in mucilage. Asci 8-spored, bitunicate, fissitunicate, 

cylindro-clavate, with a narrowed, furcate pedicel. Ascospores 

cylindrical with rounded ends, brown, 3-septate, 

deeply constricted at each septa, with sigmoid germ slit in 

each cell. 


Literature: Ahmed and Cain 1972; Ellis and Everhart 

1892; Khan and Cain 1979a, b; Luck-Allen and Cain 1975. 

Type species 

Sporormiella nigropurpurea Ellis & Everh., N. Amer. 

Pyren.: 136 (1892). (Fig. 100) 

Current name: Preussia nigropurpurea (Ellis & Everh.) 

Kruys, Syst. Biod. 7: 476. 

Ascomata 314–528 μm high×(250-)357–500 μm diam., 

solitary, scattered, or in small groups, immersed, semiimmersed 

to nearly superficial, globose, subglobose, wall 

black, coriaceous, smooth, papillate, papilla 43–115 μm long, 

72–157 μm broad, ostiolate, ostiole filled with periphyses 

(Fig. 100a and b). Peridium 20–28 μm thick laterally, up to 

40 μm thick at the apex, composed of small heavily pigmented 

cells of textura angularis, cells 5–8 μm diam., cell wall 

1–3 μm thick, apex cells smaller and walls thicker 

(Fig. 100c). Hamathecium of dense, long, septate, cellular 

pseudoparaphyses, 1.5–2 μm broad, embedded in mucilage. 

Asci (70-)110–158×9–12.5(−15) μm (x ¼ 114:3 11:1mm, n= 

10), 8-spored, bitunicate, fissitunicate, cylindrical to cylindroclavate, 

with a narrowed, furcate pedicel, 13–38 μm long, 

ocular chamber apparent (Fig. 100d and e). Ascospores 15– 

20×4–5.5 μm (x ¼ 17:3 4:9mm, n=10), obliquely uniseriate 

and partially overlapping to biseriate, shortly cylindrical 

with rounded ends, brown, 3-septate, deeply constricted at 

each septum, with sigmoid germ slit in each cell, smoothwalled 

(Fig. 100f and g). 


Material examined: USA, New field, New Jersey: 

Gloucester Co., on cow dung, Mar. 1891 (NY, holotype). 

Notes 

Morphology 

Sporormiella was formally established by Ellis and 

Everhart (1892) based on the single species, Sporormiella 

nigropurpurea, which is characterized by its “immersed to 

semi-immersed, papillate ascomata, cylindrical to cylindroclavate 

asci with a pedicel, three to multi-septate ascospores 

with elongated germ slits through the whole cell” (Ahmed 

and Cain 1972; Khan and Cain 1979a, b). Barr (1990a) has 

indicated that Sporormiella might be a synonym of 

Ohleriella, while Sporormiella is assigned to Sporormiaceae 

as a separate genus (Eriksson 2006; Lumbsch and 

Huhndorf 2007). Currently, about 90 species are included 

in this genus (http://www.mycobank.org). 


The phylogenetic analysis based on ITS-nLSU rDNA, 

mtSSU rDNA and ß-tubulin sequences indicated that Sporormiella 

nested in Preussia,andaSporormiella–Preussia complex 

is formed (Kruys and Wedin 2009). Thus, Sporormiella was 

assigned under Preussia (Kruys and Wedin 2009).


Fig. 99 Pycnidiophora dispersa (a from CBS 297.56; b-d from MSC 

133.118, type). a Ascomata scattering on the surface of the substrate. 

b Crashed ascoma. Note the numerous released asci. c Globose asci 


It is clear that the presence or absence of an ostiole cannot 

distinguish Sporormiella from Preussia according to the 

findings of Guarro et al. (1997a, b) and Kruys and Wedin 

(2009). Thus, Sporormiella should be treated as a synonym 

of Preussia (Kruys and Wedin 2009). 

Spororminula Arx & Aa, Trans. Br. Mycol. Soc. 89: 117 


Current name: Preussia Fuckel, Hedwigia 6: 175 

(1867) [1869–70]. 


Habitat terrestrial, saprobic (coprophilous). Ascomata small 

to medium, solitary, scattered, immersed to erumpent, 

globose, subglobose, to ovate, black, membraneous, papillate, 

ostiolate. Peridium thin, membraneous, composed of 

and released ascospores. d One-celled ascospores. Scale bars: a= 

200 μm, b–d=20 μm 

several layers of heavily pigmented, elongate cells of 

textura angularis. Hamathecium of dense trabeculate, 

aseptate, decomposing pseudoparaphyses. Asci bitunicate, 

broadly cylindro-clavate with a narrow furcated pedicel. 

Ascospores cylindrical to cylindro-clavate, with round ends, 

brown, multi-septate, easily breaking into partspores. 


Literature: von Arx and van der Aa 1987. 

Type species 

Spororminula tenerifae Arx & Aa, Trans. Br. Mycol. Soc. 

89: 117 (1987).(Fig. 101) 

Current name: Preussia tenerifae (Arx & Aa) Kruys, 

Syst. Biod. 7: 476. 

Ascomata 290–400 μm diam., solitary, scattered, initially 

immersed, becoming erumpent when mature, globose, 

subglobose to ovate, black, membraneous, with a cylindrical 

or somewhat conical beak, 90–150(−230) μm broad and 110–


Fig. 100 Sporormiella nigropurpurea (from NY, holotype). a 

Section of an ascoma. b Section of the papilla. Note the dense 

pseudoparaphyses. c Section of a partial peridium. d, e Eight-spored 

cylindro-clavate asci with furcate pedicels. f, g Four-celled, brown 

ascospores. Note the sigmoid germ slit in each cell. Scale bars: a= 

200 μm, b, c=50 μm, d, e=20 μm, f, g=10 μm


190 μm high (Fig. 101a). Peridium 20–33 μm thick, 1- 

layered, composed of several layers of heavily pigmented, 

elongate cells of textura angularis, cells up to 6.3×5 μm 

diam., cell wall 1–1.5 μm thick (Fig. 101b and c). 

Hamathecium of dense, long trabeculate pseudoparaphyses 

1–2 μm broad, hyaline, aseptate, decomposing when mature. 

Asci 165–220×33–42.5 μm, 8-spored, bitunicate, broadly 

clavate, with a small, thin and furcate pedicel, 35–50 μm long, 

3–5 μm broad, ocular chamber not observed (Fig. 101d and 

e). Ascospores 68–93×12.5–16 μm, 3–4 seriate to uniseriate 

near the base, cylindrical to cylindro-clavate with rounded 

ends, brown, (6-)7 transverse septa, easily breaking into 

partspores, central cells triangular in transverse section but 

rectangular in vertical section, 14–16.5×8–10 μm long, apical 

cells 12.5–15×11.5–17.5 μm long (Fig. 101f and g). 


Material examined: SPAIN, Canary Islands, Tenerifa 

Las Canadas, on rabbit? droppings, Mar. 1986, J.A. von 

Arx (HCBS 9812, holotype). 

Notes 

Morphology 

Spororminula was formally established by von Arx and van 

der Aa (1987) according to its “ostiolate ascomata, elongated 

ascospore separated into part cells by transverse septa and 

without germ slits”, and was monotypified by S. tenerifae. 

Currently, only one species was included in this genus. 


Based on a phylogenetic analysis of ITS-nLSU rDNA, mtSSU 

rDNA and ß-tubulin sequences, Spororminula tenerifae 

nested in the clade of Preussia, thus Spororminula was 

treated as a synonym of Preussia (Kruys and Wedin 2009). 


To clarify its relationship with other genera of Sporormiaceae, 

further phylogenetic study is needed, which should 

include additional related taxa. 

Excluded and doubtful genera 

Kriegeriella Höhn., Annls mycol. 16: 39 (1918). (Dothideomycetes, 

families incertae sedis, Microthyriaceae) 


Habitat terrestrial, saprobic? Ascomata small, solitary, 

scattered, superficial, subglobose, black, roughened, apex 

Fig. 101 Spororminula tenerifae (from HCBS 9812, holotype). a b 

Appearance of ascomata on the host surface. b, c Sections of the 

partial peridium. Note the elongate cells of textura angularis. d, e 

Asci with thin pedicels. f, g Ascospores, which may break into part 

spores. Scale bars: a=0.5 mm, b=100 μm, c=50 μm, d–g=20 μm 

no obvious opening. Peridium thin, composed of a single 

type of lightly pigmented thin-walled cells. Hamathecium 

long cellular pseudoparaphyses, septate. Asci 8-spored, 

bitunicate, obpyriform. Ascospores hyaline, turning brown 

when mature, multi-septate, constricted at each septum. 


Literature: von Arx and Müller 1975; Barr1975, 1987b; 

Eriksson 2006; Lumbsch and Huhndorf 2007. 

Type species 

Kriegeriella mirabilis Höhn., Annls mycol. 16: 39 (1918) 

(Fig. 102) 


scattered, superficial, with basal wall flattened on the surface of 

the substrate, subglobose, black, roughened, apex no obvious 

opening (Fig. 102a and e). Peridium thin, composed of a 

single type of lightly pigmented thin-walled cells, cells up to 

12×5 μm diam. in front view, cell wall less than 1 μm thick, 

apex cells smaller and walls thicker (Fig. 102a and f). 

Hamathecium long cellular pseudoparaphyses, 1.5–2 μm 

wide, septate. Asci 65–85×31–36 μm (x ¼ 63:1 33mm, n= 

10), 8-spored, bitunicate, fissitunicate undetermined, obpyriform, 

no pedicel, no ocular chamber was seen (Fig. 102c and 

g). Ascospores 28–37.5×8–11 μm (x ¼ 32:6 10mm, n= 

10), 8-seriate, fusoid with broadly rounded ends, hyaline, 

turning brown when mature, 5–6-septate, constricted at each 

septum, the upper first-cell longer and broader than the lower 

ones with semi-round shape, smooth (Fig. 102d, h, i). 

Anamorph: see Fig. b. 

Material examined: On the leaves of Faulenden nadeln 

von Pinus silvestris, bei Roñigstein, Sept. 1896, W. Rueges. 

(S reg. nr F12638, isolectotype). 

Notes 

Morphology 

Kriegeriella was formally established by von Höhnel (1918b) 

and was represented by two species, i.e. K. mirabilis and K. 

transiens; it was typified by K. mirabilis and assigned to 

Microthyriaceae. Subsequently, Kriegeriella was assigned to 

the subfamily of Aulographiodeae (Microthyriaceae) 

(Batista et al. 1959), Asterinaceae (Hemisphaeriales) (Luttrell 

1973) and Pseudosphaeriaceae (Dothideales) (Barr 

1975). 

After checking the original description and the type 

specimens of K. mirabilis and K. transiens, no significant

Fungal Diversity


difference could be observed, and both are described from 

rotting needles of conifers (Barr 1975; Batista et al. 1959; 

Höhnel 1918b). Morphologically, Extrawettsteinina is 

comparable with Kriegeriella. In particularly, E. pinastri 

could not be distinguished from K. transiens or K. 

mirabilis. Thus, K. transiens including Extrawettsteinina 

pinastri was treated as synonyms of K. mirabilis, andwas 

included in the section of Kriegeriella under the genus 

Kriegeriella (von Arx and Müller 1975; Barr1975). The 

other section of Kriegeriella, Extrawettsteinina, includes 

two previous Extrawettsteinina species, i.e. K. minuta and 

K. mediterranea. Barr(1987b) introduced a family, i.e. 

Kriegeriellaceae (Dothideales) to accommodate Kriegeriella 

and Extrawettsteinina. This proposal is rarely 

followed, and Kriegeriella is usually assigned to Pleosporaceae 

(Pleosporales) (Eriksson2006; Lumbsch and Huhndorf 

2007). 


None. 


Kriegeriella might belong to Microthyriaceae, although it 

would be unusual in this family in having 5-6-septate 

ascospores. Micropeltidaceae better accommodates the 

ascospores, however, the parallel arrangement of cells of 

the upper peridium are not typical. Asterinaceae may be 

most suitable as Luttrell (1973) suggested. 

Phaeotrichum Cain & M.E. Barr, Can. J. Bot. 34: 676 

(1956). (Dothideomycetes, family incertae sedis, 

Phaeotrichaceae) 



small, cleistothecial, solitary, or in small groups, 

superficial, with long straight or slightly flexed, thin, 

black appendages evenly scattered on the surface of the 

ascomata, globose, black. Peridium thin, carbonaceousmembraneous, 

1-layered, composed of dark brown thickwalled 

cells of textura angularis. Hamathecium not 

observed. Asci bitunicate form not clear, fissitunicate 

dehiscence not observed, broadly clavate, with a relatively 

thick pedicel. Ascospores oblong to ellipsoid, 

hyaline when young, turning reddish brown at maturity, 

1-septate, deeply constricted at the septum, each end with 

a subhyaline and broadly rounded germ pore, readily 

forming partspores at the septum at maturity. 


Literature: Cain 1956; Malloch and Cain 1972. 

Type species 

Phaeotrichum hystricinum Cain & M.E. Barr, Can. J. Bot. 

34: 677 (1956). (Fig. 103) 

(Some information for the following description is from 

Cain 1956) 

Ascomata 170–280 μm diam., cleistothecial, solitary, or 

in small groups, superficial, with 15–20 long straight or 

slightly flexed, thin, black appendages evenly scattered on 

the surface of the ascomata, 0.5–1 mm long, 15–25 μm 

wide at base, tapering to less than 5 μm at the blunt apex, 

with few or without septa, globose, black, smooth 

(Fig. 103a). Peridium thin, carbonaceous-membraneous, 

1-layered, composed of dark brown thick-walled cells 

of textura angularis, cells 8–16 μm diam., cell wall 

0.5–1.5 μm thick (data obtained from Cain 1956) 

(Fig. 103b). Hamathecium not observed. Asci 42–48× 

14–17 μm, 8-spored, bitunicate form not typical, lacking 

fissitunicate dehiscence, broadly clavate, with a relatively 

thick pedicel which is about 18 μm (data obtained from 

Cain 1956). Ascospores 14–16×4–5 μm, 4-seriate, oblong 

to ellipsoid, hyaline when young, turning reddish 

brown at maturity, 1-septate, deeply constricted at the 

septum, each end with a subhyaline and broadly rounded 

germ pore, smooth, readily separating into partspores at 

the septum at maturity (Fig. 103c, d, e and f). 


Material examined: CANADA, Ontario, Muskoka, 

Stoneleigh, on porcupine dung, 18 Aug. 1932, Cain (TRTC 

4361, holotype). 

Note: the ascomata of the specimen are fragile and no 

asci could be obtained. 

Notes 

Morphology 

Phaeotrichum was formally established by Cain (1956) 

to accommodate two new coprophilous fungi, i.e. P. 

hystricinum and P. circinatum Cain, and P. hystricinum 

was selected as the generic type. Phaeotrichum is 

mainly characterized by its coprophilous habitat, superficial 

cleistothecial ascocarps covered by long hairy 

appendages, reddish brown 1-septate ascospore with a 

broadly rounded germ pore at each end, readily breaking 

into partspores (Cain 1956). According to Cain (1956), 

Phaeotrichum possesses untypical bitunicate ascus, and 

the ascospore releasing is described as “simply break 

down and allow the contents to become free in the 

cavity of the ascocarp”. This ascospore releasing 

mechanism is considered as evolutionarily developed 

compared to those that “discharge the ascospores 

through an apical pore” (Cain 1956). Although lacking


Fig. 102 Kriegeriella mirabilis (from S reg. nr F12638, isolectotype). 

a Section of a superficial ascoma. b Anamorphic stage. c 

Obpyriform ascus. Note the pigmented ascospores and hyaline 

ascospores coexisted in a single ascus. d Ascospores. Scale bars: a= 

50 μm, b–d=10 μm. e Ascomata on the host surface. f, g Crashed 

ascoma. Note the peridium structure. h, i Hyaline asymmetric 

ascospores. Scale bars: e, f =100 μm, g=50 μm, h, i=10 μm 

a typical bitunicate ascus, Phaeotrichum is still 

assigned to Pleosporales, because the lack of bitunicate 

ascus does not “taken by itself, exclude a fungus from 

close relationship” (Cain 1956).

Fig. 102 (continued) 

Fungal Diversity



A single unverified isolate of Phaeotrichum benjaminii is 

placed well outside of Pleosporales in a broad phylogenetic 

study (Schoch et al. 2009). 


