Meliolales

Abstract

The order Meliolales comprises the families Armatellaceae and Meliolaceae. These are black mildews that grow on the surface of host plants, often regarded as minor plant pathogens. In this study, types or specimens of 17 genera of Armatellaceae and Meliolaceae were borrowed from herbaria and re-examined. Armatella is accepted in Armatellaceae and Amazonia, Appendiculella, Asteridiella, Cryptomeliola, Endomeliola, Irenopsis and Meliola are accepted in the family Meliolaceae. Laeviomeliola is synonymized under Meliola. Ceratospermopsis, Ectendomeliola, Haraea, Hypasteridium, Leptascospora, Metasteridium, Ophiociliomyces, Ophioirenina, Ophiomeliola, Parasteridium, Pauahia, Pleomeliola, Pleomerium, Prataprajella, Ticomyces, Urupe and Xenostigme are excluded from Meliolaceae, and are treated as doubtful genera or placed in ascomycetes genera incertae sedis. The type species of each genus is re-described and illustrated with photomicrographs. Notes are provided and comparisons made. Two new species of Meliola and one new species of Irenopsis are also introduced with molecular data and we provide the most populated phylogenetic tree of Meliolomycetidae to date. Meliola thailandicum was found on Dimocarpus longan (Sapindales) and Acacia auriculiformis (Fabales) and confirmed to be the same species in the molecular analyses. This has important implications as the several hundred Meliola species are recognized based on host associations. Thus the same species being recorded from two unrelated hosts sheds doubt on Meliola species being host-specific.

Introduction

Species of Meliolales are obligate, biotrophic, foliar pathogens, which occur as epiphytes on leaves, stems, branches, and sometimes on petioles of vascular plants (Hansford 1961; Hosagoudar 1994, 1996, 2008, 2013; Mibey and Hawksworth 1997; Old et al. 2003). They are commonly known as the “black mildews” or “dark mildews” based on their black colonies up to one cm diameter, produced on the surface of hosts, and are widely distributed especially in subtropical and tropical regions (Goos and Andersons 1974; Ainsworth 1963; Hosagoudar 1994, 1996, 2003a, 2008; Saenz and Taylor 1999; Mibey and Hawksworth 1997; Old et al. 2003; Justavino et al. 2015; Maharachchikumbura et al. 2015). They are believed to show a high degree of host specificity, and therefore it is necessary to identify the host before determining the mildew (Hansford 1961; Hirata 1971; Hosagoudar and Goos 1989). The extent of host association, however, has not yet been determined, and molecular studies are needed to establish that species are host-specific. Species of Meliolales are not pathogens in a strict sense and do not cause extensive damage to host plants (Sabulal et al. 2006; Hosagoudar et al. 1997), however, they may increase the temperature and respiration in areas covered by the fungus, and also reduce photosynthesis as in the family Asterinaceae (Hongsanan et al. 2014). Taxa of the order obtain nutrients by producing haustoria in host cells, and heavy infections result in a dirty appearance on plants and economic products (Fig. 1) and may reduce their value (Hosagoudar et al. 1997, 2012). Wellman (1972) noted that this group produces significant affects on crops, but has never been seriously studied (Hosagoudar and Riju 2013).

Fig. 1

Meliolaceae on various hosts. a, b Dimocarpus longan. c, d Citrus maxima. e Litchi chinensis. f Mangifera indica. g Citrus reticulata

Species of Meliolales are often confused with sooty moulds because they coat fruits and leaves superficially with black colonies. Sooty moulds are saprobes which develop on leaf surfaces feeding on honeydew (Chomnunti et al. 2011, 2014). Species of Meliolales produce a variety of structures which can penetrate host cells for the uptake nutrients (Mibey and Hawksworth 1997; Saenz and Taylor 1999). This order is different from powdery mildews in morphology, but can develop in the same ecological habitats (Saenz and Taylor 1999).

The order Meliolales comprises the families Armatellaceae and Meliolaceae. The family Meliolaceae was proposed by Martin (1941) based on the generic type Meliola (Martin 1941; Hosagoudar 1994, 1996, 2008; Mibey and Hawksworth 1997; Old et al. 2003; Justavino et al. 2015; Maharachchikumbura et al. 2015). Members of the family are characterized by a biotrophic habitat, superficial dark hyphae, although sometimes the hyphae may be immersed in plant cells, with lateral hyphopodia and/or phialides. Superficial ascomata develop in the black, web-like colonies and are flattened or globose to subglobose, and have a dark peridium. Setae or appendages cover the ascomata and/or black mycelia in some genera. Asci are unitunicate, usually 3–4-spored, with brown, 3–4-septate ascospores (Mibey and Hawksworth 1997; Hosagoudar and Riju 2013). The asexual states of Meliolaceae species have been reported as phialides on hyphae, however, they are morphologically relatively poorly studied. Many characters of the family, such as types of hyphopodia, nature of phialides and structure of asci, are unclear, because various studies have provided different interpretations. The family Armatellaceae was established by Hosagoudar (2003b), with generic type Armatella Theiss. & Syd. Members of the family are characterized by superficial hyphae with hyphopodia, lacking phialides, ascomata which develop in the black colonies, cylindrical to subcylindrical asci, with 4–8-spores, and ascospores hyaline to brown at maturity, and 1-septate (Hosagoudar 2003b). The asexual morph is undetermined.

Because of their biotrophic lifestyles, species of Meliolales have yet to be grown in culture. Molecular data for species of Meliolales have therefore been obtained by directly extracting DNA from ascomata. Thite (1975) and Goos (1978) succeeded in germinating the ascospores, but growth following germination was limited and later ceased. We therefore focused on obtaining DNA and sequence data directly from the ascomata.

Taxonomic review

Superficial hyphae of Meliolales species are characterized by web-like colony formation on the host surface, usually branched, septate, and brown. Hyphopodia and setae are present in the genera Cryptomeliola, Irenopsis, and Meliola (Hansford 1961; Hosagoudar 1994, 1996, 2008, 2013; Mibey and Hawksworth 1997; Old et al. 2003). Endomeliola have intercellular hyphae which extend into the mesophyll (Hughes and Pirozynski 1994).

The nature of the hyphopodia is thought to be extremely important in identifying species in Meliolales (Hansford 1961), but this has never been proven by molecular evidence. The term hyphopodia was used by Gaillard (1891). Goos and Gessner (1975) noted that the hyphopodia is one kind of appressoria. Both terms hyphopodia and appressoria have been used in the literature, so Goos and Gessner (1975) suggested retaining usage of either. However, hyphopodia and appressoria are different because appressoria are produced on germ tubes, while hyphopodia are produced on hyphae (Emmett and Parbery 1975). These terms have been used interchangeably in many studies. As hyphopodia are produced on hyphae we recommend use of the term “hyphopodia” for the structures in Meliolales species. The hyphopodia are characterized by two cells, a short basal stalk cell bearing a single capitate hyphopodium. Hyphopodia may be alternate, opposite, or mixed alternate and opposite on hyphae depending on the species. Two kinds of hyphopodia are recognized in Meliolales, capitate and mucronate (Gaillard 1891). Capitate hyphopodia comprise a short stalk cell, bearing a single hyphopodia, which is the attachment and absorption organ. This later produces a single haustorium in the epidermal layer of host plants (Hansford 1961). Different species have differently shaped hyphopodia ranging from ovate, oblong, or angular to lobate. The capitate hyphopodia on the lower surface of leaves are mostly irregular in shape, while species found on the upper surface of leaves are globose to subglobose (Hansford 1961).

