Skip to main content

Pleosporales

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.

Introduction

Historic overview of Pleosporales

Pleosporales is the largest order in the Dothideomycetes, comprising a quarter of all dothideomycetous species (Kirk et al. 2008). Species in this order occur in various habitats, and can be epiphytes, endophytes or parasites of living leaves or stems, hyperparasites on fungi or insects, lichenized, or are saprobes of dead plant stems, leaves or bark (Kruys et al. 2006; Ramesh 2003).

The Pleosporaceae was introduced by Nitschke (1869), and was assigned to Sphaeriales based on immersed ascomata and presence of pseudoparaphyses (Ellis and Everhart 1892; Lindau 1897; Wehmeyer 1975; Winter 1887). Taxa in this family were then assigned to Pseudosphaeriaceae (Theissen and Sydow 1918; Wehmeyer 1975). Pseudosphaeriales, represented by Pseudosphaeriaceae, was introduced by Theissen and Sydow (1918), and was distinguished from Dothideales by its uniloculate, perithecioid ascostromata. Subsequently, the uni- or pluri-loculate ascostromata was reported to be an invalid character to separate members of Dothideomycetes into different orders (Luttrell 1955). In addition, the familial type of Pseudosphaeriales together with its type genus, Pseudosphaeria, was transferred to Dothideales, thus Pseudosphaeriales became a synonym of Dothideales. The name “Pseudosphaeriales” has been applied in different senses, thus Pleosporales (as an invalid name due to the absence of a Latin diagnosis) was proposed by Luttrell (1955) to replace the confusing name, Pseudosphaeriales, which included seven families, i.e. Botryosphaeriaceae, Didymosphaeriaceae, Herpotrichiellaceae, Lophiostomataceae, Mesnieraceae, Pleosporaceae and Venturiaceae. Müller and von Arx (1962) however, reused Pseudosphaeriales with 12 families included, viz. Capnodiaceae, Chaetothyriaceae, Dimeriaceae, Lophiostomataceae, Mesnieraceae, Micropeltaceae, Microthyriaceae, Mycosphaerellaceae, Pleosporaceae, Sporormiaceae, Trichothyriaceae and Venturiaceae.

Familial circumscriptions of the Pleosporales were based on characters of ascomata, morphology of asci and their arrangement in locules, presence and type of hamathecium, shape of papilla or ostioles, morphology of ascospores and type of habitats (Luttrell 1973) (Table 1). Based on these characters, Luttrell (1973) included eight families, i.e. Botryosphaeriaceae, Dimeriaceae, Lophiostomataceae, Mesnieraceae, Mycoporaceae, Pleosporaceae, Sporormiaceae and Venturiaceae in Pleosporales. In their review of bitunicate ascomycetes, von Arx and Müller (1975) accepted only a single order, Dothideales, with two suborders, i.e. Dothideineae (including Atichiales, Dothiorales, Hysteriales and Myriangiales) and Pseudosphaeriineae (including Capnodiales, Chaetothyriales, Hemisphaeriales, Lophiostomatales, Microthyriales, Perisporiales, Pleosporales, Pseudosphaeriales and Trichothyriales). This proposal has however, rarely been followed. Three existing families, i.e. Lophiostomataceae, Pleosporaceae and Venturiaceae plus 11 other families were accepted in Pleosporales as arranged by Barr (1979a) (largely using Luttrell’s concepts, Table 1), and she assigned these families to six suborders. The morphology of pseudoparaphyses was given much prominence at the ordinal level in this classification (Barr 1983). In particular the Melanommatales was introduced to accommodate taxa with trabeculate pseudoparaphyses (Sporormia-type centrum development) (Barr 1983), distinguished from cellular pseudoparaphyses (Pleospora-type centrum development) possessed by members of Pleosporales sensu Barr. The order Melanommatales included Didymosphaeriaceae, Fenestellaceae, Massariaceae, Melanommataceae, Microthyriaceae, Mytilinidiaceae, Platystomaceae and Requienellaceae (Barr 1990a).

Table 1 Major circumscription changes of Pleosporales from 1955 to 2011

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; Schoch et al. 2009; Shearer et al. 2009; Suetrong et al. 2009; Tanaka et al. 2009; Zhang et al. 2009a) (Table 1). In addition, another five families, i.e. Arthopyreniaceae, Cucurbitariaceae, Diademaceae, Teichosporaceae and Zopfiaceae are tentatively included (Kruys et al. 2006; Plate 1). In the most recent issue of Myconet, 28 families were included in Pleosporales (Lumbsch and Huhndorf 2010).

Plate 1
figure 1figure 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

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. 2006 b). Thus, 26 families are temporarily accepted in Pleosporales in this study, although some such as Zopfiaceae, still require extensive DNA sequence sampling (Table 4).

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 perithecioid-like 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 can be hyaline or colored to varying degrees. They may be amerosporous (e.g. species of Semidelitschia), phragmosporous (e.g. Phaeosphaeria and Massariosphaeria), dictyosporous (e.g. most species of Pleospora and Bimuria), or scolecosporous (e.g. type species of Cochliobolus, Entodesmium or Lophionema). Although ascospore morphology had been regarded as a key factor in differentiating genera under some families, e.g. Arthopyreniaceae (Watson 1929) and Testudinaceae (Hawksworth 1979), it has been proven variable even within a single species. For instance, two types of ascospores are produced by Mamillisphaeria dimorphospora, i.e. one type is large and hyaline, and the other is comparatively smaller and brown. Numerous studies have shown the unreliability of ascospore characters above genus level classification (e.g. Phillips et al. 2008; Zhang et al. 2009a).

Asexual states of Pleosporales

Anamorphs of pleosporalean families

Anamorphs of Pleosporales are mostly coelomycetous, but may also be hyphomycetous. Phoma or Phoma-like anamorphic stages and its relatives are most common anamorphs of Pleosporales (Aveskamp et al. 2010; de Gruyter et al. 2009, 2010; Hyde et al. 2011). Some of the reported teleomorph and anamorph connections (including some listed below) are, however, based on the association rather than single ascospore isolation followed by induction of the other stage in culture (Hyde et al. 2011).

Pleosporales suborder Pleosporineae

Pleosporineae is a phylogenetically well supported suborder of Pleosporales, which temporarily includes seven families, namely Cucurbitariaceae, Didymellaceae, Didymosphaeriaceae, Dothidotthiaceae, Leptosphaeriaceae, Phaeosphaeriaceae and Pleosporaceae, and contains many important plant pathogens (de Gruyter et al. 2010; Zhang et al. 2009a). De Gruyter et al. (2009, 2010) systematically analyzed the phylogeny of Phoma and its closely related genera, and indicated that their representative species cluster in different subclades of Pleosporineae.

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, Diplodia-like and Pleurostromella (Hyde et al. 2011; Sivanesan 1984). The generic type of Cucurbitaria (C. berberidis Fuckel) is linked to Pyrenochaeta berberidis (Farr et al. 1989). Curreya has a Coniothyrium-like anamorphic stage (von Arx and van der Aa 1983; Marincowitz et al. 2008). The generic type of Curreya is C. conorum (Fuckel) Sacc., which is reported to be linked with Coniothyrium glomerulatum Sacc. (von Arx and van der Aa 1983). The generic type of Rhytidiella (R. moriformis, Cucurbitariaceae) can cause rough-bark of Populus balsamifera, and has a Phaeoseptoria anamorphic stage (Zalasky 1968). Rhytidiella baranyayi Funk & Zalasky, another species of Rhytidiella associated with the cork-bark disease of aspen is linked with Pseudosporella-like anamorphs (Funk and Zalasky 1975; Sivanesan 1984).

Didymellaceae, Didymosphaeriaceae and Dothidotthiaceae

As has been mentioned before, Phoma sensu lato species have been proved to be highly polyphyletic, and they cluster in six distinct familial clades within the Pleosporales (Aveskamp et al. 2010). Most Phoma species, including the generic type (P. herbarum), clustered in Didymellaceae (Aveskamp et al. 2010). The clade of Didymellaceae also comprises other sections, such as Ampelomyces, Boeremia, Chaetasbolisia, Dactuliochaeta, Epicoccum, Peyronellaea, Phoma-like, Piggotia, Pithoascus, as well as the type species of Ascochyta and Microsphaeropsis (Aveskamp et al. 2010; de Gruyter et al. 2009; Kirk et al. 2008; Sivanesan 1984). Leptosphaerulina is another genus of Didymellaceae, which has hyphomycetous anamorphs with pigmented and muriform conidia, such as Pithomyces (Roux 1986).

The other reported anamorphs of Didymosphaeria are Fusicladiella-like, Dendrophoma, Phoma-like (Hyde et al. 2011). Hyphomycetous Thyrostroma links to Dothidotthiaceae (Phillips et al. 2008).

Some important plant pathogens are included within Didymellaceae, such as Phoma medicaginis Malbr. & Roum., which is a necrotrophic pathogen on Medicago truncatula (Ellwood et al. 2006). Phoma herbarum is another plant pathogen, which has potential as a biocontrol agent of weeds (Neumann and Boland 2002). Ascochyta rabiei is a devastating disease of chickpea in most of the chickpea producing countries (Saxena and Singh 1987).

Leptosphaeriaceae

The anamorphic stages of Leptosphaeriaceae can be Coniothyrium, Phoma, Plenodomus and Pyrenochaeta. All are coelomycetous anamorphs, and they may have phialidic or annellidic conidiogenous cells. Phoma heteromorphospora Aa & Kesteren, the type species of Phoma sect. Heterospora and Coniothyrium palmarum, the generic type of Coniothyrium, reside in Leptosphaeriaceae (de Gruyter et al. 2009).

Pleosporaceae

Various anamorphic types can occur in Pleosporaceae, which can be coelomycetous or hyphomycetous, and the ontogeny of conidiogenous cells can be phialidic, annellidic or sympodial blastic. Both Ascochyta caulina and Phoma betae belong to Pleosporaceae (de Gruyter et al. 2009).

Some species of Bipolaris and Curvularia are anamorphs of Cochliobolus. Many species of these two genera cause plant disease or even infect human beings (Khan et al. 2000). They are hyphomycetous anamorphs with sympodial proliferating conidiogenous cells, and pigmented phragmosporous poroconidia. The generic type of Lewia (L. scrophulariae) is linked with Alternaria conjuncta E.G. Simmons (Simmons 1986), and the generic type of Pleospora (P. herbarum) is linked with Stemphylium botryosum Sacc. (Sivanesan 1984). Both Alternaria and Stemphylium are hyphomycetous anamorphs characterized by pigmented, muriform conidia that develop at a very restricted site in the apex of distinctive conidiophores (Simmons 2007).

The generic type of Pleoseptum (P. yuccaesedum) is linked with Camarosporium yuccaesedum (Ramaley and Barr 1995), the generic type of Macrospora (M. scirpicola) with Nimbya scirpicola (Fuckel) E.G. Simmons (Simmons 1989), and the generic type of Setosphaeria (S. turcica) with Drechslera turcica (Pass.) Subram. & B.L. Jain (Sivanesan 1984). Pyrenophora has the anamorphic stages of Drechslera, and the anamorphic stage of Wettsteinina can be species of Stagonospora (Farr et al. 1989).

Most common anamorphs in Pleosporaceae are Alternaria, Bipolaris, Phoma-like and Stemphylium, and they can be saprobic or parasitic on various hosts. Phoma betae A.B. Frank is a notorious pathogen on sugar beet, which causes zonate leaf spot or Phomopsis of sugar beet. Alternaria porri (Ellis) Cif., Stemphylium solani G.F. Weber, S. botryosum and S. vesicarium (Wallr.) E.G. Simmons can cause leaf blight of garlic (Zheng et al. 2009). Phoma incompta Sacc. & Martelli is a pathogen on olive, and Stemphylium botryosum, the anamorph of Pleospora herbarum, causes leaf disease of olive trees (Malathrakis 1979).

Phaeosphaeriaceae

The type species of Phoma sect. Paraphoma (Phoma radicina (McAlpine) Boerema) as well as several pathogens on Gramineae, i.e. Stagonospora foliicola (Bres.) Bubák, S. neglecta var. colorata and Wojnowicia hirta Sacc. belong to Phaeosphaeriaceae (de Gruyter et al. 2009). Other anamorphs reported for Phaeosphaeriaceae are Amarenographium, Ampelomyces, Chaetosphaeronema, Coniothyrium, Hendersonia, Neosetophoma, ?Parahendersonia, Paraphoma, Phaeoseptoria, Rhabdospora, Scolecosporiella, Setophoma, Sphaerellopsis and Tiarospora.

These anamorphic fungi can be saprobic, but mostly pathogenic on herbaceous plants. For instance, Stagonospora foliicola and Coniothyrium concentricum (Desm.) Sacc. can cause leaf spots on herbaceous plants (Zeiders 1975), and Ampelomyces quisqualis Ces. is a hyperparasite of powdery mildews.

Pleosporales suborder Massarineae

Massarineae species are mostly saprobic in terrestrial or aquatic environments. Five families are currently included within Massarineae, viz. Lentitheciaceae, Massarinaceae, Montagnulaceae, Morosphaeriaceae and Trematosphaeriaceae. Anamorphs of the five families are summarized as follows.

Lentitheciaceae

Stagonospora macropycnidia Cunnell nests within the clade of Lentitheciaceae (Plate 1). A relatively broad genus concept of Stagonospora is currently accepted, which comprises parasitic or saprobic taxa. Keissleriella cladophila (Niessl) Corbaz is another species nesting within Lentitheciaceae (Zhang et al. 2009a), and is linked with Dendrophoma sp., which has branching conidiogenous cells, and 1-celled, hyaline conidia (Bose 1961; Sivanesan 1984).

Massarinaceae

A relatively narrow concept tends to be accepted for Massarinaceae, which seems only to comprise limited species such as Byssothecium circinans, Massarina eburnea, M. cisti S.K. Bose, M. igniaria (C. Booth) Aptroot (anamorph: Periconia igniaria E.W. Mason & M.B. Ellis) and Neottiosporina paspali (G.F. Atk.) B. Sutton & Alcorn (Zhang et al. 2009a; Plate 1). Similarly, a relatively narrow generic concept of Massarina was accepted, containing only M. eburnea and M. cisti (Zhang et al. 2009b), and both species have been linked with species of Ceratophoma (Sivanesan 1984).

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, and Prosthemium betulinum is linked with the generic type of Pleomassaria (P. siparia) (Barr 1982b; Sivanesan 1984; Sutton 1980; Tanaka et al. 2010). Splanchnonema is a genus of Pleomassariaceae, the teleomorphic morphology of which is difficult to distinguish from two other genera, i.e. Asteromassaria and Pleomassaria, and the reported anamorphs of Splanchnonema are Ceuthodiplospora, Myxocyclus and Stegonsporium, which are comparable with those of Asteromassaria and Pleomassaria.

Tetraplosphaeriaceae

Tetraplosphaeriaceae was introduced to accommodate the Massarina-like bambusicolous fungi that produce Tetraploa sensu stricto anamorphs (Tanaka et al. 2009). Tetraploa aristata Berk. & Broome, the generic type of Tetraploa is widely distributed, associated with various substrates and many occur in freshwater or has been isolated from air. The polyphyletic nature of T. aristata has been well documented (Tanaka et al. 2009). Anamorphic stages can serve as a diagnostic character for this family.

Diademaceae, Massariaceae, Sporormiaceae and Teichosporaceae

The Sporormiaceae is coprophilous having Phoma or Phoma-related anamorphic states (Cannon and Kirk 2007). Comoclathris (Diademaceae) is linked with Alternaria-like anamorphs (Simmons 1952). Myxocyclus links to Massaria (Massariaceae) (Hyde et al. 2011). The anamorphic stage of Chaetomastia (Teichosporaceae) is Aposphaeria- or Coniothyrium-like (Barr 1989c).

Generally speaking, the morphologically simple conidiophores are usually considered phylogenetically uninformative (Seifert and Samuels 2000). Phoma-like anamorphs commonly occur in Pleosporales, while their colorless and unicellular conidia are also not phylogenetically informative (Seifert and Samuels 2000).

All of the above mentioned anamorphic taxa of Pleosporales have phialidic, annellidic or sympodial conidiogenous cells, representing apical wall-building type (compared to ring wall-building and diffused wall-building) (Nag Raj 1993), which may indicate that the wall-building type probably has phylogenetic significance.

Molecular phylogeny of Pleosporales

Numerous genes have been applied in phylogenetic studies of Pleosporales, mostly including LSU, SSU, mtSSU and ITS as well as the protein genes, such as RPB1, RPB2, TEF1, β-tubulin (TUB1) and actin (ACT1). A single gene such as ITS or LSU, has been used to study phylogenetic relationships between Leptosphaeria and Phaeosphaeria (Câmara et al. 2002) or Pleosporaceae and Tubeufiaceae (Kodsueb et al. 2006a, b) (Table 2). The use of these phylogenetic markers, although making important contributions, has not been successful in resolving numerous relationships in single gene dendrograms. One exception is the use of SSU sequences to demonstrate the phylogenetic significance of pseudoparaphyses (Liew et al. 2000) whilst rejecting the phylogenetic utility of pseudoparaphyses morphology (cellular or trabeculate). Analyses with combined genes have had more success. For instance combined analyses with LSU and SSU sequence data could be used to define family level classification in a few cases (Dong et al. 1998; de Gruyter et al. 2009; Lumbsch and Lindemuth 2001; Pinnoi et al. 2007; Zhang et al. 2009b) (Table 2). The addition of more than two genes has been used to determine relationships between orders. For instance, genes such as LSU, SSU and mtSSU have been used to analyze ordinal relationships in Loculoascomycetes (Lindemuth et al. 2001), and to analyze phylogenetic relationships of coprophilous families in Pleosporales (Kruys et al. 2006). Phaeocryptopus gaeumannii (T. Rohde) Petr. was shown to belong in Dothideales based on LSU, SSU and ITS sequence analysis (Winton et al. 2007), while Schoch et al. (2006) used four genes, i.e. LSU, SSU, RPB2 and TEF1 to evaluate the phylogenetic relationships among different orders of the Dothideomycetes. Five genes, viz. LSU, SSU, TEF1, RPB1 and RPB2, were used to study the phylogenetic relationships of different orders within Dothideomycetes (Schoch et al. 2009) and of different families within Pleosporales (Zhang et al. 2009a) (Table 2). It is clear that even more genes will be required to address the remaining issues and the promise of genome analyses is within reach (www.jgi.doe.gov/sequencing/why/dothideomycetes.html) for Dothideomycetes.

Table 2 List of phylogenetic studies in Pleosporales

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. 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;

  2. 2.

    To investigate the phylogeny of Pleosporales, its inter-familial relationships, and the morphological circumscription of each family;

In order to clarify morphological characters, the generic types of the majority of teleomorphic pleosporalean genera (> 60%) were studied. Most of them are from the “core families” of Pleosporales, i.e. Delitschiaceae, Lophiostomataceae, Massariaceae, Massarinaceae, Melanommataceae, Montagnulaceae, Phaeosphaeriaceae, Phaeotrichaceae, Pleomassariaceae, Pleosporaceae, Sporormiaceae and Teichosporaceae. Notes are given for those where type specimens could not be obtained during the timeframe of this study. A detailed description and illustration of each generic type is provided. Comments, notes and problems that need to be addressed are provided for each genus. Phylogenetic investigation based on five nuclear loci, viz. LSU, SSU, RPB1, RPB2 and TEF1 was carried out using available strains from numerous genera in Pleosporales. In total, 278 pleosporalean taxa are included in the phylogenetic analysis, which form 25 familial clades on the dendrogram (Plate 1). The suborder, Massarineae, is emended to accommodate Lentitheciaceae, Massarinaceae, Montagnulaceae, Morosphaeriaceae and Trematosphaeriaceae.

Materials and methods

Molecular phylogeny

Four genes were used in this analysis, the large and small subunits of the nuclear ribosomal RNA genes (LSU, SSU) and two protein coding genes, namely the second largest subunit of RNA polymerase II (RPB2) and translation elongation factor-1 alpha (TEF1). All sequences were downloaded from GenBank as listed in Table 3. Each of the individual ribosomal genes was aligned in SATé under default settings with at least 20 iterations. The protein coding genes were aligned in BioEdit (Hall 2004) and completed by manual adjustment. Introns were removed and all genes were concatenated in a single nucleotide alignment with 43% missing and gap characters out of a total set of 5081. The alignment had 100% representation for LSU, 75% for SSU, 48% for RPB2 and 65% for TEF1. The final data matrix had 280 taxa including outgroups (Table 3).

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

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 hand-cutting. 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., large-sized: > 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, 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)

Fig. 1
figure 2

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, g = 20 μm

Ascomata 1–2 mm high × 1.8–3 mm diam., scattered or gregarious, superficial, conical or semiglobose, with a flattened base not easily removed from the substrate, ostiolate, black, very brittle and carbonaceous and extremely difficult to cut (Fig. 1a and b). Peridium 250–310 μm thick, to 600 μm thick near the apex, thinner at the base, comprising three types of cells; outer cells pseudoparenchymatous, small heavily pigmented thick-walled cells of textura epidermoidea, cells 0.6–1 × 6–10 μm diam., cell wall 5–9 μm thick; cells near the substrate less pigmented, composed of cells of textura prismatica, cell walls 1–3(−5) μm thick; inner cells less pigmented, comprised of hyaline to pale brown thin-walled cells, merging with pseudoparaphyses (Fig. 1c, d and e). Hamathecium of dense, long trabeculate pseudoparaphyses, ca. 1 μm broad, embedded in mucilage, hyaline, anastomosing and sparsely septate. Asci 140220 × 13–17 μm (\( \bar{x} = { 165}.{3 } \times { 15}.{6 }\mu {\text{m}} \), 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 (\( \bar{x} = { 21}.{5 } \times { 13}.{6 }\mu {\text{m}} \), 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 thick-walled 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.

Concluding remarks

Acrocordiopsis, Astrosphaeriella sensu stricto, Mamillisphaeria, Caryospora and Caryosporella are morphologically similar as all have very thick-walled carbonaceous ascomata, narrow pseudoparaphyses in a gelatinous matrix (trabeculae) and bitunicate, fissitunicate asci. Despite their similarities, the shape of asci and ascospores differs (e.g. Mamillisphaeria has sac-like asci and two types of ascospores, brown or hyaline, Astrosphaeriella has cylindro-clavate asci and narrowly fusoid ascospores, both Acrocordiopsis and Caryosporella has cylindrical asci, but ascospores of Caryosporella are reddish brown). Therefore, the current familial placement of Acrocordiopsis cannot be determined. All generic types of Astrosphaeriella sensu stricto, Mamillisphaeria and Caryospora should be recollected and isolated for phylogenetic study.

Aigialus Kohlm. & S. Schatz, Trans. Br. Mycol. Soc. 85: 699 (1985). (Aigialaceae)

Generic description

Habitat marine, saprobic. Ascomata mostly subglobose in front view, fusoid in sagittal section, rarely subglobose, scattered, immersed to erumpent, papillate, ostiolate, ostiole rounded or slit-like, periphysate. Peridium 2-layered. Hamathecium of trabeculate pseudoparaphyses. Asci 8-spored, cylindrical, pedicellate, with an ocular chamber and conspicuous apical ring. Ascospores ellipsoidal to fusoid, muriform, yellow brown to brown, with terminal appendages.

Anamorphs reported for genus: none.

Literature: Eriksson 2006; Jones et al. 2009; Kohlmeyer and Schatz 1985; Lumbsch and Huhndorf 2007.

Type species

Aigialus grandis Kohlm. & S. Schatz, Trans. Br. Mycol. Soc. 85: 699 (1985). (Fig. 2)

Fig. 2
figure 3

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, cf = 20 μm

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 semi-immersed, 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 450640 × 22–35 μm (\( \bar{x} = 505 \times 30\mu m \), 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 7595 × 15–26 μm (\( \bar{x} = 84.3 \times 17.5\mu m \), n = 10), obliquely uniseriate and partially overlapping, broadly fusoid to fusoid with narrowly rounded ends in front view, flat on one side from side view (14–20 μm thick), yellowish brown, apical cells usually hyaline, muriform, with 14–17(−18) transversal septa, 1–3 longitudinal septa in most cells, slightly constricted at the septa, with a gelatinous cap at each end (Fig. 2c and d).

