Introduction

Phaeosphaeriaceae is a large and important family in Pleosporales and was previously reported to include more than 300 species in 35 genera (18 sexual and 17 asexual genera; Kirk et al. 2008; Zhang et al. 2009; Hyde et al. 2013; Wijayawardene et al. 2014b). Species of Phaeosphaeriaceae, especially the asexual taxa, are important plant pathogens, infecting major crops (Shoemaker and Babcock 1989b; Carson 2005; Stukenbrock et al. 2006; Zhang et al. 2009; Quaedvlieg et al. 2013; Hyde et al. 2014). For example, Parastagonospora nodorum (Berk.) Quaedvlieg et al. causes leaf and glume blotch of cereals crops, especially barley and wheat (Cunfer 2000; Stukenbrock et al. 2006; Vergnes et al. 2006; Quaedvlieg et al. 2013) while Parastagonospora avenae causes leaf blotch of barley and rye (f.sp. tritici) (Cunfer 2000; Quaedvlieg et al. 2013). Furthermore, many genera have also been reported as pathogens causing leaf spots on various hosts, such as Neosetophoma, Phaeosphaeria, Phaeosphaeriopsis, Setophoma, Wojnowicia and Xenoseptoria (Cason 2005; Arzanlou and Crous 2006; Quaedvlieg et al. 2013; Wijayawardene et al. 2013b; Crous et al. 2014; Phookamsak et al. 2014). Some species are also endophytes (Wang et al. 2005; Sánchez Márquez et al. 2007; Lawrey et al. 2012), Lautitia danica (Berl.) S. Schatz is obligate parasite on algae (Schatz 1984; Barr 1987b; Zhang et al. 2012) and many species are saprobic on monocotyledons, especially poaceous hosts, while others may be saprobes on herbaceous dicotyledonous hosts (Shoemaker 1984; Shoemaker and Babcock 1989b; Schoch et al. 2006; Zhang et al. 2009, 2012; De Gruyter et al. 2010; Hyde et al. 2013; Quaedvlieg et al. 2013).

History of Phaeosphaeriaceae

Introduction to the family

The family Phaeosphaeriaceae was introduced by Barr (1979) and typified by Phaeosphaeria with Ph. oryzae as the type species. The important family characters were described by Barr (1979) as “saprobic, pathogenic or hyperparasitic fungi typically on herbaceous stems or monocotyledonous leaves, culms, or flowers, but also on woody substrates. Ascomata were described as immersed, erumpent or superficial, globose or conical, short papillate, small to medium, asci as bitunicate, and ascospores as hyaline, yellowish or brown, narrowly or widely obovoid, aseptate or septate” Subsequent authors are in agreement with this description (Barr 1987b, 1992a, b; Zhang et al. 2012; Hyde et al. 2013). When Barr (1979) introduced the family Phaeosphaeriaceae, 15 genera were included (Table 1); i.e. Comoclathris, Didymella, Eudarluca, Heptameria, Leptosphaeria, Loculohypoxylon, Metameris, Microthelia, Nodulosphaeria, Ophiobolus, Paraphaeosphaeria, Phaeosphaeria, Rhopographus, Scirrhodothis and Teichospora (Barr 1979; Zhang et al. 2009, 2012).

Table 1 Synopsis of Phaeosphaeriaceae from 1979 to 2014
Full size table

Arrangement of genera in Phaeosphaeriaceae based on morphology (1979–1999)

Over the next 20 years the family was revised with additions and exclusions of genera and this is detailed below (Table 1). These revisions were based on morphological characters and were subjective with decisions made based on perceived importance of characters. During this period it was very difficult for researchers to develop higher classification schemes as it was not possible to confirm the importance of characters (K.D. Hyde, personal experience). Therefore, the researchers did their work to the best of their ability and have often been proven correct.

Barr (1982) included the monotypic genus Bricookea in Phaeosphaeriaceae, while Schatz (1984) introduced Lautitia, a parasite on red algae, and accommodated it in the family. Barr (1987a) transferred Leptosphaeria as the type to establish the new family Leptosphaeriaceae, because of its large scleroplectenchymatous, thick-walled peridium cells and host preference on dicotyledons (Barr 1987a, b; Khashnobish and Shearer 1996; Câmara et al. 2002). Barr (1987b) also transferred Ophiobolus and Heptameria to Leptosphaeriaceae and followed by Tanaka and Harada (2005). Eudarluca, Loculohypoxylon and Teichospora were placed in Dacampiaceae, while some species of Microthelia were regarded as synonyms of Kirschsteiniothelia, and subsequently placed in Pleosporaceae (Hawksworth 1985; Barr 1987b). Montagnula was considered as a part of Pleospora (Barr 1987b), however, the genus was re-established by Crivelli (1983). Barr (1987b) added Bricookea, Chaetoplea, Kalmusia, Lautitia and Ophiosphaerella and suggested that Amarenomyces and Entodesmium could also be accommodated in Phaeosphaeriaceae. Boise (1989) introduced Hadrospora as a new genus in Phaeosphaeriaceae and additionally, Sulcispora was established in Phaeosphaeriaceae by Shoemaker and Babcock (1989b). More recently, Mixtura (Syd.) Eriksson and Yue (1990) was placed, with some uncertainty in Phaeosphaeriaceae.

Eriksson and Hawksworth (1991) included Entodesmium and Peridiothelia in the family but this was disputed by Barr (1992a). It was suggested that Entodesmium should be placed in Lophiostomataceae because of its abrupt, beak-like apex and the periphysate ostioles which were illustrated by Shoemaker (1984) and Peridiothelia could be accommodated in Pleomassariaceae. Thus, Barr (1992a) accepted only 14 genera in Phaeosphaeriaceae: Bricookea, Chaetoplea, Didymella, Graphyllium, Hadrospora, Kalmusia, Lautitia, Metameris, Montagnula, Nodulosphaeria, Ophiosphaerella, Phaeosphaeria, Paraphaeosphaeria and Rhopographus (Table 1).

Shoemaker and Babcock (1992) revised the taxonomy of the Pleosporales and transferred Comoclathris to Diademaceae and Graphyllium was moved to Hysteriaceae based on its hysterothecia with a slit-like opening (Zhang et al. 2011; Ariyawansa et al. 2014a). The marine fungus, Carinispora K.D. Hyde (Hyde 1992), a saprobe on the intertidal Nypa palm was placed in Phaeosphaeriaceae, and it was noted that the genus was closely related to Phaeosphaeria, although differed in several important aspects. The genus was tentatively placed in Phaeosphaeriaceae due to lack of molecular data (Hyde 1992, 1994; Jones et al. 2009; Lumbsch and Huhndorf 2007, 2010).

Eriksson and Hawksworth (1993) listed 13 genera in Phaeosphaeriaceae: Bricookea, Carinispora, Eudarluca, Hadrospora, Lautitia, Metameris, Mixtura, Nodulosphaeria, Ophiosphaerella, Paraphaeosphaeria, Phaeosphaeria, Sulcispora and Teratosphaeria (Table 1). Scirrhodothis was synonymised under Metameris, while Didymella was excluded from the family. Sivanesan (1984) proposed placing Didymella in Pleosporaceae, while Reddy et al. (1998) treated it in Venturiaceae (Dianese et al. 2001; Zhang et al. 2012). Yuan (1994) established Barria as a monotypic genus from Picea within Phaeosphaeriaceae. The peridium in this genus is composed of thin-walled pseudoparenchymatous cells and it has narrow cellular pseudoparaphyses. Ramaley and Barr (1995) introduced the monotypic genus Pleoseptum from Yucca baccata and reported Camarosporium yuccaesedum Fairm. as the asexual state of the genus. Pleoseptum shares morphological characters with several Dothideomycetes genera (Ramaley and Barr 1995). However there is a lack of molecular phylogenetic evidence to place this genus in Phaeosphaeriaceae (Zhang et al. 2012; Hyde et al. 2013).

Wilmia, a monotypic genus, was introduced as a pathogen of Memora pedunculata and formed asexual morphs associated with the sexual morph. The morphology is similar to several genera such as Barria, Lautitia, and Metameris in the Phaeosphaeriaceae (Dianese et al. 2001; Zhang et al. 2012; Ariyawansa et al. 2014b).

Dothideopsella was characterized by broadly pseudoparenchymatous cells, and phragmosporous, brown ascospores. The genus was introduced by von Höhnel (1915) to accommodate non-congeneric species of Leptosphaeria (Barr and Rogerson 1999). However, the genus was reinstated under Leptosphaeria by von Arx and Müller (1975) and this was followed in Kirk et al. (2001, 2008). Barr and Rogerson (1999) treated Dothideopsella in Botryosphaeriaceae, while Lumbsch and Huhndorf (2010) placed the genus in Dothideomycetes, genera incertae sedis. Recently, Dothideopsella was reported as a genus in Leptosphaeriaceae which contained four epithet species. However, Ariyawansa et al. (2013) observed the type species and found that the genus was typical in Phaeosphaeriaceae rather than Leptosphaeriaceae. Thus the genus was transferred to Phaeosphaeriaceae and is followed by Wijayawardene et al. (2014b).

Various other genera included in Phaeosphaeriaceae by Barr (1979, 1987a, b, 1990a, 1992a), have been transferred to other families based on morphology. Comoclathris is currently accommodated in the Diademaceae (Shoemaker and Babcock 1992; Zhang et al. 2012; Hyde et al. 2013; Ariyawansa et al. 2014a), Heptameria and Rhopographus are treated in Dothideomycetes, genera incertae sedis (Lumbsch and Huhndorf 2007, 2010; Zhang et al. 2012; Hyde et al. 2013), Loculohypoxylon and Teichospora in the family Teichosporaceae (Barr 2002; Lumbsch and Huhndorf 2007, 2010; Zhang et al. 2012; Hyde et al. 2013; Wijayawardene et al. 2014b).

Initial phylogenetics for resolving Phaeosphaeriaceae

In 1995 the first phylogenetic evidence for placement of genera in Phaeosphaeriaceae started to emerge. Morales et al. (1995) attempted to determine the relationships between Leptosphaeria and Phaeosphaeria species using rRNA gene sequences. The separation of Leptosphaeria and Phaeosphaeria was not strongly supported and the asexual states were phylogenetically uninformative in their analysis. Phylogenetic analysis indicated that Phaeosphaeria was part of Leptosphaeriaceae due to the fact that Ph. nodorum and Ph. microscopica formed a clade close to L. doliolum, the type species of Leptosphaeria (Morales et al. 1995; Câmara et al. 2002.). Khashnobish and Shearer (1996) found that Phaeosphaeria formed a distinct group separate from Leptosphaeria based on a cladistic analysis of morphological and sequence data. Câmara et al. (2002) concluded that peridial characters, asexual state and host preferences are taxonomically and phylogenetically significant at the generic level, with Leptosphaeria sensu stricto and Phaeosphaeria sharing a more recent common ancestor in their analysis.

Kruys et al. (2006) investigated the relationships of families in Pleosporales and observed that Phaeosphaeriaceae is closely related to Leptosphaeriaceae and formed an unresolved monophyletic group. The main features distinguishing the two families are the ascomata wall characters. In Leptosphaeriaceae taxa have scleroplectenchymatous ascomata walls and in Phaeosphaeriaceae taxa have thin-walled, pseudoparenchymatous ascomata walls (Câmara et al. 2002; Cheng et al. 2004; Schoch et al. 2006). Shiraia, an uncertain genus grouped with Leptosphaeriaceae and Phaeosphaeriaceae in the phylogenetic analysis of Kruys et al. (2006). Cheng et al. (2004) suggested that Shiraia should be treated as a member of Phaeosphaeriaceae. The genus was unofficially placed in Shiraiaceae, by Ogawa et al. (2003) (see GenBank AB105798), but was not validly published (Kruys et al. 2006; Liu et al. 2013). Liu et al. (2013) illustrated and epitypified the type species, Shiraia bambusicola from fresh collection in Zhejiang Province, China. Based on their phylogenetic investigation, Shiraia formed a distinct clade separate from Phaeosphaeriaceae. Therefore, Liu et al. (2013) introduced a new valid family Shiraiaceae to accommodate a monotypic genus Shiraia in Pleosporales and illustrated a coelomycetous asexual state.

Zhang et al. (2009) incorporated multigene phylogenetic analysis and included 15 families in Pleosporales. They included four families in the suborder Pleosporineae viz. Didymellaceae, Leptosphaeriaceae, Phaeosphaeriaceae and Pleosporaceae. In addition, they introduced two new families, Amniculicolaceae and Lentitheciaceae to accommodate some freshwater fungi (Zhang et al. 2009, 2012). Zhang et al. (2009) re-accommodated Ophiobolus in Phaeosphaeriaceae due to its morphological characters and because Ophiobolus is often associated with the asexual genus Chaetosphaeronema on the host (Petrak 1944). However, this link has not been confirmed (Zhang et al. 2009).

Zhang et al. (2012) re-circumscribed the families Phaeosphaeriaceae in Pleosporales. These authors treated 18 sexual genera base on morphology in Phaeosphaeriaceae and accepted Entodesmium, Ophiosphaerella, Phaeosphaeria and Setomelanomma based on phylogenetic analysis (Table 1). The freshwater genera Isthmosporella Shearer and Crane (1999) and Ocala Raja et al. (2009) were also accommodated in Phaeosphaeriaceae. Zhang et al. (2012), however, removed these genera from Phaeosphaeriaceae based on morphological characters and phylogenetic analysis; Ocala formed a unique clade basal to Jahnulales (Shearer et al. 2009) and Isthmosporella was treated in Pleosporales, genera incertae sedis. Hyde et al. (2013) listed 27 sexual and asexual genera in Phaeosphaeriaceae and later, Wanasinghe et al. (2014) introduced the monotypic genus, Dematiopleospora typified by D. mariae in Phaeosphaeriaceae based on molecular data. Furthermore, Crous et al. (2014) included Setophaeosphaeria Crous & Y. Zhang ter with Sp. badalingensis Crous & Y. Zhang ter, Sp. hemerocallidis Crous & Y. Zhang ter and Sp. setosa (Leuchtm.) Crous in Phaeosphaeriaceae. Setophaeosphaeria is morphologically and phylogenetically distinct from Phaeosphaeria oryzae, the type species of Phaeosphaeriaceae, but is similar to Ph. setosa. Thus the genus was established to accommodate phaeosphaeria-like species.

Recent molecular studies place Kirschsteiniothelia in a new family Kirschsteiniotheliaceae based on molecular data (Boonmee et al. 2012) and Microthelia species are treated in synonymy of various genera of Dothideomycetes (Hyde et al. 2013). Kalmusia, Montagnula and Paraphaeosphaeria are accommodated in Montagnulaceae (Boonmee et al. 2012; Hyde et al. 2013; Lumbsch and Huhndorf 2010; Zhang et al. 2009, 2012; Ariyawansa et al. 2014b). Several genera in Phaeosphaeriaceae, such as Barria, Bricookea, Chaetoplea, Eudarluca, Hadrospora, Lautitia, Metameris, Mixtura, Nodulosphaeria, Ophiobolus, Pleoseptum, and Wilmia lack molecular data.

History of genera

Amarenomyces Eriksson was introduced as a new genus based on its multi-layered endotunica and large, thick-walled and sheathed ascospores (Eriksson 1981). Eriksson (1982) reported Amarenographium to be the asexual state of Amarenomyces. Amarenomyces was synonymized under Phaeosphaeria by Zhang et al. (2009) based on phylogenetic evidence from Amarenomyces ammophilae (Lasch) O.E. Erikss. which formed a related clade with other Phaeosphaeria species.

Didymella was treated as a member of Phaeosphaeriaceae by Barr (1979) when she introduced the new family Phaeosphaeriaceae. However, Silva-Hanlin and Hanlin (1999) suggested that Didymella belongs in Pleosporaceae rather than Phaeosphaeriaceae based on phylogenetic relationships, while Lumbsch and Huhndorf (2007) treated Didymella in Pleosporales, genera incertae sedis. Later, De Gruyter et al. (2009) introduced a new family Didymellaceae based on a multigene phylogenetic analysis with Didymella as the type genus.

Entodesmium was introduced by Reiss (1854) as typified by E. rude Riess associated with legumes (Zhang et al. 2009). The genus was previously placed in Lophiostomataceae based on its periphysate papilla (Eriksson and Hawksworth 1990; Barr 1992b; Zhang et al. 2009). Liew et al. (2000) sequenced a strain of E. rude and their phylogenetic analysis indicated that Entodesmium belongs to Phaeosphariaceae which has been upheld by subsequent authors (Schoch et al. 2009; Zhang et al. 2009, 2012; Hyde et al. 2013, 2014). However, there is only one putative strain of Entodesmium rude (CBS 650.86) available in GenBank (Liew et al. 2000), and this is not apparently linked to any herbarium material.

Eudarluca was introduced by Spegazzini (1908) as a hyperparasite in the uredinia of rust fungi from Canna (Zhang et al. 2012). Sphaerellopsis was reported to be the asexual state of this genus (Yuan et al. 1998; Nischwitz et al. 2005; Zhang et al. 2012). Many strains of Eudarluca caricis (Fr.) O.E. Erikss. are available in GenBank. Nischwitz et al. (2005) investigated host specialization and relationships of Eudarluca caricis with Ampelomyces. Based on the phylogenetic analysis, their results have shown that Eudarluca caricis formed a clade with other pleosporalean taxa and appeared to be host-specific (Nischwitz et al. 2005). The phylogenetic investigation was limited to variation between differentiated isolates and could not resolve the current placement of Eudarluca in Phaeosphaeriaceae (Bayon et al. 2006; Zhang et al. 2012).

Katumotoa was introduced as a monotypic genus in Phaeosphaeriaceae and was tentatively placed in Phaeosphaeriaceae (Tanaka and Harada 2005). Later, Tanaka et al. (2009) removed Katumotoa from Phaeosphaeriaceae and accommodated this genus in the family Lentitheciaceae (Zhang et al. 2009), based on morphological and phylogenetic evidence.

Loratospora was introduced by Kohlmeyer and Volkmann-Kohlmeyer (1993) as a monotypic genus. The type species, Loratospora aestuarii Kohlm. & Volkm.-Kohlm. is an obligate and facultative marine taxon (Kohlmeyer and Volkmann-Kohlmeyer 1993; Monkai et al. 2013). Suetrong et al. (2009) transferred Loratospora to Phaeosphaeriaceae based on phylogenetic data, whereas based on morphology it had previously been placed in Planistromellaceae (Barr 1996).

Ophiosphaerella was introduced by Spegazzini (1909) with Op. graminicola Speg. as the type species. Barr (1987b) placed Ophiosphaerella in Phaeosphaeriaceae, due to its morphological characters (Shoemaker and Babcock 1989b; Dong et al. 1998). The placement in Phaeosphaeriaceae was confirmed by Zhang et al. (2009, 2012) and Hyde et al. (2013) based on molecular phylogeny.

Phaeosphaeriopsis was introduced to accommodate Paraphaeosphaeria species that were not congeneric based on morphological characters and phylogenetic investigation and that had a Phaeostagonospora asexual state (Câmara et al. 2003; Zhang et al. 2012). The placement of the genus in Phaeosphaeriaceae was confirmed by multi-gene analysis (Quaedvlieg et al. 2013; Thambugala et al. 2014).

Setomelanomma was introduced by Morelet (1980) as a pathogen on blue and white spruce (Morelet 1980; Rossman et al. 2002). The genus was considered morphologically similar to Phaeosphaeriaceae (Rossman et al. 2002; Zhang et al. 2012). Rossman et al. (2002) confirmed the genus belongs in Phaeosphaeriaceae based on a phylogenetic analysis and this was verified with multi-gene analyses (Schoch et al. 2009; Zhang et al. 2009, 2012; De Gruyter et al. 2012; Hyde et al. 2013; Wijayawardene et al. 2014b).

Placement of asexual genera in Phaeosphaeriaceae

Before the era of molecular phylogeny it was difficult to establish if asexual genera belonged in Phaeosphaeriaceae unless single ascospore cultures produced the asexual morph or where the asexual morph was associated with the sexual state on the host (Eriksson 1982; Ramaley 1995; Ramaley and Barr 1995; Yuan et al. 1998; Câmara et al. 2003; Nischwitz et al. 2005). Previous to the compilation by Hyde et al. (2011) there were few attempts to incorporate asexual genera in sexually defined families. Perhaps the first report of an asexual state of Phaeosphaeriaceae is that of Eriksson (1982) who reported Amarenographium to be the asexual state of Amarenomyces. Barr (1987b) listed nine coelomycetous genera, Ascochyta, Chaetodiplodia, Coniothyrium, Microdiplodia, Microsphaeropsis, Phoma, Scolecosporiella, Stagonospora and Sphaerellopsis in Phaeosphaeriaceae based on their holoblastic or enteroblastic conidiogenesis and aseptate or septate conidia.

Camarosporium yuccaesedum Fairm formed conidiomata on leaves of Yucca baccata that were associated with ascomata of Pleoseptum yuccaesedum, the type species of Pleoseptum, (Ramaley and Barr 1995; Zhang et al. 2012). Based on its structurally similar and close association, Ramaley and Barr (1995) linked Camarosporium yuccaesedum as an asexual state of Pleoseptum. Camarosporium species had been reported as asexual morphs in various genera of Botryosphaeriales and Cucurbitariaceae with doubtful polyphyletic support (Crous et al. 2006; Kirk et al. 2008; Liu et al. 2012; Zhang et al. 2012; Doilom et al. 2013). However, Wijayawardene et al. (2014a, c, d) recognized Camarosporium sensu stricto as a distinct phylogenetic linage in Pleosporineae, Pleosporales hence introduced Camarosporiaceae, (Wijayawardene et al. 2014a, d)

Hyde et al. (2011) and Wijayawardene et al. (2012) also listed Hendersonia as an asexual state of Phaeosphaeriaceae. However, the genus was synonymized under Stagonospora and is currently placed in Massarinaceae (Swart and Walker 1988; Quaedvlieg et al. 2013).

Parahendersonia Ramaley (1995) was classified by its ampulliform conidiogenous cells producing thick-walled, brown, (1–)3-septate conidia. This monotypic genus is represented by P. dasylirionis. The sexual state was reported as Chaetoplea dasylirionis on Dasylirion (Agavaceae) in Leptosphaeriaceae (Ramaley 1995; Calatayud and Etayo 2001). However, Barr (1987b) accommodated Chaetoplea in Phaeosphaeriaceae and reported a microdiplodia-like asexual state for Chaetoplea (Barr 1990b; Zhang et al. 2012). Recently, Parahendersonia was treated in Leptosphaeriaceae (Index Fungorum 2014).

Rhabdospora was originally established to accommodate septoria-like species associated with stems of Nerium oleander L. (Priest 2006; Quaedvlieg et al. 2013). The genus was treated in the taxonomic studies of Sutton (1980) which 11 Rhabdospora species currently classified under Septoria in Mycosphaerellaceae (Quaedvlieg et al. 2013). Hyde et al. (2011) listed Rhabdospora as the asexual state of Phaeosphaeriaceae, while Zhang et al. (2012) treated the genus as an asexual state of Ophiobolus. Wijayawardene et al. (2014b) treated the genus in Dothideomycetes, genera incertae sedis. There are 648 epithets listed in Index Fungorum (2014) for Rhabdospora and it was accommodated in Mycosphaerellaceae. However, Rhabdospora is a poorly understood genus and is in need of modern taxonomic and molecular treatment (Quaedvlieg et al. 2013).

The type species of Scolecosporiella, Sc. typhae (Oudemans) Petr. was reported as the asexual state of Phaeosphaeria typharum (Desm.) L. Holm (Nag Raj 1989). The genus has been re-described taxonomic revision by Sutton (1968, 1980) and Nag Raj (1989). Various sexual genera, such as Ophiobolus, Ophiosphaerella, and Phaeosphaeria have been reported as Scolecosporiella sexual states (Farr et al. 1989; Nag Raj 1989; Shoemaker and Babcock 1989b; Hyde et al. 2011; Zhang et al. 2012).

Tiarospora was reported as an asexual member of Phaeosphaeriaceae by Kirk et al. (2008). However, there are no molecular data to prove that this genus belongs in Phaeosphaeriaceae. Tiarospora has four epithets in Index Fungorum (2014) with T. perforans (Sacc.) Höhn being listed as a synonym of Montagnula perforans (Aptroot 2006).

Molecular data has allowed linkage of asexual genera in Phaeosphaeriaceae

Ascochyta, Chaetodiplodia, Microdiplodia and Microsphaeropsis have been reported as asexual states of Phaeosphaeriaceae (Barr 1987b); however, using phylogenetic evidence Crous et al. (2006) treated Microdiplodia as the asexual state of Karstenula in Montagnulaceae (Ariyawansa et al. 2014b). Liu et al. (2012) and Hyde et al. (2013) reported Microdiplodia as an asexual state in Botryosphaeriaceae. Ascochyta and Microsphaeropsis were treated in Didymellaceae based on phylogenetic evidence with the sexual morph of these genera described as Didymella (De Gruyter et al. 2009; Aveskamp et al. 2010). Ascochyta was shown to be polyphyletic in Pleosporales by subsequent researchers (De Gruyter et al. 2009, 2012; Aveskamp et al. 2010). Furthermore, allied Ascochyta species were determined to belong in Pleosporaceae by De Gruyter et al. (2012). A Chaetodiplodia strain formed a clade within Leptosphaeriaceae; however, the genus is polyphyletic in Pleosporaceae (De Gruyter et al. 2009, 2012).

Based on phylogenetic evidence, Câmara et al. (2003) introduced the new genus Phaeosphaeriopsis in Phaeosphaeriaceae and transferred Paraphaeosphaeria nolinae A.W. Ramaley to Phaeosphaeriopsis. Phaeostagonospora was reported as the asexual state based on its morphology (Ramaley 1997). Sexual and asexual morph connections were studied by Quaedvlieg et al. (2013) and Thambugala et al. (2014).

De Gruyter et al. (2009) included Wojnowicia hirta (J. Schröt.) Sacc., the type species in their phylogenetic analysis and concluded that Wojnowicia hirta has an asexual state in Phaeosphaeriaceae and this phylogenetic placement was supported by Wijayawardene et al. (2013b, 2014b).

Zhang et al. (2009) listed Ampelomyces, Chaetosphaeronema, Coniothyrium, Phoma, Plenodomus, Stagonospora and Wojnowicia (asexual genera) in Phaeosphaeriaceae. Based on phylogenetic evidence, Chaetosphaeronema was placed in Phaeosphaeriaceae and often formed a sister clade with the sexual genus Entodesmium (Zhang et al. 2009, 2012; Hyde et al. 2013; Wijayawardene et al. 2014b). Phoma radicina (McAlpine) Boerema, the type species of Phoma section Paraphoma formed a sister clade with the sexual genus Setomelanomma holmii M. Morelet as basal Phaeosphaeriaceae (Zhang et al. 2009, 2012; Hyde et al. 2013; Wijayawardene et al. 2014b).

De Gruyter et al. (2010) introduced two new genera, Neosetophoma and Setophoma in Phaeosphaeriaceae and also re-introduced Paraphoma in Phaeosphaeriaceae based on phylogenetic evidence. Four Phoma species from Aveskamp et al. (2010) which clustered in Phaeosphaeriaceae were accommodated in these genera (De Gruyter et al. 2010). Hyde et al. (2011) reported 17 asexual genera in Phaeosphaeriaceae (Table 1). Later, Wijayawardene et al. (2012) updated the checklist of asexual states of Phaeosphaeriaceae and included the hyphomyceteous genera Harpophora and Mauginiella. In their compilation of families in Pleosporales, Zhang et al. (2012) accepted 17 asexual genera in Phaeosphaeriaceae. They excluded the hyphomycetous genera, based on a broad familial concept and phylogenetic support.

Hyde et al. (2013) excluded Amarenographium which formed a clade outside Phaeosphaeriaceae and this is supported by Hodhod et al. (2012). Wijayawardene et al. (2013a) accommodated Scolicosporium minkeviciusii Treigienė in Phaeosphaeriaceae as this species formed a clade with other well-established asexual genera. Furthermore, Crous et al. (2013) revealed that Xenostigmina and Mycopappus grouped in Phaeosphaeriaceae in their phylogenetic analysis.

Quaedvlieg et al. (2013) re-circumscribed the pathogenic fungi genera Septoria, and Stagonospora and related genera such as Phaeosphaeria and Phaeoseptoria. Based on their phylogenetic study, they introduced 14 new genera, 36 new species, and 19 new combinations. The new genera Neostagonospora, Parastagonospora, Xenoseptoria and Vrystaatia were placed in Phaeosphaeriaceae. Furthermore, they epitypified Phaeosphaeria oryzae I. Miyake and Phaeoseptoria papayae Speg. and confirmed sexual asexual connections. Phaeoseptoria was confirmed as the asexual morph of Phaeosphaeria based on the sequenced strain of P. papayae which formed a well-supported clade with Ph. oryzae.

Crous et al. (2014) included Septoriella oudemansii Oudem. in Phaeosphaeriaceae. Septoriella was compared with species of Phaeosphaeriaceae and a megablast search of NCBI showed that Septoriella is similar to Phaeosphaeria caricis, Ph. ammophilae and Ophiosphaerella herpotricha with high percentage similarities (Crous et al. 2014).

In a new outline of the Dothideomycetes, Wijayawardene et al. (2014b) proposed synonymies and suggested names for 34 genera while incorporating asexual genera into a natural classification. This publication resulted from the new ruling in article 59.1 of International Code of Nomenclature for Algae, Fungi, and Plants (ICN; Melbourne Code) that a species or genus can only have one name. They listed 29 genera (17 only known as asexual genera) in Phaeosphaeriaceae.

According to phylogenetic evidence, Ampelomyces species grouped in Didymellaceae and Phaeosphaeriaceae (De Gruyter et al. 2009; Aveskamp et al. 2010; Hyde et al. 2013; Wijayawardene et al. 2013a, 2014b). The type species of Ampelomyces, A. quisqualis Ces. formed a clade within Phaeosphaeriaceae (De Gruyter et al. 2009; Aveskamp et al. 2010; Wijayawardene et al. 2013a), while A. quercinus (Syd.) Rudakov grouped in Didymellaceae (De Gruyter et al. 2009). Therefore, Aveskamp et al. (2010) transferred all Ampelomyces species belonging in Didymellaceae, to Phoma based on phylogenetic relationships.

Coniothyrium Corda (1840) is characterized by pycnidial conidiomata, reduced conidiophores, annellidic conidiogenous cells, with brown, aseptate or septate, verrucose conidia (De Gruyter et al. 2012). Based on the phylogenetic analyses, Coniothyrium was revealed as polyphyletic in Pleosporales (Verkley et al. 2004, 2014). The type species of Coniothyrium, C. palmarum Corda clustered in Leptosphaeriaceae (De Gruyter et al. 2009). In De Gruyter et al. (2012), C. palmarum formed a distinct clade with other Phoma species in the reinstated Coniothyriaceae based on phylogenetic analysis. De Gruyter et al. (2012) indicated that Coniothyrium had convergent evolution with similar characters found in other families in Pleosporales such as Cucurbitariaceae, Didymellaceae, Phaeosphaeriaceae, Leptosphaeriaceae, Montagnulaceae and Sporormiaceae. However, Coniothyrium concentricum (Desm.) Sacc. is related to Phaeosphaeriaceae. A new genus should be designated to accommodate this coniothyrium-like species (Hyde et al. 2013).

Harpophora was introduced by Gams (2000) to accommodate phialophora-like species characterized by fast-growing, thin colonies and sickle-shaped phialoconidia (Gams 2000; Saleh and Leslie 2004). The genus contains the asexual state of Gaeumannomyces and Magnaporthe in Magnaporthaceae (Gams 2000; Saleh and Leslie 2004; Kirk et al. 2008; Yuan et al. 2010).

Based on a homologous test using ITS sequence data, Abdullah et al. (2005) indicated that Mauginiella scaettae Cavara was closely related to species of Phaeosphaeria which formed a clade with Phaeosphaeria associated with non-grass hosts. It was concluded that Mauginiella may represent the asexual state of Phaeosphaeria following the suggestion of Câmara et al. (2002). Câmara et al. (2002) mentioned that the morphological characters, asexual state and host preferences are important to clarify Phaeosphaeria species. It has been suggested that a great deal of convergent evolution had occurred in species of Leptosphaeria and Phaeosphaeria (Câmara et al. 2002; Abdullah et al. 2005).