The superficial cleistothecial ascocarps covered by long hairy 

appendages, the absence of hamathecium as well as the 

nontypical bitunicate ascus are all distinct from members of 

Fig. 103 Phaeotrichum hystricinum 

(from TRTC 4361, holotype). 

a Superficial ascomata on 

host surface. Note the long and 

black appendages. b Part of 

peridium. Note the large cells in 

surface view. c–f Released reddish 

brown ascospores with hyaline 

end cells. Note the strongly 

constricted middle septum. 

Scale bars: a=0.5 mm, 

b–f=20 μm


Pleosporales, but definite conclusions could only be obtained 

by further molecular phylogenetic analysis. In this study, we 

assign it to Dothideomycetes incertae cedis. 

Zeuctomorpha Sivan., P.M. Kirk & Govindu, Bitunicate 

Ascomycetes and their Anamorphs: 572 (1984). (Venturiaceae) 


Habitat terrestrial, hemibiotrophic. Ascomata small, gregarious, 

superficial, globose to slightly flattened, ostiolate, 

covered with setae. Peridium thin, composed of heavily 

pigmented pseudoparenchymatous cells of textura angularis. 

Hamathecium of rare, septate, branching and anastomosing 

pseudoparaphyses. Asci 8-spored, with a short thick pedicel, 

bitunicate, fissitunicate, broadly clavate to obclavate. Ascospores 

ellipsoid, dark brown, 1-septate, asymmetrical, deeply 

constricted at the septum. 

Anamorphs reported for genus: Acroconidiellina (Sivanesan 

1984). 

Literature: Sivanesan 1984. 

Type species 

Zeuctomorpha arecae Sivan., P.M. Kirk & Govindu, in 

Sivanesan, Bitunicate Ascomycetes and their Anamorphs: 

572 (1984). (Fig. 104) 

Ascomata 175–300 μm diam., gregarious, superficial, 

globose to slightly flattened, collapsed at the apex when 

dry, ostiolate, covered with numerous long setae 

(Fig. 104a). Peridium up to 25 μm wide, composed of 

heavily pigmented pseudoparenchymatous cells of textura 

angularis, to7μm diam. Hamathecium of rare, 2–5 μm 

broad, septate, branching and anastomosing pseudoparaphyses. 

Asci 83–185×29–40(−50) μm (x ¼ 134 35:3mm, n= 

10), 8-spored, bitunicate, fissitunicate, broadly clavate to 

obclavate, with a short thick pedicel, up to 40 μm long, 

apically rounded, with a small ocular chamber (to 4 μm 

wide×7 μm high) (Fig. 104b, c and d). Ascospores 35–43× 

12.5–18 μm (x ¼ 36:5 15:4mm, n=10), 2–4 seriate, ellipsoid, 

dark brown, 1-septate, deeply constricted at the septum, 

usually slightly asymmetric, smooth (Fig. 104e,f,g,handi). 

Anamorph: Acroconidiellina arecae (Sivanesan 1984). 

Material examined: INDIA, Shimogee, on Areca 

catechu L. leaf, 1 Nov. 1979, H.C. Govindu (IMI 246067, 

holotype). 

Notes 

Morphology 

Zeuctomorpha was formally established by Sivanesan 

(1984) based on its superficial setose ascomata, clavate 

asci, ellipsoid and 1-septate ascospores, and presence of 

pseudoparaphyses, and was monotypified by Z. arecae. 

Zeuctomorpha arecae is widely distributed in tropical 

regions of East South Asia exclusively on the leaves of 

Areca catechu (Sivanesan 1984). 


None. 


This taxon is unusual amongst the Pleosporaceae as it 

has hairy superficial ascomata, few pseudoparaphyses, 

broadly clavate to obclavate asci and 1-septate pigmented 

ascospores. All of these morphological characters are 

most comparable with species of Acantharia, which 

might be closely related to Venturiaceae (Zhang et al. 

data unpublished). 

Muroia I. Hino & Katum., J. Jap. Bot. 33: 79 (1958). 

(Ascomycota) 


Habitat terrestrial, saprobic or parasitic. Ascostromata erumpent 

through the host surface in linear rows parallel to the 

host fibers. Ascomata small- to medium-sized, semiimmersed 

to erumpent, subglobose to rectangular, black, 

coriaceous, cells of ascostromata pseudoparenchymatous, 

cells of peridium composed of pigmented cells of textura 

angularis. Hamathecium of rare, pseudoparaphyses. Asci 

bitunicate, clavate to cylindro-clavate. Ascospores oblong to 

elongated oblong, hyaline, 1-celled, usually slightly curved. 


Literature: Hino and Katumoto 1958. 

Type species 

Muroia nipponica I. Hino & Katum., J. Jap. Bot. 33: 79 

(1958). (Fig. 105) 

Ascostroma 1–6 mm long, 360–470 μm broad, linear 

parallel to the host fibers with several linearly arranged 

ascomata (Fig. 105a). Ascomata 250–400 μm diam., 

semi-immersed in substrate to erumpent, subglobose to 

rectangular with a furrow-shaped ostiole, black, coriaceous, 

cells of ascostromata pseudoparenchymatous. 

Peridium composed of pigmented cells of textura 

angularis. Hamathecium of rare, 3–4.5 μm broad 

pseudoparaphyses. Asci (120-)150–190×30–45 μm, 8- 

spored, bitunicate, fissitunicate dehiscence not observed, 

clavate to cylindro-clavate, with a short, thin, knob-like 

pedicel, lacking an ocular chamber (Fig. 105b). Ascospores 

43–50×13–18 μm (x ¼ 46:6 15:2mm, n=10), biseriate, ob-


Fig. 104 Zeuctomorpha arecae (from IMI 246067, holotype). a 

Gregarious ascomata on host surface. Note the numerous setae on the 

surface of ascomata. b Asci with ocular chamber and short peduncles. 

c, d Ascus with ocular chamber and knob-like pedicel. e–i One septate 

ascospores which are slightly asymmetrical. Scale bars: a=0.5 mm, 

b–i=20 μm


long to elongated oblong, hyaline, 1-celled, usually slightly 

curved (Fig. 105c,d and e). 


Material examined: JAPAN, Province Ugo. on moribund 

culm of Sasa kurilensis, 4 Aug. 1957, coll. H. Muroi, 

Det. I. Hino & K. Katumoto (TNS-F-230252, isotype). 

Notes 

Morphology 

Muroia was introduced based on M. nipponica, which is a 

parasite on the lower part of Sasa kurilensis (Hino and 

Katumoto 1958). Muroia is characterized by its 1-celled 

ascospores. Considering the perithecial structure and linear 

ostiole, it was assigned to the Lophiostomataceae, and was 

regarded as closely related to the amerosporous Lophiella 

(Hino and Katumoto 1958). 


None. 


The linear ascostroma and 1-celled, hyaline ascospores 

make it less likely to fit the concept of Lophiostomataceae. 

Because of the condition of the specimen, its bitunicate 

nature could not be confirmed. 

Genera not studied 

Aglaospora De Not., G. bot. ital. 2: 43 (1844). 

Type species: Aglaospora profusa (Fr.) De Not., G. bot. 

ital. 2: 43 (1844). 

Aglaospora, which was introduced by de Notaris 

(1844), has 35 species epithets (http://www.mycobank. 

org/mycotaxo.aspx) and was considered to be a synonym 

of Massaria (Voglmayr and Jaklitsch 2011) orseparate 

(Barr 1990a). In a recent phylogenetic study, Voglmayr and 

Jaklitsch (2011) confirmed that Aglaospora is a synonym of 

Massaria and is treated as such here. The immersed 

ascomata with short beaks, together with ascostroma under 

pseudostromatic tissues, cylindrical asci with a large and 

refractive apical ring, trabeculate pseudoparaphyses within a 

gel matrix, and distoseptate ascospores, are all similar to 

species of Massaria. The large and conspicuous apical ring 

of the ascus of Aglaospora has the appearance of being 

unitunicate, and thus Shoemaker and Kokko (1977) treatedit 

as a unitunicate taxon. Currently, its bitunicate status is 

widely accepted. 

Allewia E.G. Simmons, Mycotaxon 38: 260 (1990). 

Type species: Allewia proteae E.G. Simmons, Mycotaxon 

38: 262 (1990). 

Allewia was introduced by Simmons (1990) toaccommodate 

Lewia-like species but with Embellisia anamorphs. 

Embellisia differs from other similar genera by a combination 

of characters including the percentage of dictyoconidia, 

shape of conidia, thickness of septa, umbilicate sites of 

conidiophore geniculation, proliferating chlamydospores and 

hyphal coils in culture (Simmons 1971). Based on multigene 

phylogenetic analysis, A. eureka, which is closely related to 

A. proteae, clustered together with species of Alternaria. 

Thus, Allewia should be treated as a synonym of Lewia. 

Anteaglonium Mugambi & Huhndorf, System. Biodivers. 

7: 460 (2009). 

Type species: Anteaglonium abbreviatum (Schwein.) 

Mugambi & Huhndorf, System. Biodivers. 7: 460 (2009). 

≡ Hysterium abbreviatum Schwein., Trans. Am. phil. 

Soc., New Series 4: no. 2094 (1832). 

Anteaglonium was introduced to accommodate a monophyletic 

hysterothecial clade within Pleosporales, and four 

species (A. abbreviatum, A. globosum Mugambi & Huhndorf, 

A. parvulum (W.R. Gerard) Mugambi & Huhndorf 

and A. latirostrum Mugambi & Huhndorf) are included 

(Mugambi and Huhndorf 2009a). Anteaglonium is characterized 

by erumpent to superficial, globose to subglobose or 

elongate, fusoid to oblong ascomata, which are brown to 

shiny black, opening by a pronounced or indistinct 

longitudinal slit running entire length of fruit body or apex 

raised and laterally compressed; asci cylindrical with short 

pedicel, 8-spored, uniseriate or biseriate; ascospores fusoid 

to oblong, septate, constricted at the primary septum, 

hyaline or pigmented. A phylogenetic analysis based on 

DNA comparisons indicated that Anteaglonium resides as a 

separate clade but related to Tetraplosphaeria, Lophiotrema 

and other species without clear resolution. Therefore, the 

familial placement of Anteaglonium remains unclear 

(Mugambi and Huhndorf 2009a). 

Arthopyrenia A. Massal., Ric. auton. lich. crost. (Verona): 

165 (1852). 

Type species: Arthopyrenia rhyponta (Ach.) A. Massal., 

Ric. auton. lich. crost. (Verona): 166, fig. 329 (1852). 

≡ Verrucaria rhyponta Ach., K. Vetensk-Acad. Nya 

Handl.: 150 (1809). 

Arthopyrenia is a lichen genus with a Trentepohlia 

photobiont and is characterized by dimidiate perithecoid 

ascomata, which are scattered to irregularly confluent, and 

have an upper thick clypeate wall composed of periderm 

cells intermixed with dark hyphae. The pseudoparaphyses are 

branched and asci are obpyriform, obclavate to subcylindrical 

and 8-spored. Ascospores are oblong, ovoid, slipper-shaped, 

1-3-septate, hyaline and smooth-walled (Coppins 1988;


Fig. 105 Muroia nipponica (TNS-F-230252, isotype). a Linear ascostroma parallel to the host fibers. b Crashed ascus with ascospores released. 

c–e Released hyaline ascospores. Scale bars: a=5 mm, b–e=30 μm 

Upreti and Pant 1993). Multigene phylogenetic studies 

indicated that Arthopyrenia salicis, a typical species of 

Arthopyrenia, is located within Pleosporales in close 

proximity to bambusicolous species in the genus Roussoella, 

with its familial status remaining undetermined (Del Prado et 

al. 2006; Schochetal.2009; Zhang et al. 2009a). 

Ascocratera Kohlm., Can. J. Bot. 64: 3036 (1986). 

Type species: Ascocratera manglicola Kohlm., Can. J. Bot. 

64(12): 3036 (1986). 

Ascocratera is a monotypic obligate marine fungus and 

is characterized by conical, crater-like, erumpent to 

superficial and carbonaceous ascomata, a depressed 

ostiole, a thick peridium, trabeculate pseudoparaphyses, 

bitunicate, fissitunicate and cylindrical asci, and ellipsoidal, 

hyaline, 1-septate (3-septate when senescent) ascospores 

surrounded by a sheath (Kohlmeyer 1986). 

Ascocratera was reported to be one of the most common 

marine fungi of the upper intertidal zone of dead 

mangrove roots, trunks and branches (Kohlmeyer 1986). 

Based on a multigene phylogenetic analysis, Ascocratera 

nested within the clade of Aigialaceae (Schoch et al. 

2009; Suetrong et al. 2009). 

Atradidymella M.L. Davey & Currah, Am. J. Bot. 96: 1283 

(2009). 

Type species: Atradidymella muscivora M.L. Davey & 

Currah, Am. J. Bot. 96: 1283 (2009). 

Atradidymella was introduced as a pleosporalean genus 

parasitic on boreal bryophytes, and is characterized by


minute, unilocular, setose pseudothecia with 2–3 wall 

layers; brown, fusoid, 1-septate ascospores, and an anamorphic 

stage (Phoma muscivora M.L. Davey & Currah) 

(Davey and Currah 2009). Based on an ITS rDNA 

sequences analysis, Atradidymella nested within Didymellaceae 

(Davey and Currah 2009). 

Bertiella (Sacc.) Sacc. & P. Syd., in Saccardo, Syll. fung. 

(Abellini) 14: 19 (1899). 

≡ Bertia subgen. Bertiella Sacc., Syll. fung. (Abellini) 1: 

584 (1882). 

Type species: Bertiella macrospora (Sacc.) Sacc. & 

Traverso, Syll. fung. (Abellini) 19: 147 (1910). 

≡ Bertia macrospora Sacc., Michelia 1(no. 8): 452 (1882). 

Bertia subg. Bertiella was raised to generic rank by Saccardo 

(1899), and is typified by B. macrospora. After studying the 

type specimen of B. macrospora, Eriksson and Yue (1986) 

assigned it to Massarina (as M. macrospora (Sacc.) O.E. Erikss. 

& J.Z. Yue). Concurrently, Bertiella is treated as a synonym of 

Massarina. Hydeetal.(2002) assigned Bertia macrospora to 

Lophiostoma as (L. bertiellum Aptroot & K.D. Hyde). 

The superficial ascomata, cylindro-clavate asci and 

hyaline 1-septate ascospores which may become 3-septate 

and pale brown when senescent and, in particular, the 

woody habitat indicate that B. macrospora may be related 

to Lophiostoma sensu Holm and Holm (1988). A single 

isolate of Bertiella macrospora clusters with Byssosphaeria 

in the Melanommataceae in a recent DNA based phylogeny 

(Mugambi and Huhndorf 2009b). The relationship between 

Bertiella and Byssosphaeria needs further study. 

Byssothecium Fuckel, Bot. Ztg. 19: 251 (1861). 

Type species: Byssothecium circinans Fuckel, Bot. Ztg. 

19: 251 (1861). 

The isotype of Byssothecium circinans is in FH as 

exiccatae (Fungi rhenani 730c); it was described by Boise 

(1983) and could not be loaned. Byssothecium circinans is 

regarded as a saprobe or weak parasite of Medicago sativa 

(Semeniuk 1983), and a Pleospora-type centrum was 

observed (Boise 1983). A Chaetophoma-like anamorph 

was produced in culture, however, no culture or herbarium 

specimen is listed (Boise 1983). Boise (1983) regarded 

Byssothecium circinans as closely related to Teichospora, 

however, confirmation is required. An isolate of Byssothecium 

circinans was sequenced and a multigene phylogeny 

placed it in close proximity to members of Massarinaceae 

(Schoch et al. 2009; Zhang et al. 2009a; Plate 1). 

Caryospora De Not., Micr. Ital. Novi 9: 7 (1855). 

Type species: Caryospora putaminum (Schwein.) De Not., 

Micr. Ital., Dec. 9: 7 (1855). 

After studying the Caryospora species in North America, 

Barr (1979b) indicated that species of Caryospora may 

closely relate to Trematosphaeria. Boise (1985) distinguished 

Caryospora from Trematosphaeria based on the 

structure of ascospores. Currently, 17 taxa, from freshwater, 

marine, or terrestrial habitats (Raja and Shearer 2008), are 

included within Caryospora and might be polyphyletic. 