Anamorph: none reported.

Material examined: BELIZE, Wee-Wee Cay, on submerged wood of roots and branches of Rhizophora mangle L., Mar. 1983, leg. J. Kohlmeyer (NY, J.K. 4332b, isotype).

Notes

Morphology Aigialus was formally established by Kohlmeyer and Schatz (1985) based on its immersed or semi-immersed ascomata with periphysate ostiole, trabeculate pseudoparaphyses, cylindrical and fissitunicate asci, and distinctive muriform ascospores with gelatinous sheath or caps. There are five accepted species in the genus, namely A. grandis, A. mangrovei Borse, A. parvus S. Schatz & Kohlm., A. rhizophorae Borse and A. striatispora K.D. Hyde (Jones et al. 2009). Aigialus was first assigned to the Melanommatales, but its familial status was uncertain (Kohlmeyer and Schatz 1985). Barr (1990b) included Aigialus in Massariaceae based on its conspicuous apical ring in the asci and ascospore characters, and this has subsequently been widely followed (Eriksson 2006; Hawksworth et al. 1995; Kirk et al. 2001; Lumbsch and Huhndorf 2007).

Phylogenetic study The generic type of Aigialus (A. grandis) together with other three marine species, i.e. A. mangrovei, A. parvus as well as A. rhizophorae form a robust clade on the phylogenetic tree. Thus a new family, Aigialaceae, was introduced to accommodate Aigialus together with Ascocratera and Rimora (Suetrong et al. 2009).

Concluding remarks The pleosporalean status of Aigialus has been phylogenetically verified, and the single branch containing Aigialus, Ascocratera and Rimora represents a familial rank of Aigialaceae (Suetrong et al. 2009).

Amniculicola Yin. Zhang & K.D. Hyde, Mycol. Res. 112: 1189 (2008). (Amniculicolaceae)

Generic description

Habitat freshwater, saprobic. Ascomata solitary, scattered, or in small groups, initially immersed, becoming erumpent, 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)

Fig. 3
figure 4

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

Ascomata 350–450 μm high × 300–500 μm diam., solitary, scattered, or in small groups of 2–3, initially immersed, becoming erumpent, to nearly superficial, with basal wall remaining immersed in host tissue, globose, subglobose, broadly or narrowly conical, often laterally flattened, with a flattened base not easily removed from the substrate, wall black, roughened, often bearing remnants of wood fibers; apex well differentiated into two tuberculate flared lips surrounding a slit-like ostiole, 150–250 μm long, filled with a purplish amorphous matter, oriented in the axis of the wood fibers; underlying wood stained pale purple (Fig. 3a and b). Peridium 40–55 μm thick laterally, up to 120 μm thick at the apex, thinner at the base, coriaceous, 2-layered, outer layer composed of small heavily pigmented thick-walled cells of textura angularis, cells 4–9 μm diam., cell wall 2–3 μm thick, apex cells smaller and walls thicker, inner layer composed of hyaline thin-walled cells of textura angularis, 8–16 μm diam., in places with columns of textura prismatica, oriented perpendicular to the ascomatal surface, and larger, paler cells of textura prismatica towards the interior and at the base, 10–25 μm (Fig. 3c, d and e). Hamathecium of dense, long trabeculate pseudoparaphyses <1 μm broad, embedded in mucilage (Indian ink), anastomosing between and above the asci. Asci 140184 × 9–10 μm, 8-spored, bitunicate, fissitunicate, cylindrical to narrowly fusoid, with a short, narrowed, twisted, furcate pedicel which is 15–25 μm long, with a low truncate ocular chamber and a small inconspicuous apical apparatus barely seen in water (Fig. 3f). Ascospores (20.5-)28–32 × (6-)8(−9) μm, obliquely uniseriate and partially overlapping, broadly fusoid to fusoid with broadly to narrowly rounded ends, hyaline, 1-septate, deeply constricted at the median septum, the upper cell often shorter and broader than the lower one, smooth, containing four refractive globules, surrounded by an irregular hyaline gelatinous sheath 4–8.5 μm thick, best seen in India ink, released senescent ascospores are greyish and 3-septate, strongly constricted at all septa (Fig. 3g).

Anamorph: none reported.

Colonies slow growing, reaching 4 cm diam. after 70 d growth on Malt Extract Agar (MEA) at 25°C, flat, with irregular to rhizoidal margin, off-white to grey, reverse reddish purple to deep reddish purple, the medium is stained pale yellow.

Material examined: FRANCE, Ariège, Prat Communal, Ruisseau de Loumet, 1000 m, on partly submerged wood of Fraxinus excelsior, 8 Aug. 2006, leg. Jacques Fournier (PC 0092661, holotype); 3 Sept. 2004 (BPI 877774; CBS: H-17932); Rimont, Ruisseau de Peyrau, 400 m, on driftwood of Alnus glutinosa (L.) Gaertn., 23 Jul. 2006 (HKU(M) 17515, isotype).

Notes

Morphology

Amniculicola is a freshwater genus which stains the woody substrate purple (Zhang et al. 2008c, 2009a, c). This genus appears only to be reported from Europe. A detailed description of the generic type was provided by Zhang et al. (2008c).

Phylogenetic study

Three species of Amniculicola cluster together with Anguillospora longissima, Spirosphaera cupreorufescens and Repetophragma ontariense as well as Pleospora rubicunda Niessl (current name Murispora rubicunda (Niessl) Y. Zhang ter, J. Fourn. & K.D. Hyde) and Massariosphaeria typhicola (P. Karst.) Leuchtm. (current name Neomassariosphaeria typhicola (P. Karst.) Yin. Zhang, J. Fourn. & K.D. Hyde). A new family, i.e. Amniculicolaceae, was introduced to accommodate these taxa (Zhang et al. 2008c, 2009a, c).

Concluding remarks

All of the five teleomorphic taxa within Amniculicolaceae are from freshwater in Europe and their ascomata stain the woody substrate purple. Purple staining makes taxa of this family easily recognized in the field.

Anomalemma Sivan., Trans. Br. Mycol. Soc. 81: 328 (1983). (?Melanommataceae)

Generic description

Habitat terrestrial, fungicolous. Ascomata gregarious, superficial, papillate, ostiolate. Peridium composed cells of pseudoparenchymatous. Asci clavate, 8-spored. Hamathecium of dense, filliform pseudoparaphyses. Ascospores 1- (rarely 2- to 3-) septate, fusoid, reddish brown, constricted at the main septum.

Anamorphs reported for genus: Exosporiella (= Phanerocorynella) (Sivanesan 1983).

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)

Fig. 4
figure 5

Anomalemma epochnii (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

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 75108 × 9.5–12.5 μm (\( \bar{x} = 92.8 \times 11.1\mu m \), n = 10), 8-spored, bitunicate, fissitunicate, dehiscence not observed, cylindro-clavate to clavate, with a furcate pedicel up to 6–25 μm long, with a small ocular chamber best seen in immature asci (ca. 2 μm wide × 1 μm high) (Fig. 4b and c). Ascospores 20–25(−30) × 5–7.5 μm (\( \bar{x} = 23.1 \times 6.3\mu m \), 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) and Chaetosphaeria (as C. epochnii (Berk. & Broome) Keissl.; Keissler 1922). The deposition of Sphaeria epochnii in Chaetosphaeria is obviously unacceptable, as Chaetosphaeria has unitunicate asci. Melanomma has been reported having Aposphaeria or Pseudospiropes anamorphs, which differs from Exosporiella (Sivanesan 1983). In addition, the presence of well developed prosenchymatous stroma in Sphaeria epochnii can also readily distinguish it from Melanomma (Sivanesan 1983).

The gregarious ascomata and formation of prosenchymatous stroma of Anomalemma resembles those of Cucurbitaria, but the pleosporaceous dictyosporous ascospores of Cucurbitaria readily distinguish it from Anomalemma epochnii. In addition, the pseudoparenchymatous peridium, fungicolous habitat and brown 1-septate ascospores, which later becoming 3-septate differ from any other pleosporalean genus. Thus a new genus, Anomalemma, was introduced to accommodate it (Sivanesan 1983). Anomalemma is presently monotypic.

Phylogenetic study

None.

Concluding remarks

Anomalemma epochnii certainly resembles Byssosphaeria in its ascomata clustering together in groups on closely intertwined hyphae and brown ascospores, and may well be included in this genus. Its fungicolous habitat, however, distinguishes it from Byssosphaeria.

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

Generic description

Habitat terrestrial, saprobic. Ascomata small, clustered, immersed, subglobose or irregularly pyriform. Peridium thin. Hamathecium of dense, long trabeculate pseudoparaphyses. Asci 8-spored, bitunicate, fissitunicate, cylindrical, apical rounded with ocular chamber and faint ring, with short pedicels. Ascospores uniseriate to partially overlapping, fusoid, brown, 1-septate, slightly constricted at the septum.

Anamorphs reported for genus: none.

Literature: Hyde 1994a.

Type species

Appendispora frondicola K.D. Hyde, Sydowia 46: 30 (1994a). (Fig. 5)

Fig. 5
figure 6

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 = 0.5 mm, b = 100 μm, c–g = 10 μm

Ascomata 120–280 μm high × 180–280 μm diam., clustered, immersed with minute ostioles visible through cracks or blackened dots on the host surface, subglobose or irregularly pyriform (Fig. 5a and b). Peridium 40 μm thick, comprising two types of cells; outer cells, small heavily pigmented thick-walled cells of textura angularis, inner cells compressed, hyaline. Hamathecium of dense, very long trabeculate pseudoparaphyses, ca. 1 μm broad, embedded in mucilage, hyaline, anastomosing (Fig. 5e). Asci 130144 × 11–13 μm, 8-spored, bitunicate, fissitunicate, cylindrical, with an ocular chamber and faint ring, with short pedicels (Fig. 5c and d). Ascospores 2130 × 7–9 μm, uniseriate to partially overlapping, fusoid, brown, 1-septate, slightly constricted at the septum, with an irregular ridged ornamentation and 3–5 narrow appendages at each end (Fig. 5f and g).

Anamorph: none reported.

Material examined: BRUNEL, Jalan, Muara, Simpang 835, on dead rachis of Oncosperma horridum on forest floor, Nov. 1992, K.D. Hyde 1652 (BRIP 21354, holotype).

Notes

Morphology

Appendispora was described as a saprobe of palm, and is characterized by small, immersed ascomata, bitunicate, fissitunicate asci, trabeculate pseudoparaphyses, brown, 1-septate, appendaged ascospores with irregular wall striations (Hyde 1994a). Based on its trabeculate pseudoparaphyses embedded within gel matrix and its brown ascospores, Appendispora was assigned to Didymosphaeriaceae (Barr 1987b; Hyde 1994a).

Phylogenetic study

None.

Concluding remarks

The saprobic habitat and association with monocots, cylindrical asci, trabeculate pseudoparaphyses as well as its brown, 1-septate ascospores make it difficult to determine a better phylogenetic position than Didymellaceae.

Ascorhombispora L. Cai & K.D. Hyde, Cryptog. Mycol. 28: 294 (2007). (Pleosporales, genera incertae sedis)

Generic description

Habitat freshwater, saprobic. Ascomata solitary or gregarious, superficial, globose to subglobose, dark brown to black, short papillate, ostiolate, coriaceous. Peridium relatively thin, textura angularis in longitudinal section, 2-layered. Hamathecium not observed. Asci 8-spored, obpyriform, broadly clavate to saccate, pedicellate, bitunicate, apex rounded, persistent. Ascospores overlapping 2-3-seriate, broadly fusoid to rhomboid, thick-walled, surrounded by mucilaginous sheath, 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.

Anamorphs reported for genus: none.

Literature: Cai and Hyde 2007.

Type species

Ascorhombispora aquatica L. Cai & K.D. Hyde, Cryptog. Mycol. 28: 295 (2007). (Fig. 6)

Fig. 6
figure 7

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 = 10 μm (figures referred to Cai and Hyde 2007)

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 thick-walled, inner layer hyaline, relatively thin-walled (Fig. 6a and b). Hamathecium not observed. Asci 100–198 × 72–102 μm (\( \bar{x} = 186 \times 88\mu m \), 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 (\( \bar{x} = 38.5 \times 21\mu m \), n = 25), overlapping 2-3-seriate, broadly fusoid to rhomboid, thick-walled, surrounded by mucilaginous sheath, 3-euseptate, not constricted at septa, median septum wide, forming a darker band, central cells large, trapezoid, 11–18 μm long, dark brown to black, verruculose, polar end cells small, hemispherical, 3.5–4 μm long, subhyaline to pale brown, smooth (Fig. 6f).

Anamorph: none reported.

Material examined: CHINA, Yunnan, Jinghong, on submerged bamboo in a small forest stream, 26 Jan. 2003, leg. det. L. Cai, CAI-1H31 (HKU(M) 10859, holotype).

Notes

Morphology

Ascorhombispora was introduced as a monotypic genus from freshwater by Cai and Hyde (2007), and is characterized by superficial, coriaceous, non-stromatic ascomata, large, saccate asci; lack of interascal filaments and trapezoid (rhombic), 3-septate, dark brown to black ascospores with smaller end cells which are subhyaline to pale brown. Ascorhombispora is most comparable with Caryospora and Zopfia. But the globose to subglobose ascomata and thin peridium, saccate asci lacking interascal pseudoparaphyses, and the 3-septate, rhomboid ascospores with the paler end cells of Ascorhombispora differs from those of Caryospora (Cai and Hyde 2007).

Phylogenetic study

Phylogenetic analysis based on either SSU or LSU rDNA sequences indicated that Ascorhombispora aquatica belongs to Pleosporales, but its familial placement was left undetermined (Cai and Hyde 2007).

Concluding remarks

The sac-shaped asci and absence of pseudoparaphyses are uncommon in Pleosporales, especially among those from freshwater.

Asteromassaria Höhn., Sber. Akad. Wiss. Wien, Math.-naturw. Kl., Abt. I 126: 368 (1917). (?Morosphaeriaceae)

Generic description

Habitat terrestrial, saprobic. Ascomata medium-sized, clustered, at first immersed and then breaking through the host surface and becoming superficial, globose, subglobose, coriaceous. Peridium 2-layered, thicker near the base. Hamathecium of dense, septate, cellular pseudoparaphyses which branch and anastomosing frequently between and above asci. Asci (4-)8-spored, bitunicate, cylindro-clavate to clavate, with a short truncated pedicel and a small ocular chamber. Ascospores obliquely uniseriate and partially overlapping to biseriate, fusoid to fusoid-ellipsoidal, pale brown when mature, 1-septate, some becoming 3-septate when old, constricted at the median septum.

Anamorphs reported for genus: Scolicosporium (Sivanesan 1984).

Literature: Barr 1982a; b; 1993a; Boise 1985; Shoemaker and LeClair 1975; Sivanesan 1987; Tanaka et al. 2005.

Type species

Asteromassaria macrospora (Desm.) Höhn., F. von, Sber. Akad. Wiss. Wien, Math.-naturw. Kl., Abt. I 126: 368 (1917). (Fig. 7)

Fig. 7
figure 8

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

≡ 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, 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 (\( \bar{x} = 230 \times 35\mu m \), n = 10), 8-spored (sometimes 4-spored), bitunicate, fissitunicate dehiscence not observed, cylindro-clavate to clavate, with a short truncated pedicel up to 30 μm, with a small ocular chamber (ca. 3 μm wide × 3 μm high) (Fig. 7e and f). Ascospores 5058 × (14-)18–21 μm (\( \bar{x} = 55.3 \times 18.2\mu m \), n = 10), obliquely uniseriate and partially overlapping to biseriate, fusoid to fusoid-ellipsoidal, with narrowly rounded ends, lightly brown when mature, 1-septate, some becoming 3-septate when old, constricted at the median septum, the upper cell often broader and longer than the lower one, minutely verrucose (Fig. 7g, h, i and j).

Anamorph: Scolicosporium macrosporium (Berk.) B. Sutton.

Acervuli immersed in bark, brown, discrete, up to 250 μm diam., opening by irregular rupture of the overlaying tissues. Peridium of thin-walled angular cells. Conidiophores cylindrical, 1-2-septate, up to 30 μm long and 3–5 μm wide. Conidiogenous cells holoblastic, 1-2-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).

Phylogenetic study

Asteromassaria pulchra (Harkn.) Shoemaker & P.M. LeClair is basal to Morosphaeriaceae in the phylogenetic tree based on four genes, but its placement is influenced by taxon sampling that was different in several analyses.

Concluding remarks Asteromassaria can be distinguished from other comparable genera, i.e. Pleomassaria and Splanchnonema by 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)

Generic description

Habitat terrestrial, saprobic. Ascomata densely scattered or in small groups, erumpent through the outer layers of the host tissues to nearly superficial, reflexed pieces of the ruptured host tissue usually persisting around the base of the ascomata, often star-like, conical to semiglobose, with a central papilla. Peridium upper wall usually comprising a thick dark brittle pseudoparenchymatous layer, base usually flattened and thin-walled. Hamathecium of dense, filliform, trabeculate pseudoparaphyses, embedded in mucilage. Asci 8-spored, bitunicate, fissitunicate, cylindro-clavate to narrowly fusoid. Ascospores narrowly fusoid with acute ends, hyaline, pale brown or brown, 1-3-septate.

Anamorphs reported for genus: Pleurophomopsis (Hyde et al. 2011).

Literature: von Arx and Müller 1975; Barr 1990a; Chen and Hsieh 2004; Hawksworth 1981; Hawksworth and Boise 1985; Hyde and Fröhlich 1998; Hyde et al. 2000; Kirk et al. 2001; Sydow and Sydow 1913; Tanaka and Harada 2005a; b; Tanaka et al. 2009.

Type species

Astrosphaeriella stellata Syd. & P. Syd., Annls mycol. 11: 260 (1913). (Fig. 8)

Fig. 8
figure 9

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–f = 10 μm

Ascomata 360–570 μm high × 860–1150 μm diam., densely scattered or in small groups, erumpent through the outer layers of the host tissues to nearly superficial, reflexed pieces of the ruptured host tissue usually persisting around the base of the ascomata, forming star-like flanges around the ascomata from the surface view; ascomata broadly conical, with a flattened base not easily removed from the substrate, wall black; apex with a central papilla which is black and shiny at maturity, scarcely projecting (Fig. 8a). Peridium 40–70 μm thick, carbonaceous and crisp, 1-layered, composed of very small dark brown thick-walled pseudoparenchymatous cells, cells 2–5 μm diam., cell wall 2–6 μm thick, in places at the base composed of hyaline cells of textura prismatica, cells 5 × 8 μm diam. (Fig. 8b). Hamathecium of dense, very long trabeculate pseudoparaphyses, <1 μm broad, embedded in mucilage (Indian ink), anastomosing between and above the asci. Asci 130190 × 11.5–15 μm (\( \bar{x} = 161.5 \times 12.8\mu m \), n = 10), 8-spored, bitunicate, fissitunicate, cylindro-clavate to narrowly fusoid, with a short, narrowed pedicel which is 10–35 μm long, with a large ocular chamber (Fig. 8c). Ascospores 3550 × 5–7.5 μm (\( \bar{x} = 43.4 \times 6\mu m \), n = 10), biseriate, elongate- fusoid, gradually tapering towards the ends, hyaline, turning pale brown when mature, 1(−3)-septate, constricted at the median septum (Fig. 8d,e and f).

Anamorph: none reported.

Material examined: USA, Hawaii, Kapano Gulch, in bamboo culms, 5 Jun. 1947, leg. Kopf & Rogers, det. Miller (BISH 145726, as Astrosphaeriella fusispora Syd. & P. Syd.).

Notes

Morphology

Astrosphaeriella has been treated as a synonym of Microthelia (von Arx and Müller 1975), but the large conical ascomata, numerous trabeculate pseudoparaphyses and 1-septate and elongated ascospores of Astrosphaeriella all disagree with those of Microthelia (Hawksworth 1981). It was assigned to Platystomaceae by Barr (1990a) in Pleosporales or Melanommataceae by Kirk et al. (2001). Following a systematic study of Astrosphaeriella, only four species were accepted, i.e. A. aosimensis I. Hino & Katum., A. stellata, A. trochus (Penz. & Sacc.) D. Hawksw. and A. venezuelensis M.E. Barr & D. Hawksw. (Hawksworth 1981), and it was defined as a tropical genus, occurring exclusively on palms or bamboo. Astrosphaeriella stellata was selected as the type of Astrosphaeriella, and A. fusispora was regarded 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 semi-immersed to superficial ascomata with flattened base, cylindro-clavate asci with fusoid ascospores and trabeculate pseudoparaphyses, i.e. Astrosphaeriella sensu stricto (e.g. A. fusispora and A. vesuvius (Berk. & Broome) D. Hawksw. & Boise); 2. Trematosphaeria-like with rounded ascomata (e.g. A. africana D. Hawksw.); and 3. Massarina-like species with immersed ascomata (e.g. A. bakeriana (Sacc.) K.D. Hyde & J. Fröhl.) (Chen and Hsieh 2004; Tanaka and Harada 2005a; b). Currently, a broad generic concept of Astrosphaeriella is accepted, and 47 taxa are included in Astrosphaeriella.

Phylogenetic study

Phylogenetic analysis based on LSU and SSU nurDNA sequence data indicates that Astrosphaeriella is polyphyletic, and located in the basal region of the Pleosporales between Testudinaceae and Zopfiaceae/Delitschiaceae (Tanaka et al. 2009), or basal to Aigialaceae (Schoch et al. 2009). The genus is, however, clearly not related to Trematosphaeria as previously understood (Boise 1985).

Concluding remarks Astrosphaeriella is currently polyphyletic and new collections of the different elements listed above are needed in order to understand the placement of various species. We suggest that some immersed bambusicolous species may belong in Tetraplospheariaceae.

Asymmetricospora J. Fröhl. & K.D. Hyde, Sydowia 50: 183 (1998). (?Melanommataceae)

Generic description

Habitat terrestrial, saprobic. Ascomata solitary or in small groups, immersed, black, lenticular in section, uni- or often multi-locular, with a central ostiole without tissue differentiation. Upper peridium carbonaceous, thicker at sides and apex. Lower peridium composed of irregular-shaped, hyaline cells. Hamathecium of trabeculate pseudoparaphyses, branching and anastomosing between and above asci, embedded in mucilage. Asci 8-spored, bitunicate, fissitunicate unknown, clavate, short pedicellate. Ascospores 1-septate, hyaline, constricted at the septum, with a broad, spreading mucilaginous sheath.

Anamorphs reported for genus: none.

Literature: Fröhlich and Hyde 1998.

Type species

Asymmetricospora calamicola J. Fröhl. & K.D. Hyde, Sydowia 50: 184 (1998). (Fig. 9)

Fig. 9
figure 10

Asymmetricospora calamicola (from HKU(M) 7794, holotype). a Ascomata immersed in the substrate. b Section of the peridium. c Mature and immature asci in pseudoparaphyses (in cotton blue). d Clavate ascus with a small ocular chamber. e–g Ascospores with sheath. Scale bars: a, b = 0.5 mm, c = 50 μm, dg = 20 μm

Ascomata 675–950 μm high × 875–1500 μm diam., solitary or in small groups of 2–10, immersed and forming slightly protruding domes on the substrate surface, with near-white 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 (\( \bar{x} = 172.8 \times 31.5\mu m \), n = 20), 8-spored, bitunicate, fissitunicate dehiscence not observed, clavate, with short pedicel (to 25 μm), with ocular chambers (ca. 3 μm wide × 4 μm high) (Fig. 9c and d). Ascospores 35–55 × 10.5–15 μm (\( \bar{x} = 44.7 \times 12.4\mu m \), n = 50), biseriate, navicular to obovoid, hyaline, becoming pale brown when senescent, straight or usually curved, smooth, asymmetric, 1-septate, the upper cell larger with a rounded end, basal cell with a tapering end, constricted at the septum, with spreading mucilaginous sheath (Fig. 9e, f and g) (data from Fröhlich and Hyde 1998).