Phoma species have been described as the asexual state of various genera of Dothideomycetes i.e. Didymella, Mycosphaerella, Leptosphaeria and Pleospora (Boerema 1997; De Gruyter et al. 2009). De Gruyter et al. (2009) carried out a re-classification of the Phoma complex based on molecular phylogeny. Their results showed that Phoma was polyphyletic and was subsequently classified in nine sections: Phoma, Heterospora, Paraphoma, Peyronellaea, Phyllostictoides, Sclerophomella, Plenodomus, Macrospora, and Pilosa (Boerema 1997; Boerema et al. 2004; De Gruyter et al. 2009). Based on phylogenetic evidence, they concluded that Phoma section Paraphoma, as typified by Ph. radicina (McAlpine) Boerema can be assigned in Phaeosphaeriaceae, while Ph. heteromorphospora, the type species of Phoma section Heterospora should be treated in Leptosphaeriaceae. Additionally, the family Didymellaceae was proposed to accommodate others Phoma sections viz. Macrospora, Peyronellaea, Phoma, Phyllostictoides and Sclerophomella (De Gruyter et al. 2009). Aveskamp et al. (2010) indicated that the current Boeremaean subdivision was incorrect as Phoma is polyphyletic comprising six distinct clades in Pleosporales and appeared to be common in different families (Aveskamp et al. 2010). Therefore, De Gruyter et al. (2012) concluded that Phoma should be restricted only to Didymellaceae, while other Phoma sections were transferred to related families. However, Lawrey et al. (2012) treated lichenicolous Phoma species in Phaeosphaeriaceae based on their multigene phylogenetic analysis. Therefore, a new genus should be introduced to accommodate all lichenicolous Phoma species in Phaeosphaeriaceae (Hyde et al. 2013).

Plenodomus Preuss is typified by Pl. rabenhorstii Preuss to accommodate the phoma-like species which Boerema (1982) synonymized under Phoma section Plenodomus (Boerema et al. 2004; Torres et al. 2005). Torres et al. (2005) mentioned that Reddy et al. (1998) treated phoma-like species with the sexual state known as Leptosphaeria in Plenodomus in Leptosphaeriaceae and concluded that Phoma should be restricted to the species relating to Didymella in Phaeosphaeriaceae. Zhang et al. (2009) listed Plenodomus in Phaeosphaeriaceae; however based on phylogenetic evidence, De Gruyter et al. (2012) reinstated the genus Plenodomus to accommodate Phoma section Plenodomus within Leptosphaeriaceae.

Sclerostagonospora was typified by S. heraclei (Sacc.) Höhn. (1917) and characterized with pycnidial conidiomata with holoblastic, determinate, discrete, ampulliform to irregular conidiogenous cells, and subcylindrical, 3-septate, pale brown, verruculose conidia (Sutton 1980; Quaedvlieg et al. 2013). Sclerostagonospora differs from Stagonospora in its pigmented conidia (Quaedvlieg et al. 2013). Crous and Palm (1999) reported Sclerostagonospora leucadendri Crous & M.E. Palm as the asexual state of Leptosphaeria leucadendri Crous & M.E. Palm which occurs on leaves of Leucadendron. Based on phylogenetic analysis, Sclerostagonospora opuntiae (Ellis & Everh.) Huhndorf (CBS 118224) formed a clade with Phaeosphaeria and Phaeosphaeriopsis (Crous et al. 2008). Thus, Kirk et al. (2008) treated Sclerostagonospora as an asexual state in the Pleosporales. In Quaedvlieg et al. (2013), multigene phylogenetic analysis has shown that Sclerostagonospora phragmiticola Quaedvlieg et al. (CBS 338.86) is related to Phaeosphaeriaceae.

Stagonospora is characterized by holoblastic or annellidic conidiogenesis and produces phragmoconidia (Schoch et al. 2006; Quaedvlieg et al. 2013). The genus was reported as an asexual genus in Phaeosphaeriaceae, which often clusters with Phaeosphaeria and Phoma in phylogenetic analyses (Cunfer and Ueng 1999; Câmara et al. 2002; Solomon et al. 2006; Zhang et al. 2009, 2012; Hyde et al. 2013). Stagonospora has often been confused with Septoria; however, the genus was clarified by Cunfer and Ueng (1999) and Solomon et al. (2006). Based on phylogenetic evidence, Stagonospora is more closely related to other genera in Pleosporales, while Septoria clustered in Mycosphaerellaceae, Capnodiales (Quaedvlieg et al. 2013). Quaedvlieg et al. (2013) re-collected the type species of Stagonospora, St. paludosa (Sacc. & Speg.) Sacc. and their phylogenetic analysis showed that this species formed a separate clade with other Stagonospora species which occurred on Poaceae (i.e. St. avenae (A.B.. Frank) Bissett and St. nodorum (Berk.) E. Castell. & Germano). Therefore, Quaedvlieg et al. (2013) concluded that stagonospora-like species belonged in different genera and introduced the new genera, Neostagonospora and Parastagonospora to accommodate the stagonospora-like species in Phaeosphaeriaceae. The type species of Stagonospora, St. paludosa and other Stagonospora species grouped in Massarinaceae (Quaedvlieg et al. 2013).

The aim of this study is to revisit all genera of Phaeosphaeriaceae and provide revised descriptions of the sexual and asexual morphs of the genera. Available generic type specimens were examined, described and illustrated. We have also collected fresh specimens from Thailand and Italy. Phylogenetic investigation of available sequence data reveals the current placement of genera in Phaeosphaeriaceae.

Material and methods

Herbarium examination, fungal isolation, morphology and asexual state study

Type specimens of each genus in Phaeosphaeriaceae were obtained from BPI, BR, NY, S, and UB herbaria. Morphology of generic types were examined and re-described. Generic descriptions were provided based on the original descriptions and non type herbarium specimens if generic type specimens could not be located or loaned. Some types were redrawn from iconotype. Ascomata on herbarium material were rehydrated by adding 5 % KOH. Free hand sections and squash mounts were prepared and morphological characters were observed under stereo microscope and compound microscope.

Collections of Phaeosphaeriaceae were made from Italy and Thailand and returned to the laboratory for examination. If the specimens were dry, they were incubated in a moist chamber for 2–3 days at 25 °C. Ascomata were sectioned by hand and examined under an Olympus SZH10 stereomicroscope. Macro-morphological characters were photographed using a Sony DSC-T110 digital camera. The micro-morphological characters were captured by a Cannon 550D digital camera, using a Nikon ECLIPSE 80i compound microscope with DIC objectives. Indian ink was added to show mucilaginous sheaths surrounding ascospores. Cotton blue and congo red reagents were used as stains for checking ascospore septation and internal elements of the hamathecium. Permanent slides were prepared by mounting material on lactoglycerol and sealing with clear nail polish. Measurements were obtained using a Tarosoft (R) Image Frame Work version 0.9.7. Photographic plates were prepared in Adobe Photoshop version CS3 (Adobe Systems Inc., The United States).

Pure cultures were obtained from single spores following the methods described in Chomnunti et al. (2014) and Phookamsak et al. (2013). Ascomata were cut at the apex and sides walls by a razor blade. The contents inside ascoma was picked and soaked in sterile water and broken up further mechanically with the sterile needle until a spore suspension was obtained. The ascospore suspension was dropped on water agar (WA; 15.0 g/L sterile distilled water) and incubated at room temperature overnight. The germinating spores were transferred to malt extract agar (MEA; 33.6 g/L sterile distilled water, Difco malt extract) and cultivated in corn meal agar (CMA, 17 g/l sterile distilled water, Difco corn meal agar) and potato dextrose agar (PDA; 39 g/L distilled water, Difco potato dextrose) for recording growth rates and culture characters. Sterile bamboo pieces on water agar (WA; 15 g/l sterile distilled water) were used to encourage production of asexual morphs. Cultures are deposited in Mae Fah Luang University Culture Collection (MFLUCC), and duplicated in the International Collection of Microorganisms from Plants (ICMP), Landcare Research, New Zealand. Herbarium specimens were prepared by drying the specimens with silica gel at room temperature for 7–10 days depending on the humidity and kept in envelopes or small boxes. The herbarium specimens are deposited in the herbarium of Mae Fah Luang University (MFLU), Chiang Rai, Thailand and duplicates in the New Zealand Fungal Herbarium (PDD), Landcare Research, New Zealand or Université Catholique de Louvain, Belgium (MUCL).

Molecular and phylogenetic analysis

Fungal DNA extraction, PCR amplification and sequencing

The fungal colonies were grown on MEA/PDA at 25–30 °C for 2–4 weeks. After 4 weeks, the fungal mycelium was scraped and transferred to micro centrifuge tube. The fungal genomic DNA was extracted by Biospin Fungus Genomic DNA Extraction Kit (BioFlux®, China) following the manufacturer’s instructions (Hangzhou, P.R. China).

DNA amplifications were performed by Polymerase Chain Reaction (PCR). The internal transcribed spacer (ITS) was amplified by using primers ITS4 and ITS5 as defined by White et al. (1990). The partial small subunit nuclear rDNA (SSU) was amplified by using primers NS1 and NS4 (White et al. 1990). The partial large subunit nuclear rDNA (LSU) was amplified by using primers LROR and LR5 (Vilgalys and Hester 1990). The partial RNA polymerase second largest subunit (RPB2) was amplified by using primers fRPB2-5F and fRPB2-7cR (Liu et al. 1999). The translation elongation factor 1-alpha gene (TEF1) was amplified by using primers EF1-983F and EF1-2218R (Rehner 2001). The amplification reaction were carried out following protocol: The final volume of the PCR reaction was 50 μl which contained 2.0 μl of DNA template, 1.5 μl of each forward and reverse primers, 25 μl of 2× Easy Taq PCR SuperMix (mixture of EasyTaqTM DNA Polymerase, dNTPs, and optimized buffer, Beijing TransGen Biotech Co., Ltd., Chaoyang District, Beijing, PR China) and 20 μl sterilized water (Phookamsak et al. 2013).

The PCR thermal cycle program of ITS, nuSSU, nuLSU and TEF1 genes amplification were provided as: initially 94 °C for 3 min, followed by 40 cycles of denaturation at 94 °C for 30 s, annealing at 55 °C for 50 s, elongation at 72 °C for 1 min, and final extension at 72 °C for 10 min. The PCR thermal cycle program for the partial RNA polymerase second largest subunit (RPB2) was obtained by step as initially 95 °C for 5 min, followed by 40 cycles of denaturation at 95 °C for 1 min, annealing at 52 °C for 2 min, elongation at 72 °C for 90 s, and final extension at 72 °C for 10 min. The quality of PCR products were checked on 1 % agarose gel electrophoresis strained with ethidium bromide. The PCR products were send for sequencing at Shanghai Sangon Biological Engineering Technology & Services Co., Ltd (Shanghai, P.R. China) (Phookamsak et al. 2013).

Phylogenetic analyses

Combined ITS, LSU, SSU, RPB2 and TEF1 sequence data were used for phylogenetic analysis in this study. The reference nucleotide sequences of Phaeosphaeriaceae and other selected families in Dothideomycetes were obtained from the GenBank database and included in the analysis (see Supplementary table). The fungal sequence strains were combined and aligned in MEGA6 (Tamura et al. 2013) and improved manually where necessary in BioEdit v. 7.2 (Hall 1999). The combined sequences alignment was analyzed by Bayesian, maximum likelihood and maximum parsimony methods.

Bayesian analyses was carried out using MrBayes v. 3.0b4 (Ronquist and Huelsenbeck 2003). The best-fit model of sequences evolution was estimated by using MrModeltest 2.2 (Nylander 2004). Markov Chain Monte Carlo sampling (BMCMC) in MrBayes v. 3.0b4 (Huelsenbeck and Ronquist 2001) were used to determine the Posterior probabilities (PP) (Rannala and Yang 1996; Zhaxybayeva and Gogarten 2002). Phylogenetic trees were sampled every 100th generation (resulting in 10,000 total trees) in 1,000,000 generations from the running of six simultaneous Markov chains. The first 2,000 trees which contained the burn-in phase of the analyses were discarded. The remaining 8,000 trees were used to calculate the posterior probabilities (PP) in the majority rule consensus tree (Liu et al. 2011).

A maximum likelihood analysis (ML) was performed in RaxmlGUI v.1.0 (Silvestro and Michalak 2011). Alignments in PHYLIP format were loaded and automatically checked their compatibility in raxmlGUI. The available substitution models consisting of a generalized time reversible (GTR) was applied with four rate classes and a discrete gamma distribution (Yang 1994) for nucleotides, symmetric, Markovian, and ordered models for morphological characters (Liu et al. 2011; Silvestro and Michalak 2011). The best scoring tree from a separate randomized tree under the same model was selected. Rapid bootstrap with nonparametric bootstrap iterations (Stamatakis et al. 2008) was run in 1,000 replicates with the GTR model and a discrete gamma distribution (Liu et al. 2011).

A maximum parsimony (MP) analysis was conducted by using PAUP v. 4.0b10 (Swofford 2002). The heuristic search option with 1,000 random sequences addition and tree-bisection reconnection (TBR) of branch-swapping algorithm were performed. Maxtrees were setup at 1,000 and a zero of maximum branch length was collapsed. Gaps were treated as missing data and all characters were unordered and of equal weight. All multiple, equally parsimonious trees were saved. Descriptive tree statistics for parsimony consistency index (CI), retention index (RI), rescaled consistency index (RC) and homoplasy index (HI) was calculated. The robustness of the best parsimonious tree was estimated by a bootstrap (BT) value with 1,000 replicates, each with 10 replicates of random stepwise addition of taxa (Liu et al. 2011; Phookamsak et al. 2013). The best scoring phylogram from Bayesian analysis was selected for representing phylogenetic relationships and drawn in Treeview (Page 1996) with bootstrap values above and below the branches. All the sequences generated in this study have been deposited in GenBank (Liu et al. 2011; Phookamsak et al. 2013).

Results and discussion

Three phylogenetic reconstructions were generated to clarify the phylogenetic relationships of genera and species that have been accommodated in Phaeosphaeriaceae. Figure 1 deals with the genus and family level placement within the Phaeosphaeriaceae and Pleosporales. The dataset includes LSU, SSU, RPB2, and TEF1 sequences and were analyzed using maximum likelihood (ML), maximum parsimony (MP) and Bayesian analyses. The dataset comprised 85 taxa, with Massaria inquinans (M 19) selected as the outgroup taxon. Twenty-nine Phaeosphaeriaceae taxa formed a well-supported clade (74 % ML/ 1.00 PP) within the suborder Pleosporinae, while Mycopappus and Xenostigmina form a single clade close to Melanommataceae. Therefore, we exclude these later genera from Phaeosphaeriaceae in this study. Wilmia was synonymized under Letendraea by Ariyawansa et al. (2014b). Phylogenetic evidence shows Letendraea cordylinicola is related with Didymosphaeriaceae which form a strongly-supported clade (75 % ML/ 1.00 PP) with Paraphaeosphaeria michotii (CBS 652.86), Deniquelata barringtoniae (MFLUCC 11-0422) and Kalmusia brevispora (KT 2313) in Didymosphaeriaceae. Thus, Letendraea is excluded from Phaeosphariaceae in this study and transferred to Didymosphaeriaceae. Phaeosphaeriopsis, Parastagonospora, Ophiosphaerella, Phaeosphaeria and Setophoma are well-resolved genera and formed distinct clades, while the other clades comprised groups of genera without clear combining features or comprising more than one genus (Fig. 2).

Fig 1
figure 1

A Bayesian 50 % majority rule consensus tree based on combined dataset of LSU, SSU, TEF1 and RPB2 alignment. Bootstrap support values for maximum likelihood (ML, red) and maximum parsimony (MP, green) equal or greater than 60 % are given above the nodes. Bayesian posterior probabilities (BYPP, blue) equal or higher than 0.95 are given below the nodes. Massaria inquinans (M 19). Ex-type and ex-epitype strains are in bold. Newly generated sequences are in red and the type species are indicated in blue

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Fig 2
figure 2

A Bayesian 50 % majority rule consensus tree based on combined dataset of ITS, LSU, SSU and RPB2 alignment. Bootstrap support values for maximum likelihood (ML, blue) and maximum parsimony (MP, green) equal to or greater than 60 % are given above the nodes. Bayesian posterior probabilities (BYPP, red) equal or higher than 0.95 are given below the nodes. The tree is rooted to Didymella exigua. Ex-type and ex-epitype strains are in bold. Newly generated sequences are in red and the type species are indicated in blue

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Figure 2 focuses on the phylogenetic relationships of genera in Phaeosphaeriaceae. The multigene combined dataset (ITS, LSU, SSU and RPB2) were obtained from maximum likelihood (ML), maximum parsimony (MP) and Bayesian analysis. Didymella exigua (CBS 183.55) was selected as the outgroup taxon. The dataset consists of 58 taxa, with 57 taxa belonging to the Phaeosphaeriaceae. Eleven clades (A–K) can be recognized in Fig. 2. Clade A comprises the genera Chaetosphaeronema, Dermatiopleospora, Entodesmium, Loratospora. Nodulosphaeria and Ophiobolus. Dermatiopleospora, Entodesmium and Loratospora are represented by their ex-type strains and type species. This clade needs better populating with many more species from the respective genera as it is presently not well-resolved. Ophiobolus cirsii (MFLUCC 13-0218) formed a robust clade (99 % ML/ 97 % MP/ 1.00 PP) with the asexual genus Chaetosphaeronema, while O. erythrosporus (MFLU 12-2225) forms a weakly supported clade with Entodesmium rude. However, Ophiobolus is not represented by its type species and the strains used have not been morphologically clearly defined (see Ariyawansa et al. 2014c). Clade B represents the genus Phaeosphaeriopsis and is well resolved in Phaeosphaeriaceae (97 % ML/ 93 % MP/ 1.00 PP). It is represented by the type species Ps. glaucopunctata (ex-type strain), plus other strains and a new species Ps. dracaenicola is also introduced here (see details in “Taxonomy” section). Clade C represents Phaeosphaeria sensu stricto with five species, including the type species and two new species; detailed phylogenetic relationships among these species are shown in Fig. 3. Clade D represents two isolates of the genus Ophiosphaerella, which form a strongly supported clade (100 % ML/ 100 % MP/ 1.00 PP) separate from Ophiosphaerella herpotricha. The two strains of Ophiosphaerella agrostidis are clearly defined based on morphology (see under Ophiosphaerella agrostidis). However, the genus Ophiosphaerella is not represented by its type species. Further isolates and type strains from Ophiobolus and Ophiosphaerella are required to better resolve and naturally place these genera. Clade E comprises the genera Amarenomyces, Sclerostagonospora, and Wojnowicia plus a putative strain of Ophiosphaerella herpotricha and four strains of Phaeosphaeria. Phaeosphaeria ammophilae is not closely related to Phaeosphaeria sensu stricto, the species is reinstated as Amarenomyces, while other Phaeosphaeria taxa in this clade may need removing from Phaeosphaeria. However, the tree needs to be better populated with strains that are morphologically well-defined before adjustments can be made. Clade F comprises two strains, Phaeosphaeria alpina and Ph. typharum which are also unrelated to Phaeosphaeria sensu stricto and need further study. Clade G comprises the type species and ex-type strains of Ampelomyces and Neosetophoma, these genera also need to be better populated with strains that are morphologically well-defined. Clade H comprises the ex-types strains and type species of Parastagonospora and, Neostagonospora, and ex-type strains Sclerostagonospora cycadis, Scolicosporium minkeviciusii and Wojnowicia viburni (the latter three species are not generic types). Three Phaeosphaeria sensu lato strains also cluster here indicating the polyphyletic nature of the genus. Clade I comprises Xenoseptoria, Setomelanomma and Paraphoma, with Xenoseptoria and Paraphoma represented by ex-type strains of their type species. Setomelanomma is closely related to Xenoseptoria which formed a robust clade (97 % ML/ 71 % MP) in our multigene phylogenetic analyses. Clade J represents Vrystaatia with single species, V. aloeicola. Clade K represents the asexual genus Setophoma with five strains, including the type species, S. terrestris. The genus is well-resolved in our phylogenetic analyses which form a strongly supported clade (100 % ML/ 64 % MP/ 1.00 PP) at the basal of Phaeosphariaceae.

Fig 3
figure 3

Maximum parsimony tree based on a combined dataset of ITS, LSU, RPB2 and TEF1. Bootstrap support values for maximum parsimony (MP, green) (equal to or greater than 50 %) and the values of the Bayesian posterior probabilities (BYPP, blue) from MCMC analyses provided significant (equal or higher than 95 %) are given above the nodes. The tree was rooted to Phaeosphaeria juncicola. Ex-type and ex-epitype strains are in bold. Newly generated sequences are in blue

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Figure 3 deals with the species complex of Phaeosphaeria sensu stricto and Parastagonospora. Phaeosphaeria juncicola (CBS 110108) was selected as the outgroup taxon. The newly generated strains which were collected from Thailand form a robust clade with the ex-type species, Phaeosphaeria oryzae and with Ph. papayae and Ph. musae in the Phaeosphaeria sensu stricto clade. Phaeosphaeria sensu stricto is distinguished from Parastagonospora, the species having been accommodated in Phaeosphaeria and Phaeosphaeria sensu lato (Phaeosphaeria alpina and Ph. typharum).

Phaeosphaeria sensu stricto taxa (MFLUCC 11-0133; MFLUCC 11-0151) cluster with Ph. musae (CBS 120026), a pathogen of banana. An ITS pair wise comparison shows that these three isolates differ in seven base positions (Fig. 4) while there are only one to two base differentiated in TEF1 gene comparisons. Isolate MFLUCC 11-0133 is slightly morphologically different from the other two isolates (MFLUCC 11-0151 and CBS 120026) in its ascus and ascospore sizes. However, these three isolates are generally similar in morphology and phylogeny. Therefore, we treat the isolates causing leaf spot disease on Calathea sp. and Cordyline sp. from Thailand as conspecific with Phaeosphaeria musae. Phaeosphaeria thysanolaenicola (MFLUCC 10-0563), a pathogen on Thysanolaena maxima, forms a clade with. Ph. chiangraina, which is saprobic on rice. These two species form strongly-supported clades which are well-resolved from the ex-type species of Phaeosphaeria oryzae.

Fig 4
figure 4

ITS pair wise comparison of Phaeosphaeria oryzae (CBS 110110), P. oryzae (RP0086; MFLUCC 11-0170), P. papayae (CBS 135416), P. thysanolaenicola (RP0036; MFLUCC 10-0563), P. chiangraina (RP0131; MFLUCC 13-0231), P. musae (CBS 120026) and P. musae (RP0049; MFLUCC 11-0133 and RP0067; MFLUCC 11-0151)

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Phaeosphaeria avenaria, Ph. eustoma and Ph. nigrans form a clade with Parastagonospora species. Phaeosphaeria avenaria was synonymized under Parastagonospora avenae by Quaedvlieg et al. (2013), and the other two species may need to be synonymized under Parastagonospora based on the phylogenetic relationships. However, it was not possible to observe the types of these species or confirm if they are correctly identified.

Phaeosphaeria alpina and Ph. typharum form a distinct clade from the Phaeosphaeria sensu stricto species which is separate with other Phaeosphaeria sensu lato taxa. The type species need to re-visited and a new genus may need to be introduced to accommodate these species.

Taxonomy

Phaeosphaeriaceae M.E. Barr Mycologia 71(5): 948 (1979).

Faces of Fungi number: FoF00232

Endophytic, hyperparasitic, pathogenic, or saprobic on monocotyledonous and some dicotyledonous hosts. Sexual state: Ascomata scattered, clustered, solitary or gregarious, immersed to semi-immersed or erumpent through host tissue, small to large, uniloculate, globose to subglobose, glabrous or setose, brown to dark brown or black, ostiolate, with short to long papilla. Peridium with thin to thick walls, composed of brown to dark brown or black, pseudoparenchymatous cells, mostly arranged in a textura angularis to textura prismatica. Hamathecium composed of sparse to numerous, filamentous, septate, rarely anastomosing, mostly cellular pseudoparaphyses, or pseudoparaphyses occasionally lacking. Asci 8-spored, bitunicate, fissitunicate, cylindrical to cylindric-clavate, or clavate, sessile to subsessile, or usually with short pedicels, apically rounded with typically well-developed ocular chamber. Ascospores overlapping muriform, phragmosporous, or scolecosporous, varying in shape from ellipsoidal, fusiform to broadly fusiform, clavate, and cylindrical to filiform, mostly brown to dark brown or reddish-brown, sometimes hyaline, 1- to multi-septate, constricted or not constricted at the septum, walls smooth or rough, echinulate, punctuate or verrucose, some with appendage or mucilaginous sheath. Asexual state: Coelomycetous. Conidiomata pycnidial, scattered, solitary or gregarious, immersed to superficial, uni- to multi-loculate, globose to subglobose, glabrous or setose, light brown to dark brown or black, ostiole central, papillate. Conidiomata walls thin, composed of pseudoparenchymatous cells, arranged in a textura angularis to textura prismatica. Conidiophores lining inner cavity of the conidioma, typically reduced to conidiogenous cells, hyaline to brown, aseptate or septate. Conidiogenous cells enteroblastic or holoblastic, annellidic or phialidic, discrete, oblong to ampulliform, hyaline to brown. Conidia varied in shape, hyaline to brown, aseptate or septate, walls smooth or rough, echinulate, punctuate or verrucose, some with appendage or mucilaginous sheath.

Notes: Phaeosphaeriaceae is one of the largest families in Pleosporales, and demonstrates variability in morphological characters. Previously, many genera with brown spores, associated with monocotyledonous hosts, having a thin-walled peridium and producing coelomycetous asexual states were accommodated in Phaeosphaeriaceae (Barr 1979, 1987b, 1990a, 1992a; Crivelli 1983; Schatz 1984; Shoemaker 1984; Shoemaker and Babcock 1989b; Yuan 1994; Ramaley and Barr 1995; Dianese et al. 2001; Zhang et al. 2009, 2012). Most of genera in Phaeosphaeriaceae lack molecular data or only limited sequence data is available in some genera such as Entodesmium and Phaeosphaeria. Several genera in Phaeosphaeriaceae share morphological characters with genera in other families (e.g. Didymosphaeriaceae, Leptosphaeriaceae and Pleosporaceae) and thus their placement is confused. Therefore, re-visiting all type genera of Phaeosphaeriaceae to establish their morphology and to obtain molecular sequence data from fresh collections are essential to provide modern descriptions and to determine the placement of genera in this family.

Type: Phaeosphaeria I. Miyake, Bot. Mag., Tokyo 23: 93 (1909).

Notes: The following genera are placed in Phaeosphaeriaceae. Genera are listed in alphabetical order and provided with genus description and notes, descriptions of type or authentic species, notes on species, material examined and illustrations. If the genus is monotypic, only a single description which also represents the genus is provided.

Key to genera of Phaeosphaeriaceae

1. Sexual state . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

1. Asexual state . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14

2. Ascospores phragmosporous or scolecosporous . . . . . . 3

2. Ascospores dictyosporous, muriform, on Ononis spinosa (dicotyledon) . . . . . . . . . . . . . Dematiopleospora

3. Ascospores phragmosporous . . . . . . . . . . . . . . . . . . . . . . . . . 4

3. Ascospores scolecosporous . . . . . . . . . . . . . . . . . . . . . . . . . 12

4. Ascospores 2–3-septate . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

4. Ascospores mostly more than 3-septate . . . . . . . . . . . . . 10

5. Ascospores mostly 2-septate, associated with rust fungi . . . . . . . . . . . . . . . . . . . . . . . . Eudarluca

5. Ascospores 3-septate, saprobic or pathogenic on monocots or dicots . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6

6. Ascomata in groups or rows with slit-like ostioles; ascospores hyaline . . . . . . . . . . . . . . . . . . . . . . . Bricookea

6. Ascomata solitary, scattered, opening ostiolate to papillate; ascospores pale brown, yellowish brown to brown . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .7

7. Asci ovoid to ampulliform, aparaphysate, on Juncus roemerianus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Loratospora

7. Asci broadly cylindrical, cellular pseudoparaphyses . . . . . 8

8. Ascomata typically glabrous . . . . . . . . . . . . . . . . . . . . . . . 9

8. Ascomata typically setose . . . . . . . . . . . Setomelanomma

9. Ascomata typically immersed, ostioles with minute papilla, asexual state: Phaeoseptoria . . . . . . . . . . . . . Phaeosphaeria

9. Ascomata semi-immersed to erumpent, near superficial, ostiole with conical, knob-like, asexual: unknown . . . . . . . . . . . . . . . . . . . Dothideopsella

10. Ascospores cylindrical, with typically one enlarged cell . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11

10. Ascospores broadly fusiform, widest at the middle cell, saprobic on Ammophila . . . . . . . . . . . . . Amarenomyce s

11. Papilla, with numerous long, large setae in the ostiole, ascospores with one enlarged cell near apex . . . . . . . . . . . . . . . . . . . . Nodulosphaeria

11. Papilla absent, opening ostiolate, ascospores typically enlarged cell near the base . . . . . . . . . . Phaeosphaeriopsis

12. Ascospores with enlarged cells . . . . . . . . . . .Ophiobolus

12. Ascospores lacking enlarged cells . . . . . . . . . . . . . . . . 13

13. Ascospores constricted at the septa, breaking into part spores, saprobic or parasitic on legumes . . . . Entodesmium

13. Ascospores non-constricted at the septum, not separating into part spores, saprobic on various monocots . . . . . . . . . . . . . . . . Ophiosphaerella

14. Conidia amerosporous, didymosporous phragmosporous or scolecosporous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15

14. Conidia dictyosporous (muriform) . . . Amarenographium

15. Conidia aseptate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .16

15. Conidia septate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18

16. Saprobic or pathogenic, conidiomata setose . . . . . . . . 17

16. Mycoparasitic on powdery mildew . . . . . Ampelomyces

17. Conidiomata dark brown, no sexual state reported . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Paraphoma

17. Conidiomata pale brown, olivaceous to olivaceous-black, sexual state: ascospores cylindrical, hyaline, 3-septate, with enlarged in second cell from apex . . . . . . Setophoma

18. Conidiomata setose or covered by mycelium . . . . . . . 19

18. Conidiomata glabrous . . . . . . . . . . . . . . . . . . . . . . . . . . 21

19. Conidia 1–3-septate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20

19. Conidia more than 3-septate . . . . . . . . . . . . . . . Wojnowicia

20. Conidiophores present, hyaline, 1-septate conidia . . . . . . . . . . . . Chaetosphaeronema

20. Conidiophores absent, yellowish, typically 0–1(−3)-septate conidia . . . . . . . . . . . . . . . . . . . . . Neosetophoma

21. Conidia 1-septate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22

21. Conidia more than 1-septate . . . . . . . . . . . . . . . . . . . . . . . . 23

22. Conidia with appendage . . . . . . . . . . . . . . . . . Tiarospora

22. Conidia without appendage . . . . . . . . Neostagonospora

23. Conidia hyaline . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24

23. Conidia pigmented . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26

24. One type of conidia, cylindrical to obclavate . . . . . . . 25

24. Two types of conidia (macroconidia subcylindrical to narrowly obclavate; microconidia aseptate, pear-shaped to globose or ellipsoid) . . . . . . . . . . . . Vrystaatia

25. Conidiomata exuding a creamy conidial mass, conidiophores absent; sexual state: phaeosphaeria-like . . . . . . . . . . . . . . . . . . . Parastagonospora

25. Conidiomata exuding a pink to orange conidial mass, conidiophores branched; sexual state: ??Setomelanomma . . . . . . . . . . . . . . Xenoseptoria

26. Conidia usually 3-septate . . . . . . . . . . . . . . . . . . . . . . . . 27

26. Conidia usually more than 3-septate . . . . . . . . . . . . . . 29

27. Conidia produced one type of conidia . . . . . . . . . . . . . . . . 28

27. Conidia produced two types of conidia (microconidia globose to ellipsoidal, hyaline) . . . . . Phaeostagonospora

28. Conidia subcylindrical, pale brown, minutely verruculose . . . . . . . . . . . . . . . . . . . Sclerostagonospora

28. Conidia subcylindrical, wider in the middle, yellowish-brown, smooth-walled . . . . . . . Septoriella oudemansii

29. Conidia fusiform, 5–6-septate, apical cell attenuated into a short, beak-like, appendage . . . . . . . . . . . . Scolecosporiella

29. Conidia curved to sigmoid, 6–7-septate, with hyaline end cells . . . . . . . . . . . . . . . . . . . . Scolicosporium minkeviciusii

Amarenographium O.E. Erikss., Mycotaxon 15: 199 (1982), Faces of Fungi number: FoF00237.