Celtidia J.D. Janse, Ann. Jard. Bot. Buitenzorg 14: 202 (1897). 

Type species: Celtidia duplicispora J.D. Janse, Ann. Jard. 

Bot. Buitenzorg 14: 202 (1897). 

Celtidia is a monotypic genus, which is characterized by 

its echinulate ascospores (Hawksworth 1979). It is only 

known from an illustration accompanying the original 

description from root nodules of Celtis in Java. A new 

collection is needed for further study of this genus. 

Chaetopreussia Locq.-Lin., Revue Mycol., Paris 41: 185 

(1977). 

Type species: Chaetopreussia chadefaudii Locq.-Lin., 

Revue Mycol., Paris 41: 187 (1977). 

Chaetopreussia is a monotypic genus characterized by 

cleistothecioid ascomata with seta, and 3-septate ascospores 

without germ slits. Recent molecular analysis has shown that 

cleistothecioid ascomata and the presence of germ slits lack 

significance at the generic rank (Kruys and Wedin 2009). 

Chaetopreussia is possibly another synonym of Preussia. 

Clathrospora Rabenh., Hedwigia 1(18): 116 (1857). 

Type species: Clathrospora elynae Rabenh., Hedwigia 1: 

116 (1857). 

The most striking character of Clathrospora is its 

ascomata opening with an intraepidermal discoid lid and 

muriform applanate ascospores with more than one row of 

longitudinal septa (Shoemaker and Babcock 1992). The 

form of opening and applanate ascospores, however, might 

have limited significance at generic rank and thus, 

Clathrospora may be closely related to Pleosporaceae. 

Phylogenetic analysis based on nLSU, nSSU and mtSSU 

indicate that C. diplospora (Ellis & Everh.) Sacc. & 

Traverso nests in Pleosporaceae (Kruys et al. 2006). 

Clathrospora elynae is saprobic on monocots (Shoemaker 

and Babcock 1992). 

Cochliobolus Drechsler, Phytopathology 24: 973 (1934). 

Type species: Cochliobolus heterostrophus (Drechsler) 

Drechsler, Phytopathology 24: 973 (1934). 

Cochliobolus and its asexual relatives are well studied 

taxa in Pleosporales because of their economic importance. 

Cochliobolus includes both saprobic and pathogenic species 

that are significant monocot pathogens worldwide, which 

attack corn, rice, barley, sugarcane, wheat, and oats, all 

major cereal crops. Cochliobolus is characterized by globose 

or subglobose ascomata with a well defined long ostiolar 

papilla or cylindrical neck, a peridium composed of


pseudoparenchymatous cells, filliform, septate and branched 

pseudoparaphyses, and thin-walled cylindrical or broadly 

clavate asci. Ascospores are distinctively hyaline or pale 

brown, filliform, and strongly helicoid to loosely coiled in 

the asci (Sivanesan 1984). The anamorphs of Cochliobolus 

belong to Bipolaris and Curvularia (Sivanesan 1984). 

Bipolaris and Curvularia can be distinguished by characters 

of conidial morphology, conidial germination, hilum structure, 

conidial septum and wall structure, conidial septum 

ontogeny (Sivanesan 1987). Multigene phylogenetic analysis 

indicated that Cochliobolus heterostrophus and C. sativus (S. 

Ito & Kurib.) Drechsler ex Dastur nested within the clade of 

Pleosporaceae (Zhang et al. 2009a; Plate 1). Thus, its 

familial placement is confirmed. 

Comoclathris Clem., Gen. fung. (Minneapolis): 37, 173 

(1909). 

Type species: Comoclathris lanata Clem. [as ‘Comochlatris’], 

Gen. fung. (Minneapolis) (1909). 

Comoclathris is temporarily placed in Diademaceae, and 

its pivotal characters are the circular lid-like opening and 

applanate reddish-brown to dark reddish-brown muriform 

ascospores with single longitudinal septa (versus two or 

more rows of longitudinal septa of Clathrospora) (Shoemaker 

and Babcock 1992). Barr (1990b) treated it as a synonym of 

Graphyllium. Comoclathris has been linked with an Alternaria-like 

anamorphs (Simmons 1952), which may suggest 

its close relationship with Pleosporaceae. 

Coronopapilla Kohlm. & Volkm.-Kohlm., Mycol. Res. 94: 

686 (1990). 

Type species: Coronopapilla avellina Kohlm. & Volkm.- 

Kohlm., Mycol. Res. 94: 687 (1990). 

Coronopapilla is characterized by immersed ascomata 

with a conical papilla, thin peridium, 8-spored and thickwalled, 

cylindrical and fissitunicate asci. Ascospores are 

ellipsoidal, 1-3-septate, brown and distoseptate. Coronopapilla 

avellina is an obligate marine species, and was originally 

assigned to Didymosphaeriaceae (Kohlmeyer and Volkmann- 

Kohlmeyer 1990). The marine habitat of Coronopapilla 

makes it readily distinguishable from Didymosphaeria futilis 

(the generic type of Didymosphaeria). Thus, the familial 

placement of Coronopapilla is yet to be determined. 

Cucurbitaria Gray, Nat. Arr. Brit. Pl. (London) 1: 508, 519 

(1821). 

Type species: Cucurbitaria berberidis (Pers.) Gray, Nat. 

Arr. Brit. Pl. (London) 1: 508, 519 (1821). 

≡ Sphaeria berberidis Pers., Neues Mag. Bot. 1: 83 

(1794). 

A narrow generic concept of Cucurbitaria was accepted 

by Welch (1926), who restricted Cucurbitaria to five 

closely related species, which have turbinate ascomata that 

develop cespitosely in a massive subiculum or over 

compressed stromatic tissues and have a thick and obconoid 

base. A broader generic concept was accepted by Mirza 

(1968), who also included species with globose or ovoid to 

pyriform ascomata that are gregarious on the substrate with 

only sparse subiculum and lack an obconoid region in the 

base of the locule. Barr (1990b) accepted an intermediate 

concept, and described 11 related species from North 

America. Currently, 450 species are accepted in Cucurbitaria 

(http://www.mycobank.org/mycotaxo.aspx), and the 

genus was assigned to Cucurbitariaceae. In this study, an 

isolate of C. berberidis clustered with some species of 

Pyrenochaeta and Didymosphaeria futilis, and they get 

moderate bootstrap support (Plate 1). Cucurbitariaceae 

may be another family within Pleosporineae. 

Curreya Sacc., Syll. fung. (Abellini) 2: 651 (1883). 

Type species: Curreya conorum (Fuckel) Sacc., Syll. fung. 

(Abellini) 2: 651 (1883). 

Curreya is a contentious genus which had been assigned to 

Pleospora (Barr 1981). von Arx and van der Aa (1983), 

however, maintained it as distinct, because of its Coniothyrium 

anamorph, and considered Curreya should be closely 

related to Didymosphaeria, Melanomma, Paraphaeosphaeria 

or Massarina. Because of the small sclerotial cells of its 

peridium, the narrower, thinner-walled asci and its Coniothyrium-like 

anamorph, Barr (1990b) assigned it to the 

Leptosphaeriaceae. Previous phylogenetic studies indicated 

that a strain of Curreya pityophila (J.C. Schmidt & Kunze) 

Petr. nested within Massarineae (Kruys et al. 2006). 

Decorospora Inderb., Kohlm. & Volkm.-Kohlm., Mycologia 

94: 657 (2002). 

Type species: Decorospora gaudefroyi (Pat.) Inderb., 

Kohlm. & Volkm.-Kohlm., Mycologia 94: 657 (2002). 

≡ Pleospora gaudefroyi Pat., Tabl. analyt. Fung. France 

(Paris) 10: 40 (no. 602) (1886). 

Decorospora gaudefroyi (as Pleospora gaudefroyi) had 

been considered a synonym of Pleospora herbarum, 

despite its striking sheath of ascospores (Wehmeyer 

1961). Molecular phylogenetic analysis based on partial 

SSU and ITS rDNA sequences indicated that Decorospora 

gaudefroyi was a sister taxon in the Pleosporaceae 

represented by Alternaria alternata (Fr.) Keissl., Cochliobolus 

sativus, Pleospora herbarum, Pyrenophora triticirepentis 

(Died.) Drechsler and Setosphaeria rostrata K.J. 

Leonard (Inderbitzin et al. 2002). Decorospora was 

introduced as a monotypic genus represented by Decorospora 

gaudefroyi, which is characterized by black ascomata 

becoming superficial on the substrate at maturity, 

septate and branched pseudoparaphyses, fissitunicate, clavate 

asci, as well as yellowish brown ascospores with seven 

transverse septa and one to three longitudinal septa in each


segment, enclosed in a sheath with 4–5 apical extensions 

(Inderbitzin et al. 2002). Decorospora gaudefroyi is an 

obligate marine fungus, growing at or above the high water 

mark (Inderbitzin et al. 2002). 

Diadema Shoemaker & C.E. Babc., Can. J. Bot. 67: 1349 

(1989). 

Type species: Diadema tetramerum Shoemaker & C.E. 

Babc. [as ‘tetramera’], Can. J. Bot. 67: 1354 (1989). 

During their study of Leptosphaeria and Phaeosphaeria, 

Shoemaker and Babcock (1989c) found some alpine fungi 

with typical pleosporalean characters (such as perithecoid 

ascomata, bitunicate asci and presence of pseudoparaphyses) 

having relatively large, very dark brown ascospores, mostly 

with a peculiar disc-like opening (as reported in some 

species of Wettsteinina, Shoemaker and Babcock 1987). 

Thus, they introduced a new genus Diadema (typified by D. 

tetramerum) to accommodate them (Shoemaker and 

Babcock 1989c). Currently, Diadema is assigned to Diademaceae, 

and differs from other genera in the family in having 

ascospores which lack longitudinal septa (Shoemaker and 

Babcock 1992). The large, dark brown ascospores and the 

disc-like opening, however, may be an adaptation to 

environmental factors. 

Diademosa Shoemaker & C.E. Babc., Can. J. Bot. 70: 1641 

(1992). 

Type species: Diademosa californiana (M.E. Barr) Shoemaker 

& C.E. Babc. [as ‘californianum’], Can. J. Bot. 70: 1641 (1992). 

≡ Graphyllium californianum M.E. Barr, Mem. N. Y. 

bot. Gdn 62: 40 (1990). 

Diademosa is the only genus in Diademaceae that has 

terete (cylindrical, circular in cross section) ascospores 

(Shoemaker and Babcock 1992). 

Didymella Sacc., Michelia 2(no. 6): 57 (1880). 

Type species: Didymella exigua (Niessl) Sacc., Syll. fung. 

(Abellini) 1: 553 (1882). 

≡ Didymosphaeria exigua Niessl, Öst. bot. Z.: 165 

(1875). 

The type specimen of Didymella (D. exigua) is lost and a 

neotype specimen was selected by de Gruyter et al. (2009). 

Didymella was characterized by the immersed or erumpent, 

globose or flattened and ostiolate ascomata with dense, rare 

(or lack?) of pseudoparaphyses. Asci are cylindrical, 

clavate or saccate, and 8-spored. Ascospores are hyaline, 

1-septate (symmetrical or asymmetrical) and constricted at 

the septum. Didymella has been assigned under Mycosphaerellaceae, 

Pleosporales (Sivanesan 1984), Phaeosphaeriaceae 

(Barr 1979a; Silva-Hanlin and Hanlin 1999), 

Venturiaceae (Reddy et al. 1998) orPleosporales genera 

incertae sedis (Lumbsch and Huhndorf 2007). Based on a 

multigene phylogenetic analysis, the Didymella clade forms 

a familial rank within Pleosporineae, thus the Didymellaceae 

was introduced (Aveskamp et al. 2010; de Gruyter et 

al. 2009; Zhang et al. 2009a; Plate 1). Anamorphs of 

Didymellaceae include Ascochyta, Ampelomyces, Boeremia, 

Chaetasbolisia, Dactuliochaeta, Epicoccum, Microsphaeropsis, 

Peyronellaea, Phoma, Piggotia, Pithoascus, 

Pithomyces and Stagonosporopsis (Aveskamp et al. 2010; 

de Gruyter et al. 2009; Hyde et al. 2011). 

Didymocrea Kowalski, Mycologia 57: 405 (1965). 

Type species: Didymocrea sadasivanii (T.K.R. Reddy) 

Kowalski, Mycologia 57: 405 (1965). 

≡ Didymosphaeria sadasivanii T.K.R. Reddy, Mycologia 

53: 471 (1962). 

Didymocrea is a monotypic genus, and was separated 

from Didymosphaeria based on its “unitunicate asci”, 

presence of pseudoparaphyses and absence of spermatia, 

and assigned under Hypocreales (Kowalski 1965). Following 

Kowalski (1965), Luttrell (1975) also studied the 

centrum development of Didymocrea, and concluded that it 

should be a true pleosporalean fungus with functionally 

unitunicate asci, and retained it in Didymosphaeria. After 

studying the type specimen of Didymocrea sadasivanii, 

Aptroot (1995) concluded that it should be closely related to 

the loculoascomycetous genus Zopfia. Rossman et al. (1999) 

also kept it as a unique genus in Pleosporales. Based on a 

multigene phylogenetic analysis, D. sadasivanii nests within 

Montagnulaceae (Kruys et al. 2006; Schoch et al. 2009). 

Dothivalsaria Petr., Sydowia 19: 283 (1966) [1965]. 

Type species: Dothivalsaria megalospora (Auersw.) Petr., 

Sydowia 19: 283 (1966) [1965]. 

≡ Valsaria megalospora Auersw., Leipzig. Bot. 

Tauschver. 5. (1866). 

Dothivalsaria is monotypic and is represented by D. 

megalospora (Petrak 1965). The taxon is characterized by 

immersed, medium- to large-sized ascomata which usually 

aggregate under blackened stromatic tissues and have 

trabeculate pseudoparaphyses. Asci are cylindrical, while 

ascospores are brown, ellipsoid, and 1-septate and uniseriate 

in the asci (Barr 1990a). The ascostroma of D. megalospora is 

comparable with those of Aglaospora profusa as has been 

mentioned by Barr (1990a), but their relationships are unclear. 

Epiphegia G.H. Otth, Mitt. naturf. Ges. Bern: 104 (1870). 

Type species: Epiphegia alni G.H. Otth, Mitt. naturf. Ges. 

Bern: 104 (1870). 

Epiphegia was reinstated to accommodate a species 

which has Phragmoporthe-like ascocarps and Massarinalike 

asci, pseudoparaphyses and ascospores (Aptroot 1998). 

Ascomata are grouped within stromatic tissues, pseudoparaphyses 

are cellular, asci are bitunicate and ascospores are 

hyaline and trans-septate (Aptroot 1998).


Eremodothis Arx, Kavaka 3: 34 (1976) [1975] (IMI 90223= 

CBS 610.74 type). 

Type species: Eremodothis angulata (A.C. Das) Arx, 

Kavaka 3: 34 (1976) [1975]. 

≡ Thielavia angulata A.C. Das, Trans. Br. Mycol. Soc. 

45: 545 (1962). 

The type species of Eremodothis (E. angulata) was 

originally isolated from rice field soil in Fulta, India and it 

was assigned to Sporormiaceae because of the orange 

pigmentation of the colony (von Arx 1976). The cleistothecoid 

ascomata, sphaerical asci and 1-celled ascospores of E. 

angulata are comparable with those of Pycnidiophora. 

Based on a multigene phylogenetic study, both Eremodothis 

and Pycnidiophora were treated as synonyms of Westerdykella 

(Kruys and Wedin 2009). 

Extrawettsteinina M.E. Barr, Contr. Univ. Mich. Herb. 9 

(8): 538 (1972). 

Type species: Extrawettsteinina minuta M.E. Barr, Contr. 

Univ. Mich. Herb. 9(8): 538 (1972). 

Extrawettsteinina was introduced to accommodate three 

species, i.e. E. minuta, E. pinastri M.E. Barr and E. mediterranea 

(E. Müll.) M.E. Barr, which are saprobic on the dead leaves 

of gymnosperms and angiosperms, in North America and 

Europe (Barr 1972). Subsequently, a fourth species was 

introduced, viz. E. andromedae (Auersw.) M.E. Barr (Barr 

1987a). Extrawettsteinina is characterized by superficial, conical 

ascomata, containing a few saccate bitunicate asci, ellipsoidal, 

obovate-clavate, septate, smooth and hyaline ascospores which 

turn dull brown at maturity (Barr 1972). The diagnostic 

character of Extrawettsteinina is its conic ascocarps which are 

superficial on the substrate, and radiating arrangement of wall 

cells, which makes it distinguishable from comparable genera 

such as Stomatogene and Wettsteinina. 