Anamorph: none reported.

Material examined: AUSTRALIA, North Queensland, Palmerston, Palmerston National Park, on dead rattan of Calamus caryotoides A.Cunn. ex Mart., Mar. 1994, J. Fröhlich (HKU(M) 7794, holotype).

Notes

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 multi-loculate fruiting body (Fröhlich and Hyde 1998).

Phylogenetic study

None.

Concluding remarks

The placement of Asymmetricospora under Melanommataceae remains to be confirmed.

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

Generic description

Habitat terrestrial, parasitic. Ascomata small- to medium-sized, solitary or scattered, immersed, globose, subglobose, ostiolate, coriaceous. Apex with or without papilla and with a pore-like ostiole. Peridium 2-layered. Hamathecium of dense, long cellular pseudoparaphyses, septate, embedded in mucilage. Asci bitunicate, fissitunicate, cylindrical to clavate, with a short, furcate pedicel. Ascospores ellipsoid, hyaline at first, turning brown at maturity, 1-septate, strongly constricted at the septum.

Anamorphs reported for genus: none.

Literature: Yuan 1994.

Type species

Barria piceae Z.Q. Yuan, Mycotaxon 51: 314 (1994). (Fig. 10)

Fig. 10
figure 11

Barria piceae (from NY 92003, isotype). a Ascoma on the host surface. Note the wide opening ostiole. b Section of the partial peridium with two types of cells. c, d Asci with ocular chambers and short pedicels. e, f Ellipsoid ascospores which are turning brown with thin sheath around them. Scale bars: a = 0.5 mm, b = 50 μm, c, d = 20 μm, e, f = 10 μm

Ascomata 240–370 μm high × 200–320 μm diam., solitary, scattered, immersed, globose, subglobose, coriaceous, apex with or without papilla and with a pore-like ostiole (Fig. 10a). Peridium 20–35 μm thick, comprising two cell types, the outer cells comprising 3–4 layers of brown pseudoparenchymatous cells, cells 4–5 μm diam., cell wall 2–3 μm thick, inner cells comprising 3–4 layers of pale brown compressed cells, cells 2 × 16 μm diam., cell wall 0.5–1.5 μm thick (Fig. 10b). Hamathecium of dense, long cellular pseudoparaphyses, 2–3 μm broad, septate. Asci 135–200(−220) × 14–20 μm (\( \bar{x} = 156 \times 16.6\mu m \), 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. 4 μm wide × 3 μm high) (Fig. 10c and d). Ascospores 1921.5 × 10–12 μm (\( \bar{x} = 20.4 \times 11\mu m \), n = 10), uniseriate to partially overlapping, ellipsoid, hyaline or greenish with numerous small guttules at first and olive green to smoky brown at maturity, 1-septate, strongly constricted at the septum, foveolate, surrounded with sheath (Fig. 10e and f).

Anamorph: none reported.

Material examined: CHINA, Xinjiang Province, Uygur, Urumqi, Tianshan Mountain, on needles of Picea schrenkiana, 1 Jul. 1992, Z.Q. Yuan (NY 92003, isotype).

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.

Phylogenetic study

None.

Concluding remarks

The status of Barria with its unusual verrucose ascospores and thick gel coating is uncertain. In many ways it resembles Belizeana, with its cylindrical asci, 1-septate, ellipsoid ascospores with sheath and verruculose surface (Kohlmeyer and Volkmann-Kohlmeyer 1987). However, the latter is a marine genus while Barria causes leaf blight of terrestrial Picea (Yuan 1994). The placement in Phaeosphaeriaceae seems logical based on the parasitic life style, thin and simple peridium, wide cellular pseudoparaphyses and brown ascospores. However, molecular data are needed to confirm this.

Belizeana Kohlm. & Volkm.-Kohlm., Bot. Mar. 30: 195 (1987). (Pleosporales, genera incertae sedis)

Generic description

Habitat marine, saprobic. Ascomata solitary, scattered, or in small groups, medium-sized, immersed to semi-immersed, 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)

Fig. 11
figure 12

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–i = 10 μm

Ascomata 170–300 μm high × 160–290 μm diam., solitary, scattered, or in small groups of 2–3, immersed to semi-immersed, 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.μm broad, filliform pseudoparaphyses, rarely branched, embedded in mucilage (Fig. 11g). Asci 145170 × 20–30 μm (\( \bar{x} = 163 \times 25\mu m \), 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 (\( \bar{x} = 22 \times 15\mu m \), 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 can be assigned to Pleosporaceae (Pleosporales) according to Luttrell’s (1973) treatment and keys of von Arx and Müller (1975), but cannot resolve a proper family based on Barr (1979a, 1983). The unique morphology together with obligate marine habitat makes B. tuberculata readily distinguishable from all other taxa of Pleosporaceae.

Phylogenetic study None.

Concluding remarks

The ascospores of Belizeana tuberculata are most comparable with those of Acrocordiopsis patilii, but the superficial 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)

Generic description

Habitat marine, saprobic. Ascomata large, solitary or gregarious, immersed, subglobose to pyriform, ostiolate, papillate, periphysate, black, branching, carbonaceous. Hamathecium of dense, long trabeculate pseudoparaphyses, embedded in mucilage. Asci 8-spored, bitunicate, fissitunicate, cylindrical, with apical apparatus. Ascospores uniseriate to partially overlapping, fusoid, hyaline when young, becoming brown to dark brown at maturity, multi-septate towards each end, with a hyaline, globose refractive chamber or appendage at each end, not constricted at the septum.

Anamorphs reported for genus: none.

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

Type species

Biatriospora marina K.D. Hyde & Borse, Mycotaxon 26: 264 (1986). (Fig. 12)

Fig. 12
figure 13figure 13

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)

Ascomata 650–860 μm high × 350–510 μm diam., solitary or gregarious, immersed, subglobose to pyriform, ostiolate, papillate, periphysate, black, carbonaceous (Fig. 12.2a). Hamathecium of dense, long trabeculate pseudoparaphyses, 1–1.5 μm broad, branching, embedded in mucilage. Asci 175400 × 22–40 μm, 8-spored, bitunicate, fissitunicate, cylindrical, with long pedicels and apical apparatus (Fig. 12.1a, b, 2b). Ascospores 5582 × 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).

Anamorph: none reported.

Material examined: SEYCHELLES, 2 Jan. 1984 (Herb. IMI 297768 holotype).

Notes

Morphology

Biatriospora was introduced to accommodate a marine fungus B. marina, which is characterized by horizontal ascomata and ascospores with polar, globose refractive chambers and polar septa (Hyde and Borse 1986). Polar refractive chambers can also occur in other marine fungi, such as Lulworthia and Aigialus. The chambers have been proposed as important for spore attachment to substrates in a liquid environment (Hyde and Borse 1986).

Phylogenetic study

Multigene phylogenetic analysis indicated that Biatriospora marina forms a separate branch, sister to other families of Pleosporales (Suetrong et al. 2009), and maybe related to species in Roussoella (Plate 1).

Concluding remarks The familial status of Biatriospora can not be determined.

Bicrouania Kohlm. & Volkm.-Kohlm., Mycol. Res. 94: 685 (1990). (?Melanommataceae)

Generic description

Habitat marine, saprobic. Ascomata immersed gregarious, erumpent to superficial, globose to subglobose, black, periphysate, coriaceous, epapillate or papillate, ostiolate. Peridium thin, 2-layered. Hamathecium of dense, long trabeculate pseudoparaphyses, branching and anastomosing between and above the asci. Asci 8-spored, bitunicate, fissitunicate, cylindrical, with a thick, furcate pedicel lacking ocular chamber. Ascospores obliquely uniseriate and partially overlapping, ellipsoidal with broadly rounded ends, reddish brown, 1-septate, thick-walled, constricted at the septum.

Anamorphs reported for genus: none.

Literature: Jones et al. 2009; Kohlmeyer and Volkmann-Kohlmeyer 1990.

Type species

Bicrouania maritima (P. Crouan & H. Crouan) Kohlm. & Volkm.-Kohlm., Mycol. Res. 94: 685 (1990). (Fig. 13)

Fig. 13
figure 14

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

≡ Sphaeria maritima P. Crouan & H. Crouan, Florule du Finistére, Paris: 27 (1867) non Sphaeria maritima Cooke & Plowright, Grevillia 5: 120 (1877).

Ascomata 320–440 μm high × 370–460 μm diam., gregarious, immersed, mostly erumpent to superficial, globose to subglobose, black, coriaceous, with a rough surface, papillate or epapillate, ostiolate, periphysate (Fig. 13a). Peridium 40–50 μm thick laterally, up to 75 μm thick at the apex, thinner at the base, 2-layered, outer layer composed of small heavily pigmented pseudoparenchymatous cells, inner layer very thin, composed of hyaline thin-walled small cells, merging into pseudoparaphyses (Fig. 13a and b). Hamathecium of dense, very long trabeculate pseudoparaphyses, 0.8–1.2 μm broad, branching and anastomosing between and above the asci. Asci 170225 × 17.5–22.5 μm (\( \bar{x} = 199.6 \times 20\mu m \), 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 2226 × 12–15 μm (\( \bar{x} = 24.5 \times 13.3\mu m \), 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).

Anamorph: none reported.

Material examined: FRANCE, Finistère, on Halimone portulacoides (IMI 330806, isotype, as Sphaeria maritima).

Notes

Morphology

When Kohlmeyer and Volkmann-Kohlmeyer (1990) studied the four marine Didymosphaeria species, the monotypic Bicrouania was established to accommodate B. maritima (as Didymosphaeria maritima (P. Crouan & H. Crouan) Sacc.), which could be distinguished from Didymosphaeria by its superficial ascomata lacking a clypeus, thick-walled asci and its association with algae (Kohlmeyer and 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.

Phylogenetic study None.

Concluding remarks

Besides the morphological differences, its marine and substrate habitats also differ from Didymosphaeria.

Bimuria D. Hawksw., Chea & Sheridan, N. Z. J. Bot. 17: 268 (1979). (Montagnulaceae)

Generic description

Habitat terrestrial, saprobic. Ascomata solitary, superficial, globose, dark brown, epapillate, ostiolate. Peridium thin, pseudoparenchymatous. Hamathecium of few, cellular pseudoparaphyses, embedded in mucilage, rarely anastomosing and branching. Asci bitunicate, fissitunicate, broadly clavate with short pedicels, 2-3-spored. Ascospores muriform, broadly ellipsoid, dark brown with subhyaline end cells, verrucose.

Anamorphs reported for genus: none.

Literature: Barr 1987b; Hawksworth et al. 1979; Lumbsch and Huhndorf 2007.

Type species

Bimuria novae-zelandiae Hawksworth, Chea & Sheridan, N. Z. J. Bot. 17: 268 (1979). (Fig. 14)

Fig. 14
figure 15

Bimuria novae-zelandiae (from CBS 107.79, isotype). a–c Asci with a short pedicel and small ocular chamber. d Immature ascus. 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

Ascomata (185-)200 × 310(-330) μm diam., solitary, scattered, semi-immersed or superficial, globose, hyaline when 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 (\( \bar{x} = 75.6 \times 29.4\mu m \), 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 (\( \bar{x} = 59 \times 26\mu m \), 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).

Anamorph: none reported.

Material examined: NEW ZEALAND, North Island, Wairarapa District, Nutty Farm, isolated from soil, 3 Mar. 1978, Chea Chark Yen & J.E. Sheridan (CBS 107.79, isotype).

Notes

Morphology

Bimuria novae-zelandiae was first isolated from soil of a barley field in New Zealand (Hawksworth et al. 1979). Based on B. novae-zelandiae, the genus is characterized by a very thin peridium, mostly 2-spored and fissitunicate asci as well as the muriform, dark brown, verrucose ascospores (Hawksworth et al. 1979). Because of its unique morphological characters, the familial placement of this genus has been debatable and it has been placed in Pleosporaceae (Hawksworth et al. 1979), in Phaeosphaeriaceae (Barr 1987b) and in Melanommataceae (Lumbsch and Huhndorf 2007).

Morphologically, Bimuria is most comparable with some superficially similar or allied genera, in particular Montagnula (Hawksworth et al. 1979). However, the thick carbonaceous peridium distinguishes Montagnula from that of Bimuria (Hawksworth et al. 1979). In addition, the ascospores of Montagnula are discharged forcibly through the ostiole instead of forming a mass outside of the ostiole as in Bimuria (Hawksworth et al. 1979). Ascomauritiana lignicola V.M. Ranghoo & K.D. Hyde has somewhat similar ascospores in 4-spored asci, but this taxon has unitunicate asci (Ranghoo and Hyde 1999). The morphological characters of Bimuria, such as ascospore release and large, thick-walled ascospores may be an adaptation to its soil-borne habitat (Hawksworth et al. 1979).

Phylogenetic study

Bimuria novae-zelandiae was found to be closely related to Phaeodothis winteri (Niessl) Aptroot (syn. Didymosphaerella opulenta (De Not.) Checa & M.E. Barr) and Montagnula opulenta (De Not.) Aptroot in analysis of combined sequences, i.e. SSU rDNA, LSU rDNA, RPB2 and TEF1 sequences (Schoch et al. 2006, 2009). These two species had been included by Barr (2001) in her new family Montagnulaceae.

Concluding remarks

We agree with Barr (2001) and include the genus in Montagnulaceae based on both morphological and phylogenetic characters.

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

Generic description

Habitat terrestrial, saprobic (or parasitic?). Ascomata small- to medium-sized, solitary, scattered, or in small groups, immersed, erumpent to superficial, depressed globose, papillate, ostiolate. Peridium thin. Hamathecium filliform, cellular pseudoparaphyses, embedded in mucilage, anastomosing, septate. Asci bitunicate, fissitunicate, cylindrical, cylindro-clavate or slightly obclavate, with a short knob-like pedicel, with an ocular chamber. Ascospores hyaline, ellipsoid to narrowly obovoid, 3-septate, constricted at each septum.

Anamorphs reported for genus: none.

Literature: Barr 1982a; Berlese 1896; Holm 1957; Shoemaker and Babcock 1989a.

Type species

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

Fig. 15
figure 16

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

Metasphaeria sepalorum Vleugel, Svensk bot. Tidskr. 2: 369 (1908).

Ascomata 120–250 μm high × 170–440 μm diam., solitary, scattered, or in small groups, or forming locules in massive stromatic tissues, initially immersed, becoming erumpent, to nearly superficial, depressed globose, black, membraneous, roughened; apex rounded, sometimes very short and almost inconspicuous, with a somewhat slit-like or Y-shaped ostiole (Fig. 15a). Peridium 16–30 μm wide, comprising two types of cells, outer cells heavily pigmented thick-walled textura angularis, cells 4.5–8 μm diam., cell wall 1–1.5 μm thick, inner cells of subhyaline thin-walled textura angularis, cells larger than outer cells (Fig. 15b). Hamathecium of long cellular pseudoparaphyses, 1.5–2 μm broad, embedded in mucilage, anastomosing, septate. Asci 6383 × 9.5–11 μm (\( \bar{x} = 73.8 \times 10.8\mu m \), n = 10), 8-spored, bitunicate, fissitunicate, oblong, cylindro-clavate or slightly obclavate, with a short knob-like pedicel which is 5–13 μm long, with an ocular chamber (Fig. 15c, d and e). Ascospores (14-)15.5–19 × 5–7 μm (\( \bar{x} = 16.9 \times 5.9\mu m \), n = 10), obliquely uniseriate and partially overlapping to biseriate, ellipsoid to narrowly obovoid, hyaline, 3-septate, constricted at each septum, the cells above central septum often broader than the lower ones, smooth (Fig. 15f, g, h, i and j).

Anamorph: none reported.

Material examined: SWEDEN, on Juncus filliformis, Stockholm, J. Vleugel. Jul. 1907 (S, type as Metasphaeria sepalorum Vleugel).

Notes

Morphology

Bricookea was formally established by Barr (1982a) as a monotypic genus represented by B. sepalorum based on its “globose to depressed ascomata, slit-like ostiole with labial cells, bitunicate asci, cellular pseudoparaphyses, and hyaline septate ascospores”. Bricookea was morphologically assigned to Phaeosphaeriaceae. Holm (1957) checked the authentic collections from North America and type material from Europe, and observed that the ascospores of collections from North America were significantly larger than those from the type material from Sweden. Thus, Shoemaker and Babcock (1989a) considered that the collections from North America represented a new species, which they introduced as B. barrae Shoemaker & C.E. Babc. Although the short slit-like ostiole has previously been reported (Shoemaker and Babcock 1989a), it is inconspicuous in the type specimen from Sweden. Currently, only two species are accommodated in this genus.

Phylogenetic study

None.

Concluding remarks

The knob-shaped pedicel, slit-like ostiole, hyaline ascospores as well as the herbaceous substrate all disagree with any current pleosporalean family. Thus, we temporarily retain this genus under Phaeosphaeriaceae until DNA sequence comparisons can be carried out.

Byssolophis Clem., in Clements & Shear, Gen. fung., Edn 2 (Minneapolis): 286 (1931). (Pleosporales, genera incertae sedis)

Generic description

Habitat terrestrial, saprobic. Ascomata medium-sized, gregarious, semi-immersed to erumpent, coriaceous, ovoid, with a conspicuous elongate slit-like ostiole on the top. Peridium not observed. Hamathecium of dense, long pseudoparaphyses, anastomosing and branching between and above the asci. Asci 8-spored, bitunicate, fissitunicate, cylindrical or cylindro-clavate, with a furcate pedicel. Ascospores fusoid, hyaline, turning faintly brown when old, 1-septate, with a short terminal appendage at each end.

Anamorphs reported for genus: none.

Literature: Clements and Shear 1931; Holm 1986; Müller and von Arx 1962.

Type species

Byssolophis byssiseda (Flageolet & Chenant.) Clem., Gen. Fung. (Minneapolis): 286 (1931). (Fig. 16)

Fig. 16
figure 17

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

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 80105 × (5-)7.5–10 μm (\( \bar{x} = 91 \times 8\mu m \); 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 (\( \bar{x} = 20.8 \times 5.7\mu m \), n = 10), uniseriate to biseriate, fusoid, hyaline, turning faintly brown when old, 1-septate, with 1–2 distinct oil drops in each cell and usually with a short terminal appendage at each end (Fig. 16c).

Anamorph: none reported.

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

Notes

Morphology

Byssolophis was introduced as a monotypic genus based on B. byssiseda, which is characterized by its semi-immersed, gregarious, ovoid ascomata, with a conspicuous central apical ostiolar slit (Holm 1986). Subsequently, two more species were introduced, viz. B. ampla (Berk. & Broome) L. Holm and B. sphaerioides (P. Karst.) E. Müll. (Holm 1986; Müller and von Arx 1962).

Phylogenetic study

The current phylogeny places Byssolophis sphaerioides in proximity of Hypsostromataceae without resolving any sister taxa (Plate 1).

Concluding remarks

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

Generic description

Habitat terrestrial, saprobic. Ascomata medium-sized, scattered to gregarious, superficial, globose, subglobose to turbinate, non papillate with white, orange, red or green ostiolar region, wall black. Hamathecium of dense, long trabeculate pseudoparaphyses, embedded in mucilage, anastomosing between and above the asci. Asci bitunicate, fissitunicate, clavate to nearly cylindrical, with a furcate pedicel. Ascospores fusoid with narrow 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)

Fig. 17
figure 18

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 = 10 μm

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 ostiole, ostiolar region sometimes with orange and greenish tint, wall black, roughened, coriaceous (Fig. 17a). Peridium 55–85 μm thick, peridium outside of the substrate comprising two cell types, outer layer composed of brown thick-walled cells of textura epidermoidea, cells 1–3 μm diam., inner layer composed of small hyaline cells, cells 3–5 μm diam., merging into pseudoparaphyses; peridium inside the substrate one layer, composed of large pale brown cells of textura angularis, cells 6–13 μm diam. (Fig. 17c). Hamathecium of dense, long trabeculate pseudoparaphyses, 1–2 μm broad, embedded in mucilage, anastomosing between and above the asci. Asci 90–120(−148) × 10–14 μm, 8-spored, bitunicate, fissitunicate, cylindro-clavate to clavate, biseriate above and uniseriate below, pedicel 15–20(−53) μm long, the immature asci usually with longer and furcate pedicel (−68 μm) (Fig. 17d,e and f). Ascospores 29–34(−38) × 5.5–8(−10) μm, fusoid with narrow ends, mostly straight, sometimes slightly curved, smooth, pale brown, 1-septate, becoming 3-septate after discharge, with hyaline appendages at each acute to subacute end; in some mature spores the appendage may be absent (Fig. 17b).

Anamorph: Pyrenochaeta sp. (Barr 1984; Samuels and Müller 1978).

Pycnidia 70–500 μm diam. Conidiogenous cells phialidic, lining cavity, 5–8 × 4–6 μm to 5–10 × 3–6 μm. Conidia 2.5–3.5(−4) × 1.5–2(−3) μm, hyaline, ellipsoid or subglobose (Barr 1984).

Material examined: ERIE, Dublin, Glasnevin Botanic Garden, on old rope, Jun. 1872, W. Keit (K(M):108784, holotype, as Sphaeria keitii Berk. & Broome).

Notes

Morphology

Byssosphaeria was introduced by Cooke and Plowright (1879) based on its superficial ascomata seated on a “tomentose subiculum of interwoven threads”, which includes various species in Sphaeria and Byssisedae, and was validly typified by B. keitii (Cooke 1878). Byssosphaeria keitii was treated as a synonym of B. schiedermayeriana (Fuckel) M.E. Barr by Sivanesan (1971), and B. schiedermayeriana exclusively occurs in tropical regions or greenhouse environments in temperate regions (Barr 1984). Morphologically, B. keitii is characterized by its large ascomata with orange to reddish plain apices, and is closely related to B. rhodomphala (Berk.) Cooke (Barr 1984).

For a long time, Byssosphaeria was assigned to Herpotrichia sensu lato, and Byssosphaeria schiedermayeriana was renamed as H. schiedermayeriana Fuckel (von Arx and Müller 1975; Bose 1961; Luttrell 1973; Müller and von Arx 1962; Sivanesan 1971). After studying Herpotrichia in North America, Barr (1984) accepted a relatively narrow generic concept, Herpotrichia sensu stricto, and revived Byssosphaeria; this proposal is supported by phylogenetic study (Mugambi and Huhndorf 2009b). Currently Byssosphaeria comprises 32 species (http://www.mycobank.org, 08-01-2009).

Phylogenetic study

The monophyletic nature of Byssosphaeria is well demonstrated, as well as its familial status in Melanommataceae (Mugambi and Huhndorf 2009b).

Concluding remarks

Orange and greenish plain apices exist in the specimen we examined, which is different from records as “orange, bright or dull reddish plain apices” by Barr (1984). This might be because different specimens have different colours, or there may be a variation of apical colour within a single species, as both orange and green can coexist on the same ascoma (see Fig. 17a). The coloured apical rim, together with the trabeculate pseudoparaphyses as well as the presence of subiculum make Byssosphaeria readily distinguishable from other morphologically comparable genera, e.g. Herpotrichia and Keissleriella (Hyde et al. 2000).

Calyptronectria Speg., Anal. Mus. nac. Hist. nat. B. Aires 19: 412 (1909). (Melanommataceae)

Generic description

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

Anamorphs reported for genus: none.

Literature: Barr 1983; Rossman et al. 1999; Spegazzini 1909.