Saprobic on marine grasses. Sexual state: see notes. Asexual state: Conidiomata pycnidial, scattered, solitary, immersed to erumpent through host tissue by papilla, uniloculate, ellipsoidal to subglobose, glabrous, brown to dark brown, ostiole central, with papilla. Conidiomata walls thin, composed of thick-walled, reddish-brown to dark brown cells, arranged in textura angularis to textura prismatica. Conidiophores mostly reduced to conidiogenous cells, hyaline to brown, septate, branched. Conidiogenous cells holoblastic, phialidic, discrete, determinate, ampulliform, rarely cylindrical, hyaline, smooth-walled, produced macro- and microconidia. Macroconidia dictyosporous, blastic-phialidic, broadly ellipsoidal to clavate or fusiform, brown to dark brown, paler at the ends cells, muriform, septate, smooth-walled, with mucoid appendage at apex. Microconidia amerosporous, blastic-phialidic, hyaline, smooth-walled (from Eriksson 1982; Nag Raj 1989).

Type species: Amarenographium metableticum

Phylogenetic study: Hodhod et al. (2012)

Notes: Amarenographium metableticum (Trail) O.E. Erikss. is the type of Amarenographium which was introduced as the asexual state of Amarenomyces (Eriksson 1982; Nag Raj 1989; Hodhod et al. 2012). Amarenographium metableticum had previously been identified as Camarosporium metableticum which was not congeneric with Camarosporium stephensii, the generic type of Camarosporium at that time (Sutton 1980; Eriksson 1982). Therefore, Eriksson (1982) transferred Camarosporium metableticum to a new genus based on the different morphological characters (Hodhod et al. 2012; Hyde et al. 2013). Amarenographium differs from Camarosporium by its conidiophores and conidia. In Amarenographium conidiophores are longer and branched and conidia have gelatinous appendages (Eriksson 1982; Hodhod et al. 2012). Hodhod et al. (2012) introduced a new species of Amarenographium, Am. solium Abdel-Wahab et al. from mangroves and noted that Amarenographium did not belong in Phaeosphaeriaceae but clusters with Pleosporales, genera incertae sedis, while Amarenomyces formed a clade with Phaeosphaeria in Phaeosphaeriaceae (Zhang et al. 2009, 2012). However, Hodhod et al. (2012) did not study the type species and the type species of Amarenographium lacks molecular data to confirm the relationships between these genera. Thus we presently maintain the genus in Phaeosphaeriaceae.

Amarenographium metableticum (Trail) O.E. Erikss., Mycotaxon 15: 199 (1982), Faces of Fungi number: FoF00238, Fig. 5.

Fig. 5
figure 5

Amarenographium metableticum (S-F207509). a Specimens and herbarium label of Amarenographium metableticum b Conidiomata visible on host surface. c Section through conidiomata. d Section through conidiomata wall. e–g Conidiogenous cells. h–k Conidia. Scale bars: c = 100 μm, d = 20 μm, e, h, i, j, k = 10 μm, f, g = 5 μm

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≡ Camarosporium metableticum Trail, Scott. Natural., N.S. 2 (‘8’): 267 (1886) [1885–1886]

Saprobic on Ammophila. Sexual state: see notes. Asexual state: Conidiomata 160–215.5 μm high, 150–270 μm diam., pycnidial, scattered, solitary, immersed to erumpent, visible as black spots on host surface, uniloculate, globose to subglobose, glabrous, dark brown, ostiole central, with short to long papilla. Conidiomata walls 13–19 μm wide, thin-walled, of equal thickness, composed of 3–5 layers of flattened, brown to dark brown, pseudoparenchymatous cells, arranged in a textura angularis to textura prismatica. Conidiophores arising from basal cavity of conidioma, mostly reduced to conidiogenous cells, hyaline to brown, septate. Conidiogenous cells (1.5–)3–4(−9) × 1.5–2.5(−5) μm (\( \overline{x} \) = 3.9 × 2.3 μm, n = 10), holoblastic, phialidic, discrete, oblong to ampulliform, hyaline to brown, 1-septate at the base, smooth-walled. Conidia (20–)24–27(−29) × 12–15(−17) μm (\( \overline{x} \) = 25.4 × 13.4 μm, n = 30), solitary, dictyosporous, muriform, ellipsoidal, ovoid, or lageniform, with obtuse ends or sometimes truncate at the basal cell, initially hyaline, becoming brown to yellowish-brown at maturity, mostly 3–4 transverse septa, 1–4 longitudinal septa, with several segments, smooth and thick-walled, with indistinct mucoid appendage.

Material examined: SWEDEN, Öland, Böda parish, Lugnet beach, on Ammophila arenari (L.) Link (Poaceae), 2 June 1963, B. Eriksson & O. Eriksson 2022c (S-F207509).

Amarenomyces O.E. Erikss., Op. bot. 60: 124 (1981), Faces of Fungi number: FoF00235.

Type species: Amarenomyces ammophilae

Phylogenetic study: Zhang et al. (2009, 2012), Hyde et al. (2013)

Amarenomyces ammophilae (Lasch) O.E. Erikss., Op. bot. 60: 124 (1981), Faces of Fungi number: FoF00236, Fig. 6.

Fig. 6
figure 6

Amarenomyces ammophilae (S-F77691). a Specimens and herbarium label b Ascomata visible on host surface. c Section through ascoma. d Section through peridium. e Asci with cellular pseudoparaphyses. f–i Asci. j–m Ascospores. n Ascospore stained in indian ink. Scale bars: c = 100 μm, c, d = 50 μm, e, f, g, h, i = 20 μm, j, k, l, m, n = 10 μm

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≡ Sphaeria ammophilae Lasch Flora, Jena 33: 282 (1850)

Saprobic on Ammophila. Sexual state: Ascomata 170–290 μm high, 170–285 μm diam., scattered, immersed to erumpent through host tissue, visible as small black spots on host surface, uniloculate, globose to subglobose, glabrous, dark brown, ostiole central, with short papilla (50–130 μm long). Peridium 7.5–25 μm wide, thin-walled, of equal thickness, composed of 3–5 layers of flattened, dark brown, pseudoparenchymatous cells, arranged in textura angularis to textura prismatica. Hamathecium composed of numerous, 1.5–3 μm wide, filiform, distinctly septate, frequently anastomosing, narrow cellular pseudoparaphyses, embedded in mucilaginous matrix. Asci (112–)140–160(−173) × (28–)30–35(−37) μm (\( \overline{x} \) = 140 × 32.4 μm, n = 25), 8-spored, bitunicate, fissitunicate, broadly cylindrical, subsessile with minute knob-like pedicel, apically rounded, with a well-developed ocular chamber. Ascospores 39–47(−50) × 12–14 μm (\( \overline{x} \) = 43.4 × 13.2 μm, n = 30), overlapping 1–3-seriate, phragmosporous, broadly fusiform, rarely cylindric-clavate, widest at the middle cell, pale yellowish when immature, becoming yellowish-brown to brown at maturity, 6-septate, slightly constrict at the septum, smooth and thick-walled.

Material examined: BELGIUM, West-Vlaanderen, Blankenbergh, on Ammophila arenaria L. (Poaceae), Westendorp (S-F77691); ITALY, Sardegna, Cagliari, La Maddelena, March 1866, Marcucci (S-F77741).

Notes: Amarenomyces was introduced by Eriksson (1981) to accommodate the single species from a marine grass, which traditionally has been identified as Sphaeria ammophilae Lasch. Eriksson (1981) mentioned that the genus should belong in Botryosphaeriaceae rather than Phaeosphaeriaceae as its morphological characters differed from Phaeosphaeria. Zhang et al. (2009) listed Amarenomyces as a synonym of Phaeosphaeria based on phylogenetic evidence. The generic type of Amarenomyces, A. ammophilae often clusters with other Phaeosphaeria species (Zhang et al. 2009, 2012; Hyde et al. 2013). Thus the earlier name as proposed by Kohlmeyer and Kohlmeyer (1965) and Leuchtmann (1984) was reinstated as Phaeosphaeria ammophilae (Suetrong et al. 2009; Zhang et al. 2009).

Several Phaeosphaeria species have previously been transferred to other genera based on the phylogenetic evidence and due to their asexual state. For example Phaeosphaeria nodorum which was transferred to Parastagonospora, as Pa. nodorum. On the other hand, Shoemaker and Babcock (1989b) divided Phaeosphaeria into six subgenera by combining Phaeosphaeria species with similar morphological characters, but differing in ascospore characters, host preference and asexual state (Câmara et al. 2002; Zhang et al. 2012; Quaedvlieg et al. 2013). The asexual state of Phaeosphaeria has been reported as Phaeoseptoria and this is supported by phylogenetic data (Quaedvlieg et al. 2013). Quaedvlieg et al. (2013) transferred Phaeosphaeria species which did not produce Phaeoseptoria asexual states to other genera. Therefore, the genus Amarenomyces is reinstated in this study based on the phylogenetic analysis as A. ammophilae forms a separate clade to the type species, Phaeosphaeria oryzae. Although A. ammophilae has often been reported to be the sexual state of Amarenographium metableticum (Trail) O.E. Erikss., they are not treated as congeneric here. The presumed link was established as both species occur on the same host in close proximity. There is, however, no sequence data for Amarenographium metableticum and it might be predicted that if both species were host-specific then it would not be surprising that they both occur on the same host. We therefore refrain from combining the taxa until molecular data confirms the link.

Ampelomyces Ces. ex Schlecht., Bot. Zeit. 10: 302 (1852), Faces of Fungi number: FoF00291.

Hyperparasitic on Erysiphales or saprobic. Sexual state: unknown. Asexual state: Conidiomata pycnidial, scattered, solitary, superficial, uniloculate, globose or elongated to pyriform, pale brown, ostiolate, sometimes papillate. Conidiomata walls thick, composed of pale brown cells, arranged in textura angularis. Conidiophores mostly reduced to conidiogenous cells. Conidiogenous cells enteroblastic, phialidic, determinate, discrete, doliiform to ampulliform, hyaline, smooth-walled, channel and collarette minute, periclinal wall towards apex thickened, formed directly from the pycnidial wall cells. Conidia amerosporous, cylindrical to fusiform, straight to curved, very pale brown, aseptate, thin and smooth-walled, with guttules. (Sutton 1980)

Type species: Ampelomyces quisqualis Ces., Bot. Ztg. 10: 301 (1852).

Phylogenetic study: Kiss and Nakasone (1998), Szentiványi et al. (2005), Liang et al. (2007), De Gruyter et al. (2009), Aveskamp et al. (2010), Hyde et al. (2013), Wijayawardene et al.(2013a)

Notes: Ampelomyces species have been reported as mycoparasites on powdery mildews (Erysiphaceae) and are worldwide in nature (de Bary 1870; Griffith 1899; Kiss 1998; Kiss et al. 2004). The genus was introduced by Cesati (1852) to accommodate taxa that formed pycnidia as intracellular mycoparasites on powdery mildews (De Bary 1870; Kiss et al. 2004). Previously, Ampelomyces had varied morphological and cultural characteristic with species differentiated based on pycnidia, cultures and conidial types in isolates of pycnidial mycoparasites from powdery mildews (Clare 1964; Belsare et al. 1980; Kiss et al. 2004). In Kiss et al. (2004), Ampelomyces species could be distinguished into slow and fast growing types. Based on the rDNA ITS region phylogenetic analysis, Kiss et al. (2004) concluded that strains from the two types of differentiated growth rate were not congeneric. The slow-growing isolates, obtained from intracellular pycnidia in powdery mildew mycelia was Ampelomyces sensu stricto. While fast growing isolates from sessile pycnidia on mildew-infected leaves, were closely related to Phoma species. Ampelomyces quercinus (Syd.) Rudakov, A. humuli (Fautrey) Rudakov and A. heraclei (Dejeva) Rudakov are example of fast growing species (Rudakov 1979; Kiss and Nakasone 1998; Sullivan and White 2000). De Gruyter et al. (2009) included two isolates of the type species; A. quisqualis in their phylogenetic analysis and showed that the type species of Ampelomyces can be accommodated in Phaeosphaeriaceae. Whereas Ampelomyces quercinus grouped in Didymellaceae; thus, Aveskamp et al. (2010) introduced A. quercinus as new combination within Phoma in Didymellaceae.

Bricookea M.E. Barr, Mycotaxon 15: 346 (1982), Faces of Fungi number: FoF00239.

Saprobic on Juncaceae. Sexual state : Ascomata scattered or clustered, immersed to semi-immersed, becoming erumpent through host tissue or superficial at maturity, uni- to multi-loculate, globose to subglobose, glabrous, brown to dark brown, ostiole central, with slit-like opening. Peridium thin-walled, composed of brown to dark brown, pseudoparenchymatous cells, arranged in a textura angularis. Hamathecium composed of numerous, filamentous, distinctly septate, broad cellular pseudoparaphyses, anastomosing at the apex. Asci 8-spored, bitunicate, fissitunicate, cylindrical to cylindric-clavate, narrow towards apex, short pedicellate, apically rounded, with well-developed ocular chamber. Ascospores overlapping, 1–3-seriate, phragmosporous, ellipsoidal to broadly fusiform, hyaline, septate, slightly curved, constricted at the median septum, smooth and thick-walled. Asexual state: Unknown.

Type species: Bricookea sepalorum

Phylogenetic study: None

Notes: Barr (1982) introduced Bricookea to accommodate B. sepalorum that was previously identified as Leptosphaeria sepalorum (Vleugel) Lind (Lind 1928). Holm (1957) observed the collections from Sweden and compared the morphological characters with specimens from North America. He mentioned that the ascospores and asci of Swedish collections were smaller than North American specimens and regarded the species as synonym of Lizonia emperigonia (Auersw.) de Not. (Barr 1982; Shoemaker and Babcock 1989a; Zhang et al. 2012). Barr (1982) did not agree with Holm (1957) and designated the new genus, Bricookea to accommodate Leptosphaeria sepalorum. Barr (1982) suggested placing Bricookea in Phaeosphaeriaceae with its uniquely opening ostioles and lack of known asexual state. Barr (1982) mentioned that Sphaerulina inquinans Rehm may be provided as earlier epithet of Bricookea sepalorum. However, Shoemaker and Babcock (1989a) studied the type specimens of Bricookea from S herbaria and mentioned that no material was found under the name Sphaerulina inquinans. Subsequently, Shoemaker and Babcock (1989a) introduced the new species, Bricookea barrae from a North American collection which had been examined by Barr (1982). Bricookea barrae [as Br. barriae] was synonymized under the name Lophiostoma barriae in Lophiostomataceae by Eriksson (2007) leaving the genus monotypic. Zhang et al. (2012) tentatively place Bricookea in Phaeosphaeriaceae. Currently, Bricookea is generally treated in Lophiostomataceae based on its slit-like ostioles, but the type species, Br. sepalorum has been synonymized under the earlier name in Leptosphaeriaceae as Leptosphaeria sepalorum (Index Fungorum 2014).

We examined the collection from North America and type specimen from S herbaria. Both collections have similar morphological characters. However, the specimens from North America differ in having larger ascoma, asci and ascospores than the type specimen from Europe. In this study, Bricookea forms a group in pseudostroma, with slit-like ostioles; characters typical of Lophiostomataceae. However, we tentatively place Bricookea in Pheaosphaeriaceae as its peridium is thin-walled, composed of pseudoparenchymatous cells, asci are clavate with a subsessile to short acute pedicel and ascospores are phragmosporous. Fresh collection, epitypification and molecular analysis are needed to confirm the natural placement.

Bricookea sepalorum (Vleugel) M.E. Barr, Mycotaxon 15: 346 (1982), Faces of Fungi number: FoF00240, Fig. 7.

Fig. 7
figure 7

Bricookea sepalorum (S-F49661, type). a Herbarium label and specimens b Ascomata on substrate. c Section through stroma d Section through peridium. e Asci with cellular pseudoparaphyses strained in cotton blue reagent. f–g Asci. h–i Asci stained in cotton blue reagent. j Ascospores. k–n Ascospores. Scale bars: b = 1000 μm, c = 100 μm, e, f, g, h, i, j = 20 μm, d, k, l, m, n = 10 μm

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≡ Metasphaeria sepalorum Vleugel, Svensk bot. Tidskr. 2(4): 369 (1908)

Saprobic on Juncus spp. Sexual state: Ascomata 130–210 μm high, 170–250 μm diam., scattered to clustered, gregarious, immersed to semi-immersed under epidermis, becoming erumpent through host tissue, to superficial at maturity, visible as black dots in rows or groups on host surface, uni- to multi-loculate, globose to subglobose, glabrous, dark brown to black, ostioles central, with slit-like opening. Peridium 10–20 μm wide, thin-walled, of equal thickness, composed of 2–3 layers of large, brown to dark brown, pseudoparenchymatous cells, arranged in a textura angularis. Hamathecium composed of numerous, 1.5–2 μm wide, filamentous, distinctly septate, cellular pseudoparaphyses, embedded in mucilaginous matrix, anastomosing at the apex. Asci (60–)65–75(−80) × 12–14(−16) μm (\( \overline{x} \) = 71.5 × 13.5 μm, n = 30), 8-spored, bitunicate, fissitunicate, cylindric-clavate, narrowing towards the apex, with subsessile to shortly acute pedicel, apically rounded with well-developed ocular chamber. Ascospores 17.5–19(−21) × 6–7.5 μm (\( \overline{x} \) = 19 × 6.4 μm, n = 30), overlapping 1–2-seriate, phragmosporous, ellipsoidal to broadly fusiform, hyaline to subhyaline, initially forming one median septate, becoming 3-septate at maturity, slightly curved, constricted at the median septum, easily breaking into part spores at the median septum, smooth and thin to thick-walled. Asexual state: Unknown.

Material examined: SWEDEN, Suecia. Holmön, Västerbotten, on dead sepals of Juncus filiformis Linn. (Juncaceae), July 1907, J. Vleugel (S-F49661, type); USA, California, Squaw Valley Creek Meadows, Mt. Shasta, Siskiyou Co. alt. 8,500 ft., on dead sepals of Juncus parryi Engelm. (Juncaceae), 10 July 1947, W. B. Cooke & V. G. Cooke (BPI 737398); Washington, Mt. Baker National Forest, Whatcom Co. Subalpine Meadows along Washington Route #1 near Baker Lodge, on dead sepals of Juncus parryi Engelm. (Juncaceae), 12 July 1948, W. B. Cooke & V. G. Cooke 24247 (BPI 737397); California, King Creek Trail, Lassen Volcanic National Park. alt. 8,000 ft., on dead sepals of Juncus parryi Engelm. (Juncaceae), 1 July 1954 W. B. Cooke & V. G. Cooke 29530 (BPI 737400); California, King Creek Trail, Lassen Volcanic National Park. alt. 8,000 ft., on dead sepals of Juncus parryi Engelm. (Juncaceae), 1 July 1954, W. B. Cooke & V. G. Cooke 29530 (BPI 737399); California, Panther Creek Meadows, Mt. Shasta, Siskiyou Co. alt. 7,600 ft., on dead sepals of Juncus parryi Engelm. (Juncaceae), 4 July 1948, W. B. Cooke 24055 (BPI 737401).

Chaetosphaeronema Moesz, Bot. Közl. 14: 152 (1915), Faces of Fungi number: FoF00241.

Saprobic or pathogenic on Malus, Euphorbia, Anthyllis and Dorycnium spp. Asexual state: Unknown Conidiomata pycnidial, scattered, immersed to semi-immersed, uniloculate, globose, setose, brown to dark brown, ostiole central, with setose papilla. Conidiomata walls thin, arranged in a textura prismatica, pale brown, outer layers comprising dark brown, thick-walled cells. Conidiophores arising from basal cavity of conidiomata mostly reduced to conidiogenous cells. Conidiogenous cells enteroblastic, phialidic, discrete or integrated, cylindrical, hyaline, smooth-walled, with minute channel or collarette, hardly distinguishable from the inner wall cells. Conidia didymosporous, oblong to cylindrical, with obtuse ends, straight or slightly curved, hyaline, 1-septate, thin and smooth-walled, with guttules (from Sutton 1980; Quaedvlieg et al. 2013)

Type species: Chaetosphaeronema hispidulum (Corda) Moesz, Bot. Közl. 14: 152 (1915)

= Sphaeronaema hispidulum Corda, Icon. fung. (Prague) 4: 39 (1840)

Phylogenetic study: Schoch et al. (2009), De Gruyter et al. (2009, 2010), Zhang et al. (2009, 2012), Hyde et al. (2013), Quaedvlieg et al. (2013).

Notes: Chaetosphaeronema was introduced by Moesz (1915) as typified by Ch. hispidulum to accommodate species of Sphaeronaema that were not congeneric. A partial morphological treatment of the genus was provided by Sutton (1980). Petrak (1944) treated Ch. herbarum (Hollós) Moesz as a synonym of Ch. hispidulum and accepted three species in the genus; Ch. collivagum (Petr.) Petr., Ch. galii (Petr.) Petr. and Ch. hispidulum (Sutton 1980). However, Chaetosphaeronema was subsequently limited to the type species (De Gruyter et al. 2009, 2010; Zhang et al. 2012; Hyde et al. 2013; Quaedvlieg et al. 2013). De Gruyter et al. (2009) showed, using molecular data that Chaetosphaeronema was related to Phaeosphaeriaceae and Pleosporaceae. De Gruyter et al. (2010) introduced a new species, Ch. coonsii (Boerema & Loer.) Gruyter et al., and treated Chaetosphaeronema sensu stricto in Phaeosphaeriaceae. This placement has been widely accepted by subsequent authors based on the phylogenetic evidence, while no sexual state is presently known (Schoch et al. 2009; Zhang et al. 2009, 2012; Hyde et al. 2013; Quaedvlieg et al. 2013).

In this study, Chaetosphaeronema clusters with various sexual genera and is closely related to Ophiobolus cirsii (MFLUCC13-0218) as propose by Petrak (1944) and Zhang et al. (2009) who suggested that Chaetosphaeronema may be asexual state of Ophiobolus. The link between these genera has not yet equivocally been proven. Epitypification with molecular data is therefore needed to confirmed the natural placement of these genera and confirm asexual-sexual connections.

Dematiopleospora Wanasinghe et al., Cryptog. Mycol. 35 (2): 105-11(2014), Faces of Fungi number: FoF00242.

Saprobic on Ononis spinosa L (Fabaceae). Sexual state: Ascomata scattered, solitary, superficial, uniloculate, globose to subglobose, glabrous, dark brown to black, coriaceous, cupulate when dry, ostiole central, minute papilla, with thick, brown, periphyses. Peridium thin-walled, composed of pseudoparenchymatous cells, outer layer comprising heavily pigmented, thick-walled, reddish to dark brown cells, arranged in textura angularis, inner layer comprising 2–3 cell layers of thin-walled, hyaline cells, arranged in a textura angularis. Hamathecium composed of numerous, filamentous, distinctly septate, broad cellular pseudoparaphyses. Asci 8-spored, bitunicate, fissitunicate, cylindrical to cylindric-clavate, short pedicellate, apically rounded with minute ocular chamber. Ascospores overlapping 1–2-seriate, dictyosporous, muriform, ellipsoidal to subfusiform, widest in the upper part, slightly curved, 5–9 transverse septa, with 3–6 longitudinal septa, constricted at the central septum, initially hyaline, becoming yellowish-brown at maturity, smooth-walled. Asexual state: Unknown (from Wanasinghe et al. 2014)

Type species: Dematiopleospora mariae Wanasinghe et al., Cryptog. Mycol. 35 (2): 105-11(2014)

Phylogenetic study: Wanasinghe et al. (2014).

Notes: Dematiopleospora was introduced by Wanasinghe et al. (2014) to accommodate D. mariae. The genus is characterized by thick, brown, periphyses in the ostiole, superficial ascomata and muriform ascospores. Dematiopleospora is similar to Nodulosphaeria in its periphysate ostiole, but they differ in peridium structure and ascospore morphology. Dematiopleospora has muriform ascospores similar to those characterized in Phaeosphaeria vagans (Niessl) O.E. Erikss., Ph. phragmiticola Leuchtm., Ph. phragmitis (Hollós) Leuchtm and Pleoseptum yuccaesedum A.W. Ramaley & M.E. Barr (Shoemaker and Babcock 1989b; Zhang et al. 2012; Wanasinghe et al. 2014). Dematiopleospora is similar to Pleoseptum, but they differ in peridium structures and papillate ostioles. Pleoseptum lacks of molecular data to confirm its natural placement. Therefore, the genus is tentatively placed in Pleosporaceae based on its muriform ascospores and thick-walled peridium that are dominant characters in Pleosporaceae. Multigene phylogenetic analyses indicated that Dematiopleospora belongs to Phaeosphaeriaceae which forms a separate clade from those Phaeosphaeria species and groups with Chaetosphaeronema, Entodesmium, Loratospora and Nodulosphaeria. No asexual morph is known for Dematiopleospora.

Dothideopsella Höhn., Sber. Akad. Wiss. Wien, Math.-naturw. Kl., Abt. 1 124: 70 (1915), Faces of Fungi number: FoF00295

Saprobic on dead wood and stems in terrestrial habitats. Sexual state: Ascomata scattered, gregarious, immersed to semi-immersed, or erumpent, nearly superficial, globose, uniloculate, subglobose or obypyriform, glabrous, dark brown to black, ostiole central, broadly or narrowly conical, with a dark brown to black papilla. Peridium thin to thick, of unequal thickness, slightly thick at the basal ascoma, composed of dark brown to black, thin-walled, pseudoparenchymatous cells, arranged in a textura angularis. Hamathecium composed of numerous, filamentous, distinctly septate, cellular pseudoparaphyses, anastomosing without a gelatinous coating. Asci 8-spored, bitunicate, fissitunicate, cylindrical to oblong, with a knob like pedicel, apically rounded with well-developed ocular chamber Ascospores phragmosporous, ellipsoidal to fusiform, brown or yellowish brown, septate, slightly constricted at the septum, smooth-walled. Asexual state: Unknown (from Ariyawansa et al. 2013).

Type species: Dothideopsella agminalis (Sacc. & Morthier) Höhn., Sber. Akad. Wiss. Wien, Math.-naturw. Kl., Abt. 1 124: 70 (1915)

Phylogenetic study: None

Notes: Dothideopsella was introduced by Höhnel (1915) and is typified by Dothideopsella agminalis (Sacc. & Morthier) Höhn. The genus was established to accommodate Dothideomycetes species characterized by a broad peridium structure, with conical, papillate ostioles, and phragmosporous, brown ascospores (Ahn and Shearer 1998; Ariyawansa et al. 2013). Lumbsch and Huhndorf (2010) treated Dothideopsella in Dothideomycetes, genera incertae sedis, while Index Fungorum (2014) lists the genus in Leptosphaeriaceae. Ariyawansa et al. (2013) tentatively placed Dothideopsella in Phaeosphaeriaceae based on its morphological characters which are typical of Chaetoplea. Dothideopsella however differs from Chaetoplea in having a peridium composed of pseudoparenchymatous cells, while the peridium in Chaetoplea is composed of scleroplectenchymatous cells. The placement of the genus in Phaeosphaeriaceae was supported by Wijayawardene et al. (2014b). Thus we tentatively place Dothideopsella in Phaeosphaeriaceae until molecular date proves otherwise.

Entodesmium Riess, Hedwigia 1(6): 28 (1854), Faces of Fungi number: FoF00243.

Saprobic or parasitic? on various dicotyledonous and some monocotyledonous hosts. Sexual state: Ascomata scattered, solitary to gregarious, immersed to erumpent through host tissue with long neck, uniloculate, globose to subglobose, glabrous, dark brown to black, ostiole central, with long neck. Neck with thick walls of textura angularis, dark brown to black, cylindrical to cylindric-clavate, carbonaceous, with periphyses. Peridium thick-walled, composed of brown to dark brown, pseudoparenchymatous cells, arranged in a textura angularis. Hamathecium composed of numerous, filamentous, distinctly septate, broad cellular pseudoparaphyses, not constricted at the septa and anastomosing at the apex. Asci 8-spored, bitunicate, cylindrical to cylindric-clavate, short pedicellate, apically rounded, with well-developed ocular chamber. Ascospores fasciculate, scolecosporous, filiform or filamentous, initially hyaline to pale brown, becoming brown or reddish-brown at maturity, multi-septate, constricted at the septa, separating into numerous part spores, smooth-walled. Asexual state: Unknown.

Type species: Entodesmium rude

Phylogenetic study: Liew et al. (2000), Schoch et al. (2009), Zhang et al. (2009, 2012)

Notes: Entodesmium was introduced by Riess (1854) and is typified by E. rude. The genus was introduced to accommodate a Dothideomycete species with long beaked ascomata and scolecosporous ascospores, which break into numerous part-spores and occur on legumes. The placement of Entodesmium was previously unclear as Von Arx and Müller (1975) disposed the genus in Pleosporaceae, while Barr (1987b) suggested that Entodesmium should be placed in Phaeosphaeriaceae and this was accepted by Eriksson and Hawksworth (1991). Barr (1992b) assigned Entodesmium to Lophiostomataceae based on its short, blackish beak and periphysate ostiole. Barr (1992b) mentioned that these characters are more typical of Lophiostomataceae. Zhang et al. (2009, 2012) disagreed with Barr (1992b) and mentioned that the morphological characters of Entodesmium best fit in Phaeosphaeriaceae.

Entodesmium was thought to be family-specific and associated only with legumes (Barr 1992b; Shoemaker 1984; Zhang et al. 2009, 2012). Holm (1957) and Shoemaker (1984) re-described Entodesmium and reported six species associated with legumes from Europe; although, Wehmeyer (1952) identified some collections from North America (Barr 1992b). Furthermore, there is a report of E. niesslianum from Dieffenbachia (monocotyledon) in West Indies by Minter et al. (2001) in Farr and Rossman (2014). Six species are presently accommodated in Entodesmium; E. eliassonii L. Holm, E. lapponicum (L. Holm) L. Holm, E. mayorii (E. Müll.) L. Holm, E. nevadense Petr., E. niesslianum (Rabenh. ex Niessl) L. Holm and E. rude (Index Fungorum 2014). Entodesmium multiseptatum (G. Winter) L. Holm was assigned to Leptosphaeriaceae and Entodesmium vermisporum (Ellis) M.E. Barr (1992b) was treated in Lophiostomataceae (Winter 1872; Barr 1992b; Zhang et al. 2009, 2012; Index Fungorum 2014). However, Zhang et al. (2009, 2012) mentioned that E. multiseptatum and E. niessleanum are more similar to Phaeosphaeria than Leptosphaeria or Entodesmium.

Zhang et al. (2009) accepted the scolecosporous genera Entodesmium and Ophiosphaerella in Phaeosphaeriaceae based on phylogenetic evidence. Multigene phylogenetic analysis based on this sequence data has indicated that Entodesmium belongs to Phaeosphaeriaceae which has been widely accepted (Schoch et al. 2009; Zhang et al. 2009, 2012; Hyde et al. 2013).

In this study, the iconotype is re-drawn while the type specimens could not be located. Based on morphology (thin-walled peridium composed of pseudoparenchymatous cells, and scolecosporous, brown, multi-septate ascospores breaking into part-spores) and multigene phylogenetic analysis, we retain Entodesmium in Phaeosphaeriaceae. Entodesmium forms a weakly-supported clade with other genera (Chaetosphaeronema, Dematiopleospora, Loratospora, Nodulosphaeria modesta, Ophiobolus cirsii and O. erythrosporus) in Phaeosphaeriaceae. However, the sequence data is from a putative strain of Entodesmium rude from CBS, thus epitypification and molecular work is needed to confirm its natural placement.

Entodesmium rude Riess, Hedwigia 1(6): 28 (1854), Faces of Fungi number: FoF00244, Fig. 8, and 9.