Mucronate hyphopodia were illustrated by Hughes (1981) and Mueller et al. (1991). Mucronate hyphopodia were interpreted as phialides and are conoid or ampuliform on hyphae, however they vary in shape in the same colony (Hansford 1961). Phialides produce conidia or phialoconidia (Hughes 1981, Mueller et al. 1991). There is however, no report to confirm that phialoconidia can develop into robust hyphae (Hughes 1981). Most species of Meliolaceae produce opposite phialides, or occasionally they are alternate or mixed with capitate hyphopodia on the hyphae. Some species in Meliolaceae lack phialides, but the apparent lack is probably due to the difficulty of observing such structures. Phialides appear to lack penetration or function (Hansford 1961; Goos 1974; Goos and Gessner 1975; Mueller et al. 1991). Species in Armatellaceae are characterized by hyphae without phialides, and the asexual morph is undetermined (Hosagoudar 2003b).

Ascomata are solitary or scattered on colonies of superficial hyphae and mostly develop at the centre of the colony. The ascomata is first flat, cells are radially arranged, and then become globose to subglobose, however a few genera (e.g., Amazonia), have flattened ascomata even at maturity. The wall of the ascomata comprises two strata, the outer strata having dark brown walls, and inner strata hyaline walls. A rounded pore is present at the apex of ascomata and is filled with hyaline periphyses and is covered by hyphal layers, and is thus hard to observe. The surface of ascomata is mostly verrucose. In Asteridiella, the surface has raised conoid cells, while conical appendages are present in Appendiculella species. The genus Irenopsis has setae on the surface of ascomata which are dark, smooth or rough, septate or aseptate, straight to coiled, with acute, rounded or hooked ends, and narrower than appendages in Appendiculella.

The asci of Meliolales have been reported differently in many studies. Müller and von Arx (1973) considered the asci of Meliolales species to be unitunicate. Eriksson (1981) introduced the term “Pseudoprototunicate” to describe the asci in Meliolales. A pseudoprototunicate ascus is characterized by thin walls, and is thought to have evolved from bitunicate asci, in which the endotunica is reduced. There is no opening mechanism, and they are evanescent at maturity (Eriksson 1981; Barr and Huhndorf 2001; Hofmann 2009). Hawksworth and Eriksson (1986) emended the description provided by Gäumann (1964) and concluded that Meliolales had bitunicate asci. However, Hosagoudar et al. (1997, 2012) used unitunicate to describe the asci of Meliolales species. We suggest using the term unitunicate asci to define the asci in Meliolales based on phylogeny, which place the species of Meliolaceae in Sordariomycetes (Maharachchikumbura et al. 2015).

The ascospores are very uniform in shape and septation in the family Meliolaceae. They are thick-walled, cylindrical to oblong, fusiform or obovoid, and slightly curved in some species. Some species have 3-septate ascospores, but in most they are 4-septate. In most species the central cells are widest and longest. The ascospores are more or less constricted at the septa, with smooth walls. The end cells are mostly rounded, but can rarely be conoid or apiculate (Hansford 1961; Mibey and Hawksworth 1997). All species in the family are hyaline when immature, and dark brown and thick-walled at maturity. However, some genera with two to multiseptate hyaline ascospores were included in Meliolaceae (Lumbsch and Huhndorf 2010). In this study, we only accept genera with 3–4-septate and brown ascospores in Meliolaceae. Ascospores in Armatellaceae are aseptate or 1-septate, hyaline to brown, and smooth or slightly rough-walled.

History of Armatellaceae and Meliolaceae

The family Meliolaceae was established by Martin (1941) without a Latin diagnosis but validated by Hansford (1946). Roger (1953) placed the Meliolaceae with nine other families in the order Hypocreales. This family has also been placed in the orders Dothideales, Erysiphales, Meliolales, Myriangiales and Hypocreales at various times (Martin 1941; Luttrell 1951, 1989; Roger 1953; Ainsworth et al. 1971; Müller and von Arx 1973; Yarwood 1973; Barr 1976; Eriksson 1981; Hawksworth et al. 1983). Monographs of the Meliolaceae were provided by Hansford (1961, 1963), with keys to the genera of the family based on morphology and host association (Hansford 1961). Ainsworth et al. (1971) and Eriksson and Hawksworth (1993) listed 50 genera in the family. This was reduced to 25 genera (Hawksworth et al. 1995) and later 22 genera (Kirk et al. 2008). A biogeographical distribution of Meliolaceae in India was provided by Hosagoudar (2006) and he included a key to the genera in the family. Lumbsch and Huhndorf (2010) placed Meliolaceae in the class Sordariomycetes with 26 genera. This was followed by Hyde et al. (2013) and Wijayawardene et al. (2014), who excluded Meliolaceae from Dothideomycetes. The family Armatellaceae was introduced by Hosagoudar (2003b). Armatellaceae contains a single monotypic genus (Table 1). Although Basavamyces Hosag. was included in Armatellaceae by Hosagoudar (2008), Hosagoudar et al. (2012) concluded that Basavamyces is not the member of Armatellaceae because of its 2-septate ascospores.

Table 1 Genera included in Armatellaceae and Meliolaceae by different researchers

Saenz and Taylor (1999) directly sequenced DNA from ascomata and their phylogenetic tree indicated that Meliolales was a member of Pyrenomycetes, which are close to Sordariales. Pinho et al. (2012a) reported that the phylogenetic placement of members of Meliolales was uncertain because the phylogenetic analyses (28S rDNA) were not strongly supported, but this order is monophyletic within Sordariomycetes. Kirk et al. (2001) introduced a new subclass, Meliolomycetidae (Sordariomycetes) for members of Meliolaceae, but without a description or diagnosis. The placement of Meliolomycetidae in Sordariomycetes was confirmed by Justavino et al. (2015) based on their phylogenetic tree and Maharachchikumbura et al. (2015) validated the subclass. Meliolales species are biotrophic and cannot be cultured, thus there are few sequences for species in GenBank.

Ecology

The Meliolales are biotrophs, or pathogens of living leaves or occasionally dead leaves, and are common on scrubs in open park-lands and on inaccessible canopies of rainforest trees (Hansford 1961; Hosagoudar 2003b, 2008). Species of Meliolales are associated with a variety of plant substrates including living leaves, and occasionally petioles, twigs, and branches (Hansford 1961; Hosagoudar 1994, 1996, 2008; Hosagoudar and Riju 2013; Mibey and Hawksworth 1997; Old et al. 2003). Species of Armatellaceae and Meliolaceae have a worldwide distribution (Kirk et al. 2008). They are especially common in the tropics and have an extended distribution to sub-temperate to temperate regions, and are generally lacking in arid regions (Hansford 1961; Hosagoudar 1994, 1996, 2008, 2013; Saenz and Taylor 1999; Mibey and Hawksworth 1997; Old et al. 2003; Justavino and Piepenbring 2007; Kirk et al. 2008; Piepenbring et al. 2011; Pinho et al. 2012b, 2013; Justavino et al. 2015). Studies on the dispersal of ascospores in Meliolales are lacking (Hansford 1961; Hosagoudar 1996; Nayar et al. 1998). Dispersal of species of Meliolaceae probably occurs by rain splash, insects, or air (Nayar et al. 1998). Meliolaceae species can survive after the leaves have been shed. When new leaves grow during the next season they are colonized by Meliolaceae with spores from leaves on the ground (Nayar et al. 1998). Meliolaceae in forests are more abundant during the cool season in tropical to subtropical regions. Even though forests are burnt leaving only the perennial root stocks; this group of fungi still occurs during the next season (Nayar et al. 1998). There is no report concerning the dispersal of ascospores in species of Armatellaceae.

Life cycle of Armatellaceae and Meliolaceae

Species of Meliolales have different infection mechanisms and the life cycle is illustrated in Fig. 2. Thimmaiah et al. (2013) stated that colonization starts when a mature ascospore attaches to the host substrate (Fig. 2a). A primitive stalk-cell bearing a single capitate, angular or lobate hyphopodium is mostly produced from the terminal cell of the ascospore; this is likely to be an intermediate step before mature hyphopodia development (Fig. 2b) (Tucker et al. 2010, Hongsanan et al. 2014). The hyphopodium then forms an apoplastic complex which is called an interaction apparatus, based on a study of Asteridiella callista (Justavino et al. 2014). At the contact zone between epidermal cells and hyphopodia, the apoplastic interaction apparatus forms a penetration pore so that the haustorium can develop into the cytoplasmic membrane and epidermal cells, and rarely in deeper tissues. Nutrient uptake begins (Fig. 2c) and the other cells of the ascospores produce superficial hyphae with frequent branches, and then alternate or opposite lateral hyphopodia are formed on short stalks (Fig. 2d). Initial infection and web-like colony formation on the leaf surface follows (Fig. 2e, f). In Meliola species, superficial hyphae produce strong, dark setae. Ascomata are formed laterally on the hyphae, at first being flat, cells are radially arranged, and then becoming globose to subglobose at maturity (Fig. 2g). The genus Amazonia is the only genus with flattened ascomata. The ascomata are filled with sterile tissue and asci which are evanescent at maturity (Fig. 2h). Ascospores are produced within the 2–4-spored asci in Meliolaceae and 4–8-spored asci in Armatellaceae. The asexual morph of Meliolaceae develops from the hyphae, and form ampuliform hyphopodia which are called “phialides” (Fig. 2i). These structures developed on the same hyphae as the capitate hyphopodia or sometimes are found on separate hyphae, but lack penetration or apparent function (Mueller et al. 1991). Conidia, which are 1-celled, thin-walled and hyaline, are produced within the neck of phialides (Fig. 2j). The conidiogenous cells are probably present at the base of the phialides (Hughes 1981). It is unlikely that these small phialoconidia can develop into robust hyphae (Hughes 1981) and their function is unclear. There has been a lack of research to establish the asexual morph cycle. There is no reported asexual morph in Armatellaceae, and phialides are lacking.

Fig. 2

Life cycle of Meliola sp. and its mucronate hyphopodia. a Ascospore. b Mature germinating ascospore. c Haustorium developing in epidermal cell. d Lateral hyphae forming superficially. e Colony formation. f Black spot of hyphae forming on the host surface. g Ascoma with hyphal setae. h Mature ascus. i Hyphopodia with phialides. j Phialoconidia

Host specificity

Species in Meliolales are considered to have a high degree of host specificity because they are biotrophic on living leaves. Numerous studies have introduced new species based only on host association, even though taxa are morphologically identical (Hosagoudar 1987). Some species on different hosts are distinct in morphology and in penetration mechanisms (Fig. 3) (Hosagoudar 2003a; Justavino and Piepenbring 2007). Hence, most species of Meliolales species have been justified based on host association, and it is essential to establish the host genus or family before identifying the fungal species. Whether Meliolales species are host specific needs verifying using molecular techniques.

Fig. 3

Host penetration by Meliolales species (a–c redrawn and modified from Yamamoto 1954, d redrawn from Hughes and Pirozynski 1994). a Hyphopodium of Meliolina octospora. penetrating epidermis cells. b Hyphopodium of Irenina rhaphiolepis Yam. penetrating epidermis and mesophyll cells. c Hyphopodium of Irenopsis coronata var. triumfettae Stev. penetrating epidermis cell. d Hyphae with hyphopodia of Endomeliola dingleyae developing in epidermis and mesophyll cells

Molecular phylogeny of Meliolales

The morphology of Meliolales species indicate that they belong to the class Sordariomycetes (Zhang et al. 2006). Pinho et al. (2012a) used sequence data of species in Meliolaceae and their phylogenetic tree confirmed the placement within Sordariomycetes. Justavino et al. (2015) mentioned a new subclass Meliolomycetidae based on their phylogenetic tree which was represented by a basal clade of Meliolaceae in Sordariomycetes. However, the phylogenetic placement of Meliolaceae members was uncertain within the clade because the phylogenetic analyses (28S rDNA) were not strongly supported, but monophyletic within Sordariomycetes (Pinho et al. 2012a). Maharachchikumbura et al. (2015) provided a backbone tree for Sordariomycetes based on LSU, SSU, TEF and RPB2 sequence data analysis, and their phylogenetic tree indicated that Meliolomycetidae is most closely related to Sordariomycetidae in Sordariomycetes as also found by Justavino et al. (2015). There are few sequences for the species of Meliolaceae as they are biotrophic and cannot be cultured (Maharachchikumbura et al. 2015). Most genera discussed here lack molecular data, however, the placements are based on morphological similarities and where possible phylogeny. No sequence data is available for any species of Armatellaceae.

Material and methods

Morphology

Type or other specimens of Appendiculella, Asteridiella, Cryptomeliola, Endomeliola, Haraea, Irenopsis, Laeviomeliola, Leptascospora, Meliola, Ophiociliomyces, Ophioirenina, Ophiomeliola, Pauahia, Pleomeliola, and Pleomerium were obtained from B, BPI, ILL, IMI, K(M), PDD, S, URM and ZT. Their morphology was observed under a stereomicroscope. Sections of ascomata were made free-hand. Morphological characters were observed and photographed under a compound microscope (Nikon 80i). Measurements were made using the Tarosoft (R) Image Frame Work v. 0.9.7.

Fresh specimens of Meliolaceae were collected in Thailand. Morphological characters were observed and photographed in the same way as herbarium specimens. Single spore isolation was performed following the method of Chomnunti et al. (2014) on PDA (potato dextrose agar) and MEA (malt extract agar), but was generally unsuccessful. Sequences were therefore obtained directly using dry fungal fruiting bodies.

Type specimens of the Meliolaceae are deposited in the Mae Fah Luang University Herbarium (MFLU), Chiang Rai, Thailand. Faces of fungi numbers and Index fungorum numbers were obtained as in Jayasiri et al. (2015) and Index fungorum (2015).

DNA isolation, amplification and sequencing

DNA extraction was made directly from dry fungal fruiting bodies to obtain sequence data. Extraction was started by placing individual ascomata in 1.5 ml sterilized tubes and leaving overnight at −20 °C. DNA extraction was performed by E.Z.N.A® Forensic Genomic DNA Extraction Kit (OMEGA Bio-tek Norcross GA 2013), GoTaq® Hot start-Promega, Lysis Buffer for Microorganism to Direct PCR (TaKaRa); following the manufacturer‘s instructions.

For GoTaq® Hot start-Promega, amplification conditions were set up for initial denaturation of 3 min at 95 °C, followed by 35 cycles of 27 s at 94 °C, 60 s at 56 °C and 90 s at 72 °C, and a final extension period of 7 min at 72 °C. PCR-products were cleaned from excessive nucleotides and primers by adding 2 μl of 1:5 diluted ExoSAP-IT® (Affymetrix, ExoSAP-IT® For PCR Product Cleanup) to 5 μl of PCR-product. The mix was incubated in the thermocycler for 30 min at 37 °C, followed by 15 min at 80 °C, and sequencing was performed on a ABI 3130xl sequencer in the sequencing service in the faculty of biochemistry at the Ruhr-Universität Bochum. For the specimens which were performed by using Lysis Buffer for Microorganism to Direct PCR (TaKaRa) Kit and E.Z.N.A® Forensic Genomic DNA Extraction Kit, amplification conditions were set up for initial denaturation of 5 min at 95 °C, followed by 35 cycles of 45 s at 94 °C, 45 s at 52 °C and 90 s at 72 °C, and a final extension period of 10 min at 72 °C. PCR-products were checked on 1 % agarose electrophoresis gels stained with ethidium bromide. The purification and sequencing of PCR products were done by Majorbio Co., China.

Phylogenetic analysis

Sequences data were downloaded from GenBank to supplement the dataset (Supplementary Table 1). Dothidea sambuci was selected as outgroup taxon. The representative sequences included taxa from Amplistromatales, Annulatascales, Boliniales, Chaetosphaeriales, Coniochaetales, Meliolales and Sordariales, as well as some from tentatively placed lineages downloaded from GenBank (Supplementary Table 1), and aligned by using Bioedit v.7.0.1 (Hall 2004) and Clustal X 2.0.11 (Thompson et al. 1997).

RAxML (Maximum likelihood) analysis was performed by using raxmlGUIv.0.9b2 (Silvestro and Michalak 2012). The search strategy was set to rapid bootstrapping and the analysis using the GTRGAMMAI model. The number of replicates was inferred using the stopping criterion (Pattengale et al. 2009). Maximum likelihood bootstrap values equal or greater than 50 % are given as the first set of numbers above the nodes (Figs. 4 and 5). The model of evolution was performed by MrModeltest 2.2 (Nylander et al. 2008). Posterior probabilities (PP) were determined by Markov Chain Monte Carlo sampling (BMCMC) (Rannala and Yang 1996; Zhaxybayeva and Gogarten 2002) in MrBayes v3.1.2 (Huelsenbeck and Ronquist 2001). Six simultaneous Markov chains were run for 1,000,000 generations and trees were sampled every 100th generation and 10,000 trees were obtained. The first 2000 trees, representing the burn-in phase were discarded, the remaining 8000 trees were used for calculating posterior probabilities (Cai et al. 2006, 2008). Bayesian posterior probabilities (BYPP) equal or greater than 0.80 are given as second set of numbers above the nodes (Figs. 4 and 5).

Fig. 4

RAxML maximum likelihood phylogenetic tree (LSU). The first set of numbers above the nodes are RAxML value expressed from 1000 repetitions with values above 50 % shown. The second set of numbers above the nodes are Bayesian posterior probabilities, with values above 0.8 shown. Strain numbers are indicated after species names. New sequence data are in red, types of the new species are in red bold, and other types are in black bold

Fig. 5

RAxML maximum likelihood phylogenetic tree (ITS). The first set of numbers above the nodes are RAxML values expressed from 1000 repetitions with values above 50 % shown. The second set of numbers above the nodes are Bayesian posterior probabilities, with values above 0.8 shown. Strain numbers are indicated after species names. New sequence data of Meliolaceae are in red, types of the new species are in red bold, and other types are in black bold

Results and discussion

Molecular phylogeny

Phylogenetic analyses used LSU (Fig. 4) and ITS sequence data (Fig. 5) and indicate that the Meliolales clade clusters in the class Sordariomycetes, subclass Meliolomycetidae. The Meliolaceae clade includes species of the family Meliolaceae with 100 % ML support and 1.0 PP support, and is sister to Rimaconus coronatus and R. jamaicensis; this is similar to the results of Maharachchikumbura et al. (2015). However, we are unable to clarify the placement of Armatellaceae because there is no sequence data from this family. Irenopsis walsurae clustered in the Meliolaceae clade and was closely related to other species of Irenopsis with relatively high bootstrap support (99 % ML, 1.0 PP support). Meliola mucunicola is placed within the genus Meliola in the clade of Meliolaceae and closely related to M. centellae (55 % ML support). Five strains of Meliola thailandicum clustered in the clade of Meliolaceae within the genus Meliola and are sister to M. variaseta (84 % ML, 0.9 PP support). Four strains of Meliola thailandicum which were found on Dimocarpus longan (MFLU15-0377, MFLU15-0379, MFLU 15-0044, MFLU 15-0047) and one from Acacia auriculiformis (MFLU15-0378), are the same species in the LSU tree (100 % ML, 0.1 PP support), and in the ITS tree (100 % ML, 0.1 PP support). Polymorphic nucleotides from sequence data of the ITS show a few base pair differences between the four strains of Meliola thailandicum from Dimocarpus longan and one strain from Acacia auriculiformis, and are not significant to separate them (Table 2). Hence, we conclude that this species is not host-specific. Appendiculella was not closely related to Asteridiella contradicting the phylogenetic tree of Justavino et al. (2015), however, these authors noted that the two species might be closely related as the conical appendages of Appendiculella could be homologous to the projecting conical cells on the ascoma wall in Asteridiella. Justavino et al. (2015), used LSU rDNA with 417 positions, and this may have resulted in a distinct separation of Appendiculella from Asteridiella. In the present study, the backbone tree used LSU sequence data with 635 positions and indicated that Appendiculella lozanellae is related to Asteridiella nitidae with 60 % ML and 0.9 PP support. Asteridiella obese, however, formed a distinct clade with Meliola species, and clustered in different subclades with Asteridiella nitidae, thus, more sequence data for Appendiculella and Asteridiella are needed to clarify their relationships. Meliola clerodendricola is sister to Endomeliola dingleyae (97 % ML, 1.0 PP support), however, these two stains are different in morphology. We suspect this may be wrong and needs to be verified in future, although host plants of Endomeliola dingleyae and Meliola clerodendricola belong to closely related orders in the euasterids (Jansen et al. 2007; Justavino et al. 2015). Some species have an uncertain placement, therefore, more gene and sequence data are needed to clarity the placement of the genera and species.

Table 2 Polymorphic nucleotides from sequence data of the ITS show a few base pair differences between the five strains of Meliola thailandicum. Polymorphisms unique to Meliola thailandicum are highlighted

Taxonomy

Key to genera of Meliolales

1. Hyphae with capitate hyphopodia, but without phialides	Armatella (Armatellaceae)
1. Hyphae with capitate hyphopodia and phialides	2 (Meliolaceae)
2. Hyphae intercellular	Endomeliola
2. Hyphae superficial	3
3. Ascomata flattened	Amazonia
3. Ascomata globose to subglobose	4
4. Hyphae with dark brown to black setae	5
4. Hyphae without setae (setae may be present on ascomata)	6
5. Hyphal setae with bulbous tips, apical part curved, covering ascomata	Cryptomeliola
5. Hyphal setae simple, straight or sometimes slightly curved	Meliola
6. Ascomata with appendages or setae	7
6. Ascomata without appendages and setae, with raised conical cells on the ascoma wall	Asteridiella
7. Ascomata with larviform to cylindrical appendages	Appendiculella
7. Ascomata with long setae, mostly curved at the apex	Irenopsis

Armatellaceae Hosag., Sydowia 55(2): 165 (2003)

Facesoffunginumber: FoF00723

Epiphytes on the surface of leaves. Superficial hyphae dense, septate, brown, with hyphopodia, hyphal setae lacking. Hyphopodia capitate, alternate or opposite on hyphae, 2-celled, brown. Sexual morph: Ascomata superficial on the surface of hosts, globose to subglobose, flattened when immature, developing on hyphae, the surface verrucose, covered with tuberculate projections, ascomatal setae and appendages lacking. Peridium thick, comprising two strata, outer stratum amorphous and black, inner stratum thick, comprising reddish to brown, scleroparenchymatous cells of textura angularis to globulosa. Hamathecium with evanescent paraphyses. Asci 4–8-spored, unitunicate, subcylindrical to ovoid, or clavate, evanescent. Ascospores hyaline to brown, ellipsoidal to oblong, aseptate when immature, 1-septate at maturity, constricted at the septum, with rounded ends. Asexual morph: Phialides absent (Hansford 1946; Hosagoudar 2003b).

Notes: The family Armatellaceae differs from Meliolaceae by having aseptate to 1-septate, hyaline ascospores and a different peridial wall type. Phialides are lacking on the hyphae in Armatellaceae, but are found in Meliolaceae.

Type: Armatella Theiss. & Syd., Annls mycol. 13(3/4): 235 (1915)

= Artallendea Bat. & H. Maia, Atas Inst. Micol. Univ. Recife 1: 221 (1960)

Facesoffunginumber: FoF00722

Epiphytes on the surface of leaves. Superficial hyphae dense, septate, brown, with hyphopodia, hyphal setae lacking. Hyphopodia irregularly capitate, alternate or opposite on hyphae, 2-celled, brown. Sexual morph: Ascomata superficial on the surface of hosts, globose to subglobose, flattened when immature, developing on hyphae, surface verrucose, covered with tuberculate projections, ascomatal setae and appendages lacking. Peridium thick, comprising two strata, outer stratum amorphous and black, inner stratum thick, comprising reddish to brown, scleroparenchymatous cells of textura angularis to globulosa. Hamathecium with evanescent paraphyses. Asci 4–8-spored, unitunicate, subcylindrical to ovoid, or clavate, evanescent. Ascospores hyaline to brown, ellipsoidal to oblong, aseptate when immature, 1-septate at maturity, constricted at the septum, with rounded ends. Asexual morph: Phialides absent (Hansford 1946; Hosagoudar 2003b).

Notes: There is no sequence data for any species of this genus. Direct sequencing from fresh specimens is needed to clarify the phylogenetic placement.

Type species: Armatella litseae (Henn.) Theiss. & Syd., Annls mycol. 13(3/4): 235 (1915)

Facesoffunginumber: FoF00721

Epiphytes on the surface of leaves. Superficial hyphae 5–7 μm diam., ($\overline{x}=7\ \mu m$, n = 10), dense, branched, septate, brown to reddish, with hyphopodia, hyphal setae lacking. Hyphopodia 15–17 μm diam. ($\overline{x}=16\ \mu m$ , n = 10), single stellate to sublobate on stalk cells, alternate on hyphae, 2-celled, brown to reddish. Sexual morph: Ascomata 210–235 × 174–192 μm ($\overline{x}=226 \times 180\ \mu m$, n = 10), superficial on surface of hosts, scattered, initially flattened, and becoming globose to subglobose, flattened when immature, developing on hyphae, surface verrucose, covered with tuberculate projections, ascomatal setae and appendages lacking. Peridium 43–59 um, thick, comprising two strata, outer stratum amorphous and black, inner stratum 29–37 um, thick, comprising reddish to brown, scleroparenchymatous cells of textura angularis to globulosa. Hamathecium with evanescent paraphyses. Asci 51–61 × 26–29 μm ($\overline{x}=54 \times 27\ \mu m$, n = 10), 4–8-spored, unitunicate, ovoid to clavate. Ascospores 33–35 × 10–11 μm ($\overline{x}=34 \times 10\ \mu m$, n = 10), 2–3-seriate, hyaline to light brown, ellipsoidal to oblong, aseptate when immature, 1-septate at maturity, constricted at the septum, rounded ends. Asexual morph: Undetermined.

Material examined

JAPAN, Province Awa, Tokushima, Kigasumi, on Litsea glauca (Thunb.) Siebold (Lauraceae), 25 December 1897, S. Kusano (SF70331, holotype).

Notes: We re-examined the type specimen of Armatella litseae, and accept this as a separate family from Meliolaceae characterized by aggregated colonies on the surface of host, stellate hyphopodia, and ascomata covered with tuberculate projections which are amorphous and black in section, and a thick inner stratum, comprising reddish to brown scleroparenchymatous cells of textura angularis to globulosa, aseptate to 1-septate ascospores, and lacking phialides. However, the phylogenetic placement needs clarification (Fig. 6).

Fig. 6

Armatella litseae (holotype). a Herbarium packet. b–d Colonies on leaves. e Section through ascoma. f Upper cell walls of ascomata when viewed in squash mounts. g, h Hyphopodia on hyphae. i Immature ascus in Melzers’ reagent. j, k Mature 1-septate ascospores in Melzer’s reagent. Scale bars: e = 100 μm, i–k = 20 μm, g, h = 10 μm

Meliolaceae W. Martin ex Hansf., Mycol. Pap. 15: 23 (1946)

Facesoffunginumber: FoF00741

Epiphytes or pathogens on leaves, occasionally on stems or branches, often forming web-like colonies. Superficial hyphae branched, septate, brown to dark brown, hyphal setae present or lacking, with hyphopodia. Hyphal setae developing from hyphae in Cryptomeliola and Meliola, septate, branched or unbranched at apex, or with bulbous apices or apical part in Cryptomeliola, brown to dark brown. Hyphopodia capitate on hyphae, variously shaped. Sexual morph: Ascomata superficial on surface of black, web-like colonies on host, mostly globose to subglobose, or flattened in the genus Amazonia. Peridium comprising dark brown cells of textura angularis when viewed in squash mounts, with two strata, outer stratum of brown to dark brown cells, or with raised conoid cells, appendages or setae, inner stratum of hyaline to pale brown cells. Hamathecium with evanescent paraphyses. Asci 2–4-spored, unitunicate, subglobose to broadly clavate, lacking an opening mechanism. Ascospores 2–3-seriate, hyaline when immature, brown to dark brown at maturity, ellipsoid or cylindrical to ovoid, 3–4-septate. Asexual morph: Phialides, ampuliform or flask-shaped on hyphae. Conidiogenous cells formed directly from vegetative hyphae. Conidia unicellular, small and hyaline (from Cannon and Kirk 2007).

Type species: Meliola Fr., Syst. orb. veg. (Lundae) 1: 111 (1825)

Notes: Phylogenetic analyses place Meliolaceae species in the class Sordariomycetes, and they were therefore accommodated in a new subclass Meliolomycetidae (Liu et al. 2015; Justavino et al. 2015; Maharachchikumbura et al. 2015). In our phylogenies (Fig. 4), the subclass Meliolomycetidae is clearly supported.

Genera included in Meliolaceae are Amazonia, Appendiculella, Asteridiella, Cryptomeliola, Endomeliola, Irenopsis and Meliola. These genera are characterized by superficial hyphae with hyphopodia or intracellular hyphae with hyphopodia in Endomeliola, and forming lateral haustoria that may penetrate the epidermis or mesophyll cells. Hyphal setae are found in Cryptomeliola and Meliola. Ascomata are globose to subglobose except in Amazonia where they are flattened. Ascomatal setae are found in Irenopsis and ascomatal appendages are found in Appendiculella. Asci are unitunicate and ascospores brown to dark brown at maturity with 3–4 septa. Basavamyces was transferred to Armatellaceae based on having superficial hyphae without phialides, cylindrical to subcylindrical asci and 1–2-septa ascospores (Hosagoudar et al. 2012), but we treat the genus in Sordariomycetes genera, incertae sedis base on its morphology. Ceratospermopsis, Ectendomeliola, Hypasteridium, Leptascospora, Metasteridium, Ophiociliomyces, Ophioirenina, Ophiomeliola, Prataprajella, Parasteridium, Pauahia, Pleomeliola, Pleomerium, Ticomyces, Urupe and Xenostigme are excluded from Meliolaceae. This is based on morphology or because taxa are doubtful due to lack of good type specimens. Hypasteridium, Metasteridium and Parasteridium were introduced without species name and their application is uncertain (Eriksson and Hawksworth 1988). Laeviomeliola is synonymized under Meliola based on morphology.

Amazonia Theiss., Annls mycol. 11(6): 499 (1913)

Facesoffunginumber: FoF00718

Epiphytes on the surface of leaves. Superficial hyphae branched, septate, darker at the septa, brown, with hyphopodia, hyphal setae lacking. Hyphopodia capitate, alternate on hyphae, near to hyphal septa, 2-celled, brown to dark brown. Sexual morph: Ascomata superficial on surface of host, mostly gregarious, rarely solitary, circular, flattened, borne under radiating hyphae, branching at the rim, producing lateral hyphopodia, ascomatal setae and appendages absent. Peridium arranged radially when viewed in squash mounts, poorly developed at the base, with three strata, outer stratum comprising a dark brown to black amorphous layer, central stratum comprising flattened, thick-walled, brown cells of textura angularis and inner stratum of hyaline to reddish brown flattened cells. Hamathecium with evanescent paraphyses. Asci 2-spored, unitunicate, ellipsoid to obovoid, lacking an opening mechanism, short pedicellate or apedicellate, evanescent. Ascospores 2-seriate, hyaline to brown, ellipsoid to subcylindrical, 4-septate, constricted and darker at the septa, smooth-walled. Asexual morph: Phialides ampuliform, sometimes curved, alternate or opposite on hyphae, mixed with capitate hyphopodia, pale brown to brown. Phialoconidia rarely observed, hyaline.

Notes: The genus Amazonia was introduced by Theissen (1913), with the type species Amazonia psychotriae (Henn.) Theiss. Amazonia is typical of other members of Meliolaceae based on its unitunicate asci, and brown to dark brown, 4-septate ascospores. Höhnel (1918) noted this genus as transitional between Meliola and Microthyriaceae based on its flattened ascomata typical of species in Microthyriaceae (Wu et al. 2011), while asci and ascospores are typical of species in Meliolaceae. Molecular analyses place Meliolaceae in the class Sordariomycetes, while the family Microthyriaceae is placed in Dothideomycetes (Hyde et al. 2013; Maharachchikumbura et al. 2015).

Type species: Amazonia psychotriae (Henn.) Theiss., Annls mycol. 11(6): 499 (1913)

≡ Meliola asterinoides var. psychotriae Henn., Hedwigia 43: 361 (1904)

≡ Amazonia psychotriae (Henn.) Theiss., Annls mycol. 11(6): 499 (1913) var. psychotriae

= Amazonia psychotriae var. major Hansf., Reinwardtia 3: 102 (1954)

Facesoffunginumber: FoF00716

Epiphytes on the surface of leaves. Superficial hyphae 4–5 μm diam., branched, septate, darker at the septa, brown, with hyphopodia, hyphal setae lacking. Hyphopodia 10–11 μm diam. ($\overline{x} = 10\mu m$, n = 10), capitate, alternate on hyphae, near to hyphal septa, 2-celled, brown to dark brown. Sexual morph: Ascomata 440–495 μm diam. ($\overline{x} = 455\ \mu m$, n = 5), superficial on surface of host, mostly gregarious, rarely solitary, circular, flattened, borne under radiating hyphae, branching at the rim, producing lateral hyphopodia, ascomatal setae and appendages absent. Peridium 20–26 μm thick, arranged radially when viewed in squash mounts, poorly developed at the base, with three strata, outer stratum comprising a dark brown to black amorphous layer, central stratum comprising flattened, thick-walled, brown cells of textura angularis and inner stratum of hyaline to reddish brown flattened cells. Hamathecium with evanescent paraphyses. Asci 45–50 × 23–28 μm ($\overline{x}=47 \times 26\ \mu m$, n = 10), 2-spored, unitunicate, ellipsoid to obovoid, lacking an opening mechanism, short pedicellate or apedicellate, evanescent. Ascospores 37–40 × 15–20 μm ($\overline{x}=39 \times 19\ \mu m$, n = 10), 2-seriate, hyaline to brown, ellipsoid to subcylindrical, 4-septate, constricted and darker at the septa, smooth-walled, with an evanescent sheath. Asexual morph: Phialides 7–8 μm ($\overline{x}=8\ \mu m$, n = 5), ampuliform, sometimes curved, alternate or opposite on hyphae, mixed with capitate hyphopodia, sometimes occurring on separate hyphae, pale brown to brown. Phialoconidia rarely observed, hyaline.

Material examined

BRAZIL, Amazonas, Rio Negro, Manaus, on leaves of Phychotria sp. (Rubiaceae), March 1901, E. Ule No. 3152 (B700014752, syntype); BRAZIL, Amazonas, Rio Negro, Manaus, on leave of Phychotria sp. (Rubiaceae), 1901, E. Ule (B70001475).

Notes: Amazonia was introduced based on its flattened ascomata, with the type species as A. psychotriae. Meliola asterinoides var. psychotriae Henn. is a synonym of A. psychotriae Theissen (1913). We examined and illustrated the syntype of A. psychotriae from B (Figs. 7 and 8). Molecular data is lacking for this genus.

Fig. 7

Amazonia psychotriae (syntype). a, b Herbarium packet and specimens. c Colony on substrate. d Ascoma viewed in squash mount. e, f Hyphopodia at rim of ascoma. g Section through ascoma. h Peridium. i Phialides. j Ascus in cotton blue reagent. k, l Immature asci. m, n Mature asci. o, p Ascospores with 4 septa. Note the thin sheath in p. Scale bars: d, g = 100 μm, e, f, i = 10 μm, h, j, l–p = 20 μm, k = 50 μm

Fig. 8

Amazonia psychotriae (B70001475). a, b Herbarium packet and specimen. c Colony on substrate. d Ascoma borne under radiating hyphae viewed in squash mounts. e Wall comprising radiating cells. f Hyphae with capitate hyphopodia. g Hyphopodia at rim of ascoma. h, i Immature asci. j Mature ascus. k–m Mature ascospores. Scale bars: d = 100 μm, e = 50 μm, f, g = 10 μm, h–m = 20 μm

Appendiculella Höhn., Sber. Akad. Wiss. Wien, Math.-naturw. Kl., Abt. 1 128: 556 (1919)

Possibly synonyms (from Index Fungorum 2015):

≡ Irenina F. Stevens, Annls mycol. 25(5/6): 411 (1927)

≡ Meliola subgen. Irenina (F. Stevens) Cif., Annls mycol. 36(2/3): 203 (1938)

Facesoffunginumber: FoF00720

Epiphytes on the surface of leaves or stems. Superficial hyphae branched, septate, darker at the septa, with hyphopodia, hyphal setae lacking. Hyphopodia capitate, angular or lobate, alternate or opposite on hyphae, 2-celled, brown. Sexual morph: Ascomata superficial on surface of host, mostly gregarious on superficial hyphae, subglobose to globose, thick-walled, lacking ascomatal setae, but with raised conoid cells, which may extend to form larviform to cylindrical appendages. Peridium comprising dark brown cells of textura angularis when viewed in squash mounts, with two strata, outer stratum of brown to dark brown cells of textura angularis, inner stratum of hyaline to pale brown flattened cells. Hamathecium with evanescent paraphyses. Asci 2–4-spored, unitunicate, oblong to obovoid, lacking an opening mechanism, short pedicellate or apedicellate, evanescent at maturity. Ascospores 2–3-seriate, hyaline to brown, fusiform to ellipsoid, 3–4-septate, slightly constricted and darker at the septa, smooth-walled. Asexual morph: Phialides ampuliform, alternate or opposite, mixed with capitate hyphopodia on hyphae, sometimes curved, brown. Phialoconidia not seen (Justavino and Piepenbring 2007).

Notes: The genus Appendiculella was introduced by Höhnel (1919) with the type species A. calostroma (Desm.) Höhn. and is characterized by conical appendages with transversely striate walls developing on the surface of the ascomata. This feature is similar to the conical ascomatal cells of Asteridiella (Justavino et al. 2015). Appendiculella species are distributed widely in the tropics (Kirk et al. 2008), with an 87 species epithets listed in Index Fungorum (2015). Most species were described based on host association. Justavino et al. (2015) provided the first sequence data for Appendiculella lozanellae (DQ508302), which grouped in the Meliolaceae clade in their phylogenetic tree. However, this relationship was weakly supported in the phylogenetic tree of Justavino et al. (2015). In this study, the phylogenies indicate a close relationship between Appendiculella lozanellae and Asteridiella nitidae (60 % ML support). However, the morphology of these two genera differs, and therefore we maintain these genera as distinct.

Type species: Appendiculella calostroma (Desm.) Höhn., Sber. Akad. Wiss. Wien, Math.-naturw. Kl., Abt. 1 128: 556 (1919)

≡ Sphaeria calostroma Desm., Bull. Soc. bot. Fr. 4: 22 (1857)

= Chaetosphaeria calostroma (Desm.) Sacc., Syll. fung. (Abellini) 2: 95 (1883)

= Meliola calostroma (Desm.) Höhn., Annls mycol. 15(5): 363 (1917)

Facesoffunginumber: FoF00719

Epiphytes on the surface of leaves. Superficial hyphae 5–6 μm diam. ($\overline{x}=5\ \mu m$, n = 10), branched, septate, darker at the septa, with hyphopodia, hyphal setae lacking. Hyphopodia 13–15 μm diam. ($\overline{x}=14\ \mu m$, n = 10), capitate, angular or lobate, alternate or opposite on hyphae, 2-celled, brown. Sexual morph: Ascomata 125–140 high × 140–150 μm diam. ($\overline{x}=125 \times 149\ \mu m$, n = 5), superficial on surface of host, mostly gregarious on superficial hyphae, subglobose to globose, thick-walled, lacking ascomatal setae, but with raised conoid cells, which may extend to form larviform to cylindrical appendages. Peridium 28–34 μm ($\overline{x} = 30\mu m$, n = 5), comprising dark brown cells of textura angularis when viewed in squash mounts, with two strata, outer stratum of brown to dark brown cells of textura angularis, inner stratum of hyaline to pale brown flattened cells. Hamathecium with evanescent paraphyses. Asci 42–57 × 24–33 μm ($\overline{x}=45\times 28\mu m$, n = 10), 2–4-spored, unitunicate, oblong to obovoid, lacking an opening mechanism, with short pedicellate or apedicellate, evanescent. Ascospores 35–37 × 12–13 μm ($\overline{x}=36 \times 12\ \mu m$, n = 10), 2–3-seriate, hyaline to brown, fusiform to ellipsoid, 3-septate, slightly constricted and darker at the septa, smooth-walled. Asexual morph: Undetermined.

Material examined

CHILE, Bío-Bío, Concepción, on surface of leaves of Geum chilense Balb. (Rosaceae), 5 June 1895, F.W. Neger (SF5748).

Notes: Appendiculella calostroma was established by Höhnel (1919) and infects plant species in Rosaceae (Hansford 1961; Justavino et al 2015) (Figs. 9 and 10).

Fig. 9

Appendiculella calostroma (S-F5748). a, b Herbarium packet and specimen. c Ascomata on surface of leaves. d Transverse section through ascoma. e Ascoma viewed in squash mount. f Peridium comprising cells of textura angularis. g Conical appendage on ascomata. h Hyphae with hyphopodia. i Immature ascus in Melzer’s reagent. j Mature ascus. k, l Mature ascospores with 3 septa. Scale bars: d, e = 50 μm, f, g, i–l = 20 μm, h = 10 μm

Fig. 10

Appendiculella lozanellae (redrawn from Justavino and Piepenbring 2007). a Transverse section through ascoma showing appendages on ascoma. b Ascus. c Ascospores with 4 septa

Asteridiella McAlpine, Proc. Linn. Soc. N.S.W. 22(1): 38 (1897)

Possible synonymy (from Index Fungorum 2015):

= Irene Theiss et al., Annls mycol. 15(3/4): 194 (1917)

= Meliola subgen. Irene (Theiss et al.) Cif., Annls mycol. 36(2/3): 203 (1938)

= Parasteridiella H. Maia, Publicações Inst. Micol. Recife 267: 25 (1960)

Facesoffunginumber: FoF00724

Epiphytes on the surface of leaves and stems. Superficial hyphae branched, septate, darker at the septa, brown, with hyphopodia, hyphal setae lacking. Hyphopodia capitate, alternate or opposite on hyphae, near to hyphal septa, 2-celled, brown. Sexual morph: Ascomata superficial on surface of host, mostly gregarious, rarely solitary on superficial hyphae, subglobose to globose, thick-walled, ascomatal setae and appendages absent. Peridium comprising dark brown cells of textura angularis when viewed in squash mounts, with two strata, outer stratum of dark brown cells of textura angularis, with raised conical cells, acute or rounded at the apex, and inner stratum of hyaline cells of textura porrecta. Hamathecium with evanescent paraphyses. Asci 2–4-spored, unitunicate, oblong to cylindrical. Ascospores 2–4-seriate, hyaline to brown, subcylindrical to oblong, 4-septate, slightly constricted and darker at the septa, not constricted in some species, smooth-walled (description modified from Hansford 1961, 1963; Mibey and Hawksworth 1997 and own observations). Asexual morph: Phialides rarely observed on hyphae, ampuliform, alternate or opposite, sometimes curved, pale brown to brown. Conidia hyaline (Hansford 1961; Hosagoudar 2013).

Notes: Asteridiella was introduced by McAlpine (1897) and differs from Meliola species in lacking hyphal setae. Most of the genera in Meliolaceae have been separated previously by setae and appendages. These characters seem hardly sufficient to separate Asteridiella from other genera, molecular data however, show Asteridiella as a distinct clade in Meliolaceae, sister to Appendiculella. Asteridiella may well be a distinct genus in having conical cells raised from the ascomata and in lacking ascomatal setae or appendages (Justavino et al. 2015). Species in Asteridiella are differentiated based on host as they are thought to be host-specific (Hansford 1961; Hosagoudar 2013). Less significant differences can be seen in the size of ascomata, hyphopodia shape and arrangement, however this needs testing at the molecular level (Hansford 1961; Hosagoudar 2013). Asteridiella species are commonly found in the tropics (Mibey and Hawksworth 1997; Kirk et al. 2008), with around 300 species estimated in Kirk et al. (2008), and almost 450 species epithets listed in Index Fungorum (2015). Molecular evidence placed Asteridiella nitidae Rodr. Just. (EF094839) and A. obesa (Speg.) Hansf. (DQ508302) in Meliolaceae as a distinct clade, however, the two species clustered in different subclades (Justavino et al. 2015). Therefore, molecular data is needed to clarify the distinctiveness of this and other genera.

Type species: Asteridiella solani McAlpine, Proc. Linn. Soc. N.S.W. 22(1): 38 (1897)

= Asteridiella solani var. kodaikanalensis Hosagoudar et al., in Nithyatharani et al., Scientific Transactions in Environment and Technovation 4(4): 165 (2011)

Facesoffunginumber: FoF00725

Epiphytes on the surface of leaves and stems. Superficial hyphae branched, septate, darker at the septa, brown, with hyphopodia, hyphal setae lacking. Hyphopodia 10–14 μm diam. ($\overline{x}=13\ \mu m$, n = 10), capitate, alternate or opposite on hyphae, near to hyphal septa, 2-celled, brown. Sexual morph: Ascomata 120–207 μm diam. ($\overline{x}=198\ \mu m$, n = 3), superficial on surface of host, mostly gregarious, rarely solitary on superficial hyphae, subglobose to globose, thick-walled, ascomatal setae and appendages absent. Peridium 35–46 μm thick, comprising dark brown cells of textura angularis when viewed in squash mounts, with two strata, outer stratum of dark brown cells of textura angularis, with raised conical cells 35–70 μm long, acute or rounded at the apex, and inner stratum of hyaline cells of textura porrecta. Hamathecium with evanescent paraphyses. Asci 45–50 × 21–26 μm ($\overline{x}=46 \times 23\ \mu m$, n = 3), 2–4-spored, unitunicate, oblong to cylindrical. Ascospores 37–42 × 15–19 μm ($\overline{x}=38 \times 18\ \mu m$, n = 3), 2–4-seriate, hyaline to brown, oblong, 4-septate, slightly constricted and darker at the septa, sometimes not constricted, smooth-walled. Asexual morph: Phialides rarely observed on hyphae, ampuliform, alternate or opposite, sometimes curved, pale brown to brown. Conidia not seen (Hansford 1961; Hosagoudar 2013).

Material examined

AUSTRALIA, on Solanum viride G. Forst. ex Spreng. (Solanaceae), 1896, V.H. Maiden (IMI 73840, glass slide from ex-holotype); PHILIPPINES, Benguet Province, Luzon, Mt. Santo Tomas, on leaf of Solanum inaequilaterale Merrill (Solanaceae), February 1925, Mary Strong Clemens 6410 (BPI 697764)

Notes: Part of the holotype specimen of this species is preserved as a glass slide at K (as IMI 73840), but we could not observe many characters (Fig. 11). However, a specimen (BPI 697764), and a redrawing of Asteridiella solani var. kodaikanalensis Hosag. et al. published by Hosagoudar (2013), and a redrawing from Hansford (1963) are provided here (Figs. 12, 13 and 14). Molecular data is lacking for A. solani.

Fig. 11

Asteridiella solani (holotype). a, b Herbarium packet and extype slide. c Ascomata viewed in squash mount. d Hyphae with capitate hyphopodia. e Immature ascomata viewed in squash mount. f–g Ascospores with 4 septa. Scale bars: c = 100 μm, e = 50 μm, f, g =20 μm, d = 5 μm

Fig. 12

Asteridiella solani (BPI 697764). a, b Herbarium packet and specimen. c, d Ascomata on host surface. e Section through ascomata. f, g Raised conical projections on ascomata. h Upper walled of ascoma viewed in squash mount. i Hyphae with capitate hyphopodia. j Hyphae with phialides. k Ascus when immature. l, m Ascospores with 4 septa. Scale bars: e = 100 μm, f–h = 50 μm, i, j =10 μm, k–m = 20 μm

Fig. 13

Asteridiella solani (redrawn from Hansford 1963). a Ascospores. b Capitate hyphopodia on hyphae. c Raised conical projections on ascomata

Fig. 14

Asteridiella solani var. kodaikanalensis (redrawn from Hosagoudar 2013). a Capitate hyphopodia on hyphae. b Phialides on hyphae. c Raised conical projections on ascoma. d Ascospore. Scale Bar: a = 8 μm

Cryptomeliola S. Hughes & Piroz., Mycol. Pap. 174: 14 (1997)

Facesoffunginumber: FoF00727

Epiphytes on the surface of leaves, twigs, or petioles. Superficial hyphae branched, brown or dark-brown, with or without hyphopodia, hyphal setae present. Hyphopodia capitate, alternate or opposite on hyphae, near to hyphal septa, 2-celled, brown. Hyphal setae arising from hyphae, dense, septate, dark brown, curved, rounded or hooked at the apex. Sexual morph: Ascomata superficial on surface of host, solitary or gregarious, sometimes hidden amongst setae, subglobose to globose, carbonaceous, verrucose, ascomatal setae and appendages absent. Peridium comprising dark brown cells of textura angularis when viewed in squash mounts, with two strata, outer stratum with dark brown to reddish cells of textura globulosa to angularis, and inner stratum with hyaline, flattened cells. Hamathecium with evanescent paraphyses. Asci