Type species

Calyptronectria platensis Speg., Anal. Mus. nac. Hist. nat. B. Aires 19: 412 (1909). (Fig. 18)

Fig. 18
figure 19

Calyptronectria platensis (from LPS 1209, holotype). a Appearance of ascomata scattered in the substrate (after removing the out layer of the substrate). Note the protruding papilla. b Section of an ascoma. c Section of the partial peridium. Note the lightly pigmented pseudoparenchymatous cells. d Released ascospores with mucilaginous sheath. e Eight-spored asci in hamathecium and embedded in gel matrix. f Ascus with a short pedicel. Scale bars: a = 0.5 mm, b = 100 μm, c = 50 μm, d–f = 10 μm

Ascomata 120–270 μm high × 170–400 μm diam., solitary, scattered, immersed, lenticular to subglobose, papillate, ostiolate (Fig. 18a and b). Apex with a small and slightly protruding papilla. Peridium 18–30 μm wide, comprising two types of cells, outer layer composed of pseudoparenchymatous cells, cells 3–6 μm diam., cell wall 1–2 μm thick, inner layer comprising less pigmented cells, merging with pseudoparaphyses (Fig. 18b and c). Hamathecium of long, filliform pseudoparaphyses, 1–2 μm broad, branching and anastomosing, embedded in mucilage. Asci 98140 × 12.5–20 μm (\( \bar{x} = 107 \times 15.4\mu m \), n = 10), 8-spored, sometimes 4-spored, bitunicate, fissitunicate, cylindrical to cylindro-clavate, with a short, furcate pedicel, 12–20 μm long, with an ocular chamber (to 4 μm wide × 3 μm high) (Fig. 18e and f). Ascospores 1722.5 μm × (6.3-)7.5–10 μm (\( \bar{x} = 19.8 \times 7.6\mu m \), n = 10), biseriate above and uniseriate below, ellipsoid to broadly fusoid with broadly to narrowly rounded ends, hyaline, usually with (3-)5 transverse septa, with or without 1–3 longitudinal septa in the central cells, constricted at the median septum, the upper cell often broader than the lower one, smooth, surrounded by an irregular hyaline gelatinous sheath up to 3 μm thick (in dry specimen) (Fig. 18d).

Anamorph: none reported.

Material examined: ARGENTINA, La Plata, on decaying branches of Manihot carthaginensis (Jacq.) Müll., Sept. 1906, Spegazzini (LPS 1209, holotype).

Notes

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, trabeculate pseudoparaphyses and hyaline, muriform ascospores as well as its peridium that turns reddish brown in KOH (Rossman et al. 1999) (not shown here). Subsequently, C. indica Dhaware was introduced from India, and Barr (1983) transferred Teichospora ohiensis Ellis & Everh. to Calyptronectria as C. ohiensis (Ellis & Everh.) M.E. Barr. However, this proposal is inappropriate as the type specimen of T. ohiensis is “unitunicate” (Barr 1983; Rossman et al. 1999). Subsequently, Rossman et al. (1999) transferred Calyptronectria ohiensis to Thyridium (as T. ohiense (Ellis & Everh.) Rossman & Samuels).

Phylogenetic study

None.

Concluding remarks

The immersed ascomata, trabeculate pseudoparaphyses, bitunicate asci, hyaline and muriform ascospores as well as the reaction of peridium to KOH (turns reddish brown) make it distinguishable from all other reported genera (Rossman et al. 1999). Thus Calyptronectria is a morphologically well defined genus.

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

Generic description

Habitat marine, saprobic. One or two ascomata per stroma. Ascomata scattered or in small groups, developing beneath the host epidermis, erumpent, lenticular, ostiolate, lacking periphyses. Peridium pale brown, composed of thin-walled elongated cells at the sides and thick-walled cells of textura epidermoidea at the base. Hamathecium of dense, long filliform pseudoparaphyses, embedded in mucilage, anastomosing between and above the asci, rarely septate. Asci 8-spored, bitunicate, fissitunicate, clavate to cylindrical, with a short furcate pedicel, apex with an ocular chamber and apical ring. Ascospores biseriate, narrowly fusoid, yellow to pale brown, multi-septate, constricted at the septa, the two central cells being the largest, surrounded by a gelatinous sheath.

Anamorphs reported for genus: none.

Literature: Hyde 1992a, 1994b.

Type species

Carinispora nypae K.D. Hyde, J. Linn. Soc., Bot. 110: 99 (1992). (Fig. 19)

Fig. 19
figure 20

Carinispora nypae (from BRIP 17106, holotype). a Ascomata on the host surface. b Section of an ascoma. c Section of a partial peridium. d, e, g, h Asci with ocular chambers and short pedicels. f The ocular chamber and apical ring of ascus. i–j Narrowly fusoid ascospores. Scale bars: a = 1 mm, b, c = 100 μm, d, h, i = 50 μm, f = 20 μm, g, e, j =10 μm

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 142207 × 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 4266 × 7–10.6 μm, biseriate, narrowly fusoid with broadly to narrowly rounded ends, somewhat curved, yellow to pale brown, yellow in mass, 7-8-septate, constricted at the septa, the two central cells being the largest, surrounded by a gelatinous sheath; the sheath has a central “spine” and curved polar extrusions (Fig. 19i and j).

Anamorph: none reported.

Material examined: BRUNEI DARUSSALAM, Tungit Api Api mangrove, from decaying intertidal fronds of Nypa fruticans Wurmb., 14 Apr. 1987, K.D. Hyde (BRIP 17106, holotype).

Notes

Morphology

Carinispora is distinguished from Phaeosphaeria by its saprobic life style and lenticular ascomata formed under the host epidermis, peridium structure and sheath surrounding the ascospores (Hyde 1992a, 1994b). Two species were reported, i.e. C. nypae and C. velatispora K.D. Hyde.

Phylogenetic study

Suetrong et al. (2009) could not resolve Carinispora nypae in a phylogeny based on four genes.

Concluding remarks

Both Carinispora nypae and C. velatispora are reported as marine fungi, which should be taken into consideration for their familial placement.

Caryosporella Kohlm., Proc. Indian Acad. Sci., Pl. Sci. 94: 355 (1985). (?Melanommataceae)

Generic description

Habitat marine, saprobic. Ascomata densely scattered or gregarious, superficial, subglobose, black, papillate, ostiolate, periphysate, carbonaceous. Peridium carbonaceous. Hamathecium of dense, long trabeculate pseudoparaphyses, anastomosing and branching above the asci. Asci 8-spored, bitunicate, fissitunicate, cylindrical. Ascospores ellipsoidal to broadly fusoid with narrowly hyaline rounded ends, deep reddish brown, thick-walled, 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)

Fig. 20
figure 21

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

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 thick-walled cells of textura epidermoidea, mixed with subglobose, large cells near the surface, cells 7–15 μm diam., middle layer composed of deep brown, very thick-walled 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 (\( \bar{x} = 137 \times 16.3\mu m \), 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 (\( \bar{x} = 26.8 \times 11\mu m \), n = 10), uniseriate to partially overlapping, ellipsoidal to broadly fusoid with narrow hyaline rounded ends, deep reddish brown, thick-walled, 1-septate with hyaline germ pore at each end, slightly constricted at the septum, verruculose, sometimes with “net”-like ridged ornamentations (Fig. 20f and g).

Anamorph: suspected spermatia (Kohlmeyer 1985).

Material examined: BELIZE, Twin Cays, tip of prop root of Rhizophora mangle, 18 Mar. 1984, J. Kohlmeyer (NY. Herb. J. Kohlmeyer No. 4532a, holotype).

Notes

Morphology

Caryosporella was formally established by Kohlmeyer (1985) based on the obligate marine fungus, C. rhizophorae, which is characterized by its superficial ascomata, 3-layered peridium, filliform trabeculate pseudoparaphyses, and brown, 1-septate ascospores. Caryosporella was originally assigned to Massariaceae despite several major differences, such as the superficial ascomata, reddish brown ascospores (Kohlmeyer 1985). Subsequently, Caryosporella was assigned to Melanommataceae (Eriksson 2006; Lumbsch and Huhndorf 2007).

Phylogenetic study

Suetrong et al. (2009) showed that a single isolate of Caryosporella rhizophorae does not reside in Pleosporales, but is related to Lineolata rhizophorae (Kohlm. & E. Kohlm.) Kohlm. & Volkm.-Kohlm. and placed in Dothideomycetidae incertae sedis.

Concluding remarks

As an obligate marine fungus, the familial placement of Caryosporella rhizophorae is uncertain but it may not belong to Pleosporales.

Chaetomastia (Sacc.) Berl., Icon. fung. (Abellini) 1: 38 (1890). (Teichosporaceae)

Melanomma subgen. Chaetomastia Sacc., Syll. fung. (Abellini) 2: 113 (1883).

Generic description

Habitat terrestrial, saprobic. Ascomata relatively small, scattered, or in small groups, superficial, globose or subglobose, black, papillate, ostiolate, coriaceous. Peridium relatively thin, 1-layered, composed of heavily pigmented cells of textura angularis. Hamathecium of dense, long cellular pseudoparaphyses, embedded in mucilage. Asci mostly 4-spored, bitunicate, fissitunicate, broadly cylindrical with a furcate pedicel, with a large ocular chamber, especially apparent in immature asci. Ascospores ellipsoid to broadly fusoid with broadly to narrowly rounded ends, brown, 3-septate, constricted at all septa.

Anamorphs reported for genus: coelomycetous where known: conidia hyaline or brown, aseptate or 1-septate (Aposphaeria- or Coniothyrium-like) (Barr 1989c).

Literature: Barr 1987b, 1989c; 1993a; b; 2002; Berlese 1890; Clements and Shear 1931; Eriksson 1999; Eriksson and Hawksworth 1987, 1998; Holm 1957; Leuchtmann 1985; Saccardo 1883.

Type species

Chaetomastia hirtula (P. Karst.) Berl., Icon. fung. (Abellini) 1: 38 (1890). (Fig. 21)

Fig. 21
figure 22

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 = 10 μm

Sphaeria hirtula P. Karst., Fungi Fenn. Exs. N. 825 (1869).

Ascomata 214–286 μm high × 210–258 μm diam., scattered or in groups, superficial, globose, wall black; apex often opening with a broad pore within slightly raised papilla, up to 30 μm diam., coriaceous (Fig. 21a). Peridium 20–26 μm thick, 1-layered, composed of heavily pigmented cells of textura angularis, cells up to 5 × 15 μm diam., cell wall up to 3.5 μm thick (Fig. 21b). Hamathecium of dense, long cellular pseudoparaphyses, embedded in mucilage. Asci 90130 × 12.5–17.5(−22.5) μm (\( \bar{x} = 111 \times 16.3\mu m \), 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 (\( \bar{x} = 23.5 \times 8.2\mu m \), 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).

Anamorph: none reported.

Material examined: FINLAND, ETELÄ-HÄME (EH/Ta), Tammela, Mustiala, På Rub. id., 8 May 1866. P.A. Karsten (H, FFE 825, kleptotype).

Notes

Morphology

Chaetomastia was introduced by Saccardo (1883) as a 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) promoted it to genus rank. Subsequently, Chaetomastia hirtula (P. Karst.) Berl. was selected as the lectotype species of the genus (Clements and Shear 1931). Chaetomastia has been regarded as having unitunicate asci (Eriksson and Hawksworth 1986, 1998; Eriksson 1999). However its bitunicate status was confirmed by Holm (1957). Holm (1957) treated C. hirtula as Melanomma hirtulum (P. Karst.) Sacc., and Leuchtmann (1985) transferred this species to Montagnula sensu lato based on the ascospore morphology and the hyphae surrounding the ascomata. Barr (1987b) suggested that ascoma, peridium structure and ascospore characters pointed Montagnula sensu stricto to Phaeosphaeriaceae, while the characters of ascomata and peridium structure of Chaetomastia were thought to fit the definition of Dacampiaceae (Barr 1987b). In particular, the peridium and ascospore characters of C. hirtula are comparable with those of the generic type of Massariosphaeria (M. phaeospora). Thus, Barr (1989c) accepted Massariosphaeria sensu stricto and assigned the phragmosporous species of Massariosphaeria sensu lato to Chaetomastia.

Barr (2002) later assigned Chaetomastia to Teichosporaceae based on its saprobic or hypersaprobic lifestyle, occurring on woody stems and peridium structure, and this is widely followed (Eriksson 2006; Lumbsch and Huhndorf 2007). Currently, 11 species are accepted in this genus (http://www.indexfungorum.org/).

Phylogenetic study

None.

Concluding remarks

Familial placement of Chaetomastia is undetermined currently but has been included in the Teichosporaceae by authoritative sources (Eriksson 2006; Lumbsch and Huhndorf 2007) or the Dacampiaceae (http://www.indexfungorum.org/).

Chaetoplea (Sacc.) Clem., Gen. Fung. (Minneapolis): 275 (1931). (?Phaeosphaeriaceae)

Pyrenophora subgen. Chaetoplea Sacc., Syll. fung. (Abellini) 2: 279 (1883).

Generic description

Habitat terrestrial, saprobic. Ascomata small to medium, immersed, erumpent to superficial, globose to subglobose, papillate, ostiolate. Peridium not examined. Hamathecium of dense, long, narrowly cellular pseudoparaphyses. Asci 8-spored or 4-spored, bitunicate, fissitunicate, cylindro-clavate, 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.

Type species

Chaetoplea calvescens (Fr.) Clem., Gen. Fung. (Minneapolis): 275 (1931). (Fig. 22)

Fig. 22
figure 23

Chaetoplea calvescens (from FH-81113, isotype). a, b Four-spored and 8-spored asci. c Released ascospores. Scale bars: a–c = 10 μm

≡ 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 90110 × 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 1318 × 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 401, nor did Webster & Lucas in the taxonomic and life-history study (Trans. Brit. Myc. Soc. 42, 332–342. 1959) of this species.

The specimen has most of the features described by Webster & Lucas including the presence of the conidial state Microdiplodia henningsii Staritz. I did not see vertical septa in the ascospores. Webster & Lucas note that vertical septa may be occasionally be lacking. The fungus is otherwise as they describe it although some perithecia collapse and appear cupulate.”—by R.A. Shoemaker.

Phylogenetic study

None.

Concluding remarks

The substrate of Chaetoplea sensu Barr (1990b) can be herbaceous stalks, decorticated wood or periderm, or old cotton cloth and string, which may indicate its heterogeneous nature. The ascospores seem very much like Phaeosphaeria which may be an earlier name; more details concerning the ascomatal, peridial and hamathecial structures are needed to make any conclusion.

Cilioplea Munk, Dansk botanisk Arkiv 15: 113 (1953). (Pleosporales, genera incertae sedis)

Generic description

Habitat terrestrial, saprobic. Ascomata small- to medium-sized, solitary, scattered or in small groups, immersed, globose or subglobose, papilla covered with short and blackish setae, coriaceous. Peridium thin, comprising small heavily pigmented thick-walled cells of textura angularis. Hamathecium of cellular pseudoparaphyses. Asci 8-spored, bitunicate, fissitunicate, broadly clavate, with a short, furcate pedicel, and small ocular chamber. Ascospores fusoid to narrowly fusoid with narrowly rounded ends, pale brown to reddish brown, multi-transverse septa, usually with one longitudinal septum in some central cells, constricted at the primary septum.

Anamorphs reported for genus: none.

Literature: Barr 1990b, 1992b; Crivelli 1983; Lumbsch and Huhndorf 2007; Müller 1951; Munk 1953, 1957.

Type species

Cilioplea coronata (Niessl) Munk, Dansk botanisk Arkiv 15: 113 (1953). (Fig. 23)

Fig. 23
figure 24

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

≡ Pleospora coronata Niessl, Notiz. Pyr.: 16 (1876).

Ascomata 170–290 μm high × 200–410 μm diam., solitary, scattered, or in small groups, immersed, globose or subglobose, wall black, papilla raised, 50–80 μm high, with short and blackish setae, coriaceous (Fig. 23a). Peridium 9–15 μm thick laterally, up to 28 μm thick at the apex, thinner at the base, 1-layered, composed of small heavily pigmented thick-walled 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 (\( \bar{x} = 85.3 \times 12.1\mu m \), n = 10), 8-spored, bitunicate, fissitunicate, broadly clavate, with a short, thick, furcate pedicel, 5–15 μm long, and a small ocular chamber (to 3 μm wide × 2 μm high) (Fig. 23c and d). Ascospores 2127.5 × 5.5–7.5 μm (\( \bar{x} = 24 \times 6.7\mu m \), n = 10), biseriate to uniseriate at base, fusoid to narrowly fusoid with narrowly rounded ends, pale reddish brown, 5–7 transverse septa (mostly 5), usually with one longitudinal septum in some central cells, deeply constricted at the median septum, the part above the primary septum shorter and broader, smooth-walled.

Anamorph: none reported.

Material examined: GERMANY, Hadiberg. on Reseda lutea Hadiberg, 20 Sept. 1875, Niessl (M 175-89-290, lectotype; M 175-89-291, type).

Notes

Morphology

Cilioplea was introduced by Müller (1951) as a subgenus of Pleospora, and this was followed by Munk (1957), who had earlier proposed it as a separate genus typified by C. coronata based on its hairy papilla, clavate asci as well as its “perfectly paraphysoid” (see Munk 1953). A relatively narrow concept of Pleospora was accepted by Crivelli (1983), and four species was assigned under the separate genus Cilioplea, viz. C. coronata, C. genisticola (Fautrey & Lambotte) Crivelli, C. kansensis (Ellis & Everh.) Crivelli and C. nivalis (Niessl) Crivelli. Subsequently, another six species were added (Barr 1990b, 1992b). Currently, ten species are included under Cilioplea.

Phylogenetic study

None.

Concluding remarks

The most striking character of Cilioplea is its setose papilla, which has been shown to have no phylogenetic significance in Lentitheciaceae (Zhang et al. 2009a). Cilioplea was assigned under Lophiostomataceae (Lumbsch and Huhndorf 2007), but there is little morphological similarity with the Lophiostomataceae sensu stricto (Zhang et al. 2009a). Thus its familial placement needs further study.

Crivellia Shoemaker & Inderb., in Inderbitzin, Shoemaker, O’Neill, Turgeon & Berbee, Can. J. Bot. 84: 1308 (2006). (Pleosporaceae)

Generic description

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)

Fig. 24
figure 25

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 = 20 μm

≡ Cucurbitaria papaveracea De Not., Sfer. Ital.: 62 (1863).

Ascomata 210–260 μm high × 300–380 μm diam., densely scattered, immersed, erumpent to nearly superficial, flattened globose, dark brown, papillate, ostiolate (Fig. 24a). Peridium 25–30 μm thick, thicker near the apex and thinner at the base, composed of two cell types, outer cells of thick-walled and textura angularis, cells up to 10 × 5 μm diam., cell wall 2–4 μm thick, inner cells thin-walled, yellow (Fig. 24b). Hamathecium of dense, long, 1–2 μm broad, rarely septate pseudoparaphyses. Asci 85125 × 10–13 μm (\( \bar{x} = {1}0{6} \times {11}\mu {\text{m}} \), 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 (\( \bar{x} = 20.4 \times 6.3\mu m \), 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 with one or two vertical septa in central cells. Its Brachycladium anamorphic stage with phragmosporous conidia also differs from that of Stemphylium, which is the anamorphic stage of Pleospora (Inderbitzin et al. 2006). Currently, two species are included within Crivellia, i.e. C. homothallica Inderb. & Shoemaker and C. papaveracea.

Phylogenetic study

Crivellia papaveracea was shown to be closely related to some species of Alternaria, and its pleosporaceous status was confirmed following molecular studies (Inderbitzin et al. 2006).

Concluding remarks

Crivellia seems to belong to Pleosporaceae, and may be closely related to Pleospora.

Decaisnella Fabre, Annls Sci. Nat., Bot., sér. 6 9:112 (1878). (Pleosporales, genera incertae sedis)

Generic description

Habitat terrestrial, saprobic. Ascomata medium to large, immersed to erumpent, clypeate, papillate, ostiolate. Hamathecium of dense, long, cellular pseudoparaphyses, rarely septate, embedded in mucilage. Asci mostly 4- or 8-spored, rarely 2-spored, cylindrical to cylindro-clavate, with a furcate pedicel. Ascospores muriform, dark brown, oblong with broadly rounded ends.

Anamorphs reported for genus: none.

Literature: Barr 1986; 1990a; b; Fabre 1878; Saccardo 1883.

Type species

Decaisnella spectabilis Fabre, Annls Sci. Nat., Bot., sér. 6 9: 112 (1879). (Fig. 25)

Fig. 25
figure 26

Decaisnella spectabilis (NY2082, syntype). a Appearance of ascomata on the host surface. b Section of a partial peridium (immersed in the substrate). Note the pseudoparenchymatous out layer. c, d Muriform ascospores. Note the minuitely verrucose ornamentation. e Ascus with a short pedicel. Scale bars: a = 0.5 mm, b = 100 μm, c–e = 20 μm

Ascomata 520–680 μm high × 430–600 μm diam., solitary, scattered, or in small groups of 2–3, immersed to erumpent, clypeate, globose or subglobose, black, roughened, with a blunt papilla up to 170 μm high, apex with a round ostiole, coriaceous (Fig. 25a). Peridium 70–90 μm thick at sides, thicker near the apex, comprising two types of cells; part immersed in host tissue, outer layer pseudoparenchymatous, 55–65 μm thick, pigmented, inner layer composed of lightly pigmented to hyaline thin-walled compressed cells, 15–23 μm thick, cells 3.5–7 μm diam., part above host tissue heavily pigmented covered by clypeus tissues (Fig. 25b). Hamathecium of dense, long, cellular pseudoparaphyses, 1.5–3 μm broad, rarely septate, embedded in mucilage. Asci 150200 × 15–25(−33) μm (\( \bar{x} = 181 \times 20.6\mu m \), 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 3745 × 12–17 μm (\( \bar{x} = 43 \times 15\mu m \), n = 10), uniseriate and sometimes slightly overlapping, oblong with broadly rounded ends, dark brown, verrucose or smooth, 7–9 transverse septa and 1–3 longitudinal septa in some of the cells, no constriction at the septa (Fig. 25c and d).

Anamorph: none reported.

Material examined: GERMANY, Valsalpe in der Ramsau, Bayer, Alpen, on Rhamnus pumila Turra., Jul. 1913, Karl Arnold (NY2082, syntype as Teichospora megalocarpa Rehm).

Notes

Morphology

Decaisnella was formally established by Fabre (1879), but was treated as a synonym of Teichospora by Saccardo (1883). This was followed by several mycologists over a long time. The main morphological differences between Decaisnella and Teichospora include the size and septation of ascospores, shape of ascomata, structure of peridium and type of pseudoparaphyses (Barr 1986). Thus Barr (1986) revived Decaisnella and assigned it to Massariaceae based on the shape of ascomata and large, distoseptate ascospores. Currently, 15 species are accepted under Decaisnella (http://www.mycobank.org/MycoTaxo.aspx). Neither the size of ascomata nor the ascospore characters have proven sufficient to place taxa at the family level in Pleosporales (Zhang et al. 2009a), and therefore familial placement of Decaisnella remains uncertain.

Phylogenetic study

Decaisnella formosa resided in the clade of Lophiostomataceae and in proximity to Lophiostoma macrostomoides De Not. (Plate 1).

Concluding remarks

The muriform ascospores, saprobic life style and 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)

Generic description

Habitat terrestrial, saprobic (coprophilous). Ascomata medium- to large-sized, solitary or scattered, immersed to erumpent, globose or subglobose, apex with or without papilla, ostiolate. Peridium thin, composed of compressed cells. Hamathecium of dense, long pseudoparaphyses, anastomosing and branching. Asci 8-spored, cylindrical to cylindro-clavate, with short pedicel. Ascospores uni- to triseriate, pale to dark brown, ellipsoid, 1-septate, usually constricted at the septum, smooth, with a full length germ slit in each cell.

Anamorphs reported for genus: none.