Fig. 8
figure 8

Entodesmium rude (BR 5020100367693). a Herbarium packet and specimens. b Ascomata on substrate. c Section through ascomata. d Section through neck. e Section through peridium. f Asci with pseudoparaphyses. g–i Asci. Scale bars: c = 100 μm, d = 50 μm, e, f, g, h, i = 20 μm

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Fig. 9
figure 9

Entodesmium rude (iconotype) redrawn from Riess (1854). a Ascomata on host surface. b Section through ascomata. c Ascus with pseudoparaphyses. d Ascospores

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Saprobic on Astragalus, Coronilla and Lathyrus spp. Sexual state: Ascomata 160–240 μm high (excluding neck), 160–255 μm diam., scattered, solitary to gregarious, immersed to erumpent through host tissue with long neck, visible as black dots, in rows or groups on host surface, uniloculate, globose to subglobose, glabrous, dark brown to black, ostiole oblique, with long carbonaceous neck. Neck 175.5–295 μm high, 90–150 μm diam., thick-walled, cylindrical to cylindric-clavate, arranged in a textura angularis, dark brown to black, carbonaceous, with periphyses. Peridium 15–30 μm wide, thick-walled, of equal thickness, comprising two layers, outer layer comprising 3–5 cell layers of brown to dark brown, thick-walled cells, arranged in textura angularis to textura globulosa, inner layers comprising 1–2 cell layers of thin-walled, hyaline, flattened cells, arranged in textura angularis to textura prismatica. Hamathecium composed of numerous, 2–3 μm wide, filiform, distinctly septate, broad cellular pseudoparaphyses, embedded in mucilaginous matrix, anastomosing at the apex. Asci (124–)130–145(−156) × (11–)13–15(−18) μm (\( \overline{x} \) = 137.4 × 14.4 μm, n = 25), 8-spored, bitunicate, broadly cylindrical to cylindric-clavate, short pedicellate, apically rounded with well-developed ocular chamber. Ascospores (100–)110–120(−135) × 3–4 μm (\( \overline{x} \) = 115.5 × 3.5 μm, n = 25), fasciculate, scolecosporous, filiform, initially hyaline to pale brown, becoming brown to reddish-brown at maturity, multi-septate, with 18–20 septa, constricted at the septa, easily separating into part spores, smooth-walled.

Material examined: GERMANY, on stem of Astragalus glycyphyllos, December 1853, H. Riess, Hedwigia 1(6): Tab IV, (Fig 4., iconotype) (1854); CZECHOSLOVAKIA, Usti, on Astragalus glycyphyllos L. (Fabaceae), 6 May 1922, F. Petrak (BR 5020100367693); CZECHOSLOVAKIA, Moravia, Hranice (Maehr.-weisskirchen), on branches of Astragalus glycyphyllos L. (Fabaceae), July 1924, F. Petrak (BPI 626664).

Eudarluca Speg., Revta Mus. La Plata 15: 22 (1908), Faces of Fungi number: FoF00245. Hyperparasitic associated with rust species (but not necessarily parasitic) on various hosts. Sexual state: Ascomata scattered, solitary to gregarious, immersed to semi-immersed under pseudoclypeus, uni- to multi-loculate, subglobose to irregular in shape, glabrous, dark brown to black, ostiole central, with minute papilla. Peridium thin-walled, composed of brown to dark brown, flattened, pseudoparenchymatous cells, arranged in a textura angularis. Hamathecium composed of numerous, filamentous, distinctly septate, frequently anastomosing, broadly cellular pseudoparaphyses. Asci 8-spored, bitunicate, fissitunicate, cylindrical, short pedicellate, apically rounded, with well-developed ocular chamber. Ascospores phragmosporous, overlapping 2-seriate, fusiform or oblong with rounded ends, pale brown, septate, smooth-walled. Asexual state: see notes.

Type species: Eudarluca australis

Phylogenetic study: None

Notes: Eudarluca was introduced by Spegazzini (1908) and typified by Eu. australis which is associated with a rust on leaves of Canna. Eriksson (1966) treated Eu. australis as a synonym of Eudarluca caricis (Fr.) O.E. Erikss. Eudarluca caricis has been reported from various hosts, mostly in its presumed asexual state, Sphaerellopsis (Yuan et al. 1998; Nischwitz et al. 2005; Zhang et al. 2012). The asexual and sexual state connection between Eudarluca caricis and Sphaerellopsis filum was derived from ascospores growing on Puccinia extensicola-oenotherae (Mont.) Arthur on Carex sp. in Pennsylvania, USA (Keener 1951) and confirmed by Yuan et al. (1998). Yuan et al. (1998) grew ascospores of Eudarluca caricis on PDA and derived Sphaerellopsis conidia and conidiomata after 12 days. Zhang et al. (2012) compared the ITS sequence of Eudarluca caricis strain MullMK in GenBank with Leptosphaeria species and mentioned that Eudarluca should be treated in Leptosphaeriaceae due to comparable sequences and also morphological characters. However, both the generic type of Eudarluca and Sphaerellopsis have never been linked (Wijayawardene et al. 2014b).

Molecular studies on Eudarluca are limited to species variation and are based on Sphaerellopsis strains (Nischwitz et al. 2005; Bayon et al. 2006; Zhang et al. 2012). The ITS gene phylogenetic analysis showed that Eudarluca ciricis (strains MosID6 and MullMK) cluster in Leptosphaeriaceae. However, the generic type of Sphaerellopsis has found on Quercus, while the type of Eudarluca is from Canna. These two generic types have not been linked by molecular data. We suggest that caution should be followed when treating these taxa, as Eu. australis is immersed in the Canna leaves and although associated with the rust fungi (Sphaerellopsis) on Canna, it does not seem to grow on the rust.

In this study, we tentatively place Eudarluca in Phaeosphaeriaceae based on its morphological characters which are typical of Phaeosphaeria as the peridium is thin-walled, and composed of pseudoparenchymatous cells, asci are cylindrical, subsessile to short pedicellate and ascospores are fusiform and phragmosporous. Fresh collections of Eu. australis are needed for epitypification and molecular study. Sphaerellopsis may belong in Leptosphaeriaceae where it is tentatively assigned.

Eudarluca australis Speg., Revta Mus. La Plata 15: 22 (1908), Faces of Fungi number: FoF00246, Fig. 10.

Fig. 10
figure 10

Eudarluca australis (S-F11420, isotype). a Herbarium label and specimens. b Fruiting bodies on host surface. c Section through ascoma. d Section through peridium. e Pseudoparaphyses. f Asci with pseudoparaphyses. g–j Asci. k–o Ascospores. Scale bars: c = 100 μm, d, f, g, h, i, j = 20 μm, e = 10 μm, k, l, m, n, o, = 5 μm

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Hyperparasitic associated with rust fungi on leaves of Canna indica L. (Cannaceae). Sexual state: Ascomata 110–180 μm high, 170–280 μm diam., scattered, solitary to gregarious, immersed to semi-immersed, visible as raised, black tiny spots in black pseudostromata on leaf lesions, uniloculate, subglobose to irregular, glabrous, dark brown to black, ostiole central, with minute papilla. Peridium 5–13 μm wide, thin-walled, of unequal thickness, slightly thickened at the apex, composed of 1–3 layers of brown to dark brown, flattened, pseudoparenchymatous cells, arranged in textura angularis. Hamathecium composed of numerous, 1.5–3 μm wide, filiform, distinctly septate, frequently anastomosing, broadly cellular pseudoparaphyses, embedded in mucilaginous matrix. Asci (44–)50–65 × 8–10 μm (\( \overline{x} \) = 57 × 9 μm, n = 25), 8-spored, bitunicate, fissitunicate, cylindrical, short pedicellate or subsessile, apically rounded, with indistinct ocular chamber. Ascospores 15–17(−18) × 3–4.5 μm (\( \overline{x} \) = 16.7 × 4.1 μm, n = 30), phragmosporous, overlapping or 2-seriate, fusiform, or oblong with rounded ends, pale brown, 2-septate, rarely 1-septate, slightly curved, not constricted at the septa, smooth-walled. Asexual state: see notes.

Material examined: BRAZIL, São Paulo, Botanical Gardens, associated with rust (Pucciniaceae) on leaves of Canna indica L., 10 August 1905, A. Usteri (S-F11420, isotype).

Loratospora Kohlm. & Volkm.-Kohlm., Syst. Ascom. 12(1–2): 10 (1993), Faces of Fungi number: FoF00247.

Type species: Loratospora aestuarii

Phylogenetic study: Schoch et al. (2009), Suetrong et al. (2009), Zhang et al. (2012), Hyde et al. (2013).

Loratospora aestuarii Kohlm. & Volkm.-Kohlm., Syst. Ascom. 12(1–2): 10 (1993), Faces of Fungi number: FoF00248, Fig. 11.

Fig. 11
figure 11

Loratospora aestuarii a Herbarium label and specimens (Kohlmeyer_5505). b, c Ascomata on host surface (Kohlmeyer_5505, Kohlmeyer_5520). d Section through ascoma (Kohlmeyer_5505). e Section through peridium (Kohlmeyer_5505) f–g Asci (Kohlmeyer _5505). h Asci embedded in a gelatinous matrix (Kohlmeyer_5520). i–j Asci (Kohlmeyer_5520). k Ascospores (Kohlmeyer_5520) l–n Ascospores (Kohlmeyer_5520). o Ascospore (Kohlmeyer_5505). Scale bars: d = 100 μm, e = 50 μm, f, g, h, I, j, k = 20 μm, l, m, n, o = 10 μm

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Saprobic on Juncus roemerianus Scheele. Sexual state: Ascomata 130–250 μm high, 160–310 μm diam., scattered to clustered, solitary, immersed to erumpent through host surface, visible as small black spots on host surface, uni to biloculate, globose to subglobose, glabrous, brown to dark brown, ostiole central, with minute papilla, with brown periphyses. Peridium 18–42 μm wide, thin-walled, of unequal thickness, thickened at the apex, composed of several layers of brown to dark brown, pseudoparenchymatous cells, arranged in textura angularis to textura globulosa. Hamathecium absent, visible as mucilaginous matrix between asci. Asci (60–)70–85(−110) × (23–) 25–30(−35) μm (\( \overline{x} \) = 80.9 × 28.2 μm, n = 20), 8-spored, bitunicate, fissitunicate, ovoid to ampulliform, thick-walled at the apex, sessile to subsessile, apically rounded with indistinct ocular chamber, sparse in mucilaginous matrix. Ascospores 40–50(−55) × 5–7(−8) μm (\( \overline{x} \) = 46.7 × 6.7 μm, n = 30), overlapping or parallel, 4–5-seriate, phragmosporous, cylindrical, or narrowly elongate fusiform with rounded ends, initially hyaline to subhyaline, pale yellowish at maturity, 3-septate, slightly curved, not constricted at the septa, rough-walled, surrounded by a thin, distinct sheath. Asexual state: Unknown.

Material examined: USA, North Carolina, on dead culms of Juncus roemerianus (Juncaceae), Kohlmeyer_5505 (holotype), 5520, 5523, 5525, 5535 (NY_Slide box: 17, Tray no. 21); ibid. Broad Creek (Atlantic Ocean) 34° 43′ 00″N, 76° 55′ 07″W (34.717, −76.919), standing dead culms of Juncus roemerianus, J. Kohlmeyer 5520 with B. Kohlmeyer, 13 October 1993 (NY 01276343).

Notes: The monotypic genus Loratospora was established by Kohlmeyer and Volkmann-Kohlmeyer (1993) to accommodate L. aestuarii. The taxon occurs on dead culms of Juncus roemerianus (Jones et al. 2009; Suetrong et al. 2009; Monkai et al. 2013). Kohlmeyer and Volkmann-Kohlmeyer (1993) mentioned that the species often developed on culms which were not regularly submerged in salt water. Loratospora is similar to Leptosphaerulina (Didymellaceae), Monascostroma (Didymellaceae) and Wettsteinina (Dothideomycetes genera incertae sedis), but Leptosphaerulina and Monascostroma have smaller ascomata, lack periphyses, and have a thin-walled peridium. Ascospores in Leptosphaerulina are muriform and didymosporous in Monascostroma (Kohlmeyer and Volkmann-Kohlmeyer 1993; Jones et al. 2009). Loratospora forms large ascomata with brown ostiolar periphyses, a thick-walled peridium at the apex and phragmosporous, cylindrical ascospores (Kohlmeyer and Volkmann-Kohlmeyer 1993; Jones et al. 2009; Suetrong et al. 2009). Wettsteinina differs from Loratospora in lacking ostiolar periphyses, in having a thick-walled peridium of equal width and ellipsoidal to broadly fusiform ascospores (Kohlmeyer and Volkmann-Kohlmeyer 1993)

Loratospora was re-circumscribed by Jones et al. (2009) and Monkai et al. (2013). The genus was accommodated in Planistromellaceae by Barr (1996) due to its periphysate ostiole (Jones et al. 2009; Suetrong et al. 2009; Monkai et al. 2013). Based on multigene phylogenetic analysis, Loratospora was placed in Phaeosphaeriaceae (Schoch et al. 2009; Zhang et al. 2012; Hyde et al. 2013).

In the multigene phylogenetic analyses (Fig. 2), Loratospora forms a strongly supported clade with Chaetosphaeronema, Dematiopleospora, Entodesmium, Nodulosphaeria and Ophiobolus. However, the genus has a differentiated centrum type as compared with other genera. Loratospora lacks of pseudoparaphyses but has an internal mucilaginous hamathecium, while most genera in Phaeosphaeriaceae have pseudoparaphyses. It seem like the centrum-type is not significant to distinguish genera at the family level. There is only one species listed in Loratospora in Index Fungorum (2014) and no asexual state is known. The sequences data of Loratospora is limited to only one strain.

Neosetophoma Gruyter et al., in Gruyter et al. Mycologia 102(5): 1075 (2010), Faces of Fungi number: FoF00249.

Pathogenic on the grasses, Phlox paniculata L. and Urtica dioica L., or associated with soil. Sexual state: Unknown. Asexual state: Conidiomata pycnidial, scattered, solitary or gregarious, superficial on agar, uniloculate, globose or irregular, setose, olivaceous to dark olivaceous, ostiole central, with papilla or long neck. Conidiomata walls composed of pseudoparenchymatous cell. Conidiophores reduced to conidiogenous cells. Conidiogenous cells phialidic, discrete, hyaline. Conidia ellipsoidal to cylindrical, attenuate at one end, yellowish, aseptate or septate, not constricted at the septum, smooth-walled, with guttules (from De Gruyter et al. 2010; Quaedvlieg et al. 2013).

Type species: Neosetophoma samararum (Desm.) Gruyter et al. [as ‘samarorum’], in De Gruyter et al. Mycologia 102(5): 1075 (2010).

= Phoma samararum Desm., Pl. Crypt. Nord France, Edn 1 7: no. 349 (1828)

Phylogenetic study: De Gruyter et al. (2010, 2012), Zhang et al. (2012), Hyde et al. (2013), Quaedvlieg et al. (2013).

Notes: Neosetophoma was introduced by De Gruyter et al. (2010) to accommodate N. samararum. Neosetophoma often forms a single clade in Phaeosphaeriaceae. The genus has been widely accepted in Phaeosphaeriaceae by various authors (De Gruyter et al. 2010, 2012; Zhang et al. 2012; Hyde et al. 2013; Quaedvlieg et al. 2013).

Neostagonospora Quaedvlieg et al., Stud. Mycol. 75: 364 (2013), Faces of Fungi number: FoF00250.

Pathogenic on Carex acutiformis Ehrh and Elegia cuspidate Mast., or soil borne. Sexual state: Unknown. Asexual state: Conidiomata pycnidial, scattered, semi-immersed to superficial, uniloculate, globose, glabrous, producing pale luteous to creamy conidial mass on host surface. Conidiomata walls thin, composed of 2–3 layers of pale brown cells, arranged in a textura angularis. Conidiophores reduced to conidiogenous cells. Conidiogenous cells phialidic, integrated, ampulliform to doliiform, hyaline, smooth-walled, lining the inner cavity, with prominent periclinal thickening. Conidia narrow fusoid-ellipsoidal to subcylindrical, apically subobtuse to rounded, truncate at the base, hyaline, septate, initially non-constricted at the septum, becoming constricted at the septa when mature, smooth-walled, with granules (from Quaedvlieg et al. 2013).

Type species: Neostagonospora caricis Quaedvlieg et al., Stud. Mycol. 75: 364 (2013).

Phylogenetic study: Quaedvlieg et al. (2013).

Notes: The asexual genus Neostagonospora was introduced by Quaedvlieg et al. (2013) to accommodate the stagonospora-like species in Phaeosphaeriaceae. Two species were placed in this genus; Neostagonospora caricis and Ne. elegiae Quaedvlieg et al. with Ne. caricis being the type species. Neostagonospora is similar to Stagonospora but differs in its conidiogenous cells. Neostagonospora has determinate phialidic conidiogenous cells, with prominent periclinal thickening while Stagonospora has percurrent proliferation of conidiogenous cells. Based on phylogenetic analysis, Neostagonospora is related to Phaeosphaeriaceae, while the type species of Stagonospora, St. paludosa clusters in Massarinaceae (Quaedvlieg et al. 2013). In the phylogeny in Fig. 2, Neostagonospora forms a sister clade with Parastagonospora for which no sexual state is known. Phaeosphaeria sensu lato taxa are polyphyletic and therefore, it is likely that Neostagonospora may be an asexual state of phaeosphaeria-like species.

Nodulosphaeria Rabenh., Klotzschii Herb. Viv. Mycol., Edn 2: no. 725 (in sched.) (1858), Faces of Fungi number: FoF00251.

Saprobic or hemibiotrophic mostly on herbaceous dicotyledons. Sexual state: Ascomata scattered, solitary to gregarious, immersed to semi-immersed, or erumpent through host epidermis at maturity, visible as black dots on host surface, uniloculate, globose to subglobose, setose, covered by hyphae, dark brown to black, ostiole central, papillate, with broad setae in ostiolar canal, towards the apex. Peridium thin- to thick-walled, of equal thickness, composed of two types of brown to dark brown, pseudoparenchymatous cells, arranged in a textura angularis to textura globulosa. Hamathecium composed of numerous, filamentous, anastomosing, broad cellular pseudoparaphyses, with distinct septa. Asci 8-spored, bitunicate, fissitunicate, clavate to cylindric-clavate, subsessile to short pedicellate, apically rounded with indistinct ocular chamber. Ascospores overlapping 2–4-seriate, phragmosporous, cylindrical or elongated fusiform, brown to yellowish-brown, septate, with an enlarged cell near apex, smooth-walled. Asexual state: Unknown.

Type species: Nodulosphaeria hirta

Phylogenetic study: this study.

Notes: Nodulosphaeria was introduced by Rabenhorst (1858) to accommodate Dothideomycete species forming phragmosporous, brown, septate ascospores with an enlarged cell, and setose ascomata with broad setae in the papilla apex, mostly found on herbaceous dicotyledonous especially Asteraceae (Shoemaker 1984; Zhang et al. 2012). Nodulosphaeria has been reported from various hosts and may be host-specific with more than 60 species named, however it is lacking in molecular data (Shoemaker 1984). The genus was previously treated as a synonym of Leptosphaeria, but later reinstated by Holm (1957) as typified by N. hirta (Clements and Shear 1931; Zhang et al. 2012). Nodulosphaeria was treated in Phaeosphaeriaceae by Barr (1979) based on its morphological characters which are similar to Ophiobolus as the ascospores form an enlarged cell(s). The genera have often been confused and the identification of species can be subjective (Zhang et al. 2012). Shoemaker (1976) re-circumscribed the genus Nodulosphaeria and transferred various species to Ophiobolus. Later many species were added to Nodulosphaeria based on morphology by various authors, but no asexual morph is known (Shoemaker 1984; Shoemaker and Babcock 1987; Zhang et al. 2012). There are 64 epithets listed under Nodulosphaeria and unfortunately no sequences available for this genus in GenBank.

Multigene phylogenetic analyses (Figs. 1 and 2) indicate that Nodulosphaeria belongs in Phaeosphaeriaceae. The genus clusters with Chaetosphaeronema, Dematiopleospora, Entodesmium, Loratospora and Ophiobolus.

Nodulosphaeria hirta Rabenh., Klotzschii Herb. Viv. Mycol., Edn 2: no. 725 (in sched.) (1858), Faces of Fungi number: FoF00252, Fig. 12.

Fig. 12
figure 12

Nodulosphaeria hirta (S-F70410, type). a Herbarium label and specimen of Nodulosphaeria hirta. b Ascostromata on host surface. c Section through ascostroma. d Section through peridium. e Asci with cellular pseudoparaphyses. f–i Asci. j–n Ascospores. Scale bars: c = 100 μm, d, e, f, g, h, i = 20 μm, j, k, l, m, n = 10 μm

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Saprobic on herbaceous dicotyledons. Sexual state: Ascomata 230–380 μm high (including papilla), 225–395 μm diam., scattered, solitary to gregarious, semi-immersed, or erumpent through host epidermis at maturity, visible as black dots on host surface, uniloculate, globose to subglobose, setose, covered by hyphae, dark brown to black, ostiole central, papillate, with broad setae in ostiolar canal, towards the apex. Peridium 21–44 μm wide, thin, of equal thickness, composed of several layers of brown to dark brown, pseudoparenchymatous cells, outer layer comprising 1–2 layers of brown to dark brown, thickened cells, arranged in textura angularis to textura globulosa, inner layers comprised 2–3 layers of brown to dark brown, flattened cells, arranged in textura angularis. Hamathecium composed of numerous, 3–5 μm wide, filamentous, broad cellular pseudoparaphyses, with distinct septa, embedded in mucilaginous matrix, anastomosing at the apex. Asci (83–)90–110(−115) × (13.5–)15–20 μm (\( \overline{x} \) = 100 × 16.8 μm, n = 25), 8-spored, bitunicate, fissitunicate, clavate to cylindric-clavate, subsessile or short pedicellate, apically rounded with well-developed ocular chamber. Ascospores 48–55 × 5–7(−7.5) μm (\( \overline{x} \) = 51.2 × 6.5 μm, n = 30), overlapping 3–4-seriate, phragmosporous, cylindrical or elongated fusiform, with narrow ends, brown to yellowish-brown, 8-septate, slightly curved, smooth-walled, enlarged at the forth cell from the apex. Asexual state: Unknown.

Material examined: GERMANY, Sachsen, Dresdae, on dead stems, May 1858, G.L. Rabenhorst (S-F70410, type).

Nodulosphaeria modesta (Desm.) Munk ex L. Holm, in Holm, Symb. bot. upsal. 14(no. 3): 80 (1957), Faces of Fungi number: FoF00253, Fig. 13.

Fig. 13
figure 13

Nodulosphaeria modesta (MFLU 12-2214). a Ascomata on host substrate. b Close up of ascoma. c − d Section of ascoma. e Close up of the peridium. f Hamathecium of dense long cellular pseudoparaphyses. g − j Asci with short, broad pedicel bearing 8 spores. k − n Mature and immature ascospores. Scale bars: c = 100 μm, e = 20 μm, d, f, g, h, i, j, k, l, m, n, o = 10 μm

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≡ Sphaeria modesta Desm., Ann. Mag. nat. Hist., Ser. 1: 1786 (1840)

Saprobic on dead stems. Sexual state: Ascomata 130–200 μm high, 150–250 μm diam., scattered, solitary to gregarious, immersed, erumpent to superficial, base not easy to remove from the substrate, visible as small, black dots on the host surface, uniloculate, coriaceous, globose to subglobose, setose, covered by hyphae, dark brown to black, ostiole central, with blunt brown hairs around the ostiole, papillate. Peridium 7–10 μm wide, thin-walled, of equal thickness, composed of two layers of pseudoparenchymatous cells, outer layer comprising 2–3 cell layers of dark brown to black, thickened cells, arranged in a textura angularis, inner layer comprising 3–5 cell layers of thin-walled cells, hyaline, flattened cells, arranged in textura angularis to textura prismatica. Hamathecium composed of numerous, 2–3 μm wide, filamentous, broad cellular pseudoparaphyses, with distinct septa, embedded in a mucilaginous matrix, anastomosing at the apex. Asci 60–70 × 13–17 μm (\( \overline{x} \) = 67 × 15 μm, n = 20), 8-spored, bitunicate, fissitunicate, cylindrical to cylindric-clavate, subsessile to short pedicellate, apically rounded, with well-developed ocular chamber. Ascospores 29–40 × 5–6 μm (\( \overline{x} \) = 35 × 6 μm, n = 40), partially overlapping 1–2 seriate, phragmosporous, cylindrical with slightly narrow ends, initially hyaline to subhyaline with small guttules, becoming yellowish-brown to brown at maturity, 4-septate, slightly curved, constricted at the septa, smooth-walled, with small, globose appendages, without a refractive sphere near the apices, enlarged at the second cell below the apex. Asexual state: Unknown.

Material examined: ITALY, Forlì-Cesena, Montevescovo, on dead stem, 8 February 2012, E. Camporesi (MFLU 14-0304), living culture = MFLUCC 11-0461 = MUCL.

Notes: Based on the morphological comparison, our isolate is similar to Nodulosphaeria modesta due to its small ascomata covered by hairs and short, broad setae at the papilla apex, the asci are bitunicate and clavate and the ascospores are cylindrical, brown, 4-septate with narrow apices, and an enlarged cell at the second cell below the apex. The multigene phylogenetic analysis indicates that Nodulosphaeria belongs to Phaeosphaeriaceae (Figs. 1 and 2).

Ophiobolus Riess, Hedwigia 1(6): 27 (1854), Faces of Fungi number: FoF00254.

Saprobic or hemibiotrophic on various hosts. Ascomata solitary to gregarious, scattered, semi-immersed to erumpent though host tissue with short to long beak, uniloculate, subglobose or ovoid or ampulliform, glabrous or covered by hyphae, dark brown to black, ostiolate, papillate. Peridium thick-walled, composed of dark brown to black, scleroplectenchymatous cells, arrange in textura angularis to textura globulosa. Hamathecium composed of numerous, filiform, cellular pseudoparaphyses, with distinct septa, anastomosing at the apex. Asci 8-spored, bitunicate, cylindrical to cylindric-clavate, short pedicellate, apically rounded with indistinct ocular chamber. Ascospores fasciculate, 3–4-seriate, scolecosporous, filiform or filamentous, with enlarged central cells, yellowish-brown to brown, multi-septate, non-constricted at the septa, separating into two part spores, smooth-walled, with or without appendages.

Type species: Ophiobolus disseminans

Phylogenetic study: Dong et al. 1998; Ariyawansa et al. (2014c)

Notes: Ophiobolus was introduced by Riess (1854) and typified by O. disseminans. Ascomata were described as discrete with distinct ostioles and ascospores as filiform and multi-septate (Zhang et al. 2012). Ophiobolus has been found on a wide range of hosts, mostly reported from Europe and North America (Farr and Rossman 2014). Holm (1948) and Müller (1952) used a broad concept of the genus Ophiobolus; however, Holm (1957) treated Ophiobolus in a much narrower sense, based on European collections (Shoemaker 1976). Three species were accepted by Holm (1957) and most Ophiobolus species were transferred to Nodulosphaeria and Leptospora (Holm 1957; Shoemaker 1976; Zhang et al. 2012). Shoemaker (1976) re-circumscribed species of Ophiobolus from Canada and some North American collections and accepted 31 species with two new species and five new combinations. Walker (1980) provided various illustrations of scolecosporous taxa from type material which had previously been accommodated in Ophiobolus (Saccardo 1883; Zhang et al. 2012). Ophiobolus species were also transferred to Acanthophiobolus, Entodesmium, Gaeumannomyces, Leptosphaeria and Leptospora (Shoemaker 1976; Walker 1980; Zhang et al. 2012). Barr (1979) initially accommodated Ophiobolus in Phaeosphaeriaceae and later transferred it to Leptosphaeriaceae (Barr 1987b). However, Zhang et al. (2012) reinstated the genus in Phaeosphaeriaceae based on morphological characters which are similar to Nodulosphaeria and Ophiosphaerella. There are 357 epithets listed in Ophiobolus in Index Fungorum (2014).

There is very little sequence data for Ophiobolus available in GenBank with two sequences of O. fulgidus. Zhang et al. (2012) mentioned that the genus should be placed in Phaeosphaeriaceae based on morphology and the phylogenetic investigation of Dong et al. (1998). However, Dong et al. (1998) could not resolve the placement of Ophiobolus as too few strains were used in their molecular analysis. Thus the genus needs to re-circumscribed using fresh collections (Zhang et al. 2012).

The asexual state of Ophiobolus has been reported in several genera such as coniothyrium-like, Rhabdospora, phoma-like and Scolecosporiella (Shoemaker 1976; Sivanesan 1984; Hyde et al. 2011; Wijayawardene et al. 2012; Zhang et al. 2012). However, these need confirmation as Ophiobolus is probably polyphyletic.

In this study, two species of Ophiobolus are included in the phylogenetic analysis. The detailed descriptions are provided in Ariyawansa et al. (2014c) and a description of the generic type is provided below. Based on multigene phylogenetic analysis (Figs. 1 and 2), Ophiobolus clusters in Phaeosphaeriaceae. However, there are limited sequences in GenBank and the type species lacks molecular data to confirm its natural placement. Thus Ophiobolus is tentatively placed in Phaeosphariaceae until the generic type strain is sequenced to confirm the natural placement (Table 2).

Table 2 Synopsis of Ophiobolus species discussed in this study
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Ophiobolus disseminans Riess, Hedwigia 1(6): 27 (1854), Faces of Fungi number: FoF00255, Figs. 14, and 15.

Fig. 14
figure 14

Ophiobolus disseminans (BPI 629021, isotype). a Herbarium label and specimen. b Ascomata on substrate. c Section through ascoma. d Section through peridium. e Papilla. f Asci with cellular pseudoparaphyses. g–i Ascus. j Ascospores in ascus. Note the swollen part of the ascospores (arrowed). Scale bars: c = 100 μm, d, e, f, g, h, i, j = 20 μm

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Fig. 15
figure 15

Ophiobolus disseminans (iconotype) redrawn from Riess (1854). a Section through ascoma. b Ascus and pseudoparaphyses. c Ascospores

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Saprobic on Cirsium (Asteraceae). Sexual state: Ascomata 140–390 μm high (excluding papilla), 160–340 μm diam., scattered or sometimes clustered, solitary to gregarious, semi-immersed, or erumpent through host tissue, visible as slightly raised, small, black dots on the host surface, uniloculate, subglobose to ovoid or ampulliform, glabrous, dark brown to black, ostiole central, with periphyses, short papillate. Necks 60–150 μm high, 85–140 μm diam, with thick-walled cells, arranged in a textura angularis, subglobose to oblong, dark brown to black, apically rounded or obtuse, coriaceous. Peridium 30–50 μm wide, thick-walled, of unequal thickness, slightly thick at the base, composed of several layers of broadly scleroplectenchymatous cells, outer layers consisting of several layers of brown to dark brown, thickened cells, arranged in a textura angularis, inner layers comprising several layers of hyaline, flattened cells, arranged in a textura epidermoidea to textura porrecta. Hamathecium composed of numerous, 2–3 μm wide, filamentous, broad cellular pseudoparaphyses, with distinct septa, embedded in mucilaginous matrix, anastomosing at the apex. Asci (156–)170–190(−203) × 10–12(−13) μm (\( \overline{x} \) = 178.9 × 11.4 μm, n = 20), 8-spored, bitunicate, cylindrical to cylindric-clavate, short pedicellate, apically rounded with indistinct ocular chamber. Ascospores (125–)150–160(−163) × 2–3 μm (\( \overline{x} \) = 152.8 × 2.9 μm, n = 15), fasciculate, in parallel or spiral, scolecosporous, filiform or filamentous, with two enlarged cells near the centre, but with 3–4 narrow cells in between the enlarged cells, brown, multi-septate, with up to 15 septa, not constricted at the septa, smooth-walled, separating into two part spores in the centre of the spores between the enlarged cells. Asexual state: Unknown.

Material examined: ITALY, Cassellas Vicinity, on dead stem of Cirsium arvense, 1853, Riess 1823 (BPI 629021, isotype); ibid on dead stem of Cirsium arvense, 1853, H. Riess, Hedwigia 1(6): Tab IV, (Fig. 8, iconotype) (1854).