Graphyllium Clem., Botanical Survey of Nebraska 5: 6 (1901). 

Type species: Graphyllium chloës Clem., Botanical Survey 

of Nebraska 5: 6 (1901). 

Graphyllium was first described as a hysteriaceous 

fungus with elongate ascomata, but von Höhnel (1918b, 

1919) recognized its similarity to Clathrospora. Petrak 

(1952) transferred Graphyllium to Pleospora, andnoted 

that the elongate ascomata and closely grouped rows of 

small ascomata are not sufficient to recognize the genus. 

Barr (1987b, 1990b) supported this proposal and considered 

Graphyllium differs from Clathrospora by shape, 

septation and pigmentation of ascospores. A narrow 

generic concept of Graphyllium was adapted by Shoemaker 

and Babcock (1992), which is characterized by hysterothecia, 

applanate ascospores that are at least 3-septate in side view 

and have some longitudinal septa in front view, and it was 

assigned under Hysteriaceae (order Pleosporales, Shoemaker 

and Babcock 1992). But subsequent classification systems 

tend to assign it to Diademaceae (e.g. Lumbsch and 

Huhndorf 2007, 2010). This seems unlikely as pointed out 

by Zhang et al. (2011) and the genus could be placed in one 

of five families containing hysteriotheciod ascomata. Recollection 

and molecular studies are needed. 

Halomassarina Suetrong, Sakay., E.B.G. Jones, Kohlm., 

Volkm.-Kohlm. & C.L. Schoch, Stud. Mycol. 64: 161 (2009). 

Type species: Halomassarina thalassiae (Kohlm. & Volkm.- 

Kohlm.) Suetrong, Sakay., E.B.G. Jones, Kohlm., Volkm.- 

Kohlm. & C.L. Schoch, Stud. Mycol. 64: 161 (2009). 

≡ Massarina thalassiae Kohlm. & Volkm.-Kohlm., Can. 

J. Bot. 65: 575 (1987). 

Halomassarina is another marine genus which morphologically 

fits Massarina sensu lato, and is typified by H. 

thalassiae, which is characterized by subglobose to pyriform, 

immersed or erumpent, ostiolate, periphysate, papillate 

or epapillate, coriaceous ascomata, simple, rarely anastomosing 

pseudoparaphyses, 8-spored, cylindrical to clavate, 

pedunculate, thick-walled, fissitunicate asci, and ellipsoidal, 

(1-)3-septate, hyaline ascospores. Based on a multigene 

phylogenetic analysis, Halomassarina thalassiae clustered 

together with Trematosphaeria pertusa and another marine 

fungus Falciformispora lignatilis, and they are all assigned 

to Trematosphaeriaceae (Suetrong et al. data unpublished). 

Hypsostroma Huhndorf, Mycologia 84: 750 (1992). 

Type species: Hypsostroma saxicola Huhndorf, Mycologia 

84: 750 (1992). 

Hypsostroma was introduced as a tropical woodinhabiting 

genus by Huhndorf (1992). Hypsostroma has 

several striking characters including the large superficial 

ascomata which form on a subiculum, pseudoparenchymatous 

peridial cells, trabeculate pseudoparaphyses, clavate 

asci with long pedicels and a conspicuous apical apparatus, 

and ascospores that separate into partspores with a germ slit 

in each partspore (Huhndorf 1992). Phylogenetic study 

indicated that Hypsostroma should be a new genus and the 

Hypsostromataceae was reinstated to accommodate Hypsostroma 

(Mugambi and Huhndorf 2009b; Plate 1). 

Julella Fabre, Annls Sci. Nat., Bot., sér. 6 9: 113 (1879) [1878]. 

Type species: Julella buxi Fabre, Annls Sci. Nat., Bot., sér. 

6 9: 113 (1879) [1878]. 

Julella has been assigned to Thelenellaceae, a family of 

Ostropomycetidae (Lumbsch and Huhndorf 2007), and 

Arthopyreniaceae (= Xanthopyreniaceae sensu O. Eriksson, 

Pleosporales) (Barr 1985). Julella is characterized by its 

immersed, medium-sized ascomata with pseudoparenchymatous 

peridial cells, cellular pseudoparaphyses, and 

hyaline and muriform ascospores (Barr 1985). With the 

exception of hyaline ascospores, these characters are typical 

of Montagnulaceae. The taxonomic affinity of the generic


type of Julella needs confirmation following recollection. 

Julella avicenniae (Borse) K.D. Hyde is a marine fungus. A 

DNA based phylogeny containing most currently accepted 

families placed two isolates of J. avicenniae as sister to the 

families in the Pleosporineae with good support, which might 

suggest a novel family within Pleosporales (Suetrong et al. 

2009). However, J. avicenniae is not the generic type and 

therefore this conclusion must be treated with caution as only 

J. avicenniae can be considered pleosporalean. 

Lautitia S. Schatz, Can. J. Bot. 62: 31 (1984). 

Type species: Lautitia danica (Berl.) S. Schatz, Can. J. Bot. 

62: 31 (1984). 

≡ Leptosphaeria danica Berl., Icon. fung. (Abellini) 1: 

87 (1892). 

Lautitia is monotypified by L. danica, which is characterized 

by subglobose, immersed, ostiolate ascomata with a 

pseudoclypeus, a thin peridium, broad, cellular pseudoparaphyses, 

and 8-spored, bitunicate, cylindrical to clavate asci. 

Ascospores are hyaline, 1-septate, and obovate and the fungus 

is parasitic on algae (Schatz 1984). Marine or maritime fungi 

have been reported in Phaeosphaeria, suchasP. spartinae 

(Ellis & Everh.) Shoemaker & C.E. Babc. and P. ammophilae 

(Lasch) Kohlm. & E. Kohlm. (Zhang et al. 2009a). In 

addition, the prosenchymatous peridium of L. danica agrees 

with that of Phaeosphaeriaceae (Schatz 1984). 

Lepidosphaeria Parg.-Leduc, C. r. hebd. Séanc. Acad. Sci., 

Paris, Sér. D 270: 2786 (1970). 

Type species: Lepidosphaeria nicotiae Parg.-Leduc, Pubbl. 

Staz. Zool. Napoli, 1 270: 2786 (1970). 

Lepidosphaeria is a genus likely in Testudinaceae, which 

is distinguished from other genera of this family by its 

smaller ascospores, which lack furrows, and have minute 

granulate ornamentation (Hawksworth 1979). In DNA 

sequence-based phylogenies, L. nicotiae clustered with 

species of Ulospora and Verruculina (Schoch et al. 2009; 

Zhang et al. 2009a), but more recent work including 

species of Platystomaceae lacks support (Plate 1). 

Letendraea Sacc., Michelia 2: 73 (1880). 

Type species: Letendraea eurotioides Sacc., Michelia 2: 73 

(1880). 

Letendraea was introduced for L. eurotioides, which is 

characterized by superficial, globose to conical ascomata, 

filliform pseudoparaphyses, obclavate to cylindrical, 8- 

spored asci, and fusoid to oblong, 1-septate ascospores 

(Saccardo 1880). Because L. helminthicola (Berk. & 

Broome) Weese clustered with Karstenula rhodostoma, 

Letendraea was assigned to Melanommataceae (Kodsueb 

et al. 2006b). But subsequent multigene phylogenetic 

analysis indicated that both L. helminthicola and L. padouk 

Nicot & Parg.-Leduc nested within Montagnulaceae 

(Schoch et al. 2009; Zhang et al. 2009a; Plate 1), and its 

familial status seems confirmed. 

Lindgomyces K. Hirayama, Kaz. Tanaka & Shearer, 

Mycologia 102: 133 (2010). 

Type species: Lindgomyces ingoldianus (Shearer & K.D. 

Hyde) K. Hirayama, Kaz. Tanaka & Shearer, Mycologia 

102: 733 (2010). 

≡ Massarina ingoldiana Shearer & K.D. Hyde, Mycologia 

89: 114 (1997). 

Lindgomyces was introduced to accommodate a freshwater 

lineage, which belongs to Massarina ingoldiana sensu lato, 

and is characterized by scattered, subglobose to globose, 

erumpent, papillate, ostiolate ascomata, cellular pseudoparaphyses, 

and 8-spored, fissitunicate, cylindrical to clavate 

asci. Ascospores are fusoid to narrowly fusoid, hyaline and 1- 

septate but become 3–5-septate when senescent (Hirayama et 

al. 2010). A new family, Lindgomycetaceae, wasintroduced 

to accommodate Lindgomyces (Hirayama et al. 2010). 

Lophiella Sacc., Michelia 1: 337 (1878). 

Type species: Lophiella cristata (Pers.) Sacc., Michelia 1: 

337 (1878). 

≡ Sphaeria cristata Pers., Syn. meth. fung. (Göttingen) 

1: 54 (1801). 

The generic type of Lophiella, L. cristata, was treated as 

a synonym of Lophiostoma angustilabrum var. crenatum 

(Pers.) Chesters & A.E. Bell (see http://www.indexfungorum. 

org/names/Names.asp). 

Loratospora Kohlm. & Volkm.-Kohlm., Syst. Ascom. 12: 

10 (1993). 

Type species: Loratospora aestuarii Kohlm. & Volkm.- 

Kohlm., Syst. Ascom. 12: 10 (1993). 

Loratospora was introduced as a marine genus and is 

monotypified by L. aestuarii (Kohlmeyer and Volkmann- 

Kohlmeyer 1993). The generic type is characterized by 

ellipsoid, immersed to erumpent, carbonaceous ascomata, 

which are ostiolate, and with or without a papilla. Pseudoparaphyses 

comprise small subglobose cells forming irregular 

chains and finally breaking apart, and asci are 8-spored, 

clavate to ellipsoidal, and fissitunicate. Ascospores are 

hyaline, cylindrical, 3-septate and surrounded by a mucilaginous 

sheath (Kohlmeyer and Volkmann-Kohlmeyer 1993). 

The distinctive pseudoparaphyses of Loratospora aestuarii 

makes it readily distinguishable from other taxa. Based on a 

multigene phylogenetic analysis, Loratospora aestuarii nested 

within the clade of Phaeosphaeriaceae (Schoch et al. 2009; 

Suetrong et al. 2009; Plate 1), and ascospores of L. aestuarii 

are in agreement with those of Phaeosphaeria as has been 

mentioned by Kohlmeyer and Volkmann-Kohlmeyer (1993). 

Macrospora Fuckel, Jb. nassau. Ver. Naturk. 23–24: 139 

(1870) [1869–70].


Type species: Macrospora scirpicola (DC.) Fuckel, Jb. 

nassau. Ver. Naturk. 23–24: 139 (1870) [1869–70]. 

≡ Sphaeria scirpicola DC., in Lamarck & de Candolle, 

Fl. franç., Edn 3 (Paris) 2: 300 (1805). 

Macrospora had been assigned to Diademaceae based 

on its applanate and muriform ascospores with 1-row of 

longitudinal septa, with a sheath, 2–3 μm wide and 

constricted at first septum and ascospores are paler and 

larger than those of Comoclathris (Shoemaker and Babcock 

1992). Macrospora was however, considered as a synonym 

of Pyrenophora by Eriksson and Hawksworth (1991) which 

was assigned in Pleosporaceae, and this proposal was 

widely followed (Eriksson 2006; Lumbsch and Huhndorf 

2010). Nimbya anamorphs were reported for Macrospora 

(Johnson et al. 2002). 

Massaria De Not., G. bot. ital. 1: 333 (1844). 

Type species: Massaria inquinans (Tode) De Not., G. bot. 

ital. 1: 333 (1844). 

≡ Sphaeria inquinans Tode, Fung. mecklenb. sel. 

(Lüneburg) 1: Fig. 85 (1790). 

Colonies on MEA erumpent, not spreading; surface 

irregular, folded; margins even, feathery; surface olivaceous 

grey, with thin, umber margin; reverse olivaceous-grey. On 

PDA similar; surface olivaceous grey, margin dirty white; 

reverse smoke-grey to olivaceous grey; colonies reaching 

1 cm diam. On OA similar, surface olivaceous grey in 

centre, margins wide, dirty white; colonies reaching 12 mm 

diam. on all media tested; colonies sterile (based on CBS 

125591). 

Massaria was formally established by de Notaris (1844), 

and is typified by M. inquinans. Numerous fungi with 

brown septate ascospores surrounded by gelatinous sheath 

were included in the genus (Barr 1979b; Shoemaker and 

LeClair 1975). Shoemaker and LeClair (1975) accepted a 

narrow concept for Massaria, with only a few species 

characterized by large, symmetric, 4-celled ascospores 

surrounded by a massive gelatinous sheath. Barr (1979b, 

1990a) had considered Aglaospora a separate genus, but this 

subsequently proved congeneric with Massaria (Voglmayr 

and Jaklitsch 2011). Based on intensive sample collection 

and multi-gene phylogenetic analysis, Voglmayr and 

Jaklitsch (2011) acceptedMassaria as the sole genus within 

Massariaceae, which is characterized by a set of well 

defined morphological and ecological characters; Europe is 

regarded as the centre of diversity. 

Misturatosphaeria Mugambi & Huhndorf, Stud. Mycol. 

64: 108 (2009). 

Type species: Misturatosphaeria aurantonotata Mugambi & 

Huhndorf, Stud. Mycol. 64: 108 (2009). 

Misturatosphaeria was introduced to accommodate a 

group of fungi which are phylogenetically closely related to 

Amniculicolaceae, Lophiostomataceae sensu stricto and 

Sporormiaceae (Mugambi and Huhndorf 2009b; Zhang et 

al. 2009a). Species of Misturatosphaeria are characterized 

by erumpent to superficial ascomata which are scattered or 

in groups, with or without papilla; asci cylindrical or 

clavate, 8-spored; pseudoparaphyses numerous, septate, 

ascospores brown or hyaline, phragmosporous or dictyosporous, 

with or without sheath. The terrestrial saprobic 

habitat on wood, as well as its distinct morphological 

characters may indicate that this genus belongs to an 

undescribed family. A close relationship with the marine 

anamorphic species Floricola striata is unexpected and 

may suggest that some of the species in this genus could 

have marine affinities (Plate 1). 

Navicella Fabre, Annls Sci. Nat., Bot., sér. 6 9: 96 (1879) 

[1878]. 

Type species: Navicella julii Fabre, Annls Sci. Nat., Bot., 

sér. 6 9: 96 (1879) [1878]. 

Navicella is characterized by medium- to large-sized, 

immersed to erumpent, globose ascomata, apex elongated or 

rarely rounded, asci clavate or cylindrical, pseudoparaphyses 

trabeculate, ascospores reddish to dark brown, ellipsoid to 

fusoid, multi-septate, the primary septum is euseptate, and 

others distoseptate, obliquely uniseriate or biseriate (Barr 

1990a). Navicella is saprobic on bark, and was considered 

closely related to the Lophiostomataceae (Holm and Holm 

1988). Based on the wide endotunica, thin apical ring and 

distoseptate ascospores, Barr (1990a) transferred it to the 

Massariaceae. The morphological characters of Navicella do 

not match the Massariaceae sensu stricto (Voglmayr and 

Jaklitsch 2011). 

Neotestudina Segretain & Destombes, C. r. hebd. Séanc. 

Acad. Sci., Paris 253: 2579 (1961). 

Type species: Neotestudina rosatii Segretain &Destombes, 

C. r. hebd. Séanc. Acad. Sci., Paris 253: 2579 (1961). 

Neotestudina is characterized by medium- to large-sized, 

superficial, gregarious, cleistothecioid and globose ascomata 

which split on opening. Asci are 4- or 8-spored, and cylindrical 

or oblong, pseudoparaphyses are sparse and trabeculate, and 

ascospores are dark brown, ellipsoid, 1-septate, with a small 

germ pore at each end, and uniseriate or crowded in the asci 

(Barr 1990a). Based on the cleistothecioid ascomata, Neotestudina 

was assigned under Zopfiaceae (von Arx and Müller 

1975) or Testudinaceae (Hawksworth 1979). Barr (1990a) 

assigned it to Didymosphaeriaceae basedonitsascospore 

morphology. A DNA based phylogeny showed that sequence 

obtained from Neotestudina rosatii resides as sister to 

Ulospora bilgramii (D. Hawksw., C. Booth & Morgan- 

Jones) D. Hawksw., Malloch & Sivan. and other species that 

may represent Testudinaceae or Platystomaceae (Kruys et al. 