Literature: Auerswald 1866; Barr 2000; Cain 1934; Dennis 1968; Eriksson 2006; Griffiths 1901; Hyde and Steinke 1996; Kirschstein 1911; Kruys et al. 2006; 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)

Fig. 26
figure 27

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 = 50 μm

Ascomata 400–800 μm diam., solitary or scattered, immersed, globose or subglobose, black, papilla short, 70–130 μm broad, central, with a wide opening, coriaceous (Fig. 26a). Peridium ca. 15 μm thick laterally, up to 35 μm thick at the apex, up to 30 μm at the base, comprising a single layer of small lightly pigmented thin-walled cells of textura angularis, cells 4–10 μm diam., cell wall <1 μm thick, apex cells smaller and wall thicker (Fig. 26b). Hamathecium of dense, very long pseudoparaphyses, 1.5–2 μm broad, anastomosing and branching. Asci 290380 × 35–45 μm (\( \bar{x} = 357.5 \times 40.6\mu m \), n = 10), 8-spored, bitunicate, fissitunicate, cylindrical to cylindro-clavate, with short, narrowed pedicels which are rounded at the base, 25–60 μm long, apex with a wide ocular chamber (Fig. 26d and e). Ascospores 50–58 × 20–22.5 μm (\( \bar{x} = 54 \times 21.3\mu m \), n = 10), obliquely uniseriate and partially overlapping, ellipsoid with narrowly rounded ends, reddish brown, 1-septate, slightly constricted at the septum, smooth-walled, each cell with a full length germ slit (Fig. 26c).

Anamorph: none reported.

Material examined: GERMANY, Near Königstein, in forest, rare, Oct. 1904, W. Krieger (L, 1950).

Notes

Morphology

Delitschia was established by Auerswald (1866), and assigned to Sphaeriaceae. It was considered to be closely related to Sordariaceae and Amphisphaeriaceae. Winter (1887) assigned Delitschia under Sordariaceae, and this placement is followed in several subsequent studies (Griffiths 1901; Kirschstein 1911). Cain (1934) suggested that Delitschia might belong in Pleosporaceae, and this proposal was supported by Moreau (1953) and Dennis (1968). Finally, Munk (1957) established Sporormiaceae (Pseudosphaeriales), and Delitschia was assigned therein. Luck-Allen and Cain (1975) reviewed and redefined the genus as having bitunicate asci, pigmented and 1-septate ascospores with an elongated germ slit in each cell and surrounded by a gelatinous sheath, and in particular, the coprophilous habitat. Luck-Allen and Cain (1975) accepted 46 species. Subsequently, some wood-inhabiting species were also described (Hyde and Steinke 1996; Romero and Samuels 1991). Three genera, i.e. Delitschia, Ohleriella and Semidelitschia were separated from Sporormiaceae, and a new family, Delitschiaceae, was introduced by Barr (2000) to accommodate them. Delitschiaceae is characterized by a periphysate ostiole, wide endotunicate asci with a wide ocular chamber and ascospores having cells with germ slits. Delitschiaceae has been subsequently accepted (Eriksson 2006; Lumbsch and Huhndorf 2007).

The genus comprises 83 names (Index Fungorum) and is estimated to comprise 51 species (Kirk et al. 2008). Keys to Delitschia can be found in Luck-Allen and Cain (1975) and Hyde and Steinke (1996).

Phylogenetic study

Delitschia didyma and D. winteri (W. Phillips & Plowr.) Sacc. form a robust phylogenetic clade within Delitschiaceae, which is basal to other members of Pleosporales (Kruys et al. 2006; Schoch et al. 2006) except for Massariaceae (Voglmayr and Jaklitsch 2011). This might indicate its early derivation (Zhang et al. 2009a).

Concluding remarks

Morphologically, Delitschia is a well defined genus, and each cell of the ascospore has a full length germ slit. Currently, most species of this genus are coprophilous, although a few species are reported from wood (Hyde and Steinke 1996; Luck-Allen and Cain 1975). Whether the lignicolous habitat is an important character that might separate these taxa from the main coprophilous group, needs to be addressed, however, the morphological characters are similar.

Didymosphaeria Fuckel, Jb. nassau. Ver. Naturk. 22–23: 140 (1870). (Didymosphaeriaceae)

Generic description

Habitat terrestrial, saprobic or parasitic. Ascomata solitary, scattered, or in small groups, immersed to erumpent, globose to ovoid, papillate, ostiolate, periphysate. Ostiole with a pore-like opening. Peridium 1-layered, thin, composed of brown pseudoparenchymatous cells of textura angularis. Hamathecium of dense, trabeculate, anastomosing mostly above the asci. Asci (2-)4-spored or 8-spored, bitunicate, cylindrical, with a furcate pedicel. Ascospores uniseriate, ellipsoid, brown, 1-distoseptate.

Anamorphs reported for genus: Dendrophoma, Fusicladiella and Phoma (Aptroot 1995).

Literature: Aptroot 1995; Barr 1989a, b, 1990a, 1992a, b; 1993a; b; Fuckel 1870; Hawksworth 1985a, b; Hawksworth and Boise 1985; Hawksworth and Diederich 1988; Hyde et al. 2000; Lumbsch and Huhndorf 2007; Saccardo 1882; Scheinpflug 1958; Sivanesan 1984.

Type species

Didymosphaeria futilis (Berk. & Broome) Rehm, Hedwigia 18: 167 (1879). (Fig. 27)

Fig. 27
figure 28

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 = 20 μm

≡ 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 90110 × 7.5–10 μm (\( \bar{x} = 97 \times 9\mu m \), 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 1415.5 × (5.5-) 6–7.5 μm (\( \bar{x} = 14.8 \times 6.9\mu m \), 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; Hawksworth 1985a, b; Hawksworth and Boise 1985; Hawksworth and Diederich 1988; Scheinpflug 1958). Over 400 epithets of Didymosphaeria were included until the monograph of Aptroot (1995).

Aptroot (1995) examined more than 3000 specimens under the name Didymosphaeria. The type specimen of Didymosphaeria (Fungi Rhenani 1770) represents the widespread and common D. futilis (Aptroot 1995). In this study, we did not get the lectotype specimen, but described the type of D. futilis (Sphaeria futilis). Using a narrow concept (ignoring differences of host or country of origin), Aptroot (1995) accepted only seven species, which were closely related with the generic type of Didymosphaeria with over 100 synonyms distributed among them. Many taxa were found to belong to other groups, i.e. Aaosphaeria, Amphisphaeria, Astrosphaeriella, Dothidotthia, Flagellosphaeria, Kirschsteiniothelia, Megalotremis, Montagnula, Munkovalsaria, Mycomicrothelia, Parapyrenis or Phaeodothis. Didymosphaeria is mainly characterized by a peridium consisting of flattened or irregular cells or completely hyphae; a hamathecium consisting of narrow, trabeculate paraphysoids or paraphyses, richly anastomosing above the asci; and brown thinly distoseptate ascospores. Didymosphaeriaceae was maintained as a separated family within Pleosporales by Aptroot (1995) because of the distoseptate ascospores and trabeculate pseudoparaphyses mainly anastomosing above the asci. This proposal, however, has not received much support (Lumbsch and Huhndorf 2007).

Phylogenetic study

There have been few molecular investigations of Didymosphaeria when compared to the morphological studies. Didymosphaeria futilis resided in the clade of Cucurbitariaceae (or Didymosphaeriaceae) (Plate 1). The correct identification of the Didymosphaeria strain used for sequencing, however, has not been verified.

Concluding remarks

Didymosphaeria is a well established genus represented by D. futilis. Of particular significance are the narrow pseudoparaphyses which anastomose above the asci and brown 1-septate ascospores with indistinct distosepta. Familial placement of Didymosphaeria is unclear yet because of insufficient molecular data.

Dothidotthia Höhn., Ber. Deutsch. Bot. Ges. 36: 312 (1918). (Didymellaceae)

Generic description

Habitat terrestrial, saprobic. Ascomata medium-sized, solitary, clustered or somewhat gregarious, erumpent, subglobose, apex somewhat papillate to depressed, coriaceous. Peridium composed of a few layers of dark brown cells of textura angularis, and giving rise dark brown, thick-walled hyphae in the basal region, 2-layered. Hamathecium septate 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: Barr 1989b; Phillips et al. 2008.

Type species

Dothidotthia symphoricarpi (Rehm) Höhn., Ber. Deutsch. Bot. Ges. 36: 312 (1918). (Fig. 28)

Fig. 28
figure 29

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)

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 70120 × 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).

Phylogenetic study

Based on a multi-gene phylogenetic analysis, Dothidotthia formed a separate familial clade (Phillips et al. 2008). Thus Dothidotthiaceae was introduced to accommodate it (Phillips et al. 2008).

Concluding remarks

By comparing the morphological characters and phylogenetic dendrograms by Phillips et al. (2008) and de Gruyter et al. (2009), Dothidotthia seems closely related to Didymellaceae, but Dothidotthiaceae should still be treated as a separate family.

Dubitatio Speg., Anal. Soc. cient. argent. 12: 212 (1881). (Arthopyreniaceae (or Massariaceae))

Generic description

Habitat terrestrial, saprobic. Ascomata medium-sized, solitary, densely scattered, or in small groups of 2–4, immersed, covered with white crystaline rim, papillate, ostiolate. Hamathecium of dense pseudoparaphyses, long, 2–3 μm broad, branching and anastomosing. Asci cylindrical, pedicellate, with furcate pedicel. Ascospores 1-septate, asymmetrical, reddish to dark brown.

Anamorphs reported for genus: Aplosporella-like (Rossman et al. 1999).

Literature: Barr 1979b, 1987b; Müller and von Arx 1962; Rossman et al. 1999; Spegazzini 1881.

Type species

Dubitatio dubitationum Speg., Anal. Soc. cient. argent. 12: 212 (1881). (Fig. 29)

Fig. 29
figure 30

Dubitatio dubitationum (from NY, isotype; LPS, holotype). a Appearance of ascomata scattered on the host surface. Note the exposed white covering around the ostioles. b, c Section of an ascoma. Note the white covering (see arrow). d–f Cylindrical asci with short furcate pedicels. g–i Asymmetrical, 1-septate reddish-brown ascospores. Scale bars: a = 1 mm, b = 100 μm, c = 50 μm, d–i = 20 μm

Ascomata 350–530 μm high × 550–700 μm diam., solitary, densely scattered, or in small groups of 2–4, immersed, with a protruding papilla, 110–160 μm high, 160–250 μm diam., globose or subglobose, black, covered with white crystalline material which becomes hyaline and gel-like in water, ostiolate (Fig. 29a and b). Peridium 18–25 μm thick laterally (excluding the rim), up to 35 μm thick at the apex, thinner at the base, 1-layered, composed of small pale brown thin-walled cells of textura prismatica, cells 5–12 × 3–5 μm diam., cell wall up to 1 μm thick, apex cells smaller and walls thicker (Fig. 29b). Hamathecium of dense, long pseudoparaphyses, 2–3 μm broad, branching and anastomosing between and above the asci. Asci 150–190(−230) × 12.5–15 μm (\( \bar{x} = 172.5 \times 13.4\mu m \), 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 1922.5 × 10–12 μm (\( \bar{x} = 20.2 \times 11.4\mu m \), n = 10), uniseriate to obliquely uniseriate and partially overlapping, broadly ellipsoid with broadly to narrowly rounded ends, reddish brown, 1-septate, constricted at septum, asymmetric with a larger upper cell, thick-walled, possibly distoseptate (Fig. 29f, g and h).

Anamorph: Aplosporella-like (for detailed description see Rossman et al. 1999).

Conidiomata globose, ca. 300 μm diam. Conidia holoblastic, broadly fusoid, 13–15 × 7–10 μm, dark brown, finely spinulose (Rossman et al. 1999).

Material examined: ARGENTINA, Buenos Aires, Tuyu, on Celtis tala Gill., Jan. 1881, leg. det. C. Spegazzini (NY, isotype; LPS, holotype).

Notes

Morphology

When established Dubitatio, Spegazzini (1881) considered it as intermediate between Sphaeriaceae and Nectriaceae as has been mentioned by Rossman et al. (1999). Müller and von Arx (1962) treated Dubitatio as a synonym of Passerinula, while the differences of ascomata and ascospores could easily distinguish these two genera (Rossman et al. 1999). After checking the type specimen, Dubitatio was assigned to Dothideomycetes, and considered closely related to Dothivalsaria in the Massariaceae (Barr 1979b, 1987b). Dubitatio chondrospora was assigned to Pseudomassaria (as P. chondrospora (Ces.) Jacz.) (Barr 1964; Müller and von Arx 1962).

Phylogenetic study

None.

Concluding remarks

The black ascomata with white crystalline covering and central white ostiolar region as well as the asymmetrical reddish brown ascospores are striking characters of Dubitatio dubitationum. The genus cannot be assigned to any family with certainty based on morphological characters and fresh collections are needed for sequencing.

Entodesmium Reiss, Hedwigia 1: 28 (1854). (Phaeosphaeriaceae)

Generic description

Habitat terrestrial, saprobic (or parasitic?). Ascomata scattered or in small groups, immersed, papillate, ostiolate, periphysate. Peridium thin, comprising one cell type of pigmented pseudoparenchymatous cells. Hamathecium of dense, long pseudoparaphyses, septate, embedded in mucilage. Asci 8-spored, bitunicate, fissitunicate, cylindrical, with furcate pedicel. Ascospores ellipsoid to filliform, multi-septate, deeply constricted at the primary septum (usually near apex), breaking into partspores.

Anamorphs reported for genus: none.

Literature: von Arx and Müller 1975; Barr 1992b; Eriksson 1967a; b; Holm 1957; Liew et al. 2000; Shoemaker 1984a, b.

Type species

Entodesmium rude Reiss, Hedwigia 1: 28 (1854). (Fig. 30)

Fig. 30
figure 31

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, dh = 10 μm

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 100220 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. 30b and c). Hamathecium of dense, long pseudoparaphyses, septate, 2–3 μm wide, embedded in mucilage. Asci 100175 × 8–13 μm (\( \bar{x} = 147.5 \times 11.3\mu m \), n = 10), 8-spored, bitunicate, fissitunicate, cylindrical, with a furcate pedicel which is 18–50 μm long, and with a low ocular chamber (ca. 1 μm wide × 1 μm high) (Fig. 30e,f, g and h). Ascospores 108138 × 3–3.5 μm (\( \bar{x} = 123 \times 3.2\mu m \), n = 10), filliform, brown, multi-septate, breaking into 22–28 partspores, 5–7 × 3–3.5 μm diam. (Fig. 30d).

Anamorph: none reported.

Material examined: GERMANY, Königstein, on stems of Coronilla varia L., 20 May 1895, W. Krieger (H, Krieger 1070).

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) assigned Entodesmium to the Pleosporaceae sensu lato, and Shoemaker (1976) assigned E. rude (as Ophiobolus rudis) to Ophiobolus sensu lato based on the fragmenting filliform ascospores. According to the short, blackish beak and periphysate ostiole, Barr (1992b) assigned Entodesmium to Lophiostomataceae. The hosts of Entodesmium are restricted to stems of legumes (Barr 1992b; Shoemaker 1984b).

Phylogenetic study

Limited phylogenetic studies indicate that Entodesmium rude may have affinities to Phaeosphaeriaceae (Liew et al. 2000; Plate 1).

Concluding remarks

Species of Entodesmium share several morphological characters, such as immersed, papillate ascomata, periphysate ostioles, pale yellow to light yellowish brown, multi-septate (≥ 3), narrowly fusoid to filliform ascospores, and are specific to legumes. All of the above similarities indicate a close relationship among members of Entodesmium. We do not agree with Barr (1992b) who assigned Entodesmium to Lophiostomataceae because the ascomata are immersed, the papilla are not laterally compressed and the peridium comprises a single type of cells of textura angularis. These characters plus multi-septate, lightly pigmented ascospores, which break up into partspores and host specificity to legumes support inclusion in Phaeosphaeriaceae. Entodesmium multiseptatum (G. Winter) L. Holm and E. niessleanum were originally described as Leptosphaeria species (Shoemaker 1984b) indicating their similarity to Phaeosphaeria with which Leptosphaeria is commonly confused (Shoemaker 1984a; Shoemaker and Babcock 1989b). Phylogenetic study has also shown that Entodesmium rude is related to members of Phaeosphaeriaceae (Liew et al. 2000). Thus we assign Entodesmium to Phaeosphaeriaceae as a separate genus until further phylogenetic analysis is carried out on verified specimens.

Eudarluca Speg., Revta Mus. La Plata 15: 22 (1908). (?Phaeosphaeriaceae)

Generic description

Habitat terrestrial, parasitic. Ascomata small, solitary, scattered, immersed to erumpent, subglobose, ostiolate, papillate. Peridium thin, composed of a few layers cells of textura prismatica. Hamathecium of dense, cellular pseudoparaphyses, septate. Asci 8-spored, bitunicate, fissitunicate, cylindrical to fusoid, with a furcate pedicel. Ascospores broadly fusoid to fusoid, hyaline to pale yellow, rarely 1- or 3- septate, mostly 2-septate, constricted at the primary septum.

Anamorphs reported for genus: Sphaerellopsis (Sivanesan 1984).

Literature: Bayon et al. 2006; Eriksson 1966; Katumoto 1986; Ramakrishnan 1951; Spegazzini 1908.

Type species

Eudarluca australis Speg., Revta Mus. La Plata 15: 22 (1908). (Fig. 31)

Fig. 31
figure 32

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

Ascomata 160–190 μm high × 180–290 μm diam., solitary, scattered, or in small groups, semi-immersed to erumpent, subglobose to broadly ellipsoid, wall black, ostiolate, apex with a short papilla, 40–70 μm broad (Fig. 31a and b). Peridium < 10 μm wide laterally, up to 25 μm thick at the apex, thinner at the base, composed of lightly pigmented thin-walled cells of textura prismatica, cells up to 12 × 4 μm diam., cell wall <1 μm thick, apex cells heavily pigmented, smaller and walls thicker (Fig. 31b and c). Hamathecium of dense, long cellular pseudoparaphyses, 1.5–2.5 μm broad, septate. Asci 5070 × 7.5–10 μm (\( \bar{x} = 61.4 \times 8.4\mu m \), n = 10), 8-spored, with a short, thick, furcate pedicel, up to 12.5 μm long, bitunicate, fissitunicate, cylindrical to fusoid, no obvious ocular chamber (Fig. 31d, e, f and g). Ascospores 1620 × 4–6 μm (\( \bar{x} = 17.3 \times 5\mu m \), n = 10), obliquely uniseriate and partially overlapping to biseriate, broadly fusoid to fusoid, hyaline to pale yellow, 2-septate, sometimes 1- or 3-septate, constricted at the two main septa, the medium cell often broader than the others, smooth (Fig. 31h).

Anamorph: Sphaerellopsis filum (Biv.) B. Sutton (Sivanesan 1984).

Material examined: BRAZIL, Sao Paulo, on leaves of Canna sp., 1905, leg. Usteri, nro; det. Ove Eriksson (LPS 5.415, type).

Notes

Morphology

Eudarluca was introduced based on E. australis (Spegazzini 1908), and E. australis was subsequently treated as a synonym of E. caricis (Biv.) O.E. Erikss. (Eriksson 1966). The most striking character of E. australis is its 2-septate ascospores, which is quite rare in Pleosporales. Sphaerellopsis filum, anamorph of E. caricis, is a cosmopolitan hyperparasite associated with a large number of rust species (Płachecka 2005).

Phylogenetic study

A detailed phylogenetic study was conducted on Sphaerellopsis filum, the anamorphic stage of Eudarluca australis based on both AFLP and ITS sequences, and only limited variation between different isolates was detected (Bayon et al. 2006).

Concluding remarks

By blasting within GenBank, ITS sequences of E. caricis (= E. australis, strain MullMK, GB, access AY836374) are most comparable with species in Leptosphaeria and Phoma. Thus Eudarluca appears to be related to Leptosphaeriaceae pending further study.

Falciformispora K.D. Hyde, Mycol. Res. 96: 26 (1992). (Trematosphaeriaceae)

Generic description

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.

Anamorphs reported for genus: none.

Literature: Hyde 1992b; Raja and Shearer 2008.

Type species

Falciformispora lignatilis K.D. Hyde, Mycol. Res. 96: 27 (1992). (Fig. 32)

Fig. 32
figure 33

Falciformispora lignatilis (from BRIP 16972, holotype). a Section of a superficial ascoma. The peridium comprises two layers. b, c Squash mounts showing asci with wide pseudoparaphyses. The asci are cylindro-clavate with very short pedicels. d–f Hyaline multiseptate ascospores. Note the elongated appendage at the base (arrow head). Scale bars: a, b =100 μm, c = 50 μm, df = 10 μm

Ascomata 180–270 μm high × 250–340 μm diam., scattered to gregarious, erumpent and eventually superficial, depressed globose to ovoid, black, ostiolate, epapillate, coriaceous (Fig. 32a). Peridium up to 35 μm wide, comprising two cell types, outer layer composed of thick-walled cells of textura angularis, up to 8 μm diam., cell wall up to 5 μm thick, inner layer composed of hyaline compressed cells, cells 12 × 3 μm diam., cell wall 1–1.5 μm thick (Fig. 32a). Hamathecium long and cellular pseudoparaphyses, 2–3 μm broad, septate, embedded in mucilage. Asci 115130 × 23–31 μm, 8-spored, bitunicate, fissitunicate, broadly clavate to fusoid, with a short, thick pedicel, 8–15 μm long, with an ocular chamber (to 5 μm wide × 3 μm high) (Fig. 32b and c). Ascospores 4250 × 8–10 μm, 2–3 seriate, fusoid to somewhat clavate, hyaline, usually slightly curved, 6–8-septate, mostly 7-septate, slightly constricted at all septa, smooth-walled, surrounded by a thin mucilaginous sheath which is longer at the base (up to 20–30 μm) (Fig. 32d, e and f).

Anamorph: none reported.

Material examined: MEXICO, Nova Hispania, mangrove near Boca de Pascuales, saprobic on immersed intertidal mangrove wood, Mar. 1988, K.D. Hyde (BRIP 16972, holotype).

Notes

Morphology

Falciformispora was formally established by Hyde (1992b) as a monotypic genus and was assigned to Pleosporaceae by comparing with Setosphaeria, but Setosphaeria has the anamorphic stage of Exserohilum and is exclusively parasitic on Gramineae unlike Falciformispora. The setae on the ascomata of Setosphaeria could also serve as a distinguishing character from Falciformispora. Raja and Shearer (2008) also collected this species from freshwater in Florida. They considered that the species was more closely related to Chaetomastia than Setosphaeria, but that Falciformispora differed in having hyaline ascospores.

Phylogenetic study

Phylogenetic analyses in Schoch et al. (2009) and Suetrong et al. (2009) placed Falciformispora lignatilis in Trematosphaeriaceae in proximity to another marine species associated with mangroves, Halomassarina thalassiae.

Concluding remarks

Phylogenetic work confirmed that the saprobic habitat of Falciformispora is inconsistent with most other members of Pleosporaceae. The hyaline multi-septate ascospores with a mucilaginous sheath indicate affinities to Lophiostomataceae but this is not supported in DNA sequence comparisons. Carinispora is also similar and may be related.

Hadrospora Boise, Mem. N. Y. bot. Gdn 49: 310 (1989). (?Phaeosphaeriaceae)

Generic description

Habitat terrestrial (or freshwater?), saprobic. Ascomata small- to medium-sized, solitary, scattered, or in groups, immersed to nearly superficial, globose to subglobose, papillate. Peridium thin, comprising pseudoparenchymatous cells. Hamathecium dense, narrowly cellular, embedded in mucilage. Asci bitunicate, fissitunicate, oblong to ovoid, with a short pedicel. Ascospores ellipsoid to broadly fusoid with narrow ends, reddish brown, multi-septate, constricted at the primary septum.

Anamorphs reported for genus: Zalerion (Tanaka and Harada 2003a).

Literature: Boise 1984, 1989; Fisher and Webster 1992; Shearer and Crane 1971; Tanaka and Harada 2003a; Webster 1993.

Type species

Hadrospora fallax (Mouton) Boise, Mem. N. Y. bot. Gdn 49: 310 (1989). (Fig. 33)

Fig. 33
figure 34

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

Trematosphaeria fallax Mouton, Bull. Soc. R. Bot. Belg. 25: 155, (1886).