Ophiosphaerella Speg., Anal. Mus. nac. B. Aires, Ser. 3 12: 401 (1909), Faces of Fungi number: FoF00256.

Pathogenic or saprobic mostly on various monocotyledons and some flowering shrubs or dicotyledons especially Poaceae, Asteraceae and Fagaceae (Farr and Rossman 2014). Sexual state: Ascomata scattered, solitary, semi-immersed, or erumpent through host tissue with papilla visible as raised, small, black dots on host surface, uniloculate, globose to subglobose, glabrous, dark brown to black, ostiole central, with periphyses, papillate. Peridium thick-walled, composed of hyaline to brown or dark brown, pseudoparenchymatous cells, arranged in a textura angularis. Hamathecium composed of numerous, filamentous, broad cellular pseudoparaphyses, with distinct septa, embedded in mucilaginous matrix, anastomosing at the apex. Asci 8-spored, bitunicate, cylindrical to cylindric-clavate, short pedicellate, apically rounded with indistinct ocular chamber. Ascospores fasciculate, lying parallel or spiral, scolecosporous, filiform or filamentous, pale brown to brown, multi-septate, not constricted at the septum, smooth-walled. Asexual state: see under Scolecosporiella.

Type species: Ophiosphaerella graminicola

Phylogenetic study: Wetzel et al. (1999), Câmara et al. (2000), Hsiang et al. (2003), Schoch et al. (2006, 2009), Zhang et al. (2009, 2012), Hyde et al. (2013).

Notes: Ophiosphaerella was introduced by Spegazzini (1909) as typified by Op. graminicola. Most species of Ophiosphaerella are often found as pathogens or saprobes worldwide on Poaceae and Cyperaceae (Câmara et al. 2000). Spegazzini (1909) did not introduce the genus separately but described and illustrated a new species, and later the genus was validly published by Stafleu et al. (1972) (Walker 1980; Zhang et al. 2012). However, Petrak and Sydow (1936) treated Ophiosphaerella graminicola as a synonym of Ophiobolus graminicolus (Speg.) after observing the type specimens which was accepted by Ainsworth et al. (1971) and von Arx and Müller (1975) (Walker 1980; Zhang et al. 2012). Walker (1980) examined the type specimens of Ophiosphaerella and confirmed that the genus belongs to Pleosporales, but differs from Ophiobolus. Ophiosphaerella is most similar to Ophiobolus and subsequently often confused in morphological characters. However Ophiosphaerella differs from Ophiobolus by the lack of central swollen ascospore cells which do not separate into part spores (Zhang et al. 2012).

The phylogenetic relationships of Ophiosphaerella were first investigated by Wetzel et al. (1999) who treated Ophiosphaerella as monophyletic genus distinguished from Phaeosphaeria and other pleosporaceous genera. Various authors subsequently accepted the genus in Phaeosphaeriaceae based on the multigene phylogenetic investigation with Ophiosphaerella forming a monophyletic clade as a sister group of Phaeosphaeria (Schoch et al. 2006, 2009; Zhang et al. 2009, 2012; Hyde et al. 2013). However, the phylogenetic analysis was limited to one strain (Ophiosphaerella herpotricha CBS 620.86).

In the present multigene phylogenetic analyses, two strains of Ophiosphaerella agrostidis (MFLUCC 11-0152 and MFLUCC 12-0007) and Op. herpotricha (CBS 620.86) are included. The phylogenetic relationships show that Op. agrostidis (MFLUCC 11-0152 and MFLUCC 12-0007) forms a distinct clade separated from Op. herpotricha (CBS 620.86) in Phaeosphaeriaceae. Thus Ophiosphaerella seem to be polyphyletic genus. However, there are few sequences available in GenBank, thus fresh collections and molecular data are needed to for more natural understanding.

Ophiosphaerella graminicola Speg., Anal. Mus. nac. B. Aires, Ser. 3 12: 401 (1909), Faces of Fungi number: FoF00257, Fig. 16, and 17.

Fig. 16
figure 16

Ophiosphaerella graminicola (BPI 748267). a Herbarium label and specimens of Ophiosphaerella graminicola. b Ascomata on host surface. c Section through ascoma d Section through peridium. e Papilla with periphyses. f Pseudoparaphyses stained in Meltzer’s reagent. g Ascus. h–j Asci stained in Meltzer’s reagent. k–l Ascospores. m–o Ascospores stained in Meltzer’s reagent. Scale bars: c = 100 μm, e = 50 μm, d, f, g, h, i, j, k, l, m, n, o = 20 μm

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Fig. 17
figure 17

Ophiosphaerella graminicola (iconotype) redrawn from Spegazzini (1909). a Ascoma. b Ascomata immersed on host tissue. c Asci. d Ascospore

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Pathogenic or saprobic on monocotyledons. Sexual state: Ascomata 230–300 μm high, 170–230 μm diam., scattered, solitary, immersed to semi immersed or erumpent through host tissue with minute papilla, visible as raised, small black dots on host surface, uniloculate, globose to subglobose, glabrous, dark brown to black, ostiole central, with periphyses, with minute papilla. Peridium 20–40 μm wide, thick-walled, of unequal thickness, broad at the apex towards sides of ascoma, composed of several layers of pseudoparenchymatous cells, outer layers comprising several layers of brown to dark brown thickened cells at sides and apex, arranged in a textura angularis, hyaline at the base of ascoma, inner layers comprising several layers of thin-walled, flattened, hyaline cells of textura prismatica. Hamathecium composed of numerous, 2–3 μm wide, filamentous, broadly cellular pseudoparaphyses, with distinct septa, embedded in a mucilaginous matrix, anastomosing at the apex. Asci (128–)133–150(−160) × 9–12(−12.5) μm (\( \overline{x} \) = 148.7 × 11.2 μm, n = 20), 8-spored, bitunicate, cylindrical to cylindric-clavate, short pedicellate, apically rounded with indistinct ocular chamber. Ascospores (118–)130–150(−160) × 3–3.5 μm (\( \overline{x} \) = 142.4 × 3.2 μm, n = 30), fasciculate, lying parallel or spiral in the centre, scolecosporous, filiform or filamentous, narrowing towards the ends, pale brown to brown, 15–18-septate, not constricted at the septa, smooth-walled. Asexual state: Unknown.

Material examined: ARGENTINA, in garden near Tucumán, on leaf sheath of Leptochloa virgata (L.) P. Beauv. (Poaceae), April 1906, C. Spegazzini (iconotype); VENEZUELA, Edo, Aragua, Parque Nacional Henri Pittier, on the Maracay-Ocumare Rd., ca. 15 km north of Maracay, 4.5 km south of Rancho Grande Biological Station where creek crosses road, alt. 950 m. ca. 10°21′N 67°41″W, on grass culm, 5 December 1990, G.J. Samuels, B. Hein, S.M. Huhndorf (BPI 748267).

Ophiosphaerella agrostidis Dern. et al. [as ‘agrostis’], in Câmara et al., Mycologia 92(2): 320 (2000), Faces of Fungi number: FoF00258, Fig. 18.

Fig. 18
figure 18

Ophiosphaerella agrostidis (MFLU 11-0188). a Ascomata on host surface. b Section through ascoma c Section through peridium. d Cellular pseudoparaphyses stained in congo red. e–i Asci. j–n Ascospores. Scale bars: b = 100 μm, c–n = 20 μm

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Pathogenic or saprobic on monocotyledonous hosts. Sexual state: Ascomata 275–355 μm high (including papilla), 175–310 μm diam., scattered, solitary, semi immersed or erumpent through host tissue with papilla, visible as raised, small black dots on host surface, uniloculate, globose to subglobose, glabrous, dark brown to black, ostiole central, oblique, papilla, carbonaceous. Peridium 15–25 μm wide, thin-walled, of equal thickness, composed of several layers of brown to dark brown, pseudoparenchymatous cells, arranged in a textura angularis to textura globulosa. Hamathecium composed of numerous, 1.5–2.5 μm wide, filamentous, broad cellular pseudoparaphyses, with distinct septa, embedded in mucilaginous matrix, anastomosing at the apex. Asci (97–)100–135 × 7–9(−11) μm (\( \overline{x} \) = 111.5 × 8.5 μm, n = 25), 8-spored, bitunicate, fissitunicate, cylindrical to cylindric-clavate, short pedicellate, apically rounded, with a well-developed ocular chamber. Ascospores (90–)98–120(−130) × 2–2.5 μm (\( \overline{x} \) = 110.3× 2.3 μm, n = 30), fasciculate, parallel or spiral, scolecosporous, filiform, narrowing towards ends, pale brown to brown, 15-septate, not constricted at the septa, smooth-walled. Asexual state: Unknown.

Material examined: THAILAND, Phrae, Rongkwang District, Maejo University Phrae Campus, on dead culms of grass, 20 August 2010, R. Phookamsak (MFLU 11-0188), living culture = MFLUCC 11-0152; Chiang Rai, Muang District, Ta Sai, on dead culm of grass, 30 June 2011, R. Phookamsak (MFLU 11-0246), living culture = MFLUCC 12-0007.

Notes: The scolecosporous specimens were collected from dead culms of grass in Thailand. These collections are morphologically similar to Ophiosphaerella agrostidis, Op. graminicola, and Op. herpotricha with these species sharing similar ascomata, asci and ascospores, including septation of ascospores. The Thailand collections are most similar to Op. agrostidis according to the size of the ascomata, asci and ascospores, and also the septation of ascospores. However, Op. agrostidis has been reported as a pathogen on grass while the collections in the present study were saprobes. Therefore these collections are named as Op. agrostidis based on the morphological characters and these can be treated as authentic specimens until epitypification. The phylogenetic tree (Figs. 1 and 2) shows that two isolates of Op. agrostidis form a single clade in Phaeosphaeriaceae (Table 3).

Table 3 Synopsis of Ophiosphaerella species discussed in this study
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Paraphoma Morgan-Jones & J.F. White, Mycotaxon 18(1): 58 (1983), Faces of Fungi number: FoF00259.

Saprobic on various hosts, or soil borne. Mycelium abundance branched, septate, subhyaline to pale brown, smooth-walled. Sexual state: Unknown. Asexual state: Conidiomata pycnidial, scattered, solitary to gregarious, in agar semi-immersed to superficial, uniloculate, globose to subglobose, setose, brown to dark brown, ostiole central, circular. Conidiomata walls thick, outer layers comprising 2 layers of scleroplectenchymatous cells, dark brown, inner layers comprising 4–5 layers of pseudoparenchymatous cells, arranged in a textura angularis, subhyaline to pale brown. Setae numerous, thick-walled, straight to slightly curved, acuminate, septate, smooth to verruculose. Conidiophores reduced to conidiogenous cells. Conidiogenous cells monophialidic, discrete, determined, lageniform, hyaline to subhyaline, lining the inner cavity. Conidia enteroblastic, amerosporous, ellipsoidal, hyaline, aseptate, smooth-walled, with guttules (from Sutton 1980; De Gruyter et al. 2009, 2010; Quaedvlieg et al. 2013).

Type species: Paraphoma radicina (McAlpine) Morgan-Jones & J.F. White, Mycotaxon 18(1): 60 (1983)

≡ Pyrenochaeta radicina McAlpine, Fungus Diseases of stone-fruit trees in Australia: 127 (1902)

Phylogenetic study: De Gruyter et al. (2009, 2010, 2012), Aveskamp et al. (2010), Zhang et al. (2012), Hyde et al. (2013), Quaedvlieg et al. (2013).

Notes: Paraphoma as typified by P. radicina was previously treated in Pyrenochaeta by McAlpine (1902) and later, Boerema and Dorenbosch (1979) transferred the species to Phoma. Morgan-Jones and White (1983) introduced a new genus, Paraphoma to accommodate the species that produced heavily setose pycnidia and transferred P. radicina to this genus. However, the genus was treated as a section of Phoma (Boerema 1997) until De Gruyter et al. (2010) reinstated the genus and placed it in Phaeosphaeriaceae based on phylogenetic evidence. Recently, four species are accommodated in this genus for which no sexual state is known.

Parastagonospora Stud. Mycol. 75: 362 (2013), Faces of Fungi number: FoF00260.

Pathogenic on various cereal crops (Poaceae). Sexual state: Ascomata scattered, solitary to gregarious, immersed, uniloculate, globose, glabrous, brown to black, ostiole central, with papilla. Peridium thick-walled, composed of 3–6 layers of brown pseudoparenchymatous cells, arranged in a textura angularis. Hamathecium composed of numerous, filiform, cellular pseudoparaphyses, with distinct septa. Asci 8-spored, bitunicate, cylindrical to clavate, short pedicellate. Ascospores phragmosporous, fusiform, subhyaline to pale brown, septate (−3 septa), constricted at the septa, swollen at penultimate cell. Asexual state: Conidiomata pycnidial, scattered, immersed to semi-immersed, uniloculate, subglobose, glabrous, black, exuding creamy conidial mass, ostiole central. Conidiomata walls thin, composed of 2–3 layers of brown cells, arranged in textura angularis. Conidiophores reduced to conidiogenous cells. Conidiogenous cells phialidic, integrate, ampulliform to subcylindrical, hyaline, smooth-walled, lining the inner cavity, with percurrent proliferation near apex. Conidia phragmosporous, cylindrical, apically obtuse with truncate base, hyaline, septate, thin and smooth-walled, with granules or multiguttules (from Quaedvlieg et al. 2013).

Type species: Parastagonospora nodorum (Berk.) Quaedvlieg et al. Stud. Mycol. 75: 363 (2013)

≡ Depazea nodorum Berk., Gard. Chron., London: 601 (1845)

Phylogenetic study: Quaedvlieg et al. (2013)

Notes: Parastagonospora was introduced by Quaedvlieg et al. (2013) to accommodate stagonospora-like species which have been reported as asexual state of phaeosphaeria-like taxa. The genus accommodated five species; Pa. avenae (A.B.. Frank) Quaedvlieg et al., Pa. caricis Quaedvlieg et al., Pa. nodorum (Berk.) Quaedvlieg et al., Pa. poae Quaedvlieg et al. and Pa. poagena Crous & Quaedvlieg with Pa. nodorum as the type (Index Fungorum, 2014). Parastagonospora is similar to Stagonospora; however, they differ in the morphology of the conidiogenous cells (Quaedvlieg et al. 2013). Based on multigene phylogenetic analysis, Parastagonospora clusters in Phaeosphaeriaceae which forms a clade with Phaeosphaeria avenaria (synonymized under Parastagonospora avenae), Ph. eustoma and Ph. nigrans. These Phaeosphaeria species may be the sexual states of Parastagonospora.

Phaeosphaeria I. Miyake, Bot. Mag., Tokyo 23: 93 (1909), Faces of Fungi number: FoF00233.

= Phaeoseptoria Speg., Revta Mus. La Plata 15: 39. 1908.

Saprobic or pathogenic mostly on monocotyledons. Sexual state: Ascomata solitary to gregarious, immersed, visible as slightly raised, small, black dots on the host surface, uniloculate, globose to subglobose, glabrous, brown, ostiole central, with minute papilla. Peridium thin-walled, of equal thickness, composed of flattened, brown, pseudoparenchymatous cells, arranged in textura angularis. Hamathecium composed of sparse, filiform, broad cellular pseudoparaphyses, with distinct septa, slightly constricted at the septa, rarely anastomosing at the apex. Asci 8-spored, bitunicate, fissitunicate, broadly cylindrical, short pedicellate or subsessile, apically rounded, with an indistinct ocular chamber. Ascospores overlapping 2–3-seriate, phragmosporous, narrowly fusiform, pale brown to brown or yellowish-brown, septate, straight or slightly curved, constricted or not constricted at the septa, walls smooth or rough, echinulate or verruculose, with or without mucilaginous sheaths. Asexual state: Conidiomata stromatic, pycnidial, scattered to gregarious, immersed, uni- to multi-loculate, globose to subglobose, brown. Conidiomata walls thin, composed of thin-walled, of brown pseudoparenchymatous cells, arranged in textura angularis. Conidiophores reduced to conidiogenous cells. Conidiogenous cells holoblastic, phialidic, flattened ampulliform to doliiform, hyaline to pale brown, hardly distinguishable from inner cells walls of conidioma. Conidia phragmosporous, oblong to cylindrical, pale brown to brown, septate, straight or slightly curved with obtuse at the base, smooth-walled with guttules.

Type species: Phaeosphaeria oryzae

Phylogenetic study: Câmara et al. (2002); Schoch et al. (2006, 2009); Zhang et al. (2009, 2012); De Gruyter et al. (2010, 2012); Quaedvlieg et al. (2013); Hyde et al. (2013, 2014); Wijayawardene et al. (2014b)

Notes: Phaeosphaeria was established by Miyake (1909). Miyake (1910) accommodated Phaeosphaeria in Mycosphaerellaceae and mentioned that the genus differs from Sphaerulina and Leptosphaeria in ascospore colour and lack of pseudoparaphyses (Eriksson 1967). Phaeosphaeria was considered as a synonym of Leptosphaeria for a long time (Müller 1950; Munk 1957; Zhang et al. 2009, 2012). The genus was however, reinstated by Holm (1957) who found pseudoparaphyses in Phaeosphaeria oryzae and 17 mostly graminicolous species were transferred to Phaeosphaeria (Eriksson 1967; Zhang et al. 2009). Barr (1979) established the new family Phaeosphaeriaceae and designated Phaeosphaeria as the generic type and later, Eriksson (1981) introduced a broad generic concept for Phaeosphaeria which included relatively small ascomata, dictyospores, as well as ascospores with several wall layers, and an occurrence on monocotyledons (Câmara et al. 2002; Zhang et al. 2012). Subsequent authors also included Phaeosphaeria species from dicotyledonous hosts (Dennis 1978; Sivanesan 1984; von Arx and Müller 1975; Eriksson 1967; Hedjaroude 1969; Shoemaker and Babcock 1989b; Zhang et al. 2012). Shoemaker and Babcock (1989b) re-circumscribed Phaeosphaeria and divided the genus into six subgenera based on the differences in ascospore shape and septation viz. Fusispora, Ovispora, Phaeosphaeria, Sicispora, Spathispora and Vagispora. Various authors subsequently indicated that Phaeosphaeria was heterogeneous and shared several characters with Leptosphaeria. Phaeosphaeria however, differs from Leptosphaeria in morphology and these genera have been distinguished in different families by phylogenetic analysis (Câmara et al. 2002; Schoch et al. 2009; Zhang et al. 2009, 2012; Hyde et al. 2013). Presently, there are 200 epithets for Phaeosphaeria in Index Fungorum (2014) with 18,983 nucleotide sequences available in GenBank. However, several species of Leptosphaeria sensu stricto and Phaeosphaeria are confused in having similar morphology and lack phylogenetic investigation.

The asexual states of Phaeosphaeria have been reported as coelomycetes and some as hyphomycetes namely hendersonia-like, Mauginiella, Microsphaeropsis, Neostagonospora, Parastagonospora, Phaeoseptoria, Scolecosporiella and stagonospora-like (Leuchtmann 1984; Shoemaker and Babcock 1989b; Câmara et al. 2002; Abdullah et al. 2005; Kirk et al. 2008; Hyde et al. 2011, 2013; Wijayawardene et al. 2012; Zhang et al. 2012; Quaedvlieg et al. 2013). However, Leuchtmann (1984) treated the asexual state of Phaeosphaeria as species of Stagonospora and mentioned that the asexual state of Phaeosphaeria often formed pycnidial conidiomata, possessed holoblastic conidiogenous cells and had elongated to filiform, hyaline to pale brown, multiseptate conidia, which was also mentioned by Câmara et al. (2002). Câmara et al. (2002) suggested that Phaeosphaeria shares common ancestors with Leptosphaeria and that multigene phylogenetic analysis was necessary to understand the boundaries of taxa. The peridium type, asexual state and host preferences can distinguish fungal taxa in these two genera (Câmara et al. 2002)

Quaedvlieg et al. (2013) studied Septoria, Stagonospora and related genera such as Phaeosphaeria and Phaeoseptoria using molecular analysis and concluded that Stagonospora (based on the type species, S. paludosa) belongs to Massarinaceae. They introduced two new genera (Neostagonospora and Parastagonospora) to accommodate stagonospora-like genera in Phaeosphaeriaceae (Quaedvlieg et al. 2013). Additionally, Phaeosphaeria was linked to the asexual state Phaeoseptoria based on the phylogenetic analysis with the type species of Phaeosphaeria, Ph. oryzae forming a well-supported clade with Phaeoseptoria papayae, the type species of Phaeoseptoria (Quaedvlieg et al. 2013). Thus Quaedvlieg et al. (2013) synonymized the earlier name Phaeoseptoria under Phaeosphaeria as the latter was well-studied and taxonomy less confused than Phaeoseptoria which have been confused with several septoria-like in pigmented conidia (Walker et al. 1992). Hendersonia, which was treated as a synonym of Stagonospora, and Microsphaeropsis were placed in Didymellaceae (Sutton 1977; Swart and Walker 1988; Kirk et al. 2008; De Gruyter et al. 2009; Aveskamp et al. 2010). Mauginiella was treated as an asexual state of Phaeosphaeria by Abdullah et al. (2005); however, this has not been confirmed by phylogenetic investigation. In this study Mauginiella could not be resolved in Phaeosphaeriaceae. The genus is currently placed in asexual Pezizomycotina (Kirk et al. 2008). Therefore we exclude Mauginiella from Phaeosphaeriaceae. Other asexual genera of Phaeosphaeria still lack phylogenetic investigation. Therefore we accept three asexual genera (Neostagonospora Quaedvlieg et al., Parastagonospora Quaedvlieg et al., and Phaeoseptoria Speg.) for Phaeosphaeria sensu lato.

Phaeosphaeria oryzae was designated as typified of Phaeosphaeria by Miyake (1909). However, Miyake (1909) did not mention the detail of the type specimen in his publication and this type has not been found in any herbarium in Japan. Eriksson (1967) translated the protologue and found Miyake’s specimen in Sydow’s herbarium as labeled “Leptosphaeria oryzae Hori = Phaeosphaeria oryzae Miy. — on Oryza sativa L. — Prov. Susuya Shizuoka — 1907.09”. He found that most characters of this specimen match with the diagnosis except the presence of pseudoparaphyses. Thus Eriksson (1967) designated Sydow’s specimen as a lectotype of Phaeosphaeria oryzae. However, the mucilaginous sheath or appendages were not seen in Eriksson’s (1967) observation (Shoemaker and Babcock 1989b). Shoemaker and Babcock (1989b) carried out a monograph of Phaeosphaeria with Phaeosphaeria oryzae treated in subgenus Phaeosphaeria. The species description included echinulate ascospores; however Miyake (1909, 1910) and Eriksson (1967) did not mention this character. Quaedvlieg et al. (2013) epitypified Phaeosphaeria oryzae and Phaeoseptoria papayae, the type species of Phaeosphaeria and Phaeoseptoria and indicated that Phaeoseptoria is the asexual state of Phaeosphaeria based on phylogenies.

In the present study the lectotype of Phaeosphaeria oryzae was examined and the morphological characters compared with the diagnosis of Miyake (1910). On examination it appears that Phaeosphaeria oryzae lectotype has smooth-walled ascospores, but this was not mentioned by Miyake (1910) either. In addition, the asci and ascospores size in the lectotype are larger than Miyake’s diagnosis (1910) (see Table 4). Quaedvlieg et al. (2013) epitypified Phaeosphaeria oryzae and described the ascospores as echinulate following the description of Shoemaker and Babcock (1989b). Further specimens that have been determined to be Phaeosphaeria oryzae from BPI have been examined in the present study and some specimens have echinulate ascospores, while others do not. It appears that Phaeosphaeria oryzae may be a species complex which has been epitypified by a specimen that is probably a different species to the lectotype.

Table 4 Synopsis of Phaeosphaeria discussed in this study
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Based on our multigene phylogenetic analysis, Phaeosphaeria species separated in several subclades in Phaeosphaeriaceae. Several Phaeosphaeria sensu stricto specimens collected from various hosts in Thailand formed a strongly supported clade with the type species, Phaeosphaeria oryzae and also Ph. papayae and Ph. musae. These Phaeosphaeria (sensu stricto) species differ morphologically (see Table 4), but the phylogenies indicate that they are closely related species. We therefore analyzed the phylogenetic relationships among species, with Phaeosphaeria sensu stricto strains forming a distinct clade with high bootstrap support. A pair wise comparison of ITS and TEF1 helped to significantly differentiate the cryptic species (ITS pair wise comparison see Fig. 4). Currently, many plant pathogens causing serious disease of economic crops are reported as the species complexes (Wikee et al. 2011; Cannon et al. 2012; Manamgoda et al. 2012; Udayanga et al. 2012; Weir et al. 2012) which seem to be the case in Phaeosphaeria. Phaeosphaeria sensu stricto species are morphologically similar, but differ in the size of ascomata, asci, and ascospores, habitat and host occurrence. The combined gene phylogenetic analysis can distinguish taxa of Phaeosphaeria sensu stricto at the species level.

The Phaeosphaeria sensu stricto taxa collected from various monocotyledons in Thailand are introduced as two novel species and Phaeosphaeria musae is reported on other hosts from Thailand with descriptions and illustrations below.

Phaeosphaeria oryzae I. Miyake, Bot. Mag., Tokyo 23: 93 (1909), Faces of Fungi number: FoF00234, Fig. 19.

Fig. 19
figure 19

Phaeosphaeria oryzae (S-F9572, lectotype) a Herbarium label and specimens. b Ascomata on host surface. c Section through ascoma. d Section through peridium. e Asci with sparse pseudoparaphyses. f–i Asci. j–o Ascospores. Scale bars: c = 50 μm, e, f, g, h, i, j = 20 μm, d, k, l, m, n, o = 10 μm

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= Phaeoseptoria oryzae I. Miyake (1910).

Saprobic on leaves of Oryza sativa. Ascomata 70–130 μm high, 90–150 μm diam., solitary, sometimes clustered, immersed, visible as slightly raised, small, black dots on the host surface, uniloculate, globose to subglobose, glabrous, brown to dark brown, ostiole central, with minute papilla. Peridium 5–10 μm wide, of equal thickness, composed of 2–3 layers of thin-walled, brown to dark brown, flattened pseudoparenchymatous cells, arranged in textura angularis. Hamathecium composed of sparse, 2.5–4 μm wide, filiform, broad cellular pseudoparaphyses, distinctly constricted at the septa, rarely anastomosing at the apex. Asci (50–)55–70(−80) × 10–13.5(−15) μm (\( \overline{x} \) = 65.4 × 11.9 μm, n = 25), 8-spored, bitunicate, fissitunicate, broadly cylindrical, short pedicellate or subsessile, apically rounded with indistinct ocular chamber. Ascospores (22–)25–27(−29) × 5–6 μm (\( \overline{x} \) = 26 × 5.2 μm, n = 30), overlapping 2-seriate, phragmosporous, narrowly fusiform, brown, mostly 3-septate, rarely 1–2 septa, indistinctly constricted at the septa, slightly curved, smooth-walled. Asexual state: Phaeoseptoria.

Material examined : JAPAN, Shizuoka, Suruya, on Oryza sativa L. (Poaceae), September 1907 (S-F9572, lectotype); EL SALVADOR, intercepted Washington Dc #26812, on Oryza sativa, 10 March 1952, F.P. Hubert (BPI 622541); JAPAN, intercepted San Diego California #4872, on Oryza sativa L., 27 December 1957, E.B. Francis (BPI 622542); JAPAN, intercepted San Francisco California #17349, 17352, 17367, on Oryza sativa, 13 December 1940, F.J. Phelan (BPI 622544); NICARAGUA, Monagua, on living leaves of Oryza sativa, 4 November 1954, C. Pineda 37, (BPI 622545); THAILAND, Chiang Mai Province, Mae Wang District, Mae Sapok Royal Project, on dead branches of Etlingera sp. (Zingiberaceae), 5 October 2010, R. Phookamsak RP0086 (MFLU 11-0206), living culture = MFLUCC 11-0170.

Phaeosphaeria chiangraina Phookamsak & K.D. Hyde, sp. nov., Index Fungorum number: IF550736; Faces of Fungi number: FoF00261, Fig. 20.

Fig. 20
figure 20

Phaeosphaeria chiangraina (MFLU 12-2472, holotype). a Ascomata of Phaeosphaeria oryzae on host surface. b Ascomata visualized under compound microscope. c Section through ascoma. d Section through peridium. e Asci with rare pseudoparaphyses. f, i Asci. g, h Asci stained in 5 % KOH. j- o Ascospores stained in 5 % of KOH. m, n Ascospores. p Ascospore germination on WA. q Section through conidioma. r Section through conidioma wall. s–u Conidiogenous cells. v–z Conidia. Scale bars: q = 100 μm, b, c = 50 μm, d, r = 20 μm, e, f, g, h, i, p = 10 μm, j, k, l, m, n, o, s, t, u, v, w, x, y, z = 5 μm

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Etymology: Referring to Province in Thailand, in which the fungus was found.

Holotypus: MFLU 12-2472

Saprobic on leaves of Oryza sativa. Sexual state: Ascomata 120–180 μm high, 130–180 μm diam., scattered, immersed, visible as slightly raised, small, black dots on the host surface, uniloculate, globose to subglobose or irregular in shape, glabrous, brown to dark brown, ostiole central with minute papilla. Peridium 7–16 μm wide, thin-walled, of equal thickness, composed of 1–3 layers of flattened, brown to dark brown, elongated, pseudoparenchymatous cells, arranged in textura angularis. Hamathecium composed of numerous, 1.5–3 μm wide, filiform, broad cellular pseudoparaphyses, with small guttules, indistinctly constricted at the septa, embedded in mucilaginous matrix, rarely anastomosing at the apex. Asci (44–)47–60(−65) × 8–9(−10) μm (\( \overline{x} \) = 52.6 × 8.9 μm, n = 25), 8-spored, bitunicate, fissitunicate, broadly cylindrical, short pedicellate or subsessile, apically rounded with ocular chamber. Ascospores (13.5–)15–17(−18) × 3.5–4.5 μm (\( \overline{x} \) = 16.4 × 4.1 μm, n = 30), overlapping 2-seriate, phragmosporous, fusiform with obtuse ends, pale brown when young, becoming yellowish-brown to brown at maturity, dark brown when stained in 5 % of KOH, 3-septate, not constricted at the septa, slightly curved, smooth-walled with small to large guttules in each cell, surrounded by a distinct mucilaginous sheath. Asexual state: produced on media agar after 8 weeks. Conidiomata 100–305 μm high, 120–290 μm diam., pycnidial, scattered to clustered, immersed, with dark brown to black stromatic tissues embedded in the media agar, uni- to multi-loculate, globose to irregular, ostioles not observed, brown to dark brown. Conidiomata walls 13–35 μm wide, composed of several layers of brown, pseudoparenchymatous cells, arranged in a textura angularis, covered by vegetative hyphae. Conidiophores reduced to conidiogenous cells. Conidiogenous cells (2–)3–6 × (2–)4–7 μm (\( \overline{x} \) = 4.4 × 4.3 μm, n = 20), holoblastic, phialidic, ampulliform or flattened, hyaline to pale brown, difficult to distinguish from the pycnidial wall. Conidia (15–)17–21.5(−26) × 2–3 μm (\( \overline{x} \) = 19.4 × 2.5 μm, n = 30), solitary, cylindrical, pale brown to brown, 1–3-septate, slightly curved, obtuse at the apex, truncate at the base, not constricted at the septa, smooth-walled with guttules.

Culture characters: Colonies on PDA 77–80 mm diam. after 4 weeks at 25–30 °C, white to pale yellowish at the edge, pale yellowish to grey in the centre, forming different colour sectors of grey to dark grey; reverse pale yellowish at the edges, becoming yellowish-brown to brown at the centre, slightly radiating outwardly, forming dark sectors; medium dense, forming black stromatic tissues immersed in media after 8 weeks, circular to irregular, flattened to slightly raised, dull with entire edge, floccose to fairly fluffy, slightly radiating at the lower part, not pigmented.

Material examined: THAILAND, Chiang Rai Province, Muang District, Nanglae Village, on dead leaves of Oryza sativa, 20 January 2013, R. Phookamsak RP0131 (MFLU 12-2472, holotype), ex-type living culture = MFLUCC 13-0231 = MUCL.