2006; Plate 1).


Paraphaeosphaeria O.E. Erikss., Ark. Bot., Ser. 2 6: 405 

(1967). 

Type species: Paraphaeosphaeria michotii (Westend.) O.E. 

Erikss., Cryptogams of the Himalayas 6: 405 (1967). 

≡ Sphaeria michotii Westend., Bull. Acad. R. Sci. Belg., 

Cl. Sci., sér. 2 7: 87 (1859). 

Paraphaeosphaeria was separated from Leptosphaeria 

(Eriksson 1967a), and it is also quite comparable with 

Phaeosphaeria. Paraphaeosphaeria can be distinguished 

from Phaeosphaeria by its ascospores. Ascospores of Paraphaeosphaeria 

michotii have two septa, and they are biseriate, 

straight, subcylindrical with broadly rounded ends, rather dark 

brown and punctate. The primary septum is laid down closer 

to the distal end than to the proximal, and the larger, proximal 

hemispore is divided by one transversal septum. There are 

more septa in the proximal hemispore of other species such as 

Par. castagnei (Durieu & Mont.) O.E. Erikss., Par. obtusispora 

(Speg.) O.E. Erikss. and Par. vectis (Berk. & Broome) 

Hedjar. Anamorphic characters can also distinguish Paraphaeosphaeria 

and Phaeosphaeria. Paraphaeosphaeria has 

Paraconiothyrium or Coniothyrium-related anamorphs, but 

Phaeosphaeria has Hendersonia-Phaeoseptoria anamorphs 

(Eriksson 1967a). Shoemaker and Babcock (1985) redescribed 

some Canadian and extralimital species, and excluded 

Par. longispora (Wegelin) Crivelli and Par. oblongata 

(Niessl) Crivelli from Paraphaeosphaeria based on their 

longitudinal septa as well as beak-like papilla and wall 

structures. Molecular phylogenetic results based on multigenes 

indicated that Paraphaeosphaeria should belong to 

Montagnulaceae (Zhang et al. 2009a; Plate 1). 

Passeriniella Berl., Icon. fung. (Abellini) 1: 51 (1890). 

Type species: Passeriniella dichroa (Pass.) Berl., Icon. 

fung. (Abellini) 1: 51 (1890). 

≡ Leptosphaeria dichroa Pass. 

Passeriniella was introduced by Berlese in 1890 based on 

the black, ostiolate and papillate ascomata, 8-spored asci, as 

well as transverse septate ascospores, with pigmented central 

cells and hyaline terminal cells. Two species were included, i. 

e. P. dichroa and P. incarcerata (Berk. & M.A. Curtis) Berl. 

(Berlese 1890). Subsequently, more species were introduced 

including some marine taxa such as P. mangrovei G.L. Maria 

&K.R.Sridhar,P. obiones (P. Crouan & H. Crouan) K.D. 

Hyde & Mouzouras and P. savoryellopsis K.D. Hyde & 

Mouzouras (Hyde and Mouzouras 1988; Maria and Sridhar 

2002). Currently, eight species are included (http://www. 

mycobank.org, Jan. 2011). Both P. dichroa and P. incarcerata 

were considered as synonyms of Leptosphaeria obiones 

(P. Crouan & H. Crouan) Sacc. (Kohlmeyer and Kohlmeyer 

1979). The familial placement of the marine species P. 

savoryellopsis could not be resolved in a DNA based 

phylogeny but it did suggest a close relationship to 

Acrocordiopsis patilii (Suetrong et al. 2009) inPleosporales. 

Peridiothelia D. Hawksw., Bull. Br. Mus. nat. Hist., Bot. 

14: 120 (1985). 

Type species: Peridiothelia fuliguncta (Norman) D. 

Hawksw., Bull. Br. Mus. nat. Hist., Bot. 14: 121 (1985). 

≡ Microthelia fuliguncta Norman, Öfvers. kongl. 

Svensk. Vetensk.-Akad. Förhandl., Stockholm 41(no. 8): 

36 (1884). 

When dealing with the names under Microthelia, Peridiothelia 

was introduced to accommodate species having nonclypeate 

peridium which composed cells of textura globulosa 

but sometimes angularis, “dark reddish brown except 

below the generative locule where the wall is poorly 

developed or almost absent at maturity, colour not changed 

significantly in potassium hydroxide, centrum turning blue in 

iodine” (Hawksworth 1985a). Three species were included, i. 

e. P. grandiuscula (Anzi) D. Hawksw., P. fuliguncta and P. 

oleae (Körb.) D. Hawksw., and Peridiothelia was referred to 

Phaeosphaeriaceae (Hawksworth 1985a, b). However, its 

familial placement is not confirmed yet. 

Phaeodothis Syd. & P. Syd., Annls mycol. 2: 166 (1904). 

Type species: Phaeodothis tricuspidis Syd. & P. Syd., 

Annls mycol. 2: 166 (1904). 

Phaeodothis is characterized by its 1-septate euseptate 

ascospores with a sparse hamathecium consisting of thin 

pseudoparaphyses and immersed to superficial ascomata 

(Aptroot 1995).Thegenushadbeenpreviouslyassignedto 

Didymosphaeria, butAptroot(1995) consideredittobeclosely 

related to Phaeosphaeriaceae. A strain named Phaeodothis 

winteri (a synonym of P. tricuspidis Syd. & P. Syd.) nested 

within the clade of Montagnulaceae (Schoch et al. 2009). 

Platychora Petr., Annls mycol. 23: 102 (1925). 

Type species: Platychora ulmi (Schleich.) Petr., Annls 

mycol. 23(1/2): 103 (1925). 

Platychora is characterized by immersed to erumpent crustlike 

ascostroma with globose locules scattered inside (Barr 

1968). Asci are oblong to saccate or nearly cylindrical and 

bitunicate, and ascospores are hyaline 1-septate apiosporous 

and turn olivaceous when old. Platychora hadbeenpreviously 

assigned to Venturiaceae by Barr (1968), but molecular 

phylogenetic analysis indicated that a strain named Platychora 

ulmi (the generic type of Platychora) belongs to Didymellaceae 

(Winton et al. 2007; Plate 1). The generic type needs 

recollecting and epitypifying to stabilize the generic name. 

Platystomum Trevis., Bull. Soc. R. Bot. Belg. 16: 16 (1877). 

Type species: Platystomum compressum (Pers.) Trevis., 

Bull. Soc. R. Bot. Belg. 16: 16 (1877). 

≡ Sphaeria compressa Pers., Syn. meth. fung. (Göttingen) 

1: 56 (1801). 

Platystomum was introduced by Trevisan in 1877, and 

has been considered a synonym of Lophidium, as the


ascospores of Platystomum have both transverse and 

vertical septa (Barr 1990a, b; ChestersandBell1970). 

However, the boundary between Lophiostoma and Platystomum 

is not clear (Chesters and Bell 1970). Holm and 

Holm (1988) treated Platystomum as a synonym of 

Lophiostoma, and concurrently, the Platystomaceae 

should be treated as a synonym of Lophiostomataceae. 

Based on a phylogenetic analysis, however, the generic 

type of Platystomum (P. compressum) separated from 

other species of Lophiostoma, and nested with the clade of 

Platystomaceae (Mugambi and Huhndorf 2009b) which 

may be closely related to species in the Testiduniaceae 

(Plate 1). 

Polyplosphaeria Kaz. Tanaka & K. Hirayama, Stud. 

Mycol. 64: 192 (2009). 

Type species: Polyplosphaeria fusca Kaz. Tanaka & K. 

Hirayama, Stud. Mycol. 64: 193 (2009). 

Polyplosphaeria is characterized by globose ascomata 

surrounded by numerous brown hyphae and a reddish 

pigment on the host surface around the ascomata (Tanaka et 

al. 2009). Asci are cylindro-clavate with fissitunicate 

dehiscence and ascospores are narrowly fusoid surrounded 

by a sheath. The anamorph is Piricauda-like (Tanaka et al. 

2009). The cylindro-clavate asci, narrowly fusoid ascospores 

as well as its thin and numerous pseudoparaphyses are 

comparable with those of Massarina sensu lato, especially 

Lentithecium (Zhang et al. 2009a). The terrestrial and 

bambusicolous habitat of Polyplosphaeria and Piricauda 

anamorph readily distinguishes the genus from Lentithecium. 

Pontoporeia Kohlm., Nova Hedwigia 6: 5 (1963). 

Type species: Pontoporeia biturbinata (Durieu & Mont.) 

Kohlm., Nova Hedwigia 6: 5 (1963) 

≡ Sphaeria biturbinata Durieu & Mont., Flora Algéricae 

1: 497 (1849). 

Pontoporeia was introduced by Kohlmeyer in 1963, 

and is monotypified by P. biturbinata. Pontoporeia was 

treated as a synonym of Zopfia (Malloch and Cain 1972), 

which is followed by Hawksworth and Booth (1974). 

Based on its asci originating at the periphery of the 

subglobose locus, filaments occupying the center of the 

ascocarps, the irregular peridial structure, the ascospores 

having 2-layered walls with a germ pore at each end and 

its marine habitat, Pontoporeia was kept as a separate 

genus within Pleosporaceae (Kohlmeyer and Kohlmeyer 

1979). A DNA based phylogeny placed an isolate on a 

long branch in relationship with other marine species, 

Halotthia posidoniae and Mauritiana rhizophorae, but a 

familial placement awaits further resolution (Suetrong et 

al. 2009). 

Pseudotrichia Kirschst., Annls mycol. 37: 125 (1939). 

Type species: Pseudotrichia stromatophila Kirschst., Annls 

mycol. 37: 125 (1939). 

Pseudotrichia can be distinguished from Byssosphaeria, 

Herpotrichia and Lojkania by its lacking of subiculum, 

larger ascomata usually with compressed apices, the 

peripheral arrangement of asci and trabeculate pseudoparaphyses 

(Barr 1984). Phylogenetic study of strains Pseudotrichia 

mutabilis and some Herpotrichia species indicated 

that these species are closely related, and both nested within 

Melanommataceae (Mugambi and Huhndorf 2009b). But in 

this study, Pseudotrichia guatopoensis nested in the 

Testudinaceae (or Platystomaceae) (Plate 1). The types of 

both Herpotrichia and Pseudotrichia need recollecting, 

redescribing and epitypifying in order to stabiles the use of 

these generic names and clarify their familial status. 

Pseudoyuconia Lar.N. Vassiljeva, Nov. sist. Niz. Rast. 20: 

71 (1983). 

Type species: Pseudoyuconia thalictri (G. Winter) Lar. N. 

Vassiljeva [as ‘thalicti’], Nov. sist. Niz. Rast. 20: 71 (1983). 

≡ Leptosphaeria thalictri G. Winter, Hedwigia 10: 40 

(1872). 

Pseudoyuconia was introduced by Vassiljeva (1983), and 

was monotypified by P. thalictri. Currently, Pseudoyuconia 

is included in Pleosporaceae (Lumbsch and Huhndorf 

2010). 

Pyrenophora Fr., Summa veg. Scand., Section Post. 

(Stockholm): 397 (1849). 

Type species: Pyrenophora phaeocomes (Rebent.) Fr., 

Summa veg. Scand., Section Post. (Stockholm): 397 

(1849). 

≡ Sphaeria phaeocomes Rebent., Prodr. fl. neomarch. 

(Berolini): 338 (1804). 

Pyrenophora is characterized by immersed, erumpent to 

nearly superficial ascomata, indefinite pseudoparaphyses, 

clavate to saccate asci usually with a large apical ring, and 

muriform terete ascospores. Morphologically, the terete 

ascospores of Pyrenophora can be readily distinguished 

from Clathrospora and Platyspora. The indefinite pseudoparaphyses 

and smaller ascospores of Pyrenophora can be 

readily distinguished from those of Pleospora (Sivanesan 

1984). Based on both morphology and molecular phylogeny, 

Pyrenophora is closely related to Pleosporaceae 


Rechingeriella Petr., in Rechinger et al. Annln naturh. Mus. 

Wien 50: 465 (1940). 

Type species: Rechingeriella insignis Petr., Annln naturh. 

Mus. Wien, Ser. B, Bot. Zool. 50: 465 (1940). 

Rechingeriella is characterized by its erumpent to 

superficial, cleistothecioid ascomata and thin, branching 

pseudoparaphyses (Hawksworth and Booth 1974). Asci are


obovate, thick-walled, bitunicate and evanescent, and 

ascospores are globose, simple, dark brown to black (based 

on the type specimen of R. insignis) (Hawksworth and 

Booth 1974). Based on these characters, R. insignis was 

treated as a species of Zopfia (as Z. insignis (Petr.) D. 

Hawksw. & C. Booth). Rechingeriella has been assigned to 

Botryosphaeriaceae by von Arx and Müller (1975). Further 

study should be conducted on the type specimen of R. 

insignis in order to clarify its taxonomic status and fresh 

collections are needed for epitypification. 

Rhytidiella Zalasky, Can. J. Bot. 46: 1383 (1968). 

Type species: Rhytidiella moriformis Zalasky, Can. J. Bot. 

46: 1383 (1968). 

Rhytidiella was introduced based on R. moriformis, which 

causes perennial rough-bark of Populus balsamifera 

(Zalasky 1968), and produces macroconidia belonging to 

Phaeoseptoria. Subsequently, three more species were 

introduced, viz. R. baranyayi A. Funk & Zalasky, R. hebes 

P.R. Johnst. and R. beloniza (Stirt.) M.B. Aguirre (Aguirre- 

Hudson 1991; Funk and Zalasky 1975;Johnston2007), Both 

R. baranyayi and R. hebes seem closely related to R. 

moriformis on both biology and morphology (Funk and 

Zalasky 1975;Johnston2007), but R. beloniza is saprobic on 

Cordyline australis bark (Aguirre-Hudson 1991). Rhytidiella 

was temporarily assigned to Cucurbitariaceae (Barr 1987b). 

Richonia Boud., Revue mycol., Toulouse 7: 224 (1885). 

Type species: Richonia variospora Boud., Revue mycol., 

Toulouse 7: 265 (1885). 

Richonia is characterized by its 1-septate, relatively large 

ascospores which are broadly rounded at both ends, and 

have a thick ornamented undulating sheath giving an 

irregularly ridged appearance to mature spores (Hawksworth 

1979). Richonia variospora has been isolated from several 

localities in France, but it is rare (Hawksworth 1979). 

Richonia was assigned under Zopfiaceae (von Arx and 

Müller 1975; Hawksworth1979), and there are presently no 

better suggestions for its familial placement. The taxon needs 

recollecting and epitypifying. 

Rimora Kohlm., Volkm.-Kohlm., Suetrong, Sakay. & E.B. 

G. Jones, Stud. Mycol. 64: 166 (2009). 

Type species: Rimora mangrovei (Kohlm. & Vittal) 

Kohlm., Volkm.-Kohlm., Suetrong, Sakay. & E.B.G. Jones, 

Stud. Mycol. 64: 166 (2009). 

≡ Lophiostoma mangrovei Kohlm. & Vittal [as ‘mangrovis’], 

Mycologia 78: 487 (1986). 

Rimora was introduced based on a marine fungus R. 

mangrovei (syn. Lophiostoma mangrovei), and is characterized 

by its erumpent ascomata with elongated flat tops, 

cellular pseudoparaphyses and cylindrical asci (Suetrong et 

al. 2009). Ascospores are fusoid, hyaline, 3-septate and 

surrounded with an evanescent sheath (Kohlmeyer and 

Vittal 1986; Suetrong et al. 2009). Rimora forms a robust 

clade with other marine fungi, such as species of Aigialus 

and Ascocratera, and a new family, Aigialaceae was 

introduced to accommodate them (Suetrong et al. 2009). 

Roussoellopsis I. Hino & Katum., J. Jap. Bot. 40: 86 (1965). 

Type species: Roussoellopsis japonica (I. Hino & Katum.) 