Ascomata 130–240 μm high × 200–330 μm diam., solitary, scattered or in groups, initially immersed, becoming erumpent to nearly superficial, with basal wall remaining immersed in host tissue, not easily removed from the substrate, globose or subglobose, roughened, papillate, coriaceous (Fig. 33a). Peridium 30–45 μm wide, comprising cells of pseudoparenchymatous, up to 12.5 × 9 μm diam. (Fig. 33b and c). Hamathecium of dense, narrowly cellular pseudoparaphyses, 1–2 μm broad, embedded in mucilage. Asci 150200 × 40–60 μm (\( \bar{x} = 171.5 \times 48\mu m \), n = 10), 8-spored, bitunicate, fissitunicate, oblong to ovoid, with a short pedicel, 10–24 μm long, with a ocular chamber (to 5 μm wide × 6 μm high) (Fig. 33d). Ascospores 5580 × 16–22 μm (\( \bar{x} = 67.1 \times 18.1\mu m \), n = 10), biseriate to 4-seriate, ellipsoid to broadly fusoid with narrow ends, reddish brown with paler end cells, 8-septate, constricted at the primary septum, smooth-walled (Fig. 33e, f, g, h and i).

Anamorph: Zalerion sp. (Tanaka and Harada 2003a).

Material examined: BELGIUM, Beaufays, on cut off, still hard wood. Oct. 1922, V. Mouton (BR, Capsa: K 7534, holotype). (Note: The specimen is not in good condition, only a few ascomata left).

Notes

Morphology

Boise (1989) formally established Hadrospora to accommodate Trematosphaeria fallax and T. clarkia (Sivan.) Boise, and Hadrospora fallax (syn. T. fallax) was selected as the generic type. Hadrospora is a widely distributed species that has been reported from Belgium, China, Italy, Japan, Switzerland and the United States (Boise 1989; Fisher and Webster 1992; Shearer and Crane 1971; Tanaka and Harada 2003a; Webster 1993). Hadrospora was temporarily assigned to Phaeosphaeriaceae based on its “small, thin-walled 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.

Phylogenetic study

None.

Concluding remarks

Hadrospora seems not closely related to Phaeosphaeriaceae.

Halotthia Kohlm., Nova Hedwigia 6: 9 (1963). (?Zopfiaceae)

Generic description

Habitat marine, saprobic. Ascomata large, solitary, gregarious or confluent, broadly conical to subglobose, flattened at the base, carbonaceous, immersed to erumpent, ostiolate, epapillate. Peridium plectenchymatous. Hamathecium of dense, long, cellular pseudoparaphyses, septate, branching. Asci 8-spored, bitunicate, cylindrical, with a short pedicel. Ascospores uniseriate, ellipsoidal, subcylindrical or obtuse-fusoid, dark brown, 1-septate, constricted at the septum.

Anamorphs reported for genus: none.

Literature: Kohlmeyer 1963; Suetrong et al. 2009.

Type species

Halotthia posidoniae (Durieu & Mont.) Kohlm., Nova Hedwigia 6: 9 (1963). (Fig. 34)

Fig. 34
figure 35

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

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 pseudoparaphyses, 1.5–2 μm broad, septate, branching. Asci 275290 × 25–35 μm, 8-spored, bitunicate, cylindrical, with a short pedicel (Fig. 34b, c and d). Ascospores 3760.5 × 16.5–26 μm, uniseriate, ellipsoidal, subcylindrical or obtuse-fusoid, dark brown, 1-septate, constricted at the septum (Fig. 34e, f, g and h) (adapted from Kohlmeyer and Kohlmeyer 1979).

Anamorph: none reported.

Material examined: ITALY, in rhizomes of Posidonia oceanica (Posidoniaceae), 1861, Caldesi (S, isotype of Sphaeria posidoniae)

Notes

Morphology

Halotthia was introduced to accommodate the marine fungus, H. posidoniae (as Sphaeria posidoniae), which is characterized by immersed to erumpent, large, carbonaceous ascomata, thick peridium, bitunicate, 8-spored, cylindrical asci, ellipsoidal, 1-septate, and dark brown ascospores (Kohlmeyer 1963). Morphologically, Halotthia is most comparable with Bicrouania maritima, but the conical ascomata with flattened base of H. posidoniae can be readily distinguished from B. maritima.

Phylogenetic study

Phylogenetically, Halotthia posidoniae, Pontoporeia biturbinata and Mauritiana rhizophorae form a robust clade, which may represent a potential family (Suetrong et al. 2009).

Concluding remarks

Currently the familial status of Halotthia is unresolved (Suetrong et al. 2009).

Helicascus Kohlm., Can. J. Bot. 47: 1471 (1969). (Morosphaeriaceae)

Generic description

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.

Anamorphs reported for genus: none.

Literature: Kohlmeyer 1969; Kohlmeyer and Kohlmeyer 1979; Suetrong et al. 2009.

Type species

Helicascus kanaloanus Kohlm., Can. J. Bot. 47: 1471 (1969). (Fig. 35)

Fig. 35
figure 36

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 thick-walled ascospores. Note the germ pore at the lower end of the ascospores. Scale bars: a = 0.5 mm, b–d = 20 μm

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 250335 × 25–30 μm, 8-spored, subcylindrical, finally oblong-clavate (400–480 μm long), with a short pedicel, bitunicate, thick-walled, physoclastic, apically multi-layered and annulate, ectoascus forming a third, thin permeable outer layer around the base, endoascus swelling in water and becoming coiled at maturity, finally stretching and pushing the ascus into the ostiolar canal (Fig. 35b). Ascospores 36.5–48.5 × 18–22.5 μm, uniseriate, obovoid, brown, 1-septate, at each end with a germ pore, surrounded with dissolving sheath, 2.7–5.4 μm thick, with funnel-shaped, apical indentations (Fig. 35c and d) (adapted from Kohlmeyer and Kohlmeyer 1979).

Anamorph: none reported.

Material examined: USA, Hawaii, Oahu, Kaneohe Bay, Heeia Swamp, on Rhizophora mangle, 4 Jun. 1968 (Herb. J. Kohlmeyer No. 2566, holotype; No. 2565, 2567, paratype).

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.

Phylogenetic study

Multigene phylogenetic analysis indicated that both Helicascus kanaloanus and H. nypae K.D. Hyde nested within Morosphaeriaceae (Suetrong et al. 2009).

Concluding remarks

Helicascus is a well defined marine genus.

Herpotrichia Fuckel, Fungi rhenani exsic.: no. 2171 (1868). (Melanommataceae)

Generic description

Habitat terrestrial, parasitic, hyperparasitic or saprobic. Ascomata medium-sized, immersed, erumpent to nearly superficial, scattered to gregarious, globose to subglobose with a broad pore. Peridium composed of pseudoparenchymatous cells. Hamathecium of dense, long pseudoparaphyses, embedded in mucilage, septate, branching. Asci cylindrical to cylindro-clavate, with a furcated pedicel. Ascospores fusoid, ellipsoid or oblong with broadly to narrowly round ends, 1-septate, constricted at the septum, uni- to biseriate.

Anamorphs reported for genus: Pyrenochaeta or Pyrenochaeta-like (Sivanesan 1984).

Literature: von Arx and Müller 1975; Barr 1984; Cannon 1982; Freyer and van der Aa 1975; Mugambi and Huhndorf 2009b; Samuels 1973; Samuels and Müller 1978; Sivanesan 1971, 1984.

Type species

Herpotrichia rubi Fuckel, Fungi rhenani exsic 2171. (1868). (Fig. 36)

Fig. 36
figure 37

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. gh 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, eh = 10 μm

Ascomata 220–430 μm high × 240–390(-530) μm diam., scattered to gregarious, immersed to erumpent, rarely superficial, globose to subglobose, wall black, coriaceous, apex with a small sometimes inconspicuous papilla, usually with a pore, lacking periphyses (Fig. 36a and b). Peridium 32–45 μm wide at the sides, up to 60 μm wide at the apex, basal wall thinner, all walls comprising cells of textura angularis, cells 2.5–4 μm diam., cell wall 2–4(−7) μm thick, exterior cells more thick-walled and pigmented, inner cells thin-walled and less pigmented, comprising thin-walled cells up to 9 μm diam., apex cells smaller and walls thicker (Fig. 36b and c). Hamathecium of dense, long pseudoparaphyses, 2–3 μm broad, embedded in mucilage, septate, branching (Fig. 36e). Asci 105150 × 12.5–15 μm (\( \bar{x} = 137.5 \times 13.8\mu m \), 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 (\( \bar{x} = 20.5 \times 7.3\mu m \), 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) designated H. rhenana as the lectotype species with H. rubi as a synonym. This proposal was followed by Müller and von Arx (1962) and Sivanesan (1971). Herpotrichia rubi was later assigned as the generic type (Holm 1979) as it was found to be validly published 2 years earlier than H. rhenana, thus having priority (Cannon 1982). However, Cannon (1982) reported that Sphaeria herpotrichoides Fuckel (1864, cited as a synonym of H. rhenana) was the earliest name. Thus he made a new combination as H. herpotrichoides (Fuckel) P.F. Cannon and cited H. rubi as the synonym. Herpotrichia rubi is maintained as the type of the genus (Holm 1979; Cannon 1982), but the current name is H. herpotrichoides.

Herpotrichia is a morphologically well studied genus (Barr 1984; Bose 1961; Müller and von Arx 1962; Pirozynski 1972; Samuels and Müller 1978; Sivanesan 1971, 1984), and Herpotrichia sensu lato is characterized by having subglobose, pyriform to obpyriform ascomata and a peridium of textura angularis or comprising thick-walled 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).

Phylogenetic study

Herpotrichia diffusa (Schwein.) Ellis & Everh., H. juniperi (Duby) Petr., H. herpotrichoides and H. macrotricha have been shown to have phylogenetic affinity with the generic types of Byssosphaeria schiedermayeriana, Melanomma pulvis-pyrius and Pleomassaria siparia, which had been assigned under Melanommataceae (Kruys et al. 2006; Mugambi and Huhndorf 2009b; Schoch et al. 2006, 2009; Zhang et al. 2009a). In this study, Pleomassaria siparia together with its closely related species of Prosthemium is kept in a separate family, viz Pleomassariaceae.

Concluding remarks

Even species under Herpotrichia sensu stricto (according to Sivanesan 1984) have diverse hosts (such as gymnosperms (H. coulteri (Peck) S.K. Bose and H. parasitica (R. Hartig) Rostr.) and angiosperms (H. diffusa and H. villosa Samuels & E. Müll.)) or substrates (like dead or living leaves, bark or decorticated wood) (Sivanesan 1984). Species of Herpotrichia sensu stricto are also reported from various locations such as Europe, Asia or America, and they have various life styles, e.g. parasitic, hyperparasitic or saprobic (Sivanesan 1984). Additional factors (like hosts or locations) may need to be considered in order to get a natural concept for Herpotrichia.

Immotthia M.E. Barr, Mycotaxon 29: 504 (1987). (Teichosporaceae)

Generic description

Habitat terrestrial, hyperparasitic. Ascomata gregarious, globose, superficial, ostiolate, periphysate. Hamathecium of cellular pseudoparaphyses. Asci 8-spored, bitunicate, cylindrical, with a short pedicel. Ascospores 1-seriate, ellipsoidal, brown to reddish brown, 1-septate, constricted at the septum, smooth.

Anamorphs reported for genus: none.

Literature: Barr 1987a, 2002; Wang et al. 2004.

Type species

Immotthia hypoxylon (Ellis & Everh.) M.E. Barr, Mycotaxon 29: 504 (1987). (Fig. 37)

Fig. 37
figure 38

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

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 6082 × 7–9 μm, 8-spored, bitunicate, cylindrical, with a short pedicel (Fig. 37b, c and d). Ascospores 1013 × 4.4–5.4 μm, 1-seriate, ellipsoidal, brown to reddish brown, 1-septate, constricted at the septum, smooth (Fig. 37f, g and h) (adapted from Wang et al. 2004).

Anamorph: none reported.

Material examined: USA, Louisiana, Pointe a la Hache, on decaying wood, a branch of Carya oliviformis (Juglandaceae) lying on the ground in grass (parasitic on 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; Wang et al. 2004).

Phylogenetic study

None.

Concluding remarks

It seems that those Amphisphaeria species with bitunicate asci should be assigned to Pleosporales. Morphologically, Immotthia is somewhat comparable with Herpotrichia.

Isthmosporella Shearer & J.L. Crane, Mycologia 91: 141 (1999). (Pleosporales, genera incertae sedis)

Generic description

Habitat freshwater, saprobic. Ascomata small- to medium-sized, 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.

Anamorphs reported for genus: none.

Literature: Shearer and Crane 1999.

Type species

Isthmosporella pulchra Shearer & J.L. Crane, Mycologia 91: 142 (1999). (Fig. 38)

Fig. 38
figure 39

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

Ascomata 240–330 μm diam., scattered on decorticated wood, immersed, erumpent to superficial, globose, black, papillate, papilla short, cylindrical, 60 μm long × 55 μm wide, ostiolate, periphysate, membranous (Fig. 38a). Peridium 2-layered, outer 3–4 cell layers composed of brown, pseudoparenchymatic, fusoid-cylindric cells, 2–6.5 μm long; inner layer composed of 5–7 rows of fusoid, subhyaline to pale brown compressed cells, 11–20 × 2–3.5 μm diam. (Fig. 38a and b). Hamathecium of rare, broad, septate, interascal pseudoparaphyses (Fig. 38f). Asci (95-)135–160(−175) × (25-)30–45(−60) μm, 8-spored, bitunicate, fissitunicate, oblong to clavate, with a short pedicel, ocular chamber not observed (Fig. 38c, d and e). Ascospores 80–105(−110) × (7-)8–10 μm, 3–4-seriate, cylindrical to fusoid, isthmoid at centre, sometimes bent at isthmus and becoming u- or v- shaped, end cells tapering, 12–17-phragmoseptate, constricted at septa, isthmus 1-septate, 2–5.5 × 2–4.5 μm diam., hyaline, frequently fragmenting to form partspores; filled with lipid droplets that merge to form large guttules; surrounded by a gelatinous sheath with a dense region near the isthmus, sheath greatly enlarging in water (Fig. 38g, h, i and j).

Anamorph: none reported.

Colonies on yeast soluble starch agar containing balsa wood sticks effuse, white. Hyphae hyaline, septate.

Material examined: USA, New York, Adirondack Park. Piercefield. Tupper Lake at public boat launch from Rt. 30, UTM Zone 18, 539840 mE, 4892100mN; 44°10″59″N, 80°31′6″W, on submerged, decorticated wood, 7 Jul. 1994, J.L. Crane & C.A. Shearer A-254-1 (ILLS 53086, holotype).

Notes

Morphology

Isthmosporella was described as a freshwater genus typified by I. pulchra, and is characterized by globose, pseudoparenchymatous ascomata, sparse, septate pseudoparaphyses, fissitunicate asci and hyaline, cylindrical to fusoid, phragmoseptate, isthmoid ascospores surrounded with a gelatinous sheath (Shearer and Crane 1999). Based on the morphological characters, i.e. small, globose ascomata, peridium with small pseudoparenchymatous cells and sparse pseudoparaphyses, Isthmosporella was assigned to the Phaeosphaeriaceae (Shearer and Crane 1999). The aquatic habitat of Isthmosporella, however, disagree with the Phaeosphaeriaceae. Isthmosporella seems less likely to belong to Pleosporineae.

Phylogenetic study

None.

Concluding remarks

Molecular phylogenetic studies should be conducted to explore its familial placement within Pleosporales.

Kalmusia Niessl, Verh. nat. Ver. Brünn 10: 204 ( 1872). (Montagnulaceae)

Generic description

Habitat terrestrial, saprobic. Ascomata small- to medium-sized, solitary, scattered or in small groups, immersed to 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).

Literature: Barr 1987b, 1990a, 1992a; Lindau 1897; von Niessl 1872.

Type species

Kalmusia ebuli Niessl, Verh. nat. Ver. Brünn 10: 204 (1872). (Fig. 39)

Fig. 39
figure 40

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

Ascomata 290–360 μm high × 300–520 μm diam., solitary, scattered, or in small groups, immersed to erumpent, globose or subglobose, coriaceous, wall black, with or without papilla, ostiolate (Fig. 39a). Papilla small, up to 100 μm high, with small ostioles (Fig. 39a). Peridium 15–40 μm wide, comprising one cell type of small, pigmented, thick-walled cells of textura prismatica to textura angularis, cells ca. 5 × 3 μm diam., cell wall 2–3 μm thick (Fig. 39b and c). Hamathecium of dense, delicate pseudoparaphyses, 1–1.5 μm broad, septate, branching and anastomosing between and above asci, embedded in mucilage. Asci 75125 × 10–15 μm (\( \bar{x} = 90.5 \times 12\mu m \), 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. 2 μm wide × 1 μm high) (Fig. 39d, e and f). Ascospores 1518 × 5.5–6.5 μm (\( \bar{x} = 16.3 \times 5.8\mu m \), n = 10), biseriate, narrowly ovoid to clavate, pale brown, 3-distoseptate, without constriction, smooth-walled (Fig. 39g, h and i).

Anamorph: none reported.

Material examined: BELGIUM, Dolembreux, on branchlets and pieces of stumps of Sarothamnus scoparius from woodland, Oct. 1922, V. Mouton (BR 101525–63, holotype).

Notes

Morphology

Kalmusia was formally established by von Niessl (1872), and is mainly characterized as “immersed, sphaeroid ascoma with central, stout papilla, surrounded by hyphae in the substrate, stipitate asci with septate pseudoparaphyses, and brown, 3-septate, inequilateral ascospores” (Barr 1992a).

The most morphologically comparable genus to Kalmusia is Thyridaria, which had been treated as a subgenus under Kalmusia (Lindau 1897), and was subsequently transferred to Platystomaceae in Melanommatales (Barr 1987b, 1990a). Compared to Thyridaria, Kalmusia has sphaeroid ascomata, a peridium of small pseudoparenchymatous cells, basal asci and very thin pseudoparaphyses, thus it was assigned to Phaeosphaeriaceae of the Pleosporales by Barr (1990a), and the genus is utilized to accommodate both K. ebuli and K. clivensis (Berk. & Broome) M.E. Barr, as well as closely related species, i.e. K. utahensis (Ellis & Everh.) Huhndorf & M.E. Barr and K. coniothyrium (Fuckel) Huhndorf (Barr 1992a). But this proposal is questionable, as the clavate, distoseptate ascospores, as well as the clavate asci with very long pedicels are uncommon in Phaeosphaeriaceae, and most recent phylogenetic study indicated that some species of Kalmusia reside outside of Phaeosphaeriaceae (Zhang et al. 2009a).

Phylogenetic study

Both Kalmusia scabrispora Teng Kaz. Tanaka, Y. Harada & M.E. Barr and K. brevispora (Nagas. & Y. Otani) Yin. Zhang, Kaz. Tanaka & C.L. Schoch reside in the clade of Montagnulaceae (Zhang et al. 2009a). Familial placement of Kalmusia can only be verified after the DNA sequences of the generic type (K. ebuli) are obtained.

Concluding remarks

Kalmusia is distinct amongst the Pleosporales as it has pale brown ascospores with indistinct distosepta and clavate asci with long pedicels. Although both K. scabrispora and K. brevispora reside in the clade of Montagnulaceae, they both lack the distoseptate ascospores that are possessed by the generic type (K. ebuli). Thus, the familial placement of Kalmusia is still undetermined.

Karstenula Speg., Decades Mycologicae Italicae ad no. 94 (in sched.) (1879). (Montagnulaceae)

Generic description

Habitat terrestrial, saprobic. Ascomata rarely small-, usually medium-sized, immersed usually under thin clypeus, scattered to gregarious, with flattened top and rounded pore-like ostiole, coriaceous. Peridium 2-layered, outer layer composed of reddish brown to dark brown small cells, inner layer of pale compressed cells. Hamathecium of dense, cellular pseudoparaphyses. Asci cylindrical to cylindro-clavate with short furcate pedicel. Ascospores muriform, ellipsoid to fusoid, reddish brown to dark brown.

Anamorphs reported for the genus: Microdiplodia (Constantinescu 1993).

Literature: Barr 1990a; Eriksson and Hawksworth 1991; Kodsueb et al. 2006a; Munk 1957; Zhang et al. 2009a.

Type species

Karstenula rhodostoma (Alb. & Schwein.) Speg., Decades Mycologicae Italicae no. 94. (1879). (Fig. 40)

Fig. 40
figure 41

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

Sphaeria rhodostoma Alb. & Schwein., Consp. fung. (Leipzig): 43 (1805).

Ascomata 250–430 μm high × 450–650 μm diam., scattered or gregarious, immersed in the subiculum which sometimes sloths off, globose or subglobose, black, flattened top often white or reddish and sometimes slightly protruding out of the substrate surface, usually with a wide opening ostiole after removing the cover, coriaceous (Fig. 40a and b). Peridium 30–40 μm wide, comprising two cell types, outer region 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 150210 × 12.5–15 μm (\( \bar{x} = 182 \times 13.1\mu m \), 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 2026 × 7.5–10 μm (\( \bar{x} = 22.4 \times 8\mu m \), 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).

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 ascospores were brown with transverse septa and sparse longitudinal septa.

The ascomata of this species are similar to those found in Byssosphaeria and Herpotrichia, especially in the paler area around the ostiole and even in peridial structure and development under a subiculum. The numerous wide cellular pseudoparaphyses and cylindrical asci (in Herpotrichia) are also similar. The main difference of Karstenula from other two genera are the 3-septate ascospores with rare longitudinal septa (1-septate in Byssosphaeria and Herpotrichia).

Phylogenetic study

Karstenula forms a robust phylogenetic clade with Phaeodothis winteri (Niessl) Aptroot, Didymocrea sadasivanii, Bimuria novae-zelandiae, Montagnula opulenta, Curreya pityophila (J.C. Schmidt & Kunze) Arx & E. Müll. and some species of Letendraea and Paraphaeosphaeria (Kodsueb et al. 2006a; Zhang et al. 2009a). Consequently, Karstenula might be included in Montagnulaceae.

Concluding remarks

The description of the type of Karstenula here clearly excludes it from Melanommataceae as it has wide pseudoparaphyses. But its Montagnulaceae status can only be confirmed by more phylogenetic work including sequencing the generic type of Karstenula (K. rhodostoma).

Katumotoa Kaz. Tanaka & Y. Harada, Mycoscience 46: 313 (2005). (Lentitheciaceae)

Generic description

Habitat terrestrial or freshwater, saprobic. Ascomata small- to medium-sized, scattered or in small groups, immersed to erumpent, with a central protruding hairy papilla, subglobose. Peridium thin, comprising several layers of thin-walled compressed cells. Hamathecium of dense, cellular, filliform, embedded in mucilage, branching and anastomosing. Asci 8-spored, bitunicate, fissitunicate, clavate with short furcate pedicels. Ascospores apiosporous and hyaline when young, becoming 2-septate with reddish brown echinate central cell at maturity, with long gelatinous terminal appendages.

Anamorphs reported for genus: none.

Literature: Tanaka and Harada 2005b; Tanaka et al. 2009; Zhang et al. 2009a.

Type species

Katumotoa bambusicola Kaz. Tanaka & Y. Harada, Mycoscience 46: 313 (2005). (Fig. 41)

Fig. 41
figure 42

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

Some information for the following description is from Tanaka and Harada (2005).