Notes: Phaeosphaeria chiangraina was collected on Oryza in Thailand. This is the same host of the generic type of Phaeosphaeria, P. oryzae. A morphological comparison based on species described in Shoemaker and Babcock (1989b) shows that Ph. chiangraina is similar to Phaeosphaeria avenaria, Ph. eustoma, Ph. oryzae and Ph. tritici. Phaeosphaeria chiangraina differs from Ph. oryzae in the size of ascomata, asci and ascospores. Phaeosphaeria chiangraina has smaller ascospores (13.5–18 × 3.5–4.5 μm) than Ph. oryzae (15–23 × 4–5 μm). Phaeosphaeria tritici has similar sized ascospores to Ph. chiangraina; however, Ph. tritici has a larger asci and smaller ascomata. Additionally, Phaeosphaeria tritici ascospores are enlarged at the second cell and slightly constricted at the first septa and produce a phoma-like asexual state, as while Ph. chiangraina lacks an enlarged cell, is not constricted at the septa and forms a septoria-like asexual state.

Multigene phylogenetic analysis using MP shows that Phaeosphaeria chiangraina can be distinguished from Phaeosphaeria avenaria and Ph. eustoma, and forms a strongly supported clade with Ph. thysanolaenicola and Ph. oryzae (CBS 110110) (Fig. 3). However, Ph. chiangraina morphologically differs from Ph. thysanolaenicola and can be separated from the epitype of Ph. oryzae due to its smooth-walled ascospores with a distinct sheath and based on phylogenetic evidence (Fig 3).

Phaeosphaeria musae Sawada, Special Publication College of Agriculture, National Taiwan University 8: 66 (1959) Faces of Fungi number: FoF00262, Fig. 21.

Fig. 21
figure 21

Phaeosphaeria musae . (MFLU 11-0169). a Ascomata on host surface. b Section through ascoma. c Section through peridium. d Pseudoparaphyses. e–h Asci. i–m Ascospores. Note sheath in m. Scale bars: b = 50 μm, c, d, e, f, g, h, = 20 μm, i, j, k, l, m = 10 μm

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Pathogen on Asparagaceae and Marantaceae. Sexual state: Ascomata 110–140.5 μm high, 120–160 μm diam., solitary, scattered, immersed, visible as slightly raised, small black dots on host surface, uniloculate, brown to dark brown, globose to subglobose, central ostiole with minute papilla. Peridium 7–24 μm wide, thin-walled, of unequal thickness, slightly thick at the apex, composed of 1–3 layers of brown to dark brown, pseudoparenchymatous cells multi-layered and elongated, at the apex, arranged in textura angularis. Hamathecium composed of numerous, 1.5–3 μm wide, filiform, broad cellular pseudoparaphyses, with distinct septa, slightly constricted at the basal septa, embedded in gelatinous matrix, rarely anastomosing at the apex. Asci (40–)65–70(−76) × (9–)13–15(−17) μm (\( \overline{x} \) = 69 × 14.2 μm, n = 25), 8-spored, bitunicate, fissitunicate, broadly cylindric-clavate to clavate, subsessile, with acute ends, apically rounded with indistinct ocular chamber. Ascospores (16–)25–30(−33.5) × 5–7.5 μm (\( \overline{x} \) = 27.6 × 5.8 μm, n = 30), overlapping 2–3-seriate, phragmosporous, fusiform, yellowish-brown to brown, becoming dark brown when stained in 5 % KOH, 3-septate, slightly constricted at the central septa, enlarged at the second cell, slightly curved, rarely straight, rough-walled, echinulate, surrounded by narrow mucilaginous sheath. Asexual state: Unknown.

Culture characters: Colonies on MEA 25–27 mm diam. after 4 weeks at 25–30 °C, white yellowish or grey in the centre and edges; reverse yellowish-brown to greyish-yellow or olive brown, with rare tufting, radiating with dark brown colouration separating the margin from the centre; medium dense, irregular, flattened to slightly raised, dull with undulate edge, floccose or velvety, forming black stromatic tissues, embedded in agar after 8 weeks, agar tinted with yellowish-brown pigmentation.

Material examined: THAILAND, Pha Yao Province, Mae Jai District, Sri Don Kaew Village, on living leaves of Calathea sp. (Marantaceae), 19 August 2010, R. Phookamsak RP0049 (MFLU 11-0169), living culture = MFLUCC 11-0133; THAILAND, Chiang Rai Province, Muang District, Pakha village, on living leaves of Cordyline sp. (Asparagaceae), 15August 2010, S. Wikee (RP0067), MFLU11-0187, living culture = MFLUCC 11-0151.

Notes: Phaeosphaeria species (MFLUCC 11-0133 and MFLUCC 11-0151) were isolated from necrotic leaf spots symptoms on Calathea sp. and Cordyline sp. from Thailand. These two collections have similar morphological characters with Phaeosphaeria musae and Ph. panici (P. Syd.) Shoemaker & C.E. Babc. However they differ from Phaeosphaeria panici in having larger ascomata and asci and being associated with a different host. Phaeosphaeria panici lacks molecular data to confirm a natural placement, while these isolates form a strongly-supported clade (70 % MP) with Ph. musae in the Phaeosphaeria sensu stricto clade. Based on the ITS and TEF1 pair wise comparisons and multigene phylogenetic evidence, it appears that these isolates are conspecific with Phaeosphaeria musae. Thus, they are identified as Phaeosphaeria musae, and the species is reported associated with leaf spot disease on Calathea sp. and Cordyline sp. from Thailand. Pathogenicity was not confirmed.

Phaeosphaeria thysanolaenicola Phookamsak & K.D. Hyde, sp. nov., Index Fungorum number: IF550735; Faces of Fungi number: FoF00263, Fig. 22.

Fig. 22
figure 22

Phaeosphaeria thysanolaenicola (MFLU 11-0157, holotype). a Ascomata on host surface. b Section through ascoma. c Section through peridium. d Cellular pseudoparaphyses. e–h Asci. i–l Ascospores. Scale bars: b = 20 μm, c, e, f, g, h, = 10 μm, d, i, j, k, l = 5 μm

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Etymology : Referring to the host on which the fungus was found.

Holotypus: MFLU 11-0157

Pathogen associated with Paraphaeosphaeria on leaves of Thysanolaena maxima. Sexual state: Ascomata 70–100 μm high, 100–130 μm diam., scattered or sometimes clustered, immersed, visible as raised, black dots on the host surface, uni- to bi-loculate, subglobose, brown to dark brown, ostiole central, with minute papilla. Peridium 5–10 μm wide, thin-walled, of equal thickness, composed of 1–3 layers of brown to dark brown, pseudoparenchymatous cells, arranged in textura angularis. Hamathecium composed of numerous, 2–5 μm wide, filiform, broad cellular pseudoparaphyses, with distinct septa, slightly constricted at the septa, embedded in a gelatinous matrix, rarely anastomosing at the apex. Asci (41.5–)43–50(−57) × 7–9 μm (\( \overline{x} \) = 49.3 × 8.4 μm, n = 25), 8-spored, bitunicate, fissitunicate, broadly cylindrical or cylindric-clavate, sessile to subsessile, apically rounded with ocular chamber. Ascospores (15–)17–19(−20) × 3–4 μm (\( \overline{x} \) = 17.8 × 3.8 μm, n = 30), overlapping 1–3-seriate, phragmosporous, narrowly fusiform, with obtuse ends, yellowish-brown to brown, 3-septate, rarely 1–2-septate, slightly constrict at the central septum, straight to curved, rough-walled, echinulate, surrounded by mucilaginous sheath. Asexual state: Unknown.

Culture characters: Colonies on PDA 33–35 mm diam. after 4 weeks at 25–30 °C, white to pale yellowish-brown at the edge, white with grey sectors in the centre; reverse white to yellowish at the edges, becoming orange brown to grey at the centre with orange radiations separating the edges from the centre; medium dense, circular to irregular, flattened to raised, or umbonate, rough with entire to slightly undulate edge, floccose to velvety or cottony, radiating in the lower part, not producing pigments.

Material examined: THAILAND, Chiang Rai Province, Phan District, Pukang Waterfall, on living leaves of Thysanolaena maxima Kuntze (Poaceae), 22 May 2010, R. Phookamsak (RP0036) (MFLU 11-0157, holotype), ex-type living culture = MFLUCC 10-0563 = MUCL.

Notes: Phaeosphaeria thysanolaenicola is most similar to Ph. oryzae as they share a similar ascoma, asci and ascospore size range. However, they differ in habitat and host preferences, as Ph. thysanolaenicola is associated with other fungi on leaf spots of Thysanolaena maxima. Whereas Ph. oryzae was found as a saprobe on Oryzae sativa. The ITS pair wise comparison shows that they differ in eight base positions. Phylogenetic analysis performed with a combination of ITS, LSU, RPB2 and TEF1 genes shows that Ph. thysanolaenicola forms a clade with Ph. chiangraina. Phaeosphaeria thysanolaenicola differs from Ph. chiangraina in its smaller asci and ascospores, habitat, host occurrence and ascospore ornamentation. Phaeosphaeria thysanolaenicola has echinulate ascospores while Ph. chiangraina has smooth-walled ascospores.

Phaeosphaeriopsis M.P.S. Câmara et al., in Câmara et al., Mycol. Res. 107(5): 519 (2003), Faces of Fungi number: FoF00264.

Saprobic or pathogenic on various monocotyledons. Sexual state: Ascomata scattered to clustered, immersed, visible as small, black spots on host surface, uni- to bi-loculate, globose to subglobose, glabrous, brown, ostiole central, penetrating through host surface. Peridium thin-walled, of equal thickness, composed of brown to dark brown, pseudoparenchymatous cells of textura angularis. Hamathecium composed of numerous, filiform, frequently anastomosing, broad cellular pseudoparaphyses, with distinct septa and constricted at the septa. Asci 8-spored, bitunicate, fissitunicate, broadly cylindrical to cylindric-clavate, subsessile to short pedicellate, apically rounded, with indistinct ocular chamber. Ascospores overlapping 1–3-seriate, phragmosporous, oblong to cylindrical, often with one enlarged cell, yellowish-brown to brown or reddish-brown, 3–5-transeptate, slightly curved, not constricted at the septa, walls smooth or rough, echinulate or verruculose, surrounded by mucilaginous sheath. Asexual state: see under Phaeostagonospora.

Type species: Phaeosphaeriopsis glaucopunctata

Phylogenetic study: Câmara et al. (2003); Schoch et al. (2009); Zhang et al. (2012); Quaedvlieg et al. (2013); Hyde et al. (2013), Thambugala et al. (2014).

Notes: Phaeosphaeriopsis was introduced by Câmara et al. (2003) and typified by Ps. glaucopunctata to accommodate non congeneric species of Paraphaeosphaeria. Phaeosphaeriopsis agavensis (A.W. Ramaley et al.) Câmara et al., Ps. amblyspora A.W. Ramaley, Ps. glaucopunctata (Grev.) Câmara et al., Ps. nolinae (A.W. Ramaley) Câmara et al., and Ps. obtusispora (Speg.) Câmara et al. were introduced in the genus based on morphological characters and phylogenetic analyses of Phaeosphaeriopsis glaucopunctata (CBS 653.86) (Câmara et al. 2003; Zhang et al. 2012; Thambugala et al. 2014). Farr et al. (2006) transferred Sphaeria phacidiomorpha to Phaeosphaeriopsis phacidiomorpha based on the type morphological characters and microsphaeropsis-like asexual state which was most similar to Phaeosphaeriopsis. However, Sphaeria phacidiomorpha differs from other Phaeosphaeriopsis species by its hyaline ascospores (Farr et al. 2006; Thambugala et al. 2014). Phaeosphaeriopsis phacidiomorpha is currently synonymized under the name Glomerella phacidiomorpha (Ces.) Petr. (Index Fungorum 2014; Thambugala et al. 2014).

Phaeosphaeriopsis musae was introduced and accommodated in Phaeosphaeriaceae by Arzanlou and Crous (2006) and is a pathogen causing leaf spots on banana. Subsequent phylogenetic analysis indicated that Ps. musae clusters with Phaeosphaeria oryzae and separate to Phaeosphaeriopsis glaucopunctata (Zhang et al. 2012; Hyde et al. 2013; Phookamsak et al. 2013, 2014). Thus, Thambugala et al. (2014) synonymized Phaeosphaeriopsis musae under the older name Phaeosphaeria musae Sawada (1959) based on morphological characters and phylogenetic evidence. Additionally, Thambugala et al. (2014) epitypified Phaeosphaeriopsis glaucopunctata and described the new species Phaeosphaeriopsis triseptata K.M. Thambugala & K.D. Hyde with sexual and asexual morphs. Six species are listed in Index Fungorum (2014) and 59 sequences are available in GenBank. Molecular analysis confirms the placement of Phaeosphaeriopsis in Phaeosphaeriaceae (Zhang et al. 2012; Hyde et al. 2013; Quaedvlieg et al. 2013; Thambugala et al. 2014; Wijayawardene et al. 2014b).

A collection of Phaeosphaeriopsis was made which causes leaf spots on leaves of Dracaena lourieri Gagnep in Thailand. Based on morphological characters and multigene phylogenetic analysis, a new species, Phaeosphaeriopsis dracaenicola, is introduced to accommodate this taxon in the present paper.

Phaeosphaeriopsis glaucopunctata (Grev.) M.P.S. Câmara et al. [as ‘glauco-punctata’], in Câmara et al., Mycol. Res. 107(5): 519 (2003), Faces of Fungi number: FoF00265, Fig. 23.

Fig. 23
figure 23

Phaeosphaeriopsis glaucopunctata (E00074283, syntype). a Herbarium label and specimens of Phaeosphaeriopsis glaucopunctata. b Ascomata on host surface. c Section through ascomata d Section through peridium. e Asci with pseudoparaphyses. f Cellular pseudoparaphyses. g Ascus. h–i Asci stained by KOH reagent. j–o Ascospores. Scale bars: c = 100 μm, d–i = 20 μm, j–o = 5 μm

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≡ Cryptosphaeria glaucopunctata Grev., Fl. Edin.: 1-478 (1824)

Saprobic on Ruscus aculeatus (Asparagaceae). Sexual state: Ascomata 110–145 μm high, 130–220 μm diam, scattered, solitary, immersed, visible as small, brown spots surrounded by pale yellowish region, uniloculate, subglobose, glabrous, with central black ostiole penetrating through host surface. Peridium 7–18 μm wide, thin-walled, of unequal thickness, composed of 3–4 layers of thickened, brown, pseudoparenchymatous cells, arranged in a textura angularis. Hamathecium composed of sparse, 1–3 μm wide, filiform, frequently anastomosing, broad cellular pseudoparaphyses, with distinct septa, constricted at the septa. Asci (55–)60–75(−76) × (11–)12–14(−15) μm (\( \overline{x} \) = 68.2 × 13.4 μm, n = 20), bitunicate, 8-spored, broadly cylindrical to cylindric-clavate, subsessile to short pedicellate, apically rounded with indistinct ocular chamber. Ascospores 18–20(−22) × (3–)4–5.5 μm (\( \overline{x} \) = 20 × 4.8 μm, n = 20), overlapping 1–3-seriate, phragmosporous, cylindrical, narrower and longer at the lower cell, brown to dark brown, 4-septate, often enlarged at the forth cell, rough-walled, echinulate, surrounded by mucilaginous sheath. Asexual state: Conidiomata pycnidial, scattered, solitary, immersed to superficial (in agar), uniloculate, globose, glabrous, brown to black, ostiolate, with black droplets. Conidiomata walls thin, comprising 2–3 layers of yellowish-brown to brown, pseudoparenchymatous cells, arranged in textura angularis. Conidiophores reduced to conidiogenous cells. Conidiogenous cells holoblastic, phialidic, annellidic, with a short collarette, ampulliform to subcylindrical, hyaline, lining the inner cavity of conidioma. Conidia amerosporous, cylindrical to oblong, with rounded ends, hyaline to yellow, becoming yellowish-brown at maturity, aseptate, smooth to finely verruculose, accumulating in a huge bluish-black slimy mass at the tip of ostiole (from Thambugala et al. 2014).

Material examined: UK, near Edinburgh, Slateford, on dead leaves of Ruscus aculeatus L. (Asparagaceae), 15 August 1824, R.K. Greville (E 00074283, syntype).

Phaeosphaeriopsis dracaenicola Phookamsak & K.D. Hyde, sp. nov., Index Fungorum number: IF550737; Faces of Fungi number: FoF00266, Fig. 24.

Fig. 24
figure 24

Phaeosphaeriopsis dracaenicola (MFLU11-0193, holotype). on living leaves of Dracaena lourieri. a Leaf lesion on host. b Ascomata visible on host surface. c Section through ascoma. d Section through peridium. e Asci with pseudoparaphyses. f–i Asci. j Ascospores stained in Indian ink. k–n Ascospores. o Conidiomata forming on bamboo pieces on water agar. p Pycnidia. q Section through pycnidium. r Section through pycnidial wall. s–u Conidiogenous cells. v Conidia. w–ab Conidia. Scale bars: c = 100 μm, p, q = 50 μm, d, e, f, g, h, i = 20 μm, j, k, l, m, n, r, s, v = 10 μm, t, u, w, x, y, z, aa, ab = 2 μm

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Etymology: referring to the host, of which the fungus was found.

Holotypus: MFLU 11-0193

Pathogenic causing necrotic leaf spot on Dracaena lourieri; lesions 3–6 cm, infecting mostly on leaf margin, oval-shaped, pale brown to yellowish-brown, separated from healthy tissue by a reddish-brown to orange-brown margin. Sexual state: Ascomata 88.5–170 μm high, 140–200 μm diam., solitary to gregarious, immersed, visible as slightly raised, small black spots on host surface, subglobose, brown to dark brown, ostiole central. Peridium 5–17.5 μm wide, thin-walled, of equal thickness, composed of 3–4 layers of dark brown, pseudoparenchymatous cells, arranged in textura angularis, outer layer comprising 2–3 layers of thick-walled cells, inner layers comprising 1–2 layers of thin-walled hyaline cells. Hamathecium composed of numerous, 1.5–4 μm wide, frequently septate, frequently anastomosing, branching, broad cellular pseudoparaphyses, embedded in mucilaginous matrix. Asci 62–78 × 9–12 μm (\( \overline{x} \) = 71.5 × 10.8 μm, n = 25), 8-spored, bitunicate, fissitunicate, cylindrical to cylindric-clavate, short pedicelate, apically rounded, with well-developed ocular chamber. Ascospores 16–19 × (3.5–)4–5 μm (\( \overline{x} \) = 17.4 × 4.5 μm, n = 30), overlapping 2–3-seriate, phragmosporous, oblong to cylindrical, initially yellowish becoming brown to reddish-brown, 4–5 septate, usually widest in the third cell, narrower at the lowest cell, smooth-walled, guttulate, surrounded by thick mucilaginous sheath. Asexual state: forming on bamboo pieces on water agar (WA) after 8 weeks. Conidiomata 42–135.5 μm high, 50–140 μm diam., pycnidial, solitary, superficial on bamboo pieces, or semi-immersed on agar, visible as black slimy or shiny spots on surface, globose to subglobose or cup-shaped when sectioned through conidiomata, centrally ostiolate. Conidiomata walls 5–12 μm wide, composed of 3–4 layers of dark brown cells, arranged in textura angularis, outer layers comprising 2–3 layers of thick-walled cells, brown to dark brown, paler at the base, inner layer comprising 2–3 layers of thin-walled cells, hyaline. Conidiophores simple, rarely branched, doliiform to cylindrical or ampulliform, septate, hyaline, mostly reduced to conidiogenous cells. Conidiogenous cells 4–12 × 3–4 μm, holoblastic, phialidic, single, discrete, sometimes integrated, ampulliform or cylindric-clavate, hyaline, arising from basal stratum. Conidia 3–4.5 × 3–4.5 μm (\( \overline{x} \) = 3.9 × 3.6 μm, n = 30), amerosporous, globose to subglobose, initially hyaline, becoming brown to dark brown, aseptate, smooth-walled.

Culture characters: Colonies on malt extract agar (MEA) 34–34.5 mm diam. after 20 days at 25–30 °C, white to cream at the margins, pale yellowish to yellowish-brown in the middle and pale brown to brown or orange-brown at the centre, with small white to grey droplets; reverse white to cream at the margins, brown to orange-brown in the middle and pale yellowish to yellowish at the centre; medium dense, irregular, flattened to slightly raised, with rough surface, undulate edge with entire to slightly radiating margins, floccose to fairly fluffy, no pigment produced in agar.

Material examined: THAILAND, Chiang Rai Province, Muang District, Nang Lae Village, on living leaves of Dracaena lourieri Gagnep (Asparagaceae), 21 September 2010, S. Wikee, RP0073 (MFLU11-0193, holotype), ex-type living culture = MFLUCC 11-0157 = MUCL; ibid. Chiang Mai Province, Mae Wang District, Doi Inthanon, Royal Project, on living Dracaena loureiri, 16 November 2010, R. Phookamsak, RP0109 (MFLU 11-0229, paratype), ex-paratype living culture = MFLUCC 11-0193 = MUCL.

Notes: Phaeosphaeriopsis dracaenicola is typical of Phaeosphaeriopsis, due to the asexual state, habitat and host preferences and it shares similar morphological characters to Ps. agavensis, Ps. nolinae, and Ps. obtusispora in its brown, 5-septate ascospores. However, these can be differentiated by size of ascomata, asci and ascospores. Phaeosphaeriopsis dracaenicola has smaller ascospores, asci and ascomata than the other species in the genus, and ascospores are smooth-walled, while the other species have rough-walled, echinulate or punctuate ascospore ornamentation. In addition, Phaeosphaeriopsis dracaenicola was found associated with lesions on leaves of Dracaena, while the other species are saprobic on Agave and Nolina (Ramaley 1997; Câmara et al. 2001, 2003; Thambugala et al. 2014). Pathogenicity was not confirmed. Phaeosphaeriopsis dracaenicola formed an asexual state similar to Ps. agavensis and Ps. obtusispora, but differed from Ps. nolinae as conidia are globose. Multigene phylogenetic analysis shows that Ps. dracaenicola formed a robust clade (100 % ML/ 100 % MP/ 1.00 PP) at the basal of Phaeosphaeriopsis species in Phaeosphaeriaceae.

Phaeostagonospora A.W. Ramaley, Mycotaxon 61: 351 (1997), Faces of Fungi number: FoF00267.

Saprobic on Nolina erumpens (Torr.) S. Wats. Sexual state: reported as Phaeosphaeriopsis. Asexual state: Conidiomata pycnidial, scattered, solitary, immersed or subepidermal, uniloculate, elongated to globose, flattened at the apex against the host epidermis, with small papilla penetrating the host epidermis. Conidiomata walls thick, composed of 3–6 layers of brown-walled cells, with 1–2 hyaline interior layers, abundant light brown hyphae from walls associated with host substrate. Conidiophores reduced to conidiogenous cells. Conidiogenous cells holoblastic, discrete, doliiform, ampulliform or irregular in shape, hyaline, smooth-walled, lining inner cavity of conidiomata. Macroconidia phragmosporous, ellipsoidal to cylindrical, with rounded ends, brown, 1–3-septate, rough-walled, foveolate. Microconidia globose or ellipsoidal, hyaline, smooth-walled, forming on the conidiomatal wall near the ostiole (from Ramaley 1997).

Type species: Phaeostagonospora nolinae A.W. Ramaley, Mycotaxon 61: 351 (1997).

Phylogenetic study: None

Notes: When Câmara et al. (2003) introduced the genus Phaeosphaeriopsis; they also mentioned the asexual state of this genus as Phaeostagonospora. This was based on co-occurrence on the host (Ramaley 1997; Câmara et al. 2001). The sexual and asexual connection was, however, not equivocally confirmed. Thambugala et al. (2014) epitypified Phaeosphaeriopsis glaucopunctata and described the asexual state from cultures. The asexual state of P. glaucopunctata differs from Phaeostagonospora nolinae as conidia are aseptate, smaller, and verruculose, while Phaeostagonospora nolinae has septate conidia and it produces larger microconidia (Ramaley 1997; Thambugala et al. 2014). Therefore the link between Phaeostagonospora and Phaeosphaeriopsis requires confirmation.

Sclerostagonospora Höhn., Hedwigia 59: 252 (1917), Faces of Fungi number: FoF00268.

Pathogenic or saprobic on various monocotyledons and dicotyledons. Sexual state: phaeosphaeria-like. Asexual state: Conidiomata pycnidial, scattered, solitary, immersed, uniloculate, globose, glabrous, dark brown to black, ostiole central, with papilla. Conidiomata walls thin, composed of thick-walled cells, arranged in textura angularis, dark brown, inner layers lined with a hyaline cells bearing conidiogenous cells. Conidiophores reduced to conidiogenous cells. Conidiogenous cells holoblastic, discrete, determinate, ampulliform to irregular, hyaline, smooth-walled, lining inner cavity of conidiomata. Conidia phragmosporous, cylindrical to tapered slightly to the base, obtuse at the apex, with truncate base, pale brown, septate, slightly constricted at the septa, thin and rough-walled, verruculose (from Quaedvlieg et al. 2013).

Type species: Sclerostagonospora heraclei (Sacc.) Höhn., Hedwigia 59: 252. 1917.

Phylogenetic study: Crous et al. (2008), Quaedvlieg et al. (2013).

Notes: Sclerostagonospora was introduced by Höhnel (1917) and typified by S. heraclei (Sacc.) Höhn. The genus is similar to Stagonospora but differs in having pigmented conidia (Sutton 1980; Quaedvlieg et al. 2013). Crous et al. (2008) included Sclerostagonospora species in their phylogenetic analysis and Sclerostagonospora form a clade with other Phaeosphaeria species. Quaedvlieg et al. (2013) introduced a new species, S. phragmiticola from Phragmites australis (Poaceae) and confirmed that Sclerostagonospora is asexual genus belongs to Phaeosphaeriaceae based on their phylogenetic investigation. Quaedvlieg et al. (2013) mentioned that the sexual state of this genus is phaeosphaeria-like.

In this study, two species of Sclerostagonospora form a distinct clade in our multigene phylogenetic analyses (Fig. 2). Sclerostagonospora phragmiticola (CBS 338.86) groups with Phaeosphaeria phragmiticola (CBS 459.84) in clade E and other genera in Phaeosphaeriaceae with weak bootstrap support. Whereas Sclerostagonospora cycadis (CBS 123538) clusters with Parastagonospora in clade H (Fig. 2). However, Sclerostagonospora is not represented by their type species. This genus needs better populating as it is presently not well-resolved.

Scolecosporiella Petr., Annls mycol. 19(1/2): 30 (1921), Faces of Fungi number: FoF00269.

Saprobic or pathogenic on various host plants. Sexual state: reported as Ophiosphaerella and Phaeosphaeria. Asexual state: Conidiomata stromatic, pycnidial, amphigenous, scattered to gregarious, immersed to subepidermal, uniloculate, glabrous, depressed-globose to globose or ovoid, pale brown to brown, ostiole central, circular, non-papillate, with dark tendrils of conidia extruding through ostiole. Conidiomata walls thin, composed of thin-walled, subhyaline to pale brown cells, arranged in textura angularis. Conidiophores reduced to conidiogenous cells. Conidiogenous cells holoblastic, annellidic, discrete, ampulliform, globose or irregular, hyaline, smooth-walled. Conidia phragmosporous, fusiform, with acute apex and a truncate base, subhyaline to pale brown, multi-septate, constricted at the septa, thick and smooth-walled, apical cell attenuated into a short, beak-like, appendage (from Nag Raj 1989).

Type species: Scolecosporiella typhae (Oudem.) Petr., Annls mycol. 19(1/2): 31 (1921).

≡ Hendersonia typhae Oudem., Ned. kruidk. Archf, 2 sér. 1: 255 (1873)

Phylogenetic study: None

Notes: Scolecosporiella was introduced by Petrak (1921) with S. typhae (Oudemans) Petr. as the type species. The genus was treated by Sutton (1968, 1980) and Sutton and Alcorn (1974) with five species being accommodated. Nag Raj (1989) re-examined the type species of Scolecosporiella, S. typhae and described the morphological characters in detail. Nag Raj (1989) distinguished Urohendersoniella mastigospora Petr. from Scolecosporiella typhae and reinstated the genus Urohendersoniella as this monospecific genus had been synonymized as Scolecosporiella mastigospora (Petr.) B. Sutton in Sutton (1980). Additionally, a new genus Orphanocoela was established to accommodate non-congeneric species of Scolecosporiella (Nag Raj 1989). Six species are listed in Index Fungorum (2014).

Scolecosporiella has been reported as the asexual state of Ophiosphaerella herpotricha (Farr et al. 1989; Câmara et al. 2000; Zhang et al. 2012). However, there are no phylogenic evidence to confirm the connection between the sexual and asexual state, thus the generic connection is still questionable.

Scolicosporium Lib. ex Roum., Fungi Selecti Galliaei Exs.: no. 676 (1880)

Type species: Scolicosporium fagi Lib. ex Roum., in Roumeguère, Revue mycol., Toulouse 2: 22, 676 (1880)

Phylogenetic study: Wijayawardene et al. (2013a)

Notes: Scolicosporium Lib. ex Roum. was introduced by Roumeguère (1880) and typified by S. fagi Lib. ex Roum. Sutton (1980) treated Scolicosporium fagi, the type species of Scolicosporium as a synonym of S. macrosporium (Berk.) B. Sutton and designated S. macrosporium as the type species of Scolicosporium. However, Scolicosporium fagi is currently reported as the type species in Index Fungorum (2014). Scolicosporium was treated as a hyphomycete by Spooner and Kirk (1982) and Seifert et al. (2011). However, Treigiene and Mel’nik (2002) introduced S. minkeviciusii Treigiene as coelomycete, while Wijayawardene et al. (2013a) confirmed the species as a coelomycete. Furthermore, Wijayawardene et al. (2013a) showed that S. minkeviciusii belongs to Phaeosphaeriaceae, Pleosporales, in their molecular analyses. However, the genus was not represented by the type species, hence further collections of the type species of Scolicosporium, S. fagi are essential to confirm the familial placement of Scolicosporium sensu stricto. Below we only discuss S. minkeviciusii.

Scolicosporium minkeviciusii Treigiene., in Treigiene.& Mel’nik, Mikol. Fitopatol. 36(6): 45 (2002), Faces of Fungi number: FoF00270.

Saprobic on bark of Quercus pubescens Willd. and Q. robur L. Sexual state: Unknown. Asexual state: Conidiomata 100–120 μm high, 120–180 μm diam., pycnidial, solitary to gregarious, uniloculate, elongate-globose, dark brown. Ostiole central, sometimes towards one side, papillate when young, opening longitudinally at maturity. Conidiomata walls comprising 3–4 layers, outer layers 8–10 μm wide, dark brown, cells of textura angularis, inner layers hyaline, 5–40 μm wide. Conidiophores 13–30 × 2–3 μm, 1–2-septate, branched at the base, hyaline, cylindrical. Conidiogenous cells annellidic, hyaline, cylindrical. Conidia 60–65 × 7–9 μm (\( \overline{x} \) = 62.2 × 7.7 μm, n = 20), phragmosporous, curved to sigmoid, pale to moderately dark brown, with hyaline end cells, 6–7-transverse eusepta, smooth-walled, tapered to the obtuse apex, base truncate (Wijayawardene et al. 2013a).

Septoriella Oudem., Ned. kruidk. Archf, 2 sér. 5: 52 (1889)

Type species: Septoriella phragmitis Oudem. [as ‘phragmitidis’], Ned. kruidk. Archf, 2 sér. 5: 54 (1889)

Phylogenetic study: Crous et al. (2014)

Notes: Septoriella was introduced by Oudemans (1889) with the generic type Septoriella phragmitis. Crous et al. (2014) introduced a new species Septoriella oudemansii based on a megablast nucleotide search in GenBank. The species was similar to Phaeosphaeria caricis, Ph. ammophilae and Ophiosphaerella herpotricha (Crous et al. 2014). The taxon clusters with the asexual species, Wojnowicia hirta and Amarenomyces ammophilae in Phaeosphaeriaceae in LSU gene phylogenetic analysis (data is not shown). However, the type species of Septoriella is not represented in GenBank. Thus, we tentatively include Septoriella in Phaeosphaeriaceae. Below we only discuss S. oudemansii.