I. Hino & Katum., J. Jap. Bot. 40: 86 (1965). 

≡ Didymosphaeria japonica I. Hino & Katum., Bulletin 

of the Faculty of Agriculture, Yamaguchi University 5: 229 

(1954). 

Roussoellopsis was introduced by Hino and Katumoto 

(1965) based on three bambusicolous fungal species, i.e. R. 

japonica, R. macrospora (I. Hino & Katum.) I. Hino & 

Katum. and R. tosaensis (I. Hino & Katum.) I. Hino & 

Katum. These three species have immersed and gregarious 

ascomata, clavate to cylindro-clavate asci, numerous and 

filliform pseudoparaphyses, and 1-septate, asymmetrical 

ascospores (Hino and Katumoto 1965). All these characters 

point Roussoellopsis to Pleosporales, but its familial 

placement cannot be determined. 

Saccothecium Fr., Fl. Scan.: 349 (1836). 

Type species: Saccothecium sepincola (Fr.) Fr. [as ‘saepincola’], 

Summa veg. Scand., Section Post. (Stockholm): 398 

(1849). 

≡ Sphaeria sepincola Fr. [as ‘saepincola’], Observ. 

mycol. (Havniae) 1: 181 (1815). 

Saccothecium is characterized by its subglobose, immersed 

to erumpent ascomata, absence of pseudoparaphyses 

and hyaline, muriform to phragmosporous ascospores. It has 

been assigned to the Dothioraceae (Barr 1987b; Müllerand 

von Arx 1950). Molecular phylogenetic analysis indicated 

that a strain named S. sepincola nested within Didymellaceae 

(Schoch et al. 2009; Plate 1). The generic type needs 

recollecting, redescribing and epitypifying. 

Setosphaeria K.J. Leonard & Suggs, Mycologia 66: 294 

(1974). 

Type species: Setosphaeria turcica (Luttr.) K.J. Leonard & 

Suggs, Mycologia 66: 295 (1974). 

≡ Trichometasphaeria turcica Luttr., Phytopathology 48: 

282 (1958). 

Setosphaeria was segregated from Keissleriella on the 

basis of lacking a clypeus, lysigenous development of the 

ostiole, occurrence of setae on the perithecial wall, the 

absence of periphyses in the ostiole, and the hyphomycetous 

conidial states, and four species were included, i.e. S. 

prolata, S. holmii, S. pedicellata (R.R. Nelson) K.J. 

Leonard & Suggs and S. turcica (Leonard and Suggs 

1974). Currently, nine species are included in Setosphaeria 

(http://www.mycobank.org, Jan/2011). Setosphaeria


monoceras Alcorn nested within Pleosporaceae based on 

multigene phylogenetic analysis (Schoch et al. 2009; 

Plate 1). 

Syncarpella Theiss. & Syd., Annls mycol. 13: 631 (1915). 

Type species: Syncarpella tumefaciens (Ellis & Harkn.) 

Theiss. & Syd., Annls mycol. 13(5/6): 633 (1915). 

≡ Sphaeria tumefaciens Ellis & Harkn., J. Mycol. 2: 41 

(1886). 

Syncarpella was introduced by Theissen and Sydow 

(1915) as a genus of Montagnellaceae within Dothideales. 

A detailed description of S. tumefaciens can be seen in Barr 

and Boise (1989). Syncarpella was considered closely 

related to Leptosphaeria, and was treated as a synonym 

(Clements and Shear 1931). Syncarpella is characterized by 

its abundant globose, ovoid to turbinate ascomata with 

minute papillae which are seated on a common basal stroma 

and which are erumpent through fissures in the host tissues 

(Barr and Boise 1989). The peridium is thicker at the base, 

pseudoparaphyses are cellular, and asci are bitunicate, 

clavate to oblong with a furcate pedicel. Ascospores are 

pale brown to brown, oblong to narrowly obovoid, ends 

obtuse, transversely septate, smooth-walled. All these 

characters fit Cucurbitariaceae, where Barr and Boise 

(1989) transferred Syncarpella. 

Teichospora Fuckel, Jb. nassau. Ver. Naturk. 23–24: 160 

(1870) [1869–70]. 

Type species: Teichospora trabicola Fuckel, Jb. nassau. 

Ver. Naturk. 23–24: 161 (1870) [1869–70]. 

Teichospora was introduced by Fuckel (1870), and was 

typified by T. trabicola, with four more species included, 

i.e. T. brevirostris Fuckel, T. dura Fuckel, T. morthieri 

Fuckel and T. obducens (Schumach.) Fuckel. Only T. 

brevirostris and T. trabicola were kept in Teichospora 

(Barr 1987b). After studying the type specimens, Barr 

(1987b) indicated that Teichospora was different from 

Strickeria with Teichospora belonging to Pleosporales, 

and Strickeria closely related to Melanomma (Melanommatales). 

Currently, more than 250 names are included 

within Teichospora (http://www.mycobank.org, Jan/2011), 

but almost no molecular phylogenetic study has been 

conductedonthisgenus. 

Testudina Bizz., Atti Inst. Veneto Sci. lett., ed Arti, Sér. 6 

3: 303 (1885). 

Type species: Testudina terrestris Bizz., Fungi venet. nov. 

vel. Crit. 3: 303 (1885). 

Testudina terrestris is characterized by its reticulately 

ridged ascospores, which readily distinguish it from other 

genera of Zopfiaceae (Hawksworth 1979). The species is 

usually associated with other fungi, or on the wood of Abies? 

and Pinus or on the fallen leaves of Taxus in Europe 

(Hawksworth and Booth 1974; Hawksworth1979). 

Tetraplosphaeria Kaz. Tanaka & K. Hirayama, Stud. 

Mycol. 64: 177 (2009). 

Type species: Tetraplosphaeria sasicola Kaz. Tanaka & K. 


Tetraplosphaeria was introduced by Tanaka et al. (2009) 

to accommodate bambusicolous fungi with immersed to 

erumpent, globose to subglobose and smaller (mostly< 

300 μm) ascomata. The peridium is thin, and is composed 

of thin-walled cells of textura angularis. The pseudoparaphyses 

are cellular, and asci are fissitunicate, 8-spored, 

cylindrical to clavate with short pedicels. Ascospores are 

narrowly fusoid, hyaline and surrounded with a sheath. 

Species of Tetraplosphaeria have Tetraploa sensu stricto 

anamorphic stage, which is quite unique in Tetraplosphaeriaceae 

(Tanaka et al. 2009). 

Tingoldiago K. Hirayama & Kaz. Tanaka, Mycologia 102: 

740 (2010). 

Type species: Tingoldiago graminicola K. Hirayama & 

Kaz. Tanaka, Mycologia 102(3): 740 (2010). 

Tingoldiago is a genus of freshwater ascomycetes characterized 

by flattened, globose, immersed to erumpent ascomata, 

and numerous cellular pseudoparaphyses (Hirayama et al. 

2010). Asci are fissitunicate and cylindrical, and ascospores 

are 1-septate, which usually turn 3-septate and pale brown 

when old, usually with a sheath (Hirayama et al. 2010). 

Based on both morphology and multigene phylogenetic 

analysis, Tingoldiago should be treated as a synonym of 

Lentithecium (Shearer et al. 2009a; Zhang et al. 2009a). 

Tremateia Kohlm., Volkm.-Kohlm. & O.E. Erikss., Bot. 

Mar. 38: 165 (1995). 

Type species: Tremateia halophila Kohlm., Volkm.-Kohlm. 

& O.E. Erikss., Bot. Mar. 38: 166 (1995). 

Tremateia was introduced as a facultative marine genus 

which is characterized by depressed globose, immersed 

ascomata, numerous and cellular pseudoparaphyses, fissitunicate 

and clavate asci, ellipsoid muriform ascospores, 

and a Phoma-like anamorph (Kohlmeyer et al. 1995). 

These characters point Tremateia to Pleosporaceae 

(Kohlmeyer et al. 1995). DNA sequence based phylogenies 

placed T. halophila as sister to Bimuria novae-zelandiae 

in Montagnulaceae (Schoch et al. 2009; Suetrong et al. 

2009). 

Triplosphaeria Kaz. Tanaka & K. Hirayama, Stud. Mycol. 

64: 186 (2009). 

Type species: Triplosphaeria maxima Kaz. Tanaka & K. 


Triplosphaeria was introduced as a bambusicolous genus 

characterized by immersed ascomata, numerous cellular


pseudoparaphyses, bitunicate, cylindrical to clavate asci with 

a short pedicel, fusoid, hyaline, 1-septate ascospores surrounded 

with a sheath, and with a Tetraploa-like anamorph 

(Tanaka et al. 2009). Together with Tetraplosphaeria, 

Pseudotetraploa, Quadricrura and Polyplosphaeria, Triplosphaeria 

was assigned to the Tetraplosphaeriaceae (Tanaka 

et al. 2009). 

Ulospora D. Hawksw., Malloch & Sivan., in Hawksworth, 

Can. J. Bot. 57: 96 (1979). 

Type species: Ulospora bilgramii (D. Hawksw., C. Booth 

& Morgan-Jones) D. Hawksw., Malloch & Sivan., Can. J. 

Bot. 57: 96 (1979). 

Ulospora was introduced as a monotypic genus to 

accommodate taxa of Testudinaceae whose ascospore has 

3–6 fissures (Hawksworth 1979). Genera of Testudinaceae 

are distinguished based on the morphology of ascospores, 

although the validity of this classification needs to be 

confirmed by molecular study. DNA sequence based 

phylogenies placed sequences from an unverified culture 

of U. bilgramii in a clade together with Verruculina enalia, 

and Lepidosphaeria nicotiae and it may have a close 

relationship to species in Platystomaceae (Mugambi and 

Huhndorf 2009b; Schoch et al. 2009; Plate 1). 

Zopfia Rabenh., Fungi europ. exsicc.: no. 1734 (1874). 

Type species: Zopfia rhizophila Rabenh., Fungi europ. 

exsicc.: no. 1734 (1874). 

Zopfia was introduced by Rabenhorst (1874) as a 

monotypic genus (typified by Z. rhizophila), and it was 

assigned to the Perisporiaceae by Saccardo (1882) and 

Winter (1884). Arnaud (1913) described the Zopfiaceae to 

accommodate Zopfia, and considered that it should be 

excluded from the Perisporiaceae. A relatively broad 

generic concept was accepted by Hawksworth and Booth 

(1974), in which they take the ascospore size and 

ornamentation variation as criteria under generic rank 

classification, and they treat Celtidia, Lepidosphaeria, 

Marchaliella, Neotestudina, Pontoporeia, Pseudophaeotrichum, 

Rechingeriella, Richonia and Testudina as synonyms 

of Zopfia. A narrow generic concept was adopted by 

Hawksworth (1979), and Zopfia is characterized by 1- 

septate ascospores, which are apiculate at both ends, 

smooth-walled by light microscope, with minute irregular 

pitting by SEM, and larger than other species of Zopfia 

sensu Hawksworth and Booth (1974). Three species were 

accepted, viz. Z. albiziae Farr, Z. biturbinata (Dur. & 

Mont.) Malloch & Cain and Z. rhizophila, and they all 

occur on roots of plants (Hawksworth 1979). DNA 

sequences from an unverified culture of Zopfia rhizophila 

placed it in close proximity to species in Delitschiaceae 

without strong statistical support (Kruys et al. 2006; Schoch 

et al. 2009; Plate 1). 

Zopfiofoveola D. Hawksw., Can. J. Bot. 57: 98 (1979). 

Type species: Zopfiofoveola punctata (D. Hawksw. & C. 

Booth) D. Hawksw., Can. J. Bot. 57: 98 (1979). 

≡ Zopfia punctata D. Hawksw. & C. Booth, Mycol. Pap. 

153: 23 (1974). 

Zopfiofoveola was hesitantly separated from Zopfia as a 

monotypic new genus based on its evenly distributed 

ornamentation with pale minute pits readily visible under 

the light microscope, and the more elongate shape and less 

pronounced apical papilla than those of Zopfia (Hawksworth 

1979). The type specimen of this species however, cannot be 

redescribed, because “the type species is only known from a 

microscopic preparation obtained from earthworm excrements 

in Sweden” as has been mentioned by Hawksworth (1979). 

General discussion 

Molecular phylogenetic studies based on four to five genes 

indicate that 20 families should be included in Pleosporales 

(Schoch et al. 2009; Shearer et al. 2009; Suetrong et al. 

2009; Tanaka et al. 2009; Zhang et al. 2009a). Together 

with five unverified families (marked with “?”), 26 families 

are currently assigned under Pleosporales (Table 4). The 

Phaeotrichaceae lacks pseudoparaphyses, has cleistothecial 

ascomata with long setae, and conspicuous ascospores with 

germ pores at each end. These characters do not agree with 

the current concept of Pleosporales (Zhang et al. 2009a), 

and therefore Phaeotrichaceae is excluded from Pleosporales 

(Table 4). 

Families in Pleosporales 

Based on LSU and SSU rDNA, RPB1, RPB2 and TEF1 

sequence analysis, Pleosporineae is emended, and in this 

study, seven families are tentatively included, i.e. Cucurbitariaceae, 

Didymellaceae, Didymosphaeriaceae, Dothidotthiaceae, 

Leptosphaeriaceae, Phaeosphaeriaceae and 

Pleosporaceae (Zhang et al. 2009a; Plate1). In this study, 

Massarineae was emended to accommodate another five 

families, viz. Lentitheciaceae, Massarinaceae, Montagnulaceae, 

Morosphaeriaceae, Trematosphaeriaceae. The subordinal 

affinity of other families remained undetermined. 

Most of the families accepted within Pleosporales received 

high bootstrap support (Plate 1). The characters used to 

define a family, however, do not appear to have clear cut 

boundaries, as the ascomatal and hamathecial characters also 

seem to be poorly defined in some families. For example, 

both trabeculate and cellular pseudoparaphyses coexist in the 

Amniculicolaceae. Pycnidiophora, a genus of Sporormiaceae, 

has cleistothecial ascomata with spherical asci irregularly 

arranged in it. Brown phragmosporous ascospores are 

reported in Amniculicolaceae, Leptosphaeriaceae, Lophios-


tomataceae, Melanommataceae, Montagnulaceae, Phaeosphaeriaceae 

and Pleosporaceae. Similarly muriform ascospores 

occur in Aigialaceae, Amniculicolaceae, 

Didymellaceae, Lophiostomataceae, Montagnulaceae, Pleosporaceae 

and Sporormiaceae. AnamorphsofPleosporales 

are also variable to a large degree at the family level. Both 

hyphomycetous and coelomycetous anamorphs co-exist in 

Didymellaceae, Melanommataceae or Pleosporaceae. 

Phoma and Phoma-like anamorphs exist in Didymellaceae, 

Leptosphaeriaceae, Phaeosphaeriaceae, Pleosporaceae and 

Melanommataceae (de Gruyter et al. 2009; Zhang et al. 

2009a). It is clear that some characters, e.g. cleistothecial or 

perithecial ascomata, shape, colour and septation of ascospores, 

shape or arrangement (regular or irregular) of asci, or 

even presence or absence of pseudoparaphyses have evolved 

on numerous occasions which make the use of morphological 

characters in segregating families complicated. It is 

therefore unclear with our present state of knowledge which 

characters are taxonomically important at the family level or 

whether a suit of characters are necessary to define a family. 

DNA sequence comparisons are essential in delineating 

these taxa in combination with other characters. It is hoped 

that additional characters, i.e. biochemical, genomic and 

subcellular will be used to further distinguish these groups 

into natural taxa. Below we discuss each of the families, 

their genera and their considered important characteristics. 

Aigialaceae Suetrong, Sakay., E.B.G. Jones, Kohlm., 

Volkm.-Kohlm. & C.L. Schoch 2010 

The Aigialaceae was introduced by Suetrong et al. 

(2009) based on its carbonaceous ascomata without papilla, 

cylindrical asci with apical apparatus, trabeculate pseudoparaphyses 

and ascospores with a sheath. The type genus 

(Aigialus) of the Aigialaceae was previously incorporated 

within the Massariaceae (Lumbsch and Huhndorf 2007). 

Currently, three genera are assigned under Aigialaceae, viz. 

Ascocratera, Aigialus and Rimora (Suetrong et al. 2009). 