Ascomata 240–330 μm high × 260–420 μm diam., scattered or in small groups, immersed, becoming erumpent, with a slightly protruding papilla covered with brown hyphae, subglobose (Fig. 41a). Peridium 13–30 μm thick, composed of a few layers of lightly pigmented, depressed cells. Hamathecium of dense, long cellular pseudoparaphyses, 1.5–3 μm broad, embedded in mucilage, branching and anastomosing. Asci 110160 × 17.5–24 μm (\( \bar{x} = 139 \times 21\mu m \), n = 10), 8-spored, bitunicate, fissitunicate, cylindro-clavate with a short furcate pedicel which is up to 25 μm long (Fig. 41b and d). Ascospores 39–50(−57) × 7–10 μm (\( \bar{x} = 45.8 \times 8.2\mu m \), n = 10), biseriate, fusoid to narrowly fusoid with acute ends, usually curved, apiosporus and hyaline when young, constricted at the primary septum, the upper cell longer and broader than the lower one, smooth, surrounded by a bipolar sheath which is up to 15 μm long, best seen in India ink, senescent ascospores yellowish brown, 2–4-septate (Fig. 41c).

Anamorph: none reported.

Material examined: JAPAN, Mt. Iwate, near Yakebashiri, Hirakasa, Nishine, Iwate, on culms of Oryza sativa L., 19 Oct. 2003, K. Tanaka (HHUF 28663, holotype).

Notes

Morphology

Katumotoa was formally established by Tanaka and Harada (2005b) to accommodate the monotypic species, K. bambusicola, which is characterized by immersed ascomata with a thin peridium comprising thin-walled compressed cells, cellular pseudoparaphyses, cylindro-clavate and fissitunicate asci and fusoid ascospores with an elongated bipolar mucilaginous sheath. Based on its immersed ascomata, psuedoparenchymatous peridium cells and cellular pseudoparaphyses, Katumotoa was assigned to Phaeosphaeriaceae (Tanaka and Harada 2005b; Tanaka et al. 2009), but this classification has been shown to be incorrect in subsequent phylogenetic studies (Tanaka et al. 2009; Zhang et al. 2009a).

Phylogenetic study

Phylogenetic analysis based on five genes (LSU, SSU, RPB1, RPB2 and EF1) indicates that Katumotoa bambusicola resides in Lentitheciaceae, and this receives high bootstrap support (Zhang et al. 2009a). In particular, K. bambusicola forms a robust clade with Ophiosphaerella sasicola (Nagas. & Y. Otani) Shoemaker & C.E. Babc., which has filliform ascospores (Shoemaker and Babcock 1989b).

Concluding remarks

The hyaline, apiosporous ascospores which become 2–4-celled with central reddish brown cells and large unraveling appendages are the most striking features of this species and readily distinguish it from other pleosporalean taxa. Both Katumotoa bambusicola and Ophiosphaerella sasicola are associated with bambusicolous hosts, which might indicate that host spectrum in this case, has greater phylogenetic significance than some morphological characters (Zhang et al. 2009a).

Keissleriella Höhn., Sber. Akad. Wiss. Wien, Math.-naturw. Kl., Abt. 1 128: 582 (1919). (Lentitheciaceae)

Generic description

Habitat terrestrial or freshwater, saprobic. Ascomata small- to medium-sized, immersed, erumpent to nearly superficial, globose, papillate, ostiolate. Papilla covered by dark setae or small blackened cells. Peridium thick, composed of cells of pseudoparenchymatous and inner layer composed of pale cells. Hamathecium of dense, long pseudoparaphyses, rarely septate, anastomosing and branching. Asci 4- or 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)

Fig. 42
figure 43

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

Pyrenochaeta aesculi Höhn., Ber. dt. bot. Ges. 35: 249 (1917).

Ascomata ca. 250 μm high × 450 μm diam., gregarious, immersed to erumpent, globose or subglobose, with a small black papilla, ca. 75 μm high and 110 μm broad, with short black external setae (Fig. 42a). Peridium ca. 25–40 μm wide laterally, up to 70 μm near the apex, thinner at the base, comprising two types of cells which merge in the middle; outer cells composed of small heavily pigmented thick-walled cells, cells ca. 4 μm diam., cell wall up to 4 μm thick, and thick near the apex and thinner laterally and absent in the immersed part of the ascoma, inner cells less pigmented, comprising lightly pigmented to hyaline cells, 5–7 μm thick (Fig. 42a). Hamathecium of dense, long pseudoparaphyses, 0.8–1.2 μm broad, rarely septate, anastomosing and branching, thicker near the base, ca. 2 μm, constricted near the septum (Fig. 42b). Asci 80120 × 6–11 μm (\( \bar{x} = 101 \times 8.5\mu m \), 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 1318 × 4–5.5 μm (\( \bar{x} = 14.5 \times 4.8\mu m \), n = 10), obliquely uniseriate and partially overlapping, fusoid with narrowly rounded ends, hyaline, 1-septate, constricted at the septum, smooth (Fig. 42c and d).

Anamorph: none reported.

Material examined: AUSTRIA, Brentenmaistal in the Viennese forest, Aesculus hippocastanum L., 1916, Höhnel (FH, holotype of Otthiella aesculi). (Note: only two slides; setae cannot be seen from the slides but could be seen from the drawings on the cover).

Notes

Morphology

Keissleriella is characterized by ascomata with setae in and over the papilla, asci are cylindrical and ascospores are hyaline, 1-septate. Based on the morphological characters, K. aesculi was regarded as conspecific with K. sambucina; as an earlier epithet, K. sambusina typifies the genus (see comments by Barr 1990a). Munk (1957) placed Trichometasphaeria and Keissleriella in Massarinaceae, and distinguished them by their substrates (Trichometasphaeria occurs on herbaceous plants and Keissleriella on woody substrates). Bose (1961) combined Trichometasphaeria under Keissleriella, which was followed by some workers (von Arx and Müller 1975; Dennis 1978; Eriksson 1967a; Luttrell 1973). Barr (1990a), however, maintained these as distinct genera based on the differences of peridium structure and pseudoparaphyses.

Phylogenetic study

The phylogeny of Keissleriella is poorly studied. Limited phylogenetic information indicates that K. cladophila forms a robust clade with other species of Lentitheciaceae (Zhang et al. 2009a).

Concluding remarks

The presence of black setae on the surface of papilla is a striking character of Keissleriella, but phylogenetic significance of setae is undetermined yet.

Lentithecium K.D. Hyde, J. Fourn. & Yin. Zhang, Fungal Divers. 38: 234 (2009). (Lentitheciaceae)

= Tingoldiago K. Hirayama & Kaz. Tanaka, Mycologia 102: 740 (2010) syn. nov.

Generic description

Habitat freshwater, saprobic. Ascomata small, scattered or gregarious, immersed, slightly erumpent, depressed spherical to lenticular, ostiolate, papillate or epapillate. Peridium thin. Hamathecium of cellular pseudoparaphyses. Asci 8-ascospored, bitunicate, fissitunicate, clavate, short-stipitate. Ascospores broadly fusoid with broadly rounded ends, 1-septate, constricted, hyaline, usually with sheath.

Anamorphs reported for genus: none.

Literature: Shearer et al. 2009; Zhang et al. 2009a, b.

Type species

Lentithecium fluviatile (Aptroot & Van Ryck.) K.D. Hyde, J. Fourn. & Yin. Zhang, Fungal Divers. 38: 234 (2009). (Fig. 43)

Fig. 43
figure 44

Lentithecium fluviatile (from IFRD 2039). a Erumpent ascomata scattering on the host surface. b Habitat section of the immersed ascomata. c, d Section of an ascoma and a partical peridium. Note the peridium cells of textura angularis. e Clavate 8-spored ascus with a short pedicel. f, g Hyaline, 1-septate broadly fusoid ascospores. Scale bars: a, b = 0.5 mm, c = 100 μm, d = 50 μm, e–g = 20 μm

Massarina fluviatilis Aptroot & Van Ryck., Nova Hedwigia 73: 162 (2001).

Ascomata 230–260 μm high × 280–325 μm diam., scattered or gregarious, immersed, slightly erumpent, subglobose to depressed spherical, under a small black pseudostroma originating from the apical part of the peridium, apex slightly papillate, ostiole rounded, 60–70 μm diam. (Fig. 43a and b). Peridium 15–20 μm thick at sides and at base, comprising 4–5 layers of angular cells more thick-walled outwards, 50–55 μm 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).

Anamorph: none reported.

Material examined: FRANCE, Haute Garonne: Avignonet, Lac de Rosel, artificial lake, on bark and wood of a submerged branch Populus sp., 23 Nov. 2006, leg. Michel Delpont, det. Jacques Fournier (IFRD 2039, holotype).

Notes

Morphology

Lentithecium was introduced to accommodate some freshwater fungi previous assigned under Massarina, such as M. arundinacea (Sowerby) Leuchtm. and M. fluviatilis (Zhang et al. 2009a). It is characterized by its immersed and lenticular ascomata, thin peridium which is almost equal in thickness, short pedicellate asci and fusoid or filliform, hyaline or rarely lightly pigmented, 1- to multi-septate ascospores (Zhang et al. 2009b). Lentitheciaceae was introduced to accommodate Lentithecium and some other related taxa (Zhang et al. 2009a).

Phylogenetic study

The clade of Lentitheciaceae comprises the generic type Lentithecium fluviatile, as well as L. arundinaceum (Sowerby) K.D. Hyde, J. Fourn. & Yin. Zhang, Stagonospora macropycnidia, Wettsteinina lacustris (Fuckel) Shoemaker & C.E. Babc., Keissleriella cladophila, and the bambusicolous species Katumotoa bambusicola and Ophiosphaerella sasicola, which receive high bootstrap support (Zhang et al. 2009a).

Concluding remarks

Tingoldiago graminicola K. Hirayama & Kaz. Tanaka form a robust clade with species of Lentithecium (Shearer et al. 2009). Tingoldiago has lenticular immersed to erumpent ascomata, numerous and septate pseudoparaphyses, cylindro-clavate asci and hyaline, 1-septate ascospores with sheath. All of these characters fit Lentithecium well. We treat Tingoldiago as a synonym of Lentithecium.

Leptosphaeria Ces. & De Not., Comm. Soc. crittog. Ital. 1: 234 (1863). (Leptosphaeriaceae)

Generic description

Habitat terrestrial, saprobic or parasitic. Ascomata small- to medium-sized, solitary, scattered or in small groups, erumpent to superficial, subglobose, broadly or narrowly conical, papillate, ostiolate. Peridium thick, comprising layers of cells of textura angularis. Hamathecium of dense cellular pseudoparaphyses, embedded in mucilage, anastomosing and branching. Asci 8-spored, bitunicate, fissitunicate unknown, cylindrical with a furcate pedicel and a large ocular chamber. Ascospores fusoid or narrowly fusoid, brown or reddish brown, 3-septate, constricted at each septum.

Anamorphs reported for genus: Coniothyrium and Phoma (Hyde et al. 2011; Sivanesan 1984).

Literature: von Arx and Müller 1975; Barr 1987a, b; Cesati and de Notaris 1863; Crane and Shearer 1991; Dong et al. 1998; Eriksson 1967a; Eriksson and Hawksworth 1986, 1991; de Greuter et al. 1988; Hedjaroude 1969; von Höhnel 1907; Holm 1957, 1975; Huhndorf et al. 1990; Luttrell 1973; Müller 1950; Munk 1957; Saccardo 1878b, 1883, 1891, 1895; Schoch et al. 2009; Shearer 1993; Shearer et al. 1990; Shoemaker 1984a; Sivanesan 1984; Zhang et al. 2009a.

Type species

Leptosphaeria doliolum Ces. & De Not., Comm. Soc. crittog. Ital. 1: 234 (1863). (Fig. 44)

Fig. 44
figure 45

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

≡ Sphaeria doliolum Pers., Icon. Desc. Fung. Min. Cognit. (Leipzig) 2: 39 (1800).

Ascomata 340–450 μm high × 380–500 μm diam., solitary, scattered or in small groups, superficial, subglobose, broadly or narrowly conical, with a flattened base on the host surface, black, usually with 2–4 ring-like ridges surrounding the ascomata surface, apex with a conical, usually shiny papilla (Fig. 44a). Peridium 85–110 μm wide at sides, thinner at the apex, comprising two types of cells, outer layer composed of small thick-walled cells of textura angularis, cells <2 μm diam., cell wall up to 8 μm thick, surface heavily pigmented and inner lightly pigmented, apex cells smaller, walls thicker, and cells more heavily pigmented, inner layer composed of subhyaline relatively thin-walled cells of textura angularis, 3–6 μm diam., wall up to 5 μm, cells near the base larger and wall thinner and paler (Fig. 44b). Hamathecium of dense, long cellular pseudoparaphyses, 1.5–3 μm broad, embedded in mucilage, anastomosing and branching. Asci 110150 × 7–9(−10) μm (\( \bar{x} = 130.6 \times 8.5\mu m \), n = 10), 8-spored, bitunicate, fissitunicate unknown, cylindrical, furcate pedicel which is usually less than 25 μm long, with a large ocular chamber (Fig. 44c, d and e). Ascospores 2531 × 4.5–6 μm (\( \bar{x} = 27.7 \times 5.3\mu m \), n = 10), uniseriate and somewhat partially overlapping, narrowly fusoid with sharp to narrowly rounded ends, reddish brown, 3-septate, constricted at each septum, smooth (Fig. 44f).

Anamorph: Phoma hoehnelii (Sivanesan 1984).

Material examined: Herb., Persoon 910270–650 (L, lectotype).

Notes

Morphology

Leptosphaeria was first established by Cesati and de Notaris (1863) with 26 species included; L. doliolum (Pers.:Fr.) Ces. & De Not. was subsequently selected as the lectotype species (de Greuter et al. 1988; Holm 1975; Shearer et al. 1990). Leptosphaeria was originally defined based mainly on the characters of ascospores being ellipsoid or fusoid, one to many septa, hyaline to dark brown. These few common characters meant that Leptosphaeria comprised many species, and some of them should be assigned to either Euascomycetes or Loculoascomycetes (Crane and Shearer 1991). Leptosphaeria had been divided based on host and habitat (Saccardo 1878b, 1891, 1895) as well as the pseudothecium (glabrous, hairy, setose) and ascospore septation (see comments by Crane and Shearer 1991). von Höhnel (1907) used centrum structure in the classification of Leptosphaeria, and divided Leptosphaeria into three genera, viz. Leptosphaeria, Scleropleella and Nodulosphaeria. Müller (1950) subdivided Leptosphaeria into four sections based on pseudothecial and centrum structure as well as ascospore characters. This classification was modified by Munk (1957), who named these four sections as section I (Eu-Leptosphaeria), section II (Para-Leptosphaeria), section III (Scleropleella) and section IV (Nodulosphaeria). Holm (1957) used a relatively narrow concept for Leptosphaeria, which included species closely related to the generic type, L. doliolum. This viewpoint was accepted by some workers (Eriksson 1967a; Hedjaroude 1969; Shoemaker 1984a). Nevertheless, it still seems a heterogeneous group of fungi (see comments by Crane and Shearer 1991). Its position among the Loculoascomycetes is also debated. It has been placed in the Pleosporaceae (von Arx and Müller 1975; Luttrell 1973; Sivanesan 1984) or Leptosphaeriaceae (Barr 1987a, b; Eriksson and Hawksworth 1991) or Phaeosphaeriaceae (Eriksson and Hawksworth 1986).

Phylogenetic study

Molecular phylogenetic analysis based on multigenes indicated that species of Leptosphaeria (including the generic type L. doliolum) and Neophaeosphaeria form a paraphyletic clade with moderate bootstrap support (Dong et al. 1998; Schoch et al. 2009; Zhang et al. 2009a), which is sister to other families of Pleosporales (Zhang et al. 2009a). Thus the familial rank of the Leptosphaeriaceae could be temporarily verified, but further molecular phylogenetic study is needed in which more related taxa should be included.

Concluding remarks

Morphologically, Leptosphaeria is mostly comparable with Amarenomyces, Bricookea, Diapleella, Entodesmium, Melanomma, Nodulosphaeria, Paraphaeosphaeria, Passeriniella, Phaeosphaeria and Trematosphaeria. While it prefers non-woody parts of dicotyledonous hosts, its cylindrical ascus with short pedicel and smooth, fusoid and multi-septate ascospores make it readily distinguishable from all other genera (Shoemaker 1984a).

Leptosphaerulina McAlpine, Fungus diseases of stone-fruit trees in Australia and their treatment: 103 (1902). (Didymellaceae)

Generic description

Habitat terrestrial, parasitic or saprobic. Ascomata small, scattered, immersed, globose to subglobose, with a small, slightly protruding papilla, ostiolate. Peridium thin. Hamathecium of rare or decomposing cellular pseudoparaphyses. Asci bitunicate, obpyriform. Ascospores broadly clavate or cylindrical, hyaline, turning pale brown when old, asymmetrical, multi-septate, smooth-walled.

Anamorphs reported for genus: Pithoascus and Pithomyces (Hyde et al. 2011).

Literature: Barr 1972; Chlebicki 2002; Crivelli 1983; Kodsueb et al. 2006a; Zhang et al. 2009a.

Type species

Leptosphaerulina australis McAlpine, Fungus diseases of stone-fruit trees in Australia and their treatment: 103 (1902). (Fig. 45)

Fig. 45
figure 46

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

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 3853 × 55–75 μm (\( \bar{x} = 67.5 \times 43.3\mu m \), 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 (\( \bar{x} = 36.5 \times 13\mu m \), 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.

Material examined: USA, Kansas, Kansas State College, on Poa pratensis L. Grass plots, 2 Jul. 1953, leg. T. Rogerson, det. L.E. Wehmeyer (NY, C.T. Rogerson 3836).

Notes

Morphology

Leptosphaerulina, introduced by McAlpine (1902), is characterized by small immersed ascomata, obpyriform asci with a large ocular chamber and apical ring as well as muriformly septate ascospores which may be hyaline or pigmented. Species of Leptosphaerulina may occur on monocotyledons or dicotyledons. Leptosphaerulina is most comparable with Pleospora, and the only difference between them is that Leptosphaerulina has smaller ascomata and hyaline ascospores that only become pigmented after discharge, whereas the ascospores of Pleospora become brown within the asci. Currently, about 60 names are accepted in this genus, and some even reported from marine environments, e.g. L. mangrovei (Inderbitzin et al. 2000).

Phylogenetic study

Based on multigene phylogenetic analysis, two putative strains of Leptosphaerulina australis, the generic type of Leptosphaerulina, from Switzerland (CBS 311.51) and Indonesia (CBS 317.83) resided within Didymellaceae (de Gruyter et al. 2009; Zhang et al. 2009a).

Concluding remarks

Because of its morphological confusion with Pleospora and the diversity of habitats within the genus, Leptosphaerulina sensu lato is likely to be polyphyletic. Fresh collections of this species are needed from Australia to epitypify this taxon and define the genus in a strict sense. The specimen described here is a collection from USA and therefore may not represent the type.

Lewia M.E. Barr & E.G. Simmons, Mycotaxon 25: 289 (1986). (Pleosporaceae)

Generic description

Habitat terrestrial, parasitic or saprobic? Ascomata small, scattered, erumpent to nearly superficial at maturity, subglobose to globose, black, smooth, papillate, ostiolate. Papilla short, blunt. Peridium thin. Hamathecium of pseudoparaphyses. Asci (4–6-)8-spored, bitunicate, fissitunicate, cylindrical to cylindro-clavate, with a short, furcate pedicel. Ascospores muriform, ellipsoid to fusoid.

Anamorphs reported for genus: Alternaria (Simmons 1986).

Literature: Kwasna and Kosiak 2003; Kwasna et al. 2006; Simmons 1986, 2007; Vieira and Barreto 2006.

Type species

Lewia scrophulariae (Desm.) M.E. Barr & E.G. Simmons, Mycotaxon 25: 294 (1986). (Fig. 46)

Fig. 46
figure 47

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

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 100140 × 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 (\( \bar{x} = 21.5 \times 9.1\mu m \), n = 10), constricted at median septum, smooth or verruculose (Fig. 46b, e and f).

Anamorph: Alternaria conjuncta (Simmons 1986).

Primary conidiophore simple with a single conidiogenous locus; conidia produced in chains, the first conidia in chain is larger, 30–45 × 10–12 μm, 7 transverse septa, 1–2 longitudinal or oblique septa in lower cells. Secondary conidiophore with 5–7 conidiogenous loci, sometimes branched; sporulation in chains, rarely branched.

Material examined: (FH, slide from lectotype).

Note: The specimen contains only a slide, so limited structures could be observed e.g. ascospores. The information about ascomata, peridium and whole asci is referred to Simmons (1986).

Notes

Morphology

Lewia has “Pleospora-like” teleomorphs, while it has Alternaria anamorphs, which are characterized by the beakless conidia connected together with secondary conidiophore (Simmons 1986). Based on these characters, more species under this genus were subsequently reported, i.e. Lewia avenicola Kosiak & Kwaśna (Kwasna and Kosiak 2003); L. chlamidosporiformans B.S. Vieira & R.W. Barreto (Vieira and Barreto 2006); L. alternarina (M.D. Whitehead & J.G. Dicks.) E.G. Simmons and L. daucicaulis E.G. Simmons (Simmons 2007). Currently Lewia comprises 15 species (http://www.mycobank.org, 24-02-2009).

Phylogenetic study

Phylogenetic analysis based either on SSU rDNA sequences or on multigenes indicated that Lewia species (Allewia eureka (E.G. Simmons) E.G. Simmons = L. eureka) form a robust clade with other members of Pleosporaceae (Schoch et al. 2006; Schoch et al. 2009; Zhang et al. 2009a).

Concluding remarks

Its position in Pleosporaceae is confirmed.

Lichenopyrenis Calat., Sanz & Aptroot, Mycol. Res. 105: 634 (2001). (?Pleomassariaceae)

Generic description

Habitat terrestrial, parasitic on lichens. Ascomata medium-sized, 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.

Type species

Lichenopyrenis galligena Calat., Sanz & Aptroot, Mycol. Res. 105: 636 (2001). (Fig. 47)

Fig. 47
figure 48

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

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 6585 × 15–20 μm (\( \bar{x} = 74 \times 18\mu m \), n = 10), 8-spored, bitunicate, fissitunicate, broadly clavate, with a short, thick, sometimes furcate pedicel, up to 13 μm long, ocular chamber not observed (Fig. 47f, g, h and k). Ascospores 1620 × 9–11 μm (\( \bar{x} = 18 \times 10\mu m \), n = 10), biseriate, ellipsoidal, pale orange-brown, 1-distoseptate, with prominent swelling at the septum, containing refractive globules, smooth (Fig. 47i, j and l).

Anamorph: The following description is from Calatayud et al. (2001).

Conidiomata pycnidial, arising in galls together with the ascomata, immersed, ca. 100200 μm diam.; wall dark brown throughout, composed of 2–5 layers of angular to laterally compressed cells; cells relatively large, ca. 816 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 57 × 1.5–2 μm. Vegetative hyphae hyaline.

Material examined: SPAIN, Andalucía, Province, Jaén, Andújar, lichenicolous on Leptochidium albociliatum (Desm.) M. Choisy on acid volcanic rock, 19 Apr. 2000, V. Calatayud (MA-Lichen 12715, holotype).

Notes

Morphology

Lichenopyrenis was formally established by Calatayud et al. (2001) based on its “perithecioid ascomata with peridium comprising compressed cells, fissitunicate and J- asci, wide hamathecium filaments, and 1-septate pale orange-brown ascospores with distoseptate thickenings at maturity”, and is monotypic with L. galligena. The genus was temporarily assigned to Pleomassariaceae. Lichenopyrenis galligena is a parasite of lichens, occurring in galls in the thallus of the host (Calatayud et al. 2001).

Phylogenetic study

None.

Concluding remarks

This is one of the few species that are parasitic on lichens. The most comparable species are Parapyrenis lichenicola Aptroot & Diederich and Lacrymospora parasitica Aptroot (both in Requienellaceae, Pyrenulales) as well as some species from Dacampiaceae. The peridium structure, cellular pseudoparaphyses, distoseptate and smooth, orange-brown ascospores as well as the anamorphic stage of Lichenopyrenis can easily distinguish from all of them (Calatayud et al. 2001).