Septoriella oudemansii Crous & Quaedvl., Persoonia, Mol. Phyl. Evol. Fungi 32: 281 (2014), Faces of Fungi number: FoF000304.

Saprobic on Phragmites australis (Cav.) Trin. ex Steud (Poaceae). Sexual state: Unknown. Asexual state: Conidiomata up to 250 μm diam., pycnidial, solitary, immersed, uniloculate, globose, glabrous, brown, ostiole central, with a brown, oozing, conidial cirrhus. Conidiomata walls composed of 3–4 layers, brown, arranged in textura angularis. Conidiophores reduced to conidiogenous cells. Conidiogenous cells 5–11 × 4–5 μm, ampulliform, hyaline, smooth-walled, apex with minute periclinal thickening or percurrent proliferation, lining the inner cavity. Conidia (17–)23–32(−35) × (3.5–)4(−6), solitary, subcylindrical, with obtuse ends, slightly wider in the middle, yellowish-brown, 3(−5)-septate, becoming slightly constricted at the septa with age, smooth-walled, with guttules and flared mucoid appendages at the ends (from Crous et al. 2014).

Setomelanomma M. Morelet, Bull. Soc. Sci. nat. Arch. Toulon et du Var 227: 15 (1980), Faces of Fungi number: FoF00271.

Type species: Setomelanomma holmii.

Phylogenetic study: Rossman et al. (2002), Schoch et al. (2009), Zhang et al. (2009, 2012), Hyde et al. (2013), Wijayawardene et al. (2014b).

Notes: Setomelanomma was introduced by Morelet (1980) as typified by S. holmii to accommodate the monotypic genus from France associated with branches of Picea. Morelet (1980) however, did not illustrate the fungus and did not provide a detailed description (Rossman et al. 2002; Zhang et al. 2012). Rossman et al. (2002) re-circumscribed the type species and compared the morphological characters with North American collections. Detailed descriptions using morphology and molecular phylogeny were given by Zhang et al. (2012). Rossman et al. (2002) placed the genus in Phaeosphaeriaceae due to the morphology and phylogenetic data and this was later accepted by various authors (Kodsueb et al. 2006, 2007; Schoch et al. 2009; Zhang et al. 2009, 2012; Hyde et al. 2013). The genus is morphologically typical of Phaeosphaeriaceae as the ascomata are relatively small, the peridium is thin-walled comprising pseudoparenchymatous cells, pseudoparaphyses are cellular; and brown to yellowish-brown, and ascospores are phragmosporous (Rossman et al. 2002; Zhang et al. 2012). Setomelanomma however, differs from other genera in Phaeosphaeriaceae by its setose ascomata. No asexual state is known for the genus. There are two species, S. holmii and S. prolata K.J. Leonard & Suggs listed in Index Fungorum (2014)

Based on the multigene phylogenetic analyses (Figs. 1 and 2), Setomelanomma forms a strongly supported clade with the asexual species, Xenoseptoria neosaccardoi and Paraphoma radicina (CBS 111.79) in Phaeosphaeriaceae and these genera may be the asexual morphs of Setomelanomma.

Setomelanomma holmii M. Morelet, Bull. Soc. Sci. nat. Arch. Toulon et du Var 36(no. 227): 15 (1980), Faces of Fungi number: FoF00272, Fig. 25.

Fig. 25
figure 25

Setomelanomma holmii (BPI 620251) a Herbarium label and specimens. b Ascomata on substrate. c Section through ascoma. d Section through peridium. e Cellular pseudoparaphyses. f–i Asci. j–n Ascospores. Scale bars: c = 50 μm, d, e, f, g, h, i = 20 μm, j, k, l, m, n = 10 μm

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Pathogenic on branches of Picea. Sexual state: Ascomata 100–150 μm high, 140–175 μm diam., scattered, gregarious semi-immersed to erumpent through host tissue, becoming superficial, visible as abundant, small black dots on the host surface, uniloculate, globose to subglobose, covered by setae, dark brown to black, with central ostiole. Peridium 12.5–26 μm wide, thin-walled, of equal thickness, composed of 5–7 layers of black, broadly pseudoparenchymatous cells, arranged in a textura angularis to textura globulosa. Hamathecium composed of numerous, 1.5–3 μm wide, filamentous, frequently anastomosing, broad cellular pseudoparaphyses, with distinct septa, embedded in mucilaginous matrix. Asci (67–)68–77(−85) × (13–)15–17(−20) μm (\( \overline{x} \) = 75.2 × 15.2 μm, n = 20), 8-spored, bitunicate, fissitunicate, broadly cylindrical to cylindric-clavate, subsessile to short pedicellate, apically rounded with well-developed ocular chamber. Ascospores (18–)21–23(−24) × (6–)7–9(−11) μm (\( \overline{x} \) = 21.3 × 8.2 μm, n = 20), overlapping 1–2-seriate, phragmosporous, ellipsoidal to broadly fusiform, initially hyaline to pale yellowish, becoming light brown to brown or yellowish-brown at maturity, initially forming one median septum, becoming 3-septate at maturity, slightly curved, often constricted at the median septum, smooth and thick-walled. Asexual state: see under Xenoseptoria neosaccardoi Quaedvlieg et al. (2013)

Material examined: USA, Kansas, Shawnee Coounty, on branches of Picea pungens Engelm (Pinaceae), 5 May 1959, B. Herbert (BPI 620251).

Setophoma Gruyter et al., in De Gruyter et al. Mycologia 102(5): 1077 (2010), Faces of Fungi number: FoF00273, Fig. 26.

Fig. 26
figure 26

Setophoma sacchari (MFLU 11-0190 and MFLU 12-2470) on living leaves of Saccharum officinarum. a Symptoms on host (MFLU 11-0190). b Ascomata immersed in host tissue (MFLU 11-0190). c Section through ascoma (MFLU 11-0190). d Section through peridium (MFLU 11-0190). e Cellular pseudoparaphyses (MFLU 11-0190). f–h Asci (MFLU 11-0190). i–k Ascospores (MFLU 11-0190). l Spore germination (MFLU 11-0190). m Immature conidiomata on living culture (MFLU 11-0190) n Conidiomata on WA (MFLU 12-2470). o Section through conidiomata (MFLU 12-2470). p Section through conidiomata walls (MFLU 12-2470). q–r Conidiogenous cells (MFLU 12-2470). s, t Conidia (MFLU 11-0190). u, v Conidia (MFLU 12-2470) Scale bars: n, o = 100 μm, c = 50 μm, d, e, f, g, h, l, p = 20 μm, q = 10 μm, i, j, k, r, s, t, u, v = 5 μm

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Pathogenic or saprobic on various hosts (Alliaceae, Asteraceae and Poaceae). Sexual state: Ascomata, scattered, gregarious, immersed, visible as raised, minute black dots on host surface, uniloculate, globose to subglobose, glabrous, brown to dark brown, ostiole central, with minute papilla. Peridium thin-walled, of equal thickness, composed of pseudoparenchymatous cells, arranged in flattened textura angularis to textura prismatica. Hamathecium composed of numerous, filiform, broad cellular, pseudoparaphyses, with distinct septa, anastomosing at the apex. Asci 8-spored, bitunicate, fissitunicate, cylindrical to cylindric-clavate, short pedicellate, apically rounded, with well-developed narrowly ocular chamber. Ascospores overlapping 2-seriate, phragmosporous, cylindrical to cylindric-clavate, hyaline, 3-septate, usually enlarge at the second cell from apex, smooth-walled with large guttules. Asexual state: produced on rice straw and bamboo pieces on water agar after four weeks. Conidiomata scattered to clustered, gregarious, superficial, globose to subglobose, setose, and sometimes glabrous when mature, initially light brown, becoming brown to dark brown. Conidiomata walls thin, composed of light brown to brown pseudoparenchymatous cells, arranged in a textura angularis. Conidiophores reduced to conidiogenous cells. Conidiogenous cells enteroblastic, phialidic, determinate, discrete, unbranched, oblong to cylindrical, or ampulliform, hyaline, slightly curved, aseptate, lining the inner cavity of conidioma. Conidia amerosporous, oblong to ellipsoidal, rarely irregular, with rounded or obtuse ends, hyaline, aseptate, smooth-walled (from De Gruyter et al. 2010; Phookamsak et al. 2014).

Type species: Setophoma terrestris (H.N. Hansen) Gruyter et al., in De Gruyter et al. Mycologia 102(5): 1077 (2010).

≡ Phoma terrestris H.N. Hansen, Phytopathology 19(8): 699 (1929)

Phylogenetic study: De Gruyter et al. (2010), Quaedvlieg et al. (2013), Phookamsak et al. (2014)

Notes: Setophoma was introduced by De Gruyter et al. (2010) to accommodate phoma-like species and is typified by S. terrestris. Two species, S. sacchari (Bitanc.) Gruyter et al. and S. terrestris (H.N. Hansen) Gruyter were accommodated in this genus. Quaedvlieg et al. (2013) introduced a new species Setophoma chromolaenae Quaedvlieg et al. on leaves of Chromolaena odorata from Brazil. No sexual state was reported. Phookamsak et al. (2014) reported the sexual state of Setophoma based on asexual-sexual connection of S. sacchari and phylogenetic data. The sexual state of Setophoma is similar to Phaeosphaeria species, but differs in ascospore colour and asexual state (Phookamsak et al. 2014). Setophoma chromolaenae, S. sacchari, S. terrestris and S. vernoniae Crous & Alfenas are presently accommodated in the genus (Index Fungorum 2014; Crous et al. 2014; Phookamsak et al. 2014). Based on multigene phylogenetic analysis, we accept Setophoma in Phaeosphaeriaceae which forms a single clade at basal of the family (Fig. 2).

Tiarospora Sacc. & Marchal, Revue mycol., Toulouse 7(no. 26): 148 (1885), Faces of Fungi number: FoF00274.

Saprobic or pathogenic on Ammophila Host, Deschampsia P. Beauv., Elymus L. (Poaceae) and Dryas grandis Jurtzev (Rosaceae). Sexual state: Unknown. Asexual state: Conidiomata scattered, gregarious, immersed to semi-immersed, uniloculate, globose, glabrous, brown, ostiole central, with short papilla. Conidiomata walls thick, composed on several layers, outer layers comprising thick-walled dark brown, flattened cells of textura angularis, inner layers comprising thin-walled paler cells. Conidiophores reduced to conidiogenous cells. Conidiogenous cells holoblastic, sympodial, discrete, doliiform, cylindrical or lageniform, hyaline, smooth-walled. Conidia didymosporous, broadly ellipsoid, with obtuse to flattened ends, hyaline to pale brown, 1-septate, continuous, thick and smooth-walled, with guttules or irregular or cap-like gelatinous appendage (from Sutton 1980).

Type species: Tiarospora westendorpii Sacc. & Marchal, Revue mycol., Toulouse 7(no. 26): 148 (1885).

Phylogenetic study: None

Notes: Tiarospora was introduced by Saccardo and Marchal (1885) and is typified by T. westendorpii Sacc. & Marchal. Four species were accommodated under Tiarospora; T. deschampsiae Krusch., T. perforans (Sacc.) Höhn., T. pirozynskii Chleb. and T. westendorpii Sacc. & Marchal. However, Tiarospora perforans was transferred to Montagnulaceae by Aptroot (2006). Tiarospora is similar to Tiarosporella Höhn.; however, they differ in the morphology of their conidia. Tiarospora has 1-septate conidia, while Tiarosporella has aseptate conidia (Abbas et al. 2012). Tiarospora was described in Sutton (1980); however, no family placement was given for the genus. Kirk et al. (2001, 2008) treated the genus under Phaeosphaeriaceae but did not mention any literature to confirm this and this was accepted by Wijayawardene et al. (2012). Oertel (2011) however, mentioned that the type species of Tiarospora, T. westendorpii is a synanamoph of Amarenographium metableticum, which is the asexual state of Amarenomyces, also in Phaeosphaeriaceae. Therefore the current placement of this genus is somewhat unclear. The genus is relatively poorly known, lacks molecular data and no sexual state is known. We tentatively place Tiarospora in Phaeosphaeriaceae until molecular data resolves this issue.

Vrystaatia Quaedvlieg et al. Stud. Mycol. 75: 372 (2013), Faces of Fungi number: FoF00275, Fig. 27.

Fig. 27
figure 27

Vrystaatia aloeicola, redrawn from Quaedvlieg et al. (2013). Scale bars = 10 μm

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Saprobic on Aloe maculata (Aloaceae). Sexual state: Unknown. Asexual state: Conidiomata pycnidial, clustered, gregarious, erumpent on culture, with substromata on host, globose, glabrous, black, ostiole central, surrounded by dark brown area exuding cirrhus of orange conidia. Conidiomata walls thin, composed of 6–8 layers, arranged in textura angularis, pale brown. Conidiophores reduced to conidiogenous cells. Conidiogenous cells enteroblastic, annellidic, discrete, globose to ampulliform, hyaline, smooth-walled, with prominent periclinal thickening, or proliferating several times percurrently near apex, producing macro- and microconidia. Macroconidia scolecosporous, subcylindrical to narrowly obclavate, apex obtuse to subobtuse, with truncate base, hyaline, (1–)3-septate, smooth-walled, with guttules, widest at the basal septum. Microconidia pear-shaped to globose or ellipsoidal, apex obtuse, with truncate base, hyaline, aseptate, smooth-walled (from Quaedvlieg et al. 2013).

Type species: Vrystaatia aloeicola Quaedvlieg et al. Stud. Mycol. 75: 372 (2013)

Phylogenetic study: Quaedvlieg et al. (2013).

Notes: Vrystaatia was introduced by Quaedvlieg et al. (2013) to accommodate a non-congeneric species of Septoria sensu stricto and typified by V. aloeicola. Vrystaatia is similar to Septoria sensu stricto, but differs in conidiogenous cells morphology and forms macro- and microconidia in culture, which was not typical of Septoria (Quaedvlieg et al. 2013). Based on multigene phylogenetic analysis (Fig. 2), Vrystaatia forms a single clade at the basal of Paraphoma, Setomelanomma and Xenoseptoria clade. Thus we accept Vrystaatia for which no sexual state is known in Phaeosphaeriaceae.

Wojnowicia Sacc., Syll. fung. (Abellini) 10: 328 (1892), Faces of Fungi number: FoF00276.

Pathogenic on monocotyledonous or dicotyledonous hosts. Sexual state: Unknown. Asexual state: Conidiomata pycnidial, scattered, solitary, immersed to superficial on host tissue, globose, glabrous or setose, dark brown, ostiole central, with minute papillate or pore-like opening. Conidiomata walls thick, composed of pseudoparenchymatous cells, outer layers thick-walled, arranged in textura angularis, dark brown, inner layers thin-walled, hyaline, thinner towards inner conidiogenous cells. Conidiophores reduced to conidiogenous cells. Conidiogenous cells enteroblastic, phialidic, discrete, determine, doliiform to ampulliform, hyaline, smooth-walled, with minute collarette and channel, lining inner cavity of conidiomata. Conidia phragmosporous, fusiform to cylindrical, with obtuse ends, pale brown to brown, continuous, septate, straight to curved, thin and smooth-walled, guttulate.

Type species: Wojnowicia hirta (J. Schröt.) Sacc., in Saccardo & Sydow, Syll. fung. (Abellini) 14(2): 960 (1899).

Phylogenetic study: De Gruyter et al. (2009), Wijayawardene et al. (2013b).

Notes: Wojnowicia was introduced by Saccardo (1892) and is typified by W. hirta. The genus was discussed by Sutton (1975, 1980) who accepted two species, W. hirta and W. ephedrae Hollós. Sutton (1980) treated W. tenella Pat. and W. graminis (McAlpine) Sacc. & D. Sacc as synonyms of W. hirta. Farr and Bills (1995) named a collection from Juniperus deppeana Steud. and Pinus edulis Engelm. As W. colluvium D.F. Farr & Bills. Eleven species are listed in Index Fungorum (2014). Molecular data is only available for two species in GenBank. De Gruter et al. (2009) included two strains of Wojnowicia hirta in their analysis and showed that W. hirta belongs to Phaeosphaeriaceae. Wijayawardene et al. (2013b) introduced a new species, W. viburni Wijayaw. et al. on leaves of Viburnum utile Hemsl. ex Forb. & Hemsl. from China and confirmed that Wojnowicia belongs in Phaeosphaeriaceae. They also suggested that Wojnowicia should be the asexual state of Ophiosphaerella based on phylogenetic evidence (Wijayawardene et al. 2013b). However, Wijayawardene et al. (2013b) and De Gruyter et al. (2009) did not include many isolates of Phaeosphaeriaceae in their analysis. In this study with several sexual and asexual genera, the multigene phylogenetic analysis (Fig. 2) indicates that Wojnowicia hirta (CBS 160.73, CBS 295.69) clusters with Amarenomyces, Ophiosphaerella herpotricha (CBS 620.86) and other Phaeosphaeria sensu lato species while W. viburni forms a single clade at the base of Neostagonospora. Therefore the findings of Wijayawardene et al. (2013b) and De Gruyter et al. (2009) that Wojnowicia may be an asexual state of Ophiosphaerella is questionable, while Ophiosphaerella is obviously polyphyletic and requires further work.

Xenoseptoria Quaedvlieg et al. Stud. Mycol. 75: 371 (2013), Faces of Fungi number: FoF00277.

Pathogenic on Lysimachia vulgaris var. davurica (Ledeb.) R. Knuth (Primulaceae). Sexual state: unknown but possibly Setomelanomma. Asexual state: Conidiomata pycnidial, scattered, solitary, immersed to erumpent, uniloculate, globose, glabrous, brown, ostiole developing 1–3 papilla or necks, exuding a pink to orange conidial mass. Conidiomata walls thin, composed of 4–8 layers of brown cells, arranged in textura angularis. Conidiophores reduced to conidiogenous cells, hyaline, septate, branched, smooth-walled. Conidiogenous cells enteroblastic, mono- to polyphialidic, with prominent periclinal thickening, or percurrent proliferation, discrete or integrated, ampulliform to doliiform or subcylindrical, hyaline, smooth-walled. Conidia scolecosporous, cylindrical to obclavate, tapering to subobtuse apex, with obtuse base, straight to irregular curved, hyaline, septate, smooth-walled, with guttules (from Quaedvlieg et al. 2013)

Type species: Xenoseptoria neosaccardoi Quaedvlieg et al., Stud. Mycol. 75: 371 (2013).

Phylogenetic study: Quaedvlieg et al. (2013)

Notes: Xenoseptoria was introduced by Quaedvlieg et al. (2013) and was typified by X. neosaccardoi. The genus was introduced to accommodate a single species associated with leaf spot disease of Lysimachia vulgaris var. davurica (Primulaceae). The genus can be distinguished from Septoria sensu stricto as Xenoseptoria has multi-papillate conidiomata and mono- to polyphialidic conidiogenous cells (Quaedvlieg et al. 2013). In multigene phylogenetic analysis, Xenoseptoria clusters with the sexual genus Setomelanomma and the asexual genus Paraphoma with strong support (91 % ML, 1.00 PP). The genus is closely related to Setomelanomma. These genera may need placing in synonymy with Setomelanomma, but it would be unwise to do so until the tree becomes better populated.

Genera transferred to other families

Botryosphaeriaceae

Recent accounts of Botryosphaeriaceae include those of Liu et al. (2012) and Hyde et al. (2013, 2014)

Metameris Theiss. & Syd., Annls mycol. 13(3/4): 342 (1915), Faces of Fungi number: FoF00289.

Saprobic or parasitic on ferns and Bougainvillea. Sexual state: Ascostromata scattered, solitary to gregarious, erumpent through host epidermis, visible as slightly raised, black lines or black spots on the host surface. Locules clustered in pseudostroma, uni to bi-loculate, globose to subglobose, glabrous, black, ostiole in centre of each locule, with minute papilla. Peridium thick-walled, composed of thickened, black, scleroplectenchymatous cells, arranged in a textura angularis. Hamathecium composed of numerous, filamentous, broad cellular pseudoparaphyses, anastomosing at the apex. Asci 8-spored, bitunicate, cylindrical to cylindric-clavate, subsessile to short pedicellate, apically rounded, with well-developed ocular chamber. Ascospores overlapping 1–2-seriate, 2-celled, oblong to cylindrical, narrow towards lower end cell, hyaline, septate, smooth-walled, cytoplasm stains blue in cotton blue reagent, the large guttules in the upper part of the upper cell not staining. Asexual state: Unknown.

Type species: Metameris japonica

Phylogenetic study: None.

Notes: Metameris was established to accommodate Monographus japonicus Syd. by Theissen and Sydow (1915). Metameris forms linear pseudostromata, and has hyaline, 1-septate ascospores and Clements and Shear (1931) treated the genus under Dothideaceae (Zhang et al. 2012). Müller and von Arx (1962) treated Metameris as a synonym of Scirrhia and placed the genus in Mycosphaerellaceae (Crous et al. 2011). Later, von Arx and Müller (1975) assigned the genus to Pleosporaceae and did not accept that Metameris was synonymous with Scirrhia (Crous et al. 2011; Zhang et al. 2012). Recently, Metameris was accommodated in Phaeosphaeriaceae due to its small ascomata, broad cylindrical to obclavate asci with a knob-like pedicel and monocotyledons host preferences (Eriksson 2006; Lumbsch and Huhndorf 2007, 2010; Crous et al. 2011; Zhang et al. 2012; Hyde et al. 2013). However, the genus lacks molecular data to confirm its familial placement (Crous et al. 2011; Zhang et al. 2012).

There are five epithets listed for Metameris in Index Fungorum (2014); however, three species are synonymized under Scirrhia. In this study, we exclude Metameris from Phaeosphaeriaceae. The large, elongated pseudostromata, with a peridium of thickened scleroplectenchymatous cells, broad, or irregularly cellular pseudoparaphyses, ascus morphology and didymosporous ascospores are characters typical of genera in Botryosphaeriaceae where it is tentatively referred. Leptoguignardia is probably most similar (Liu et al. 2012) and should be compared in future studies of fresh collections.

Metameris japonica (Syd.& P. Syd.) Syd., in Theissen & Sydow, Annls mycol. 13(3/4): 342 (1915), Faces of Fungi number: FoF00290, Fig. 28.

Fig. 28
figure 28

Metameris japonica (holotype). a Herbarium label and specimens. b Ascostromata on host surface. c Section through ascostroma. d Section through peridium. e Pseudoparaphyses. f Asci and pseudoparaphyses strained in cotton blue reagent. g Ascus stained in cotton blue reagent. h–j Asci. k–m Ascospores. n–o Ascospores stained in cotton blue reagent. Scale bars: c = 100 μm, d, e, f, g, h, i, j = 20 μm, k, l, m, n, o = 10 μm

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≡ Monographus japonicus Syd. & P. Syd., Annls mycol. 10(4): 408 (1912)

Saprobic on Osmunda regalis var. japonica L. Sexual state: Ascostromata 170–240 μm high, 370–640 μm diam., clustered, solitary to gregarious, erumpent through host epidermis, visible as slightly raised, black, elongate lining on host surface. Locules 160–240 μm high, 200–250 μm diam, clustered, immersed in pseudostroma, uni to bi-loculate, globose to subglobose, glabrous, black, ostiole in centre of each locule, with minute papilla. Peridium 23–67 μm wide, thick-walled, of unequal thickness, slightly thick at the base towards sides of pseudostroma, composed of several layers of thickened, dark brown to black, scleroplectenchymatous cells, arranged in textura angularis. Hamathecium composed of numerous, 3–7 μm wide, filamentous, broad, irregular cellular pseudoparaphyses, with distinct septa which are constricted at the septum, embedded in mucilaginous matrix, anastomosing at the apex. Asci 80–110(−118) × (12.5–)13–18 μm (\( \overline{x} \) = 99.2 × 14.9 μm, n = 25), 8-spored, bitunicate, cylindrical to cylindric-clavate, narrow towards the apex, subsessile to short pedicellate, apically rounded with well-developed ocular chamber, arising from the basal ascoma. Ascospores (22–)24–28 × 6–8 μm (\( \overline{x} \) = 26.7 × 6.8 μm, n = 30), overlapping 1–2 seriate, 2-celled, oblong to cylindrical, narrow towards lower end cell, hyaline, 1-septate, constricted at the septum, smooth-walled, mostly upper cell larger than lower cell, cytoplasm stains blue in cotton blue reagent, the large guttules in the upper part of the upper cell not staining. Asexual state: Unknown.

Material examined: Japan, Mino, Kawauye-mura, on Osmunda regalis L. var. japonica (Osmundaceae), 10 May 1912, K. Hara (S-F7166, holotype).

Didymellaceae

For a discussion of Didymellaceae see Hyde et al. (2013). Zhang et al. (2012) questioned whether the placement Mixtura in Phaeosphaeriaceae was natural. The genus is considered to belong in Didymellaceae and is transferred herein.

Mixtura O.E. Erikss. & J.Z. Yue, Mycotaxon 38: 203 (1990), Faces of Fungi number: FoF00278.

Type species: Mixtura saginata

Phylogenetic study: None.

Mixtura saginata (Syd.) O.E. Erikss. & J.Z. Yue, Mycotaxon 38: 203 (1990), Faces of Fungi number: FoF00279, Fig. 29.

Fig. 29
figure 29

Mixtura saginata (S-F8935, lectotype). a Herbarium label and specimens. b Ascomata on leaf lesion. c Section through ascoma d Section through peridium. e Pseudoparaphyses stained in cotton blue reagent. f Immature ascus. g–i Asci. j Ocular chamber of asci stained in cotton blue reagent. k Immature ascospore stained in cotton blue reagent. l–m Mature ascospores. Scale bars: c = 100 μm, d–m = 20 μm

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≡ Leptosphaeria saginata Syd., Annls mycol. 37(4/5): 377 (1939).

Pathogen causing necrotic leaf spots on Chusquea serrulata Pilger. Lesions up to 2–5 cm long, infecting mostly the leaf margin, elongate cylindrical, pale brown to yellowish-brown, dry, separated from healthy tissue by a reddish-brown to purple-brown or black margin. Sexual state: Ascomata 120–260 μm high, 180–335 μm diam., scattered, sometimes clustered, solitary, immersed, visible as raised, small black dots on leaf lesions, uniloculate, globose to subglobose, glabrous, dark brown to black, ostiole central with minute papilla. Peridium 12–40 μm wide, thin-walled, of unequal thickness, thickened at apex at sides, composed of 3 or more layers of brown to dark brown, pseudoparenchymatous cells, arranged in a textura angularis, outer layers comprising 2–5 layers of thickened cells, brown to dark brown, inner layers comprising several layers of flattened cells, hyaline to pale brown. Hamathecium composed of numerous, 3–8 μm wide, filamentous, frequently anastomosing, broad cellular pseudoparaphyses, with distinct septa which are constricted at the septa, embedded in mucilaginous matrix. Asci (105–)112–135(−142) × (57–)60–100(−127) μm (\( \overline{x} \) = 121.3 × 80.3 μm, n = 20), 8-spored, bitunicate, fissitunicate, globose to subglobose, or ovoid, sessile to subsessile, apically rounded with well-developed ocular chamber, producing a few asci, arising from the basal ascoma. Ascospores 60–90(−95) × (19–)22–25(−27) μm (\( \overline{x} \) = 81.2 × 23 μm, n = 30), irregular overlapping 4-seriate, phragmosporous, broadly fusiform with rounded, germ pore at the ends, initially hyaline to pale brown, becoming yellowish-brown or brown at maturity, paler at the end cells, distoseptate with 7-septa, slightly curved, rough, thick-walled, surrounded by thin, distinct sheath. Asexual state: Unknown.

Material examined: ECUADOR, Tungurahua, Hacienda San Antonio pr. Baños, on leaves of Chusquea serrulata Pilger (Poaceae), 9 January 1938, H. Sydow (S-F8935, lectotype).

Notes: The monotypic genus Mixtura was introduced by Eriksson and Yue (1990) to accommodate Mi. saginata which causes leaf spots on Chusquea serrulata. Mixtura is an uncertain genus with morphological characters and habitat agreeing with several genera such as Mycosphaerella, Phaeosphaeria, Teratosphaeria, Trematosphaeria and Wettsteinina (Eriksson and Yue 1990; Zhang et al. 2012). Mixtura is similar to Mycosphaerella and Phaeosphaeria species in causing a leaf spot disease on monocotyledons, but differs from Mycosphaerella species in its centrum type (Eriksson and Yue 1990). Mixtura differs from Trematosphaeria due to its fusiform, olive brown ascospores and the genus differs from Wettsteinina as its rather thick-walled peridium and ellipsoidal asci (Eriksson and Yue 1990). Eriksson and Yue (1990) treated the genus in Phaeosphaeriaceae based on the peridium structure. However, Zhang et al. (2012) disagreed with Eriksson and Yue (1990) and placed the genus in Teratosphaeriaceae with Mixtura having relatively similar morphology to the type species of Teratosphaeria.

Mixtura is similar to other genera in Phaeosphaeriaceae such as Hadrospora and Loratospora in its saccate asci and large ascospores in Hadrospora. However, these genera differ in their centrum type. Mixtura forms dense, thick, cellular pseudoparaphyses, while Hadrospora forms trabeculate pseudoparaphyses, and pseudoparaphyses are lacking in Loratospora. Hadrospora is excluded as a member of Phaeosphaeriaceae in this study, but Loratospora is confirmed based on phylogenetic evidence. Mixtura is similar to Leptosphaerulina in habitat on host and asci. However, they differ in the ascospores and centrum type as Leptosphaerulina lacks pseudoparaphyses and has muriform ascospores. Mixtura has dense pseudoparaphyses and phragmosporous ascospores with distosepta. Therefore, we agree with Zhang et al. (2012) and exclude Mixtura from Phaeosphaeriaceae. The genus is tentatively placed in Didymellaceae.

Didymosphaeriaceae

For an account of this family see Ariyawansa et al. (2014b). Barria and Wilmia were transferred from Phaeosphaeriaceae to Didymosphaeriaceae in Ariyawansa et al. (2014b and are only briefly discussed here.

Barria Z.Q. Yuan, Mycotaxon 51: 313 (1994), Faces of fungi number: FoF00031.

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

Phylogenetic study: None.

Notes: Barria is a monotypic genus introduced by Yuan (1994) based on Barria piceae. The genus was reported as pathogenic on needles of Picea schrenkiana Fisch. & C.A. Mey. Barria is excluded from Phaeosphaeriaceae in this study based on its morphology. Barria is associated with gymnosperms, asci are clavate and short pedicellate, and ascospores are didymosporous with thick walls. Whereas the generic type, Phaeosphaeria is associated mostly with angiosperms, asci are often broad cylindrical to cylindric-clavate and subsessile and ascospores are phragmosporous (Yuan 1994; Zhang et al. 2012; Hyde et al. 2013; Quaedvlieg et al. 2013). Additionally, most genera with confirmed phylogenetic placement in Phaeosphaeriaceae have phragmospores, scolecospores, or muriform ascospores, but there are no didymosporous taxa (Zhang et al. 2012; Hyde et al. 2013; Quaedvlieg et al. 2013). Barria is associated in the same host family as Setomelanomma which was confirmed as phaeosphaeriacous based on phylogenetic evidence. Barria has didymosporous ascospores, while Setomelanomma has phragmosporous ascospores. Most didymosporous genera in Phaeosphaeriaceae lack molecular phylogenetic data to confirm their current placements. Furthermore, Barria is more similar to genera in Didymosphaeriaceae rather than Phaeosphaeriaceae (Ariyawansa et al. 2014b), where is it tentatively accommodated. Zhang et al. (2012) accepted Lautitia in Phaeosphaeriaceae; in contrast, all genera of marine and freshwater fungi (Carinispora, Isthmosporella and Ocala) were excluded from Phaeosphaeriaceae in Zhang et al. (2012). However, Lautitia differs from other genera in Phaeosphaeriaceae in having clavate asci with long pedicels and forming a pseudostromata with the peridium arranged in a textura porrecta. Other genera in Phaeosphaeriaceae mostly have cylindrical asci with subsessile to short pedicels and uniloculate ascomata with the peridium arranged in a textura angularis. Metameris forms large, elongated pseudostromata, with a thick-walled peridium of scleroplectenchymatous cells which is rarely found in other genera of Phaeosphaeriaceae. A new species of Letendraea (=Wilmia) was introduced based on phylogenetic evidence and belongs in Didymosphaeriaceae (see Ariyawansa et al. 2014b).

Wilmia Dianese et al., Mycologia 93(5): 1014 (2001), Faces of fungi number: FoF00046.

Type species: Wilmia brasiliensis Dianese et al., Mycologia 93(5): 1014 (2001).

Phylogenetic study: Ariyawansa et al. (2014a).

Notes: Wilmia was introduced by Dianese et al. (2001) to accommodate a monotypic genus, Wilmia brasiliensis that caused a leaf spot disease of Memora pedunculata (Vell.) Miers. in Brazil. Dianese et al. (2001) treated Wilmia in Phaeosphaeriaceae based on the morphological characters which fitted Barr’s concept (Barr 1987b, 1992a). Wilmia is similar to other genera in Phaeosphaeriaceae viz. Barria, Lautitia and Metameris in having didymosporous ascospores and a coelomycetous asexual morph (Dianese et al. 2001; Zhang et al. 2012). Zhang et al. (2012) tentatively placed Wilmia in Phaeosphaeriaceae as the genus is poorly known and lacks molecular data, although Zhang et al. (2012) mentioned that the genus should be accommodated in Leptosphaeriaceae based on its dicotyledonous host habit.

A Dothideomycete species was collected from living and dead leaves of Cordyline sp. in Thailand. The species also caused leaf spot disease and had similar morphological characters to Wilmia brasiliensis. Ariyawansa et al. (2014b synonymized Wilmia under Letendraea based on morphology and phylogeny. Thus, the new species collected from Cordyline sp. was accommodated in Didymosphaeriaceae and named Letendraea cordylinicola Phookamsak et al. (Ariyawansa et al. 2014b).

Leptosphaeriaceae

For a discussion on Leptosphaeriaceae see Hyde et al. (2013); Chaetoplea is considered to belong in this family and is transferred herein.

Chaetoplea (Sacc.) Clem., in Clements & Shear, Gen. fung., Edn 2 (Minneapolis): 275 (1931), Faces of Fungi number: FoF00280.

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

Saprobic on woody stems or herbaceous dicotyledonous. Sexual state: Ascomata solitary, gregarious, semi-immersed to superficial, or erumpent through host tissue, uniloculate, subglobose, glabrous, brown to dark brown, ostioles central, with short papilla. Peridium thick-walled, of equal thickness, composed of dark brown to black, scleroplectenchymatous cells of textura angularis or textura globulosa. Hamathecium composed of numerous, filamentous, distinctly septate, broad cellular pseudoparaphyses, anastomosing at the apex. Asci 8-spored, bitunicate, clavate to cylindric-clavate, short pedicellate, apically rounded, with an ocular chamber. Ascospores overlapping 1–2-seriate, phragmosporous, fusiform or broadly fusiform to ellipsoidal, brown to yellowish-brown, septate, slightly constricted at the septa, smooth-walled. Asexual state: Conidiomata pycnidial, scattered, solitary to gregarious, semi-immersed to erumpent, uniloculate, ellipsoidal to broadly fusiform with rounded apex, glabrous, dark brown to black, ostioles central, with pore-like opening. Conidiomata walls thin-walled, composed of flattened, scleroplectenchymatous cells, arranged in textura angularis to textura prismatica, brown to dark brown. Conidiophores reduced to conidiogenous cells. Conidiogenous cells holoblastic, phialidic, discrete, ampulliform, hyaline, smooth-walled, hard to distinguish from the inner wall cells. Conidia oblong to ellipsoidal, with obtuse ends, brown, 1-septate, smooth-walled.

Type species: Chaetoplea calvescens

Phylogenetic study: None

Notes: Chaetoplea was introduced by Clements and Shear (1931) to accommodate Dothideomycete species from woody substrates which included species transferred from Cucurbitaria, Leptosphaeria, Pleospora, Pyrenophora, Sphaeria and Teichospora (Shoemaker 1961; Wehmeyer 1961; von Arx and Müller 1975; Barr 1990b; Yuan and Barr 1994; Zhang et al. 2012). Barr (1987b, 1990a, b) treated Chaetoplea sensu lato in Phaeosphaeriaceae. The type specimen has not been recovered from any herbaria worldwide. Zhang et al. (2012) however observed the isotype which is a slide from the Farlow Herbarium (FH) and has only asci and ascospores. Zhang et al. (2012) mentioned that Chaetoplea may be heterogeneous in nature and occurs in a wide range of substrates such as herbaceous stalks, decorticated wood and periderm tissues.

The asexual state of Chaetoplea (for Pyrenophora calvescens (Fr.) Sacc.) was first reported as Dendryphium comosum Walk. (Saccardo 1883). Later Webster and Lucas (1959) treated Microdiplodia henningsii Staritz. as the asexual state of Chaetoplea and this was accepted by Barr (1987b). However, Crous et al. (2006) treated Microdiplodia as an asexual state of Karstenula placed in the family Montagnulaceae. Ramaley (1995) reported Parahendersonia as the asexual state of Chaetoplea dasylirionis being saprobic on Dasylirion (Agavaceae) and placed it in Leptosphaeriaceae. In this study, the asexual state of Chaetoplea was observed occurring on the same host as the lectotype. The asexual state of Chaetoplea forms oblong to ellipsoidal, brown, 1-septate, smooth-walled conidia which are microdiplodia-like.

After examining Fries exsiccatus (Scler. Suecicae exsiccatus no. 401) in the US National Fungus Collections (BPI) it was determined that this should be designated as a lectotype in this study. Based on morphology, Chaetoplea is more similar to Leptosphaeriaceae than Phaeosphaeriaceae and this supports the current placement of Chaetoplea as listed in Index Fungorum (2014). Therefore Chaetoplea should currently be placed in Leptosphaeriaceae. Recollection, epitypification, and molecular data are needed to confirm the natural placement of Chaetoplea.

Chaetoplea calvescens (Fr.) Clem., in Clements & Shear, Gen. fung., Edn 2 (Minneapolis): 275 (1931), Faces of Fungi number: FoF00281, Fig. 30.

Fig. 30
figure 30

Chaetoplea calvescens (lectotype of Sphaeria calvescens). a Label and specimen of Chaetoplea calvescens. b Ascomata on host surface. c Section through ascoma. d Section through peridium. e Immature asci with pseudoparaphyses. f–i Asci. j–n Ascospores. o Section through conidioma. p Section through conidioma wall. q, s Conidiogenous cells stained in cotton blue reagent. r Conidiogenous cell. t Conidia stained in cotton blue reagent. u–x Conidia. Scale bars: c, o = 100 μm, e = 50 μm, d, f, g, h, i, p, t = 20 μm, j, k, l, m, n = 10 μm, q, r, s, u, v, w, x = 5 μm

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≡ Sphaeria calvescens Fr., Sclerom. Suec.: no. 401 (1822)

Saprobic on herbaceous dicotyledonous hosts. Sexual state: Ascomata 140–220 μm high, 280–350 μm diam., scattered, solitary to gregarious, semi-immersed to superficial, visible as small black dots on the host surface, uniloculate, subglobose, glabrous, dark brown to black, ostiole central, with short papilla. Peridium 15–40 μm wide, thick-walled, of equal thickness, composed of several layers of scleroplectenchymatous cells, outer layers comprising 2–3 cell layers of thick-walled, black, textura angularis to textura globulosa, inner layers comprising 4–5 cell layers of thin-walled, hyaline to pale brown, flattened cells of textura angularis to textura prismatica. Hamathecium composed of numerous, 2–3 μm wide, filamentous, broad cellular pseudoparaphyses, with distinct septa, not constricted at the septa, anastomosing at the apex. Asci (80–)90–115(−120) × 11–13(−15) μm (\( \overline{x} \) = 100 × 12.1 μm, n = 20), 8-spored, bitunicate, fissitunicate, clavate, short pedicellate, apically rounded, with indistinct ocular chamber. Ascospores (14–)15–17(−119) × 6–8.5 μm (\( \overline{x} \) = 16.7 × 7.4 μm, n = 30), overlapping 1–2-seriate, phragmosporous, broadly fusiform to ellipsoidal, enlarged at the second cell, brown to yellowish-brown, 3-septate, rarely 2-septate, straight or curved, constricted at the septa, smooth-walled. Asexual state: Conidiomata growing with sexual state, 170–190 μm high, 150–190 μm diam., pycnidial, solitary to gregarious, semi-immersed to erumpent, visible as black dots on the host surface, uniloculate, ellipsoidal to broadly fusiform, with rounded apex, glabrous, dark brown to black, ostiole central, with minute papilla. Conidiomata walls 13.5–25 μm wide, thick-walled, of equal thickness, composed of several layers of flattened, scleroplectenchymatous cells, outer layers comprising 2–3 cell layers of thick-walled, dark brown to black cells, arranged in a textura angularis, inner layers comprising 3–5 layers of thin-walled, hyaline to pale brown cells, arranged in textura angularis to textura prismatica. Conidiophores arising from basal cavity of conidioma, mostly reduced to conidiogenous cells, hyaline, aseptate. Conidiogenous cells (3–)5–7(−11) × (3–)4–6(−7) μm (\( \overline{x} \) = 6.2 × 4.6 μm, n = 25) holoblastic, phialidic, discrete, ampulliform or lageniform, hyaline, aseptate, smooth-walled, hardly distinguished from inner wall cells. Conidia (11–)12–14(−15) × 4–6(−7) μm (\( \overline{x} \) = 13 × 5.2 μm, n = 30), solitary, didymosporous, oblong to ellipsoidal, with rounded or obtuse ends, brown, 1-septate, smooth-walled.

Material examined: SWEDEN, on dead wood, 1822, E.M. Fries, no. 401 (Fries SBARBARO COLLECTION), BPI (lectotype of Sphaeria calvescens designated here).

Sphaerellopsis Cooke, Grevillea 12(no. 61): 23 (1883), Faces of Fungi number: FoF00292.

Type species: Sphaerellopsis quercuum Cooke, Grevillea 12(no. 61): 23 (1883).

Phylogenetic study: Nischwitz et al. (2005), Bayon et al. (2006).

Notes: Based on ITS gene phylogenetic analysis (data not shown), Sphaerellopsis forms a basal clade in Leptosphaeriaceae. For details see under Eudarluca.

Pleosporaceae

Pleoseptum is considered to belong in this family and is transferred herein. For an account of the family see Hyde et al. (2013).

Pleoseptum A.W. Ramaley & M.E. Barr, Mycotaxon 54: 76 (1995), Faces of Fungi number: FoF00282.

Type species: Pleoseptum yuccaesedum.

Phylogenetic study: None.

Pleoseptum yuccaesedum A.W. Ramaley & M.E. Barr, Mycotaxon 54: 76 (1995), Faces of Fungi number: FoF00283, Fig. 31.

Fig. 31
figure 31

Pleoseptum yuccaesedum (BPI 802381, holotype). a Herbarium label and specimens of Pleoseptum yuccaesedum. b Ascomata on host surface. c Section through ascoma. d Section through peridium. e Pseudoparaphyses. f–h Ascospores. i Ascospore stained in Indian ink. j–l Asci. m Section through conidiomata. n Pycnidial wall. o Conidiogenous cells. p–q Conidiogenous cells stained in congo red reagent (p = annellidic, q = sympodial proliferation). r–u Conidia. Scale bars: c, m = 100 μm, e, j, k, l = 50 μm, d, n, o, r, = 20 μm, f, g, h, i, p, q, s, t, u = 10 μm

Full size image

= Camarosporium yuccaesedum Fairm., Mycologia 10(5): 261 (1918)

Saprobic on leaves of Yucca baccata Torr. Sexual state: Ascomata 250–360 μm high, 230–470 μm diam., scattered, solitary, immersed, visible as numerous black spots on host surface, uniloculate, globose to ovoid or conoid, dark brown to black, glabrous, with centrally opening ostiole penetrating the host epidermis. Peridium 70–90 μm wide, thick-walled, of unequal thickness, broad at the apex, composed of two layers of thickened pseudoparenchymatous cells, outer layer comprising several layers of thick-walled cells, arranged in a textura angularis, brown to dark brown, inner layers comprising several layers of thin-walled cells, hyaline, arranged in textura angularis. Hamathecium composed of numerous, 2–3 μm wide, filiform, distinctly septate, frequently anastomosing, narrow cellular pseudoparaphyses, embedded in mucilaginous matrix. Asci (125–)130–170(−185) × 18–22(−24) μm (\( \overline{x} \) = 153.7 × 20.9 μm, n = 25), 8-spored, bitunicate, fissitunicate, cylindrical to cylindric-clavate, short pedicellate, apically rounded, with well-developed ocular chamber. Ascospores (23–)25–28(−29) × 10–12(−13.5) μm (\( \overline{x} \) = 25.9 × 11.7 μm, n = 30), overlapping 1-seriate, dictyosporous, muriform, ellipsoidal to fusiform or obovoid, brown to dark brown, with 3–5 transverse septa, and 0–2 longitudinal septa, constricted at the central septum, smooth-walled, with distinct mucilaginous sheath. Asexual state: Conidiomata growing with the sexual state, 270–355 μm high, 265–390.5 μm diam., pycnidial, scattered, solitary to gregarious, immersed, visible as black spots on host surface, uniloculate, globose to ovoid or conoid, brown to dark brown, glabrous, with ostiolar opening penetrating the host epidermis. Conidiomata walls 30–65 μm wide, thick-walled, of unequal thickness, broad at the apex, composed of several layers of thickened pseudoparenchymatous cells, arranged in textura angularis, brown to dark brown. Conidiophores (10–)15–30(−45) × 5–7(−8.5) μm (\( \overline{x} \) = 23.8 × 6.4 μm, n = 20), discrete, oblong to cylindrical, usually unbranched, hyaline to brown, septate, arising from basal cavity of conidiomata. Conidiogenous cells enteroblastic, annellidic or sympodial. Conidia (22–)24–27(−28) × 10–12(−14) μm (\( \overline{x} \) = 25.9 × 11.7 μm, n = 30), muriform, oblong to obovoid or ellipsoidal with rounded or obtuse ends, with truncate base, initially hyaline, becoming brown to reddish-brown or dark brown at maturity, mostly with 3–5 transverse septa and 1–3 longitudinal septa, with several segments, smooth-walled.

Material examined: USA, Colorado. Montezuma County, hillside near entrance to Mesa Verde National Park, on leaves of Yucca baccata Torr. (Asparagaceae), 11 October 1992, A. Ramaley 9237 (BPI 802381, holotype).

Notes: The monotypic genus Pleoseptum was established by Ramaley and Barr (1995) to accommodate P. yuccaesedum. The genus was tentatively treated in Phaeosphaeriaceae based on its morphological characters and asexual morph (Lumbsch and Huhndorf 2010; Zhang et al. 2012; Hyde et al. 2013). Pleoseptum was compared with many genera such as Chaetoplea, Curreya, Heptameria and was considered most similar to Leptosphaeria based on its peridium structure (Ramaley and Barr 1995; Zhang et al. 2012). However, these genera differ in asexual states as in Pleoseptum conidia are dictyosporous, whereas in Leptosphaeria, they are stated to be amerosporous (De Gruyter et al. 2012; Hyde et al. 2013; Wijayawardene et al. 2014b).

Pleoseptum is a poorly known genus represented by only one species in Index Fungorum (2014) that lacks molecular sequence data in GenBank. The species has muriform spores which are similar to Dematiopleospora (Wanasinghe et al. 2014). Pleoseptum differs from Dematiopleospora by its peridium structure which is very thick, composed of heavily pigmented, thick-walled cells with ostiole and typical of Pleosporaceae in the former. Whereas the peridium is thinner, composed of lighter pigmented, thin-walled cells, with thick periphyses in the ostioles and typical of Phaeosphaeriaceae in Dematiopleospora. In Pleoseptum pseudoparaphyses are narrow, and additionally Pleoseptum forms a camarosporium-like asexual state. Pleoseptum is therefore excluded for Phaeosphaeriaceae whose genera typically have a thin-walled peridium and broad cellular pseudoparaphyses and tentatively placed in Pleosporaceae. However, Pleoseptum yuccaesedum needs recollecting and subjected to molecular study to confirm this placement is correct.

Camarosporium yuccaesedum was introduced as the asexual state of Pleoseptum yuccaesedum by Ramaley and Barr (1995) as both states co-inhabited the same host. Sexual-asexual links were connected by culture comparisons with conidia produced in culture from a sexual state (Ramaley and Barr (1995). Camarosporium however, has also been reported as the asexual state in various genera in Botryosphaeriales and Cucurbitariaceae (Kirk et al. 2008; Zhang et al. 2012; Hyde et al. 2013). Recent molecular based research on Camarosporium by Crous et al. (2014) and Wijayawardene et al. (2014a, c, d) showed that Camarosporium sensu lato is highly polyphyletic in Pleosporales, while Camarosporium sensu stricto is well-established genus in Pleosporineae, Pleosporales (Wijayawardene et al. 2014a). Thus, Neocamarosporium Crous & M.J. Wingf. (Pleosporales incertae sedis fide Crous et al. 2014), Paracamarosporium Wijayaw. & K.D. Hyde (Montagnulaceae fide Wijayawardene et al. 2014a), Pseudocamarosporium Wijayaw. & K.D. Hyde (Montagnulaceae fide Wijayawardene et al. 2014c) and Eucamarosporium Wijayaw. & K.D. Hyde (Didymellaceae fide Wijayawardene et al. 2014d) have been introduced to accommodate several camarosporium-like taxa in Pleosporales. Significantly, Ramaley and Barr (1995) reported microconidia from Camarosporium yuccaesedum which have not been reported from Camarosporium sensu stricto (Crous et al. 2013; Wijayawardene et al. 2014a). However, Crous et al. (2013) reported macroconidia from Camarosporium psoraleae Crous & M.J. Wingf. (current name Paracamarosporium psoraleae (Crous & M.J. Wingf.) Wijayaw. & K.D. Hyde fide Wijayawardene et al. 2014c), hence the status of Camarosporium yuccaesedum is uncertain. Camarosporium yuccaesedum is synonymized under Pleoseptum yuccaesedum herein.

Trematosphaeriaceae

For accounts of this family see Suetrong et al. (2011) and Hyde et al. (2013).

Hadrospora Boise, Mem. N. Y. bot. Gdn 49: 310 (1989), Faces of Fungi number: FoF00285.

Saprobic on woody or monocotyledonous hosts. Sexual state: Ascomata scattered, solitary, immersed to semi-immersed, or erumpent through host tissue, visible as tiny, small, black dots on the host surface, uniloculate, globose to subglobose, glabrous, dark brown to black, ostiole central with a minute papilla. Peridium thin-walled, composed of black, broad pseudoparenchymatous cells, arranged in a textura angularis. Hamathecium composed of numerous, filamentous, distinctly septate, frequently anastomosing, narrow cellular pseudoparaphyses. Asci 8-spored, bitunicate, fissitunicate, ovoid to ampulliform, sessile to minutely pedicellate, apically rounded, with indistinct ocular chamber. Ascospores overlapping 1–3-seriate, phragmosporous, broadly fusiform, vermiform, brown to reddish-brown, paler at the end cells, septate, smooth-walled. Asexual state: Unknown.

Type species: Hadrospora fallax

Phylogenetic study: None

Notes: Hadrospora was introduced to accommodate two species, originally placed in Trematosphaeria by Boise (1989), and was typified by Hadrospora fallax. Hadrospora has been reported from various hosts and habitats, both terrestrial and freshwater, from Belgium, China, Italy, Japan, Switzerland, and the United States (Boise 1989; Shearer and Crane 1971; Fisher and Webster 1992; Webster 1993; Tanaka and Harada 2003; Zhang et al. 2012). The genus was tentatively placed in Phaeosphaeriaceae due to its small ascomata, thin-walled peridium and narrow pseudoparaphyses (Boise 1989; Zhang et al. 2012). However, Hadrospora differs from other genera in Phaeosphaeriaceae by its large vermiform ascospores, ampulliform to saccate asci, trabeculate pseudoparaphyses and the hyphomycetous asexual morph with the type species which was treated as an amphibious species (Shearer 1993; Boise 1989; Tanaka and Harada 2003; Zhang et al. 2012). Other genera of Phaeosphaeriaceae are mostly terrestial and have cylindrical to cylindri-clavate asci with broad cellular pseudoparaphyses and produce coelomycetous asexual morphs. Hadrospora is similar to Mixtura in its large ascospores, and saccate asci; although Hardrospora has been reported as saprobic on various hosts, while Mixtura was found as plant pathogen of bamboo. The second species, Hadrospora clarkii (Sivan.) Boise is listed as Trematosphaeria clarkii Sivan. in Index fungorum (2014).

A zalerion-like asexual morph was reported for Hadrospora with conidia being produced on PDA after 4 months (Tanaka and Harada 2003). The genus is characterized by holoblastic, helicoid, multiseptate, brown to dark brown, smooth-walled conidia which are constricted at the septum (Tanaka and Harada 2003). Jones et al. (2009), however, treated Zalerion maritima as asexual morph of Lulwoana (Lulworthiales) and introduced a new genus, Halenospora to accommodate a non-congeneric species of Zalerion maritima.

Hardrospora is a poorly known genus which lacks sequences in GenBank. In this study, the proposal of Zhang et al. (2012) is followed excluding the genus from Phaeosphaeriaceae, based on its morphological characters, and it is tentatively placed in Trematosphaeriaceae.

Hadrospora fallax (Mouton) Boise, Mem. N. Y. bot. Gdn 49: 310 (1989), Faces of Fungi number: FoF00286, Fig. 32.

Fig. 32
figure 32

Hadrospora fallax (BR 5020009617141, holotype). a Fruiting bodies on host surface (BPI 878936). b Section through ascoma (BPI 878936). c Section through peridium (BPI 878936). d Pseudoparaphyses (BPI 878936). e–g Asci (BPI 878936). h–i Ascospores (BPI 878936). j–l Ascospores (BR5020009617141). Scale bars: B = 100 μm, d, e, f, g = 50 μm, c, h, i, j, k, l = 20 μm

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

Saprobic on woody hosts. Ascomata 200–350 μm high, 180–400 μm diam., scattered, sometimes clustered, solitary, immersed to semi-immersed, becoming erumpent through host tissue, visible as tiny, black dots on the host surface, uniloculate, globose to subglobose, glabrous, dark brown to black, ostiole central, with minute papilla, truncate at the apex. Peridium 20–50 μm wide, thick-walled, of equal thickness, composed of several layers of black, broad pseudoparenchymatous cells, arranged in a textura angularis. Hamathecium composed of numerous, 1–1.7 μm wide, filamentous, distinctly septate, anastomosing, narrow cellular pseudoparaphyses, embedded in mucilaginous matrix, with few asci. Asci 140–160(−186) × (54–)60–67 μm (\( \overline{x} \) = 157.3 × 61.8 μm, n = 10), 8-spored, bitunicate, fissitunicate, ovoid to ampulliform, sessile to minutely pedicellate, apically rounded with indistinct ocular chamber, broad at the apex. Ascospores (50–)65–75(−80) × (18–)21–23(−28) μm (\( \overline{x} \) = 72 × 21.3 μm, n = 25), irregularly overlapping 1–3-seriate, phragmosporous, broad fusiform, vermiform, widest at the median cells, reddish-brown to dark brown, with lighter end cells, 8-septate, slightly constricted at the median septum, smooth and thick-walled.

Material examined: BELGIUM, Beaufays, on recently cut wood, V. Mouton 245, (BR5 020009617141, holotype, microslide); SLOVAKIA, Kal’niste (East Slovakia), on stems of over wintered Triticum aestivum L. (Poaceae), 20 February 2009, M. Pastircak (BPI 878936).

Pleosporales, genera incertae sedis

For notes on Pleosporales, genera incertae sedis see Hyde et al. (2013)

Setophaeosphaeria Crous & Y. Zhang ter, Persoonia, Mol. Phyl. Evol. Fungi 32: 271 (2014), Faces of Fungi number: FOF000305.

Pathogenic on Hemerocallis fulva L. (Hemerocallidaceae). Sexual state: Ascomata scattered, immersed to subepidermal, uniloculate, globose, glabrous, ostiole central, somewhat papillate. Peridium composed of 2–3 layers of brown cells arranged in textura angularis. Hamathecium composed of hyaline, hyphal pseudoparaphyses. Asci 8-spored, bitunicate, narrowly ellipsoid, rostrate, short pedicellate, apically rounded with inconspicuous ocular chamber. Ascospores 2–3-seriate, fusoid to ellipsoidal, pale brown, 5-septate, enlarged at the second cell from apex, smooth-walled, with guttules and mucoid caps at each end. Asexual state: forming in culture. Conidiomata pycnidial, scattered, erumpent with papilla, uniloculate, globose, setose, brown, ostiole central, with round to ellipsoid papilla. Conidiomata walls thin, composed of 2–3 layers, of pale brown cells, arranged in textura angularis. Conidiophores reduced to conidiogenous cells. Conidiogenous cells ampulliform, hyaline, smooth, percurrently proliferating at the apex. Conidia subcylindrical, rounded or obtuse at the apex, base truncate, with marginal frill, hyaline, aseptate, smooth-walled, with guttules (from Crous et al. 2014).

Type species: Setophaeosphaeria hemerocallidis Crous & Y. Zhang ter, Persoonia, Mol. Phyl. Evol. Fungi 32: 271 (2014).

Phylogenetic study: Crous et al. (2014).

Notes: Setophaeosphaeria was introduced by Crous et al. (2014). The genus was established to accommodate non-congeneric species of Phaeosphaeria. Crous et al. (2014) introduced two new species, Setophaeosphaeria badalingensis and Sp. hemerocallidis and synonymized Phaeosphaeria setosa under the genus. Crous et al. (2014) mentioned that Setophaeosphaeria differs from the type species of Phaeosphaeria, Ph. oryzae, based on its asexual state. Phaeosphaeria oryzae forms a septoria-like asexual state, while Setophaeosphaeria species form phoma-like asexual states (Quaedvlieg et al. 2013; Crous et al. 2014). Based on phylogenetic analysis, the type species, Setophaeosphaeria hemerocallidis is closely related to Phaeosphaeria setosa, but can be distinguished by dimensions of their spores (Crous et al. 2014). Thus, Crous et al. (2014) introduced a new genus to accommodate these three species in Phaeosphaeriaceae. Phylogenetic analysis of ITS and LSU combined genes (data is not shown) indicates that the type species, Setophaeosphaeria hemerocallidis forms a clade outside Phaeosphaeriaceae, close to Cucurbitariaceae, while Sp. badalingensis clusters with Phaeosphaeria sensu stricto in Phaeosphaeriaceae. Thus, Setophaeosphaeria is polyphyletic and excluded from Phaeosphaeriaceae and tentatively placed in Pleosporales, genera incertae sedis. The second species, Setophaeosphaeria badalingensis may need to be synonymized under Phaeosphaeria based on its phylogenetic relationships.

Dothideomycetes, genera incertae sedis

One-hundred and sixteen genera are listed as genera incertae sedis in the class Dothideomycetes in Lumbsch and Huhndorf (2010) and updated to 322 genera in Wijayawardene et al. (2014b), as this included asexual genera. Lautitia previously placed in Phaeosphaeriaceae were also added here.

Lautitia S. Schatz, Can. J. Bot. 62(1): 31 (1984), Faces of Fungi number: FoF00287. Type species: Lautitia danica .

Phylogenetic study: None.

Lautitia danica (Berl.) S. Schatz, Can. J. Bot. 62(1): 31 (1984), Faces of Fungi number: FoF00288, Fig. 33.

Fig. 33
figure 33

Lautitia danica (NY 01390859). a Herbarium label and specimens of Lautitia danica. b Ascostromata on host tissue. c Section through ascostroma. d Section through peridium stained in congo red. e Asci with pseudoparaphyses strained in congo red. f–h Asci. i–j Asci stained in congo red. k–n Ascospores. o Ascospore stained in Indian ink. Scale bars: c = 100 μm, d, e = 20 μm, f–j = 10 μm, = 20 μm

Full size image

≡ Leptosphaeria danica Berl., Icon. fung. (Abellini) 1(2): 87 (1892)

Biotrophic in red algae (Chondrus crispus Stackh.). Ascostromata 600–1000 μm diam., scattered, immersed to semi-immersed, visible as raised, black circular or irregular regions on the host surface. Locules 180–240 μm high, 170–270 μm diam., uni to multi-loculate under pseudoclypeus, globose to subglobose, glabrous, pale brown, ostiole central, with pore-like opening. Peridium 10–22 μm wide, thin-walled, of unequal thickness, slightly thickened at the base, composed of several layers of prosenchymatous cells, arranged in textura epidermoidea at the apex and sides, with subhyaline to pale brown cells of textura porrecta at the base. Hamathecium composed of numerous, 1–1.5 μm wide, filamentous, anastomosing, rough-walled, narrow cellular pseudoparaphyses, with distinct septa, embedded in a mucilaginous matrix. Asci (57–)70–90(−104) × (12–)15–18(−20) μm (\( \overline{x} \) = 78.8 × 15.9 μm, n = 25), 8-spored, bitunicate, clavate, long pedicellate, apically rounded to truncate, with an indistinct ocular chamber. Ascospores (31–)35–40(−42) × 5–7 μm (\( \overline{x} \) = 35.9 × 5.8 μm, n = 30), overlapping 1–3-seriate, 2-celled, obclavate with narrow upper cell, mostly the upper cell longer than the lower cell, hyaline, 1-septate, slightly constricted at the septum, rough-walled with appendage on upper cell.

Material examined: FRANCE, Belclem near Roscoff, Finistère (Atlantic Ocean), on Chondrus crispus (Gigartinaceae), 5 October 1971 J. Kohlmeyer no. 2920 (NY 01390859); USA, Massachusetts, Gloucester, Halibus Point, 2 m deep, on Chondrus crispus, 28 September 1980, Kate van den Bosch; ibid. Rhode Island, Watch Hill Lighthouse (Atlantic Ocean), on Laminaria, 20 August 1966, J. Kohlmeyer_no. 1988 (NY 01390747); no. 1988a, (NY 01389139); Slide Box 15, Tray 20, S. Schatz (NY 01319510–NY 01319526).

Notes: The monotypic genus Lautitia was introduced by Schatz (1984) as typified by L. danica and placed in Dothideomycetes (Schatz 1984). A detailed description and taxonomic history were given by Wilson and Knoyle (1961) who treated the species then named Didymosphaeria danica (Schatz 1984). Berlese (1892) considered the species as a Leptosphaeria based on its hyaline, 1-septate, immature ascospores. Lind (1913) transferred the species to Didymosphaeria as D. marina (Rostrup) Lind, due to the ascospores turning brown and being 1-septate at maturity.Wilson and Knoyle (1961) could not find brown ascospores, but retained the species in Didymosphaeria. Schatz (1984) disagreed with Wilson and Knoyle (1961) as the species has cellular pseudoparaphyses, while genera in Didymosphaeria have trabeculate pseudoparaphyses. Thus Schatz (1984) designated a new genus, Lautitia, to accommodate the taxon. Schatz (1984) treated Lautitia as a member Phaeosphaeriaceae due to its pseudoparenchymatous to prosenchymatous peridium, and elongate, hyaline asymmetric ascospores. The genus was accepted in Phaeosphaeriaceae by Zhang et al. (2012) and Hyde et al. (2013).

In this study, we exclude Lautitia from Phaeaosphaeriaceae based on its morphology. Lautitia can be distingiushed from other genera in Phaeosphaeriaceae which differ in peridium structure, asci and ascospores. Lautitia has prosenchymatous perdial cells, clavate asci with long pedicels, and asymmetric didymosporous ascospores, while other genera in Phaeoaperiaceae have pseudoparenchymatous peridial cells, cylindrical to cylindric-clavate asci which are subsessile to short pedicellate, and phragmosporous, scolecosporous or muriform ascospores (Zhang et al. 2009, 2012; Hyde et al. 2013). It is difficult to place this species in any family or order any thus we refer it to Dothideomycetes, genera incertae sedis.