The genera included in Aigialaceae have a wide range of 

morphological variation, with very few shared features as 

mentioned above, but all are found in mangrove habitats 

(Suetrong et al. 2009). The ascospores, however, vary 

widely from having 1 to 3 transverse septa and being hyaline 

to muriformly septate and brown (Suetrong et al. 2009). It is 

still unclear which characters unify the family and therefore 

placement of unsequenced genera is difficult. Further 

molecular work is needed to better understand this family. 

Amniculicolaceae Yin. Zhang, C.L. Schoch, J. Fourn., 

Crous & K.D. Hyde 2009 

Members of Amniculicolaceae form a well supported 

clade, and all are freshwater fungi which usually stain the 

woody substrate purple (Zhang et al. 2009a, c). Genera of 

Amniculicolaceae have ascomata with compressed papilla 

Table 4 Families currently accepted in Pleosporales (syn. Melanommatales) 

with included genera 

Pleosporales subordo. Pleosporineae 

?Cucurbitariaceae 

Cucurbitaria Gray 

Curreya Sacc. 

?Rhytidiella Zalasky 

Syncarpella Theiss. & Syd. 

Didymellaceae 

?Appendispora K.D. Hyde 

Didymella Sacc. ex D. Sacc. 

Didymosphaerella Cooke 

Leptosphaerulina McAlpine 

Macroventuria Aa 

?Platychora Petr. 

Didymosphaeriaceae 

Didymosphaeria Fuckel 

Phaeodothis Syd. & P. Syd. 

Dothidotthiaceae 

Dothidotthia Höhn. 

Leptosphaeriaceae 

Leptosphaeria Ces. & De Not. 

Neophaeosphaeria Câmara, M.E. Palm & A.W. Ramaley 

Phaeosphaeriaceae 

Barria Z.Q. Yuan 

?Bricookea M.E. Barr 

?Chaetoplea (Sacc.) Clem. 

?Eudarluca Speg. 

Entodesmium Reiss 

?Hadrospora Boise 

Lautitia S. Schatz 

Loratospora Kohlm. & Volkm.-Kohlm. 

Metameris Theiss. & Syd. 

Mixtura O.E. Erikss. & J.Z. Yue 

Nodulosphaeria Rabenh. 

Ophiobolus Reiss 

Ophiosphaerella Speg. 

Phaeosphaeria I. Miyake 

Phaeosphaeriopsis Câmara, M.E. Palm & A.W. 

Ramaley 

Pleoseptum A.W. Ramaley & M.E. Barr 

Setomelanomma M. Morelet 

Wilmia Dianese, Inácio & Dornelo-Silva 

Pleosporaceae 

Cochliobolus Drechsler 

Crivellia Shoemaker & Inderbitzin 

Decorospora Inderbitzin, Kohlm. & Volkm.-Kohlm. 

Extrawettsteinina M.E. Barr 

Lewia M.E. Barr & E.G. Simmons 

Macrospora Fuckel 

Platysporoides (Wehm.) Shoemaker & C.E. Babc. 

Pleospora Rabenh. ex Ces. & De Not. 

Pseudoyuconia Lar. N. Vasiljeva 

Pyrenophora Fr. 

Setosphaeria K.J. Leonard & Suggs 

Pleosporales subordo. Massarineae 

Lentitheciaceae 

Lentithecium K.D. Hyde, J. Fourn. & Yin. Zhang



Katumotoa Kaz. Tanaka & Y. Harada 

Keissleriella Höhn. 

?Wettsteinina Höhn. 

Massarinaceae 

Byssothecium Fuckel 

Massarina Sacc. 

Saccharicola D. Hawksw. & O.E. Erikss. 

Montagnulaceae 

Bimuria D. Hawksw., Chea & Sheridan 

?Didymocrea Kowalsky 

Kalmusia Niessl 

Karstenula Speg. 

Letendraea Sacc. 

Montagnula Berl. 

Paraphaeosphaeria O.E. Erikss. 

Tremateia Kohlm., Volkm.-Kohlm. & O.E. Erikss. 

Morosphaeriaceae 

?Asteromassaria Höhn 

Helicascus Kohlm. 

Morosphaeria Suetrong, Sakay., E.B.G. Jones & C.L. Schoch 


Falciformispora K.D. Hyde 

Halomassarina Suetrong, Sakay., E.B.G. Jones, Kohlm., Volkm.-Kohlm. & C.L. 

Schoch 

Trematosphaeria Fuckel 

Other families 

Aigialaceae 

Aigialus Kohlm. & S. Schatz 

Ascocratera Kohlm. 

Rimora Kohlm., Volkm.-Kohlm., Suetrong, Sakay. & E.B.G. Jones 

Amniculicolaceae 

Amniculicola Y. Zhang & K.D. Hyde 

Murispora Yin. Zhang, C.L. Schoch, J. Fourn., Crous & K.D. Hyde 

?Massariosphaeria (E. Müll.) Crivelli 

Neomassariosphaeria Yin. Zhang, J. Fourn. & K.D. Hyde 

?Arthopyreniaceae (Massariaceae) 

Arthopyrenia A. Massal. 

Dothivalsaria Petr. 

?Dubitatio Speg. 

Massaria De Not. 

Navicella Fabre 

Roussoëlla Sacc. 

?Roussoellopsis I. Hino & Katum. 

Delitschiaceae 

Delitschia Auersw. 

Ohleriella Earle 

Semidelitschia Cain & Luck-Allen 

?Diademaceae 

Clathrospora Rabenh. 

Comoclathris Clem. 

Diadema Shoemaker & C.E. Babc. 

Diademosa Shoemaker & C.E. Babc. 

Graphyllium Clem. 

Hypsostromataceae 

Hypsostroma Huhndorf 

Lindgomycetaceae 

Lindgomyces K. Hirayama, Kaz. Tanaka & Shearer 2010 



Lophiostoma Ces. & De Not. 

Melanommataceae 

?Astrosphaeriella Syd. & P. Syd. (Syn. Javaria) 

?Anomalemma Sivan. 

?Asymmetricospora J. Fröhl. & K.D. Hyde 

Bertiella (Sacc.) Sacc. & P. Syd. 

?Bicrouania Kohlm. & Volkm.-Kohlm. 

Byssosphaeria Cooke 

Calyptronectria Speg. 

?Caryosporella Kohlm. 

Herpotrichia Fuckel 

?Mamillisphaeria K.D. Hyde, S.W. Wong & E.B.G. Jones 

Melanomma Nitschke ex Fuckel 

Ohleria Fuckel 

Pseudotrichia Kirschst. 

Pleomassariaceae 

?Lichenopyrenis Calatayud, Sanz & Aptroot 

?Splanchnonema Corda 

?Peridiothelia D. Hawksw. 

Pleomassaria Speg. 

Sporormiaceae 

Chaetopreussia Locq.-Lin. 

Eremodothis Arx 

Pleophragmia Fuckel 

Preussia Fuckel 

Sporormia De Not. 

Westerdykella Stolk 

?Teichosporaceae 

Chaetomastia (Sacc.) Berl 

Immotthia M.E. Barr 

Loculohypoxylon M.E. Barr 

Sinodidymella J.Z. Yue & O.E. Erikss. 

Teichospora Fuckel 

Tetraplosphaeriaceae 

Polyplosphaeria Kaz. Tanaka & K. Hirayama 

Tetraplosphaeria Kaz. Tanaka & K. Hirayama 

Triplosphaeria Kaz. Tanaka & K. Hirayama 

?Zopfiaceae (syn Testudinaceae) 

Caryospora De Not. 

Celtidia J.M. Janse 

?Coronopapilla Kohlm. & Volkm.-Kohlm. 

Halotthia Kohlm. 

Lepidosphaeria Parg.-Leduc 

?Mauritiana Poonyth, K.D. Hyde, Aptroot & Peerally 

Pontoporeia Kohlm. 

?Rechingeriella Petr. 

Richonia Boud. 

Testudina Bizz. 

Verruculina Kohlm. & Volkm.-Kohlm. 

Ulospora D. Hawksw., Malloch & Sivan. 

Zopfia Rabenh. 

Zopfiofoveola D. Hawksw. 

Pleosporales genera incertae sedis 

Acrocordiopsis Borse & K.D. Hyde 

Aglaospora De Not. 

Anteaglonium Mugambi & Huhndorf 

Ascorhombispora L. Cai & K.D. Hyde 

Atradidymella Davey & Currah



Belizeana Kohlm. & Volkm.-Kohlm. 

Biatriospora K.D. Hyde & Borse 

Byssolophis Clem. 

Carinispora K.D. Hyde 

Cilioplea Munk 

Decaisnella Fabre 

Epiphegia Nitschke ex G.H. Otth 

Isthmosporella Shearer & Crane 

Julella Fabre 

Lineolata Kohlm. & Volkm.-Kohlm. 

Lophiella Sacc. 

Lophionema Sacc. 

Lophiotrema Sacc. 

Moristroma A.I. Romero & Samuels 

Neotestudina Segretain & Destombes 

Ostropella (Sacc.) Höhn. 

Paraliomyces Kohlm. 

Passeriniella Berl. 

Quintaria Kohlm. & Volkm.-Kohlm. 

Saccothecium Fr. 

Salsuginea K.D. Hyde 

Shiraia P. Henn. 

Xenolophium Syd. 

Family excluded 

Phaeotrichaceae 

Echinoascotheca Matsush. 

Phaeotrichum Cain & M.E. Barr 

Trichodelitschia Munk 

Genera excluded 

Kriegeriella Höhn. 

Muroia I. Hino & Katum. 

Zeuctomorpha Sivan., P.M. Kirk & Govindu 

and cylindrical to cylindro-clavate asci. Neomassariosphaeria 

typhicola was traditionally assigned to Massariosphaeria (as 

M. typhicola), and Massariosphaeria is characterized by 

staining the woody substrate purple (Crivelli 1983; 

Leuchtmann 1984). Eriksson (1981 p. 135) had pointed out 

that “Purple-staining species of Pleospora, treated by Webster 

(1957), are not congeneric with P. herbarum (Eriksson 1967b: 

13), and certainly do not even belong to the Pleosporaceae”. 

This is mirrored in Murispora rubicunda, a 

previous Pleospora species (as P. rubicunda) staining the 

woody substrate purple, closely related to the Amniculicolaceae 

in a subsequent phylogenetic study (Zhang et al. 

2009a). The anamorphs of this family are possibly 

Anguillospora longissima, Spirosphaera cupreorufescens 

and Repetophragma ontariense (Zhang et al. 2009a). 

?Arthopyreniaceae (or Massariaceae) W. Watson 1929 

The Arthopyreniaceae was introduced as a lichenized 

family of Pyrenocarpales, which comprises Acrocordia, 

Arthopyrenia, Athrismidium, Bottaria, Celothelium, Laurera, 

Leptorhaphis, Microthelia, Microtheliopsis, Polyblastiopsis, 

Pseudosagedia, Raciborskiella and Tomasellia 

(Watson 1929). Subsequently, Arthopyreniaceae was 

assigned under Dothideales (suborder Pseudosphaeriineae) 

(von Arx and Müller 1975). The generic type of Massaria 

(M. inquinans) and Torula herbarum and Arthopyrenia 

salicis together with members of Roussoella as well as 

Roussoellopsis form a robust clade, which makes their 

familial placement uncertain (Massariaceae or Arthopyreniaceae) 

(Schoch et al. 2009; Zhang et al. 2009a). 

?Cucurbitariaceae G. Winter 1885 

The Cucurbitariaceae is characterized by its aggregated 

ascomata which form from a basal stromatic structure, 

ostiolate, fissitunicate and cylindrical asci, and pigmented, 

phragmosporous or muriform ascospores (Cannon and Kirk 

2007). Currently, no molecular study has been able to 

resolve its ordinal status, but some characters are similar to 

Leptosphaeriaceae or Phaeosphaeriaceae (Cannon and 

Kirk 2007). Cucurbitaria elongata clustered within Pleosporales 

(Schoch et al. 2006). 

Delitschiaceae M.E. Barr 2000 

The Delitschiaceae was established to accommodate 

some species of the Sporormiaceae, which is characterized 

by its ascomata with periphysate ostioles, ocular chamber 

surrounded by a dome and usually in having four refractive 

rods, ascospores with or without a septum, having a germ 

slit in each cell and being surrounded by a mucilaginous 

sheath (Barr 2000). Species of the Delitschiaceae are 

hypersaprotrophic on old dung or exposed wood (Barr 

2000). Based on a molecular phylogenetic studies, Delitschia 

didyma and D. winteri form a robust clade basal to 

other pleosporalean fungi (Schoch et al. 2009; Zhang et al. 

2009a). The familial status of two other genera, Ohleriella 

and Semidelitschia, remains undetermined. 

?Diademaceae Shoemaker & C.E. Babc. 1992 

The Diademaceae was introduced by Shoemaker and 

Babcock (1992) based on its ascomata opening as a flat 

circular lid and bitunicate asci, ascospores are fusiform, 

brown, mostly applanate, and having three or more 

transverse septate and with or lacking longitudinal septa 

and usually having a sheath. Five genera had been included 

viz. Clathrospora, Comoclathris, Diadema, Diademosa and 

Macrospora (Shoemaker and Babcock 1992). 

Didymellaceae Gruyter, Aveskamp & Verkley 2009 

The generic type of Didymella (D. exigua) together with 

some Phoma or Phoma-related species form a robust 

familial clade on the phylogenetic tree, thus the Didymellaceae 

was introduced to accommodate them (de Gruyter et 

al. 2009). Subsequently, Didymellaceae was assigned to


Pleosporineae (suborder of Pleosporales) (Zhang et al. 

2009a). A detailed study was conducted on the Didymellaceae 

based on LSU, SSU rDNA, ITS as well as β-tubulin, 

which indicated that many Phoma or Phoma-related 

species/fungi reside in this clade of the Didymellaceae 

(Aveskamp et al. 2010). 

Didymosphaeriaceae Munk 1953 

The Didymosphaeriaceae was introduced by Munk (1953), 

and was revived by Aptroot (1995) based on its distoseptate 

ascospores and trabeculate pseudoparaphyses, mainly anastomosing 

above the asci. The familial status of the Didymosphaeriaceae 

is debatable, and Lumbsch and Huhndorf (2007) 

assigned it to the Montagnulaceae, while von Arx and Müller 

(1975) treated it as a synonym of the Pleosporaceae. Inthis 

study, Didymosphaeria futilis (the generic type of Didymosphaeria) 

is closely related to the Cucurbitariaceae (Plate 1). 

Herein, we accept it as a separate family containing three genera, 

namely Appendispora, Didymosphaeria and Phaeodothis. More 

information could only be obtained by further molecular work 

based on correctly identified strains. 

Dothidotthiaceae Crous & A.J.L. Phillips 2008 

Dothidotthiaceae was introduced to accommodate the 

single genus Dothidotthia, which is characterized by gregarious, 

erumpent, globose ascomata, hyaline, septate pseudoparaphyses, 

8-spored, bitunicate, clavate asci, ellipsoid, 1- 

septate ascospores, and has anamorphic Thyrostroma 

(Phillips et al. 2008). In this study, Dothidotthiaceae is 

closely related to Didymellaceae, but it is still treated as a 

separate family (Plate 1). 

Hypsostromataceae Huhndorf 1994 

Hypsostromataceae was introduced based on two tropical 

genera (i.e. Hypsostroma and Manglicola), which have superficial, 

large, elongate ascomata with a soft-textured, pseudoparenchymatic 

wall, trabeculate pseudoparaphyses and stipitate asci 

attached in a basal arrangement in the centrum; asci with an 

apical chamber and fluorescing ring; and fusiform, septate 

ascospores (Huhndorf 1994). Hypsostromataceae was assigned 

to Melanommatales sensu Barr (Huhndorf 1994). In a 

subsequent phylogenetic study, Hypsostromataceae was recovered 

as a strongly supported monophyletic group nested within 

Pleosporales (Mugambi and Huhndorf 2009b). 

Lentitheciaceae Yin. Zhang, C.L. Schoch, J. Fourn., Crous 

& K.D. Hyde 2009 

Phylogenetic analysis based on multi-genes indicate that 

freshwater taxa, e.g. Lentithecium fluviatile, L. arundinaceum, 

Stagonospora macropycnidia, Wettsteinina lacustris, 

Keissleriella cladophila, Katumotoa bambusicola and 

Ophiosphaerella sasicola form a well supported clade, which 

most likely represent a familial rank (Zhang et al. 2009a). 

Their morphology, however, varies widely, e.g. ascomata 

small- to medium-sized, ascospores fusoid to filliform, 

hyaline to pale yellow, 1- to multi-septate (Zhang et al. 

2009a). In particular, they are saprobic on monocotyledons 

or dicotyledons. Currently, no conspicuous, unique morphological 

character has been noted in Lentitheciaceae, which 

makes it difficult to recognize based on morphology. 

Leptosphaeriaceae M.E. Barr 1987a 

The Leptosphaeriaceae was introduced by Barr (1987a) 

based on Leptosphaeria. The familial status of the Leptosphaeriaceae 

is subsequently supported by molecular phylogenetic 

studies, in which members of the Leptosphaeriaceae 

form a paraphyletic clade with moderate bootstrap support 

(Dong et al. 1998; de Gruyter et al. 2009;Schochetal.2009; 

Zhang et al. 2009a). Coniothyrium palmarum, the generic 

type of Coniothyrium nested within this family (de Gruyter 

et al. 2009). Further molecular phylogenetic study is needed, 

in which more related taxa are included. 

Lindgomycetaceae K. Hirayama, Kaz. Tanaka & Shearer 2010 

Lindgomycetaceae was introduced as a monotypic 

family represented by Lindgomyces (Hirayama et al. 

2010). Lindgomycetaceae is another freshwater family in 

Pleosporales, which is characterized by its subglobose to 

globose, ostiolate and papillate ascomata, numerous, septate, 

branching and anastomosing pseudoparaphyses, fissitunicate, 

cylindrical to clavate, 8-spored asci, fusiform to 

cylindrical, uni- to multiseptate, hyaline to brown ascospores 

usually covered with an entire sheath and/or bipolar 

mucilaginous appendages (Hirayama et al. 2010). 

Lophiostomataceae Sacc. 1883 

The Lophiostomataceae had been characterized by its 

slot-like ostiole on the top of a flattened neck (Holm and 

Holm 1988). Based on this, 11 genera were assigned under 

the Lophiostomataceae, viz. Byssolophis, Cilioplea, Entodesmium, 

Herpotrichia, Lophiella, Lophionema, Lophiostoma, 

Lophiotrema, Massariosphaeria, Muroia and 

Quintaria (Holm and Holm 1988). The Lophiostomataceae 

was thought to be heterogeneous, as the “papilla form is an 

unstable and highly adaptive character” (Holm and Holm 

1988). Most recent phylogenetic analysis support the 

monophyletic status of the Lophiostomataceae sensu stricto 

(which tends to comprise a single genus of Lophiostoma) 

(Zhang et al. 2009a, b). The familial placement of other 

genera, however, remains unresolved. 

Massarinaceae Munk 1956 

The Massarinaceae was established based on Keissleriella, 

Massarina, Metasphaeria, Pseudotrichia and Trichometasphaeria 

(Munk 1956). Subsequently, the Massarinaceae is 

sometimes treated as a synonym of Lophiostomataceae (Barr


1987b). Based on a multigene phylogenetic study, the generic 

type of Massarina (M. eburnea) together with M. cisti, 

Neottiosporina paspali and Byssothecium circinans form a 

well supported clade (Zhang et al. 2009a, b). It seems that a 

relatively narrow familial concept should be accepted. 

Melanommataceae G. Winter 1885 

The traditional circumscription of the Melanommataceae 

was based on its globose or depressed perithecial ascomata, 

bitunicate and fissitunicate asci, pigmented phragmosporous 

ascospores as well as the trabeculate pseudoparaphyses 

(Barr 1990a; Sivanesan 1984). However, the family has 

recently proved polyphyletic (Liew et al. 2000; Kodsueb et 

al. 2006a; Kruys et al. 2006; Wang et al. 2007). Bimuria, 

Ostropella, Trematosphaeria and Xenolophium occur outside 

Melanommataceae (Mugambi and Huhndorf 2009b; 

Zhang et al. 2009a). Species of Byssosphaeria, Bertiella, 

Herpotrichia, Pseudotrichia, Pleomassaria as well as 

Melanomma resided in the clade of Melanommataceae 

(Mugambi and Huhndorf 2009b; Schoch et al. 2009; Zhang 

et al. 2009a). The familial status of many genera previously 

listed under this family remains to be sorted out (Lumbsch 

and Huhndorf 2007). 

Montagnulaceae M.E. Barr 2001 

The Montagnulaceae was introduced to accommodate 

some pleosporalean genera with ascomata immersed under 

a clypeus, a pseudoparenchymatous peridium with small 

cells, cylindric or oblong asci with pedicels and brown 

ascospores (Barr 2001). Three genera were included, i.e. 

phragmosporous Kalmusia, dictyosporous Montagnula and 

didymosporous Didymosphaerella (Barr 2001). Our molecular 

phylogenetic analysis based on multi-genes indicated 

that species from Kalmusia, Phaeosphaeria, Bimuria, 

Didymocrea, Paraphaeosphaeria, Karstenula, Letendraea 

as well as Montagnula resided in the monophylogenetic 

clade of the Montagnulaceae (Schoch et al. 2009; Zhang et 

al. 2009a). 

Morosphaeriaceae Suetrong, Sakay., E.B.G. Jones & C.L. 

Schoch 2009 

Four marine species, viz. Massarina ramunculicola 

(as Morosphaeria ramunculicola), Massarina velataspora 

(Morosphaeria velataspora), Helicascus kanaloanus and 

H. nypae together with the freshwater species Kirschsteiniothelia 

elaterascus form a well supported clade, which 

most likely represent a familial rank (Suetrong et al. 

2009). Thus, Morosphaeriaceae was introduced to accommodate 

these taxa (Suetrong et al. 2009). In 

this study, Asteromassaria pulchra is basal to other 

species of Morosphaeriaceae, and gets well support 

(Plate 1). Thus we tentatively assign Asteromassaria under 

Morosphaeriaceae. 

Phaeosphaeriaceae M.E. Barr 1979a 

The Phaeosphaeriaceae was introduced to accommodate 

some pleosporalean genera which have saprobic, parasitic or 

hyperparasitic lifestyles and have small- to medium-sized, 

subglobose or conical ascomata, bitunicate asci and hyaline or 

pigmented ascospores with or without septation (Barr 1979a). 

Fourteen genera were included, viz. Comoclathris, Didymella, 

Eudarluca, Heptameria, Leptosphaeria, Loculohypoxylon, 

Metameris, Microthelia, Nodulosphaeria, Ophiobolus, Paraphaeosphaeria, 

Rhopographus, Scirrhodothis and Teichospora 

(Barr 1979a), which were subsequently assigned to 

various families, such as Loculohypoxylon and Teichospora to 

the Teichosporaceae, Paraphaeosphaeria to the Montagnulaceae, 

Leptosphaeria to the Leptosphaeriaceae, Comoclathris 

to the Diademaceae, Didymella to the Didymellaceae and 

Heptameria and Rhopographus to genera incertae sedis of 

Dothideomycetes (Aveskamp et al. 2010; de Gruyter et al. 

2009; Lumbsch and Huhndorf 2007; Zhang et al. 2009a). 

Based on multi-gene phylogenetic analysis, a relatively narrow 

familial concept is accepted, which is mostly associated with 

monocotyledons, with perithecoid, small- to medium-sized 

ascomata, and septate ascospores which are fusiform to 

filliform (Zhang et al. 2009a). Four genera were accepted, 

Ophiosphaerella, Phaeosphaeria, Entodesmium and Setomelanomma 

(Zhang et al. 2009a). Together with Cucurbitariaceae, 

Didymellaceae, Didymosphaeriaceae, Dothidotthiaceae, Leptosphaeriaceae 

and Pleosporaceae, thePhaeosphaeriaceae is 

assigned under Pleosporineae (Zhang et al. 2009a). 

Pleomassariaceae M.E. Barr 1979a 

Both Asteromassaria and Splanchnonema were designated 

as representative genera of Pleomassariaceae (Barr 

1979a). Currently, four genera are included in Pleomassariaceae, 

viz. ?Lichenopyrenis, ?Splanchnonema, ?Peridiothelia 

and Pleomassaria (Table 4). The generic type of 

Pleomassaria (P. siparia) clustered with species of Melanommataceae 

in previous and present studies (Schoch et al. 

2009; Zhang et al. 2009a; Plate 1). Zhang et al. (2009a) has 

attempted to assign Pleomassariaceae to Melanommataceae 

(Zhang et al. 2009a). Based on the distinct morphology and 

anamorphic stage of Pleomassaria siparia as well as the 

divergence of dendrogram, we hesitantely reinstate Pleomassariaceae 

as a separate family in this study. 

Pleosporaceae Nitschke 1869 

The Pleosporaceae is one of the earliest introduced 

families in Dothideomycetes. The Pleosporaceae was 

originally assigned under Sphaeriales, which accommodated 

species with paraphyses and immersed perithecia (Ellis 

and Everhart 1892; Lindau 1897; Winter 1887). Subsequently, 

many of the Pleosporaceae species were transferred 

to the Pseudosphaeriaceae, which was subsequently 

elevated to ordinal rank as Pseudosphaeriales (Theissen


and Sydow 1918). Luttrell (1955) introduced the Pleosporales 

(lacking a Latin description), which is characterized by 

its Pleospora-type of centrum development. Based on this, 

the Pleosporaceae and the Lophiostomataceae as well as 

other five families were placed in Pleosporales (Luttrell 

1955). Pleosporaceae is the largest and most typical family 

in Pleosporales. Wehmeyer (1975) stated that the Pleosporatype 

centrum development is verified in a small number of 

genera, and centrum development in the majority of genera 

is unknown; thus the placement of families or genera is quite 

arbitrary. In addition, the circumscription of Pleosporaceae 

is not clear-cut, and “……ascostromata of many different 

types, which are previously placed in various other families 

(Trichosphaeriaceae, Melanommataceae, Cucurbitariaceae, 

Amphisphaeriaceae etc.) are to be found here” (Wehmeyer 

1975). Thus, the heterogeneous nature of Pleosporales is 

obvious (Eriksson 1981), and had been confirmed by 

subsequent molecular phylogenetic studies (e.g. Kodsueb et 

al. 2006a). Based on the multi-gene phylogenetic analysis, 

some species from Lewia, Cochliobolus, Pleospora, Pyrenophora 

and Setosphaeria resided in the Pleosporaceae 


Sporormiaceae Munk 1957 

The Sporormiaceae is the largest coprophilous family in 

Pleosporales, which bears great morphological variation. 

Ascomata vary from cleistothecoid to perithecoid, asci are 

regularly or irregularly arranged, clavate or spherical, 

ascospores with or without germ slits or ornamentations. 

Based on phylogenetic analysis, Sporormiaceae is most 

likely monophyletic as currently circumscribed (Kruys et al. 

2006; Kruys and Wedin 2009). 

?Teichosporaceae M.E. Barr 2002 

The Teichosporaceae was introduced by segregating some 

non-lichenized members of the Dacampiaceae which are 

apostrophic on woody stems and periderm or hypersaprotrophic 

on other ascomycetous fungi (Barr 2002). The Dacampiaceae 

together with its synonym, Pyrenidiaceae was only 

maintained to accommodate its lichenicolous genera (Barr 

2002). This proposal does not have any molecular phylogenetic 

support. 

Tetraplosphaeriaceae Kaz. Tanaka & K. Hirayama 2009 

The Tetraplosphaeriaceae was introduced to accommodate 

five genera, i.e. Tetraplosphaeria, Triplosphaeria, 

Polyplosphaeria and the anamorphic genera Pseudotetraploa 

and Quadricrura (Tanaka et al. 2009). The Tetraplosphaeriaceae 

is characterized by its Massarina-like 

teleomorphs and its Tetraploa-like anamorphs with setaelike 

appendages, and its monophylogenetic status has been 

recently confirmed based on DNA phylogenetic studies 

(Tanaka et al. 2009). 


Three species, viz. Falciformispora lignatilis, Halomassarina 

thalassiae and Trematosphaeria pertusa form a 

robust clade, which forms a sister group with other 

pleosporalean families (Schoch et al. 2009; Suetrong et al. 

2009). Trematosphaeriaceae is waiting to be formally 

proposed (Suetrong et al. data unpublished). 

?Zopfiaceae G. Arnaud ex D. Hawksw. 1992 

The Zopfiaceae was introduced by Arnaud (1913), but 

was invalid due to the lack of a Latin diagnosis (see 

comments by Eriksson and Hawksworth 1992). The Zopfiaceae 

was formally introduced by Eriksson and Hawksworth 

(1992), and is characterized by its cleistothecial ascomata, 

thick-walled peridium, globose or saccate asci and oneseptate, 

dark brown ascospores (Cannon and Kirk 2007). 

Currently, eleven genera are included, but the family is likely 

polyphyletic (Kruys et al. 2006). 

Excluded family 

Phaeotrichaceae Cain 1956 

The cleistothecioid ascomata, ascospores with germ pore 

at each end and the absence of pseudoparaphyses indicate 

that the Phaeotrichaceae may not be closely related to 

Pleosporales. This was confirmed by DNA based phylogenies 

(Schoch et al. 2009). Thus, we exclude it from 

Pleosporales. 

Final remarks 

Problems and concerns 

Recently, many new pleosporalean lineages from freshwater 

(Shearer et al. 2009; Zhang et al. 2009a), marine 

(Suetrong et al. 2009) or from bambusicolous hosts (Tanaka 

et al. 2009) have been reported. In particular, large-scale 

phylogenetic analysis indicate that numerous unresolved 

clades still exist, which may also indicate that a large 

number of fungal lineages are not resolved. As has been 

estimated, 95% of all fungi are unreported (Hawksworth 

1991), and a large portion of them might exist only as 

hyphae (or DNA-only fungi, Taylor 1993). Under the 

influence of human activities, environmental situations are 

changing quickly, which may result in numerous fungal 

taxa losing their habitats and/or become endangered. More 

field work is urgently needed. 

A future polyphasic approach to study Pleosporales 

The use of DNA sequence comparisons have proved 

invaluable in modern concepts of fungal taxonomy. It is now 

clear many fungi do not produce reproductive structures or


only do so under very rare circumstances and many fungi 

cannot be cultured (Begerow et al. 2010). More and more 

morpho-species have proven to be cryptic taxa, and already a 

large percentage of fungal diversity is documented only by 

DNA sequences. DNA sequence analysis is an essential way 

to resolve these problems. But are they enough for fully 

informed fungal taxonomy? Each single morphological 

character may be the outcome of the expression of one to 

numerous genes, which might be composed of thousands of 

base pairs. DNA barcoding methods are “a breakthroughfor 

identification, but they will not supplant the need to 

formulate and rigorously test species hypothesis” (Wheeler 

et al. 2004). Thus, integration of classical morphological 

approaches and DNA and protein based sequence comparisons 

are critical to produce a modern taxonomy that reflects 

evolutionary similarities and differences (DeSalle et al. 2005; 

Godfray 2002). In particular, the advent of comparative 

genomics and advances in our understanding of secondary 

metabolites and host or habitat spectra allow the possibility 

to tie phylogenetic hypotheses derived from DNA and 

protein sequence to the biology of the organisms. (Bitzer et 

al. 2008; Stajich et al. 2009; Zhang et al. 2009a, b). 

Acknowledgement We are grateful to the Directors and Curators of 

the following herbaria for loan of specimens in their keeping: BAFC, 

BISH, BPI, BR, BRIP, CBS, E, ETH, FFE, FH, G, H, Herb. J. 

Kohlmeyer, HHUF, IFRD, ILLS, IMI, K(M), L, LPS, M, MA, NY, 

PAD, PC, PH, RO, S, TNS, TRTC, UB, UBC, UPS and ZT; to Dr. L. 

Cai, Dr. A.J.L. Phillips, Dr. C. Shearer and some other mycologists for 

their permission to use or refer to their published figures, to J.K. Liu, 

H. Zhang, Y.L. Yang and J. Fournier for helping me loan or collect 

specimens, to H. Leung for technical help. The third coauthor 

acknowledges the Intramural Research Program of the NIH, National 

Library of Medicine. The Global Research Network and King Saud 

University are also thanked for support. 

Open Access This article is distributed under the terms of the 

Creative Commons Attribution Noncommercial License which permits 

any noncommercial use, distribution, and reproduction in any 

medium, provided the original author(s) and source are credited. 

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Pleosporales - CBS - KNAW

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