Lineolata Kohlm. & Volkm.-Kohlm., Mycol. Res. 94: 687 (1990). (Pleosporales, genera incertae sedis)

Generic description

Habitat marine, saprobic (or perthophytic?). Ascomata medium-sized, gregarious, immersed to erumpent, obpyriform, ostiolate, papillate. Peridium thin, comprising two types of cells; outer cells thick stratum pseudostromatic, inner stratum thin, composed of a few layers of hyaline cells of textura angularis. Hamathecium of dense, long trabeculate pseudoparaphyses, embedded in mucilage, anastomosing and septate. Asci 8-spored, bitunicate, cylindrical, with short pedicels, with an ocular chamber. Ascospores uniseriate to partially overlapping, ellipsoidal, dark brown, 1-septate.

Anamorphs reported for genus: none.

Literature: Kohlmeyer and Kohlmeyer 1966; Kohlmeyer and Volkmann-Kohlmeyer 1990.

Type species

Lineolata rhizophorae (Kohlm. & E. Kohlm.) Kohlm. & Volkm.-Kohlm., Mycol. Res. 94: 688 (1990). (Fig. 48)

Fig. 48
figure 49

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–b = 20 μm, c–f = 10 μm

≡ Didymosphaeria rhizophorae Kohlm. & E. Kohlm. Icones Fungorum Maris (Lehre) 4 & 5: tab. 62a (1967).

Ascomata 300–490 μm high × 200–360 μm diam., gregarious, immersed to erumpent, obpyriform, ostiolate, papillate, subcarbonaceous to subcoriaceous, blackish brown (Fig. 48a). Peridium 37–45 μm thick, comprising two types of cells; outer cells thick stratum pseudostromatic, composed of irregular or roundish, dark brown cells, on the outside with a more or less recognizable hyphal structure, enclosing some decaying cells of the host, inner stratum thin, composed of four or five layers of hyaline, polygonal, elongate, thin-walled cells with large lumina, merging into the pseudoparaphyses. Hamathecium of dense, long trabeculate pseudoparaphyses, 1–1.5 μm broad, embedded in mucilage, anastomosing and septate. Asci 150175 × 14–17.5 μm, 8-spored, bitunicate, cylindrical, with short pedicels, with an ocular chamber (Fig. 48b). Ascospores 23–32(−33) × 9–12 μm, uniseriate to partially overlapping, ellipsoid, dark brown, 1-septate, not or slightly constricted at the septum, striate by delicate costae that run parallel or in a slight angle to the longitudinal axis of the ascospore (Fig. 48c, d, e and f) (adapted from Kohlmeyer and Kohlmeyer 1979).

Anamorph: none reported.

Material examined: US, Florida, Middle Torch Key, on Rhizophora mangle, 21 Nov. 1965, J. Kohlmeyer (Herb. J. Kohlmeyer No. 2390b, isotype); Pirate Grove Key, on R. mangle, 5 Jan. 1964 (Herb. J. Kohlmeyer No. 1721 paratype); Florida, Virginia Key, on R. mangle, 1 Jan. 1964, leg. E. Kohlmeyer (Herb. J. Kohlmeyer No. 1751 paratype); Florida, Torch Key, on R. mangle, 20 Nov. 1965, leg. J. Kohlmeyer (Herb. J. Kohlmeyer No. 2423 paratype).

Notes

Morphology

Lineolata was monotypified by L. rhizophorae, which was originally introduced by Kohlmeyer and Kohlmeyer (1966) as a species of Didymosphaeria (as D. rhizophorae). Based on the morphology of ascomata and asci, Barr (1990a) assigned it under Lojkania (as L. rhizophorae). Kohlmeyer and 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).

Phylogenetic study

Three isolates of Lineolata rhizophorae from varied geographic localities were analyzed by Suetrong et al. (2009) and shown to be related to Caryosporella rhizophorae in Dothideomycetidae and excluded from Pleosporomycetidae and Pleosporales.

Concluding remarks

Based on initial molecular work it is likely that this species does not belong to Pleosporales in spite of its dense pseudoparaphyses and other characters shared with the order.

Loculohypoxylon M.E. Barr, Mycotaxon 3: 326 (1976). (Teichosporaceae)

Generic description

Habitat terrestrial, saprobic. Ascomata relatively small, gregarious, immersed to erumpent, globose or subglobose, forming under a clypeus, papillate, ostiolate. Peridium thin, a single layer comprising hyaline thin-walled 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.

Anamorphs reported for genus: none.

Literature: von Arx and Müller 1975; Barr 1976.

Type species

Loculohypoxylon grandineum (Berk. & Rav.) Barr, Mycotaxon 3: 326 (1976). (Fig. 49)

Fig. 49
figure 50

Loculohypoxylon grandineum (from NY). a Appearance of ascomata on the host surface. b Habitat section of ascomata. c Section of an ascoma. Note the pale brown thin-walled peridium cells. d, e Uniseriate ascospores in asci. f–f Cylindro-clavate asci with ascospores. Note the ocular chamber in (g). Scale bars: a = 100 μm, b = 200 μm, c = 50 μm, dh = 10 μm

≡ 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 7090 × 10–12.5 μm (\( \bar{x} = 76.5 \times 10.9\mu m \), 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 (\( \bar{x} = 8.3 \times 5.9\mu m \), n = 10), uniseriate to partially overlapping at the upper part, broadly elliptic to subglobose, often apiculate at both ends, pale to dark brown, aseptate, with a germ slit (Fig. 49d and e).

Anamorph: none reported.

Material examined: USA, New Jersey, Newfield, on bark of Quercus coccinea, Sept. 1878, as Diatrype grandinea, Ellis N.A.F. 494 (NY, MASS); on Quercus sp. wood, Nov. 1893, as Anthostoma grandinea B. & Rav., Ellis & Everhart, N.A.F. 494 (NY); Newfield, Oct. 1881, as Diatrype grandinea (NY); Newfield, Jan. 1882, on Quercus coccinea, as Diatrype grandinea B. & Rav, Ex Herb Ellis (NY); Newfield, Nov. 1893, as Anthostoma grandinea, on bark of fallen trunks of Quercus coccinea (NY).

Notes

Morphology

Loculohypoxylon grandineum is one of the rare pleosporalean species having aseptate ascospores. When emphasis is given to ascospore morphology, Semidelitschia (monotypified by S. agasmatica Cain & Luck-Allen) is the most comparable genus. The large ascomata and ascospores, the mucilaginous sheath surrounding the ascospores as well as the coprophilous habitat of S. agasmatica differ from L. grandineum greatly. Thus Loculohypoxylon was introduced as a new genus.

Phylogenetic study

None.

Concluding remarks

Aseptate ascospores are rare in Pleosporales, and the position of this fungus needs further verification. The familial status of Loculohypoxylon in Teichosporaceae is questionable, as it is simply based on the similarity of living habitat, ascomata and asci with Immotthia and Teichospora (Barr 2002).

Lophionema Sacc., Syll. fung. (Abellini) 2: 717 (1883). (Pleosporales, genera incertae sedis)

Generic description

Habitat terrestrial, saprobic? Ascomata solitary, scattered or in small groups, immersed to erumpent, globose to subglobose, with a flattened base, wall black, papillate, ostiolate. Peridium comprising two types of cells which merge in the middle. Hamathecium of trabeculate pseudoparaphyses, septate, rarely anastomosing and branching. Asci 8-spored, bitunicate, fissitunicate unknown, clavate to cylindro-clavate, with a short and furcate pedicel and a small inconspicuous ocular chamber. Ascospores filliform, hyaline to pale yellow, multi-septate, slightly constricted at each septum.

Anamorphs reported for genus: none.

Literature: Barr 1992b; Chesters and Bell 1970; Ellis and Everhart 1892; Höhnel 1909; Solheim 1949.

Type species

Lophionema vermisporum (Ellis) Sacc., Syll. fung. (Abellini) 2: 717 (1883). (Fig. 50)

Fig. 50
figure 51

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

≡ Lophiostoma vermispora Ellis, Bull. Torrey bot. Club 9: 19 (1882).

Ascomata 320–430 μm high × 280–350 μm diam., solitary, scattered or in small groups of 2–3, immersed to erumpent, globose to subglobose, black, papillate, ostiolate. Papilla 80120 μm high, up to 150 μm broad, cylindrical to somewhat vertically flattened neck; mostly with a short slot-like ostiole, periphysate (Fig. 50a). Peridium 30–45 μm wide at the sides and slightly thicker at the apex, 2-layered, lateral walls and wall adjacent to neck comprising two types of cells which merge in the middle; outer cells small heavily pigmented thick-walled cells of textura angularis, cells 4–7 μm diam., cell wall 3.5–5 μm thick, inner cells less pigmented, comprising thin-walled compressed cells; apical wall cells smaller and walls thicker, basal wall thinner (ca. 15 μm wide), composed of lightly pigmented thin-walled compressed cells (Fig. 50b and c). Hamathecium of trabeculate pseudoparaphyses, 1–2 μm broad, septate, anastomosing and branching rarely between and mostly above the asci. Asci 105–130(−150) × 10–15 μm (\( \bar{x} = 123 \times 12\mu m \), 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 (\( \bar{x} = 95 \times 3.5\mu m \), n = 10), filliform, gradually tapering towards the base, hyaline to light yellow, (6-)7(−8)-septate, slightly constricted at each septum, smooth (Fig. 50f).

Anamorph: none reported.

Material examined: USA, New Jersey, Newfield, on dead stems of Oenothera biennis, Aug. 1881, Ellis (NY 643, holotype, NY 885, isotype).

Notes

Morphology

Lophionema is a relatively poorly studied genus, which was formally established by Saccardo (1883) as a monotypic genus represented by L. vermisporum based on its “globose ascomata, compressed ostiole, cylindrical to clavate ascus, and filamentous, septate, subhyaline to lightly pigmented ascospores”. Lophionema vermisporum was consequently listed as the generic type (Clements and Shear 1931). Berlese (1890) placed the genus in Lophiostomataceae but mentioned that the genus was similar to Ophiobolus according to the variable apex, and Shoemaker (1976) transferred Lophionema vermisporum to Ophiobolus sensu lato. Chesters and Bell (1970) however, had regarded Lophionema as related to Lophiostoma despite the distinct ascospore morphology. Barr (1992b) assigned Lophionema to Entodesmium based on the morphology of ascomata, papilla, peridium structure, pseudoparaphyses as well as the hyaline or slightly yellowish ascospores with a terminal appendage (not observed here). Species of Entodesmium, however, exclusively occur on legumes, but Lophionema vermisporum does not. We also note that the filliform ascospores, bitunicate asci, pseudoparaphyses and nature of the peridium may also be considered as typical of genera in the Tubeufiaceae (Barr 1980; Kodsueb et al. 2006b).

Phylogenetic study

None.

Concluding remarks

The immersed to erumpent ascomata, trabeculate pseudoparaphyses and laterally flattened papilla and periphysate ostioles indicate that this genus should be included in Lophiostomataceae. We do not accept the above proposals and, consider that Lophionema should be maintained as a separate genus with filliform ascospores in Lophiostomataceae until representative taxa can be sequenced and analyzed. Currently Lophionema comprises 10 species (http://www.mycobank.org, 08-01-2009). However, many of these are poorly studied and obscure.

Lophiostoma Ces. & De Not., Comm. Soc. crittog. Ital. 1: 219 (1863). (Lophiostomataceae)

Generic description

Habitat terrestrial, saprobic. Ascomata immersed to erumpent, usually with a distinct depressed papilla and a slot-like 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)

Fig. 51
figure 52

Lophiostoma macrostomum (ah, j from UPS, leptotype; i from IFRD 2005). a Appearance of ascomata on the host surface. Note the raised crest-like areas and full length germ slits. b Section of the peridium. c–e Cylindro-clavate asci with ascospores arranged in a 2-3-seriate manner. f Hamathecium comprising branching and septate pseudoparaphyses. g–j Released or unreleased ascospores. Note the smooth young ascospores with terminal sheath, and the verrucose senescent ascospores. Scale bars: a = 0.5 mm, b = 200 μm, cj = 10 μm

Sphaeria macrostoma Tode, Fung. mecklenb. sel. (Lüneburg) 2: 12 (1791).

Ascomata 400–600 μm high × 420–560 μm diam., densely scattered to gregarious, semi-immersed to erumpent, globose or subglobose, with a small to large flattened crest-like raised area above the ascomata which is variable in shape, up to 300 μm high and 480 μm wide, with a slit-like ostiole along the full length of the crest (Fig. 51a and b). Peridium 30–45 μm thick at the sides, thicker at the apex and thinner at the base, composed of one cell type of small lightly pigmented thin-walled cells of textura prismatica, cells ca. 6–9 × 3–4 μm diam., apex composed of pseudoparenchymatous cells (Fig. 51b). Hamathecium of dense, filliform, up to 3 μm near the base and less than 1.5 μm broad in the upper place, septate pseudoparaphyses, embedded in mucilage, anastomosing and branching between and above the asci (Fig. 51f). Asci 110145 × 10–15 μm (\( \bar{x} = 127.5 \times 13\mu m \), 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 (\( \bar{x} = 31.2 \times 6.4\mu m \), n = 10), biseriate, fusoid, curved, hyaline, usually 1-septate, with 3–5 septa and faintly brown when old, with (2-)3(−4) distinct oil drops in each cell and short terminal appendage at ends (Fig. 51h, i and j), and ornamented with warts when spores are senescent (Fig. 51g).

Anamorph: none reported.

Material examined: SWEDEN, Smaland, Femsjö par., Femsjö, on Prunus, 2006, Elias Fries, det. Geir Mathiassen (UPS, lectotype, as Sphaeria macrostoma Fr.). FRANCE, Ariège, Rimont, Las Muros, on dead stems of Vitis vinifera, 2 Sept. 1996 (IFRD2005).

Notes

Morphology

Lophiostoma is morphologically a well studied genus (Barr 1990a; Chesters and Bell 1970; Holm and Holm 1988; Mugambi and Huhndorf 2009b; Yuan and Zhao 1994), and currently it comprises about 30 species (Tanaka and Harada 2003b). The genus was characterized as having immersed to erumpent ascomata with a cylindrical or crest-like papilla and full length, slit-like ostiole; a peridium of unequal thickness, which was broader near the base (Lophiostoma-type); mostly clavate, bitunicate asci and 1- to several septate, hyaline to pigmented ascospores with terminal appendages or surrounded by a mucilaginous sheath (Holm and Holm 1988). This definition was followed by Barr (1990a), Yuan and Zhao (1994) and Hyde et al. (2002).

The crest-like papilla has been regarded as a prominent morphological character of Lophiostoma macrostomum (Chesters and Bell 1970; Holm and Holm 1988). In the lectotype specimen, the raised area above the ascomata is up to 300 μm high and 480 μm long, and seen as a flattened or even Y-shaped crest (Fig. 51a). In Lophiostoma curtum (Fr.) De Not. and Lophiotrema boreale Math. the raised area above the ascomata varies considerably in height or is even lacking (Holm and Holm 1988). Thus the variable “crest-like raised area in Lophiostomataceae” was explained as an evolutionarily adaptation to the hard substrate within which the ascomata develop (Holm and Holm 1988). The ascospores of L. macrostomum usually turn reddish brown when mature, and minutely verrucose ornamentation was also found on the surface of the pigmented ascospores. Hyaline ascospores that became pigmented with age are common in Lophiostoma, such as in L. appendiculatum Fuckel, L. massarioides (Sacc.) L. Holm & K. Holm, L. semiliberum, L. subcorticale Fuckel and L. winteri (Holm and Holm 1988; Tanaka and Harada 2003b). The phylogenetic significance of this character should be observed carefully in the future but at present its phylogenetic significance is unclear as this also occurs in some Lophiotrema species.

Phylogenetic study

Phylogenetic affinity with some Massarina species has been reported by Liew et al. (2002), and several Massarina species were transferred into Lophiostoma. In a systematic study of Lophiostoma- and Massarina-related fungi conducted by Zhang et al. (2009b), Lophiostoma taxa clustered into two groups; one includes the type species L. macrostomum with crest-like ostioles, L. rugulosum Yin. Zhang, J. Fourn. & K.D. Hyde with a wide, umbilicate pore surrounded by 4–6 radial ridges, and L. glabro-tunicatus with small ostiolar pores; the other cluster comprises Lophiostoma-like taxa with slot-like ostioles lacking raised crests, which includes L. arundinis (Pers.) Ces. & De Not., L. caulium, L. compressum (Pers.) Ces. & De Not., L. crenatum (Pers.) Fuckel, L. fuckelii (Sacc.) Sacc., L. macrostomoides, L. semiliberum and L. viridarium Cooke, which seems to represent a natural group at the family level. This conclusion is tentative until verified sequences of L. macrostomum are included in analyses (see comments of Zhang et al. 2009a).

Concluding remarks

We tend to accept a narrow concept of Lophiostomataceae, which only comprises species of Lophiostoma sensu stricto (Zhang et al. 2009a).

Lophiotrema Sacc., Michelia 1: 338 (1878). (Pleosporales, genera incertae sedis)

Generic description

Habitat terrestrial, saprobic. Ascomata small- to medium-sized, with or without short papilla. Hamathecium of dense, long, septate pseudoparaphyses, anastomosing and branching between and above asci. Asci cylindrical to cylindro-clavate. Ascospores hyaline, 1–3-septate, usually with mucilaginous sheath.

Anamorphs reported for genus: none.

Literature: Barr 1990a; Chesters and Bell 1970; Holm and Holm 1988; Saccardo 1878a; Tanaka and Harada 2003c; Tang et al. 2003; Yuan and Zhao 1994.

Type species

Lophiotrema nucula (Fr.) Sacc., Michelia 1: 338 (1878). (Fig. 52)

Fig. 52
figure 53

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

Sphaeria nucula Fr., Syst. mycol. (Lundae) 2: 466 (1823).

Ascomata 200–240 μm high × 200–280 μm diam., scattered, erumpent to nearly superficial, with basal wall remaining immersed in host tissue, globose to subglobose, often laterally flattened, with a flattened base not easily removed from the substrate, black, roughened; with a cylindrical or slightly compressed papilla. Papilla to 120 μm long and 150 μm high, protruding, with a pore-like ostiole (Fig. 52a). Peridium 25–30 μm wide, very thin at the base, composed of heavily pigmented pseudoparenchymatous cells near the apex, cells 2–2 × 6 μm diam., wall 1–3(−4) μm thick, lower sides composed of pigmented cells of textura angularis, 3–5 μm diam., wall 0.8–1.5 μm thick, ostiole wall composed of heavily pigmented and thick-walled small cells (Fig. 52b and c). Hamathecium of dense, long, septate pseudoparaphyses, 1–2 μm broad, anastomosing and branching between and above asci, embedded in mucilage (Fig. 52i). Asci 90115 × 9–11.5 μm (\( \bar{x} = 99.5 \times 11.5\mu m \); n = 10), 8-spored, bitunicate, fissitunicate, cylindrical, with a short, narrowed, furcate pedicel which is up to 10 μm long, with a small ocular chamber (ca. 1.5 μm wide × 1 μm high) (J-) (Fig. 52d, e, f and h). Ascospores 17–21(−25) × (4-)5–6.5 μm (\( \bar{x} = 19.5 \times 5.5\mu m \), n = 10), obliquely uniseriate and partially overlapping to biseriate, broadly fusoid to fusoid, with narrowly rounded ends, hyaline and lightly pigmented on very rare occasions when senescent, 1-septate, 3-septate when old, constricted at the median septum, the upper cell often broader than the lower one (Fig. 52g).

Anamorph: none reported.

Material examined: on decaying wood (UPS, lectotype as Sphaeria nucula Fr.).

Notes

Morphology

Holm and Holm (1988) provided a relatively strict definition for Lophiotrema after they examined several specimens including the type material which they lectotypified. Lophiotrema was mainly defined on the unique characters of small to medium ascomata, a “Lophiotrema-type” peridium and 1-septate ascospores. In Lophiotrema, Holm and Holm (1988) considered the ascomata to be small- to medium-sized, ca. pyriform but neck often reduced, even lacking and sometimes cylindrical. The peridium was of approximately equal thickness, 20–30 μm, composed of an outer textura angularis of uniformly pigmented cells, up to 12 μm, and an inner layer of very small hyaline cells, with somewhat thickened walls. Asci are cylindrical, spores hyaline, at first 1-septate, becoming 3-septate, with distinct guttules, often with a mucilaginous sheath. Much emphasis was given to the 1-septate ascospores by Holm and Holm (1988) when they described and distinguished the three Lophiotrema species: L. boreale, L. nucula, L. vagabundum (Sacc.) Sacc. and two other unnamed species. This concept was widely accepted by later workers (Kirk et al. 2001; Yuan and Zhao 1994). Tanaka and Harada (2003c) considered the peridium and asci to distinguish Lophiotrema from Lophiostoma, while Tang et al. (2003) introduced a new Lophiotrema species with elongated slit-like ostiole stating that the main difference between Lophiotrema and Lophiostoma were size of ascomata, structure of peridium, shape of asci and sheath of ascospores. This peridium concept however, is not supported by the lectotype specimen we examined here, which has a flattened thin-walled base. Thus the “Lophiotrema-like peridium” sensu Holm and Holm (1988) should not serve as a diagnostic character of Lophiotrema, while the ostiole, asci and ascospores might have some phylogenetic significance (Zhang et al. 2009b). No anamorph is yet known for Lophiotrema. Although the ascospores was reported by Holm and Holm (1988) to be verruculose this could not be observed in the lectotype examined under light microscope (1000 ×) in the present study.

Phylogenetic study

In the phylogenetic study of Lophiostoma, Massarina and related genera (Zhang et al. 2009b), Lophiotrema nucula formed a consistent and robust clade with three other Lophiotrema species: L. lignicola Yin. Zhang, J. Fourn. & K.D. Hyde, L. brunneosporum Yin. Zhang, J. Fourn. & K.D. Hyde and L. vagabundum, separate from other members of Lophiostoma and Massarina sensu stricto. This clade might represent Lophiotrema sensu stricto, however, the correctness of strains of L. vagabundum (CBS 628.86) and L. nucula (CBS 627.86) used in the phylogenetic study are not verified and warrant further study.

Concluding remarks

Holm and Holm (1988) distinguished Lophiostoma from Lophiotrema based on the smaller ascomata, 1-septate versus multi-septate ascospores, and peridial wall structure. However, we doubt that these distinguishing characters (size of ascomata, number of septa of ascospores) can be confidently used to separate these genera and we could not establish any characters that could reliably distinguish between these two genera. The molecular data, however, does separate Lophiostoma macrostomum and Lophiotrema nucula into separate clades and provides some support that these are separate genera. Although the strain of L. nucula (CBS 627.86) was isolated by K. & L. Holm, who had examined the type specimen of L. nucula (Holm and Holm 1988), the culture of Lophiostoma macrostomum used in the analysis are unverified (see comment by Zhang et al. 2009b). For the purpose of this monograph we tentatively maintain Lophiotrema as distinct from Lophiostoma.

Macroventuria Aa, Persoonia 6: 359 (1971). (Didymellaceae)

Generic description

Habitat terrestrial, saprobic. Ascomata small, solitary, scattered, or in groups, initially immersed, becoming erumpent, to nearly superficial, globose to subglobose, roughened with cylindrical setae erect from apex. Peridium thin, membranous. Hamathecium of cellular pseudoparaphyses, seems to easily disappear when mature. Asci bitunicate, somewhat obclavate to fusoid. Ascospores fusoid with broadly to narrowly rounded ends, hyaline, 1-septate, constricted at the septum.

Anamorphs reported for genus: none.

Literature: van der Aa 1971; von Arx and Müller 1975; Barr 1987a.

Type species

Macroventuria wentii Aa, Persoonia 6: 361 (1971). (Fig. 53)

Fig. 53
figure 54

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)

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 7593 × 24–30 μm, 8-spored, without pedicel, bitunicate, somewhat obclavate to fusoid (Fig. 53b). Ascospores 2232 × 8–14 μm, 1–3 seriate, fusoid with broadly to narrowly rounded ends, hyaline, 1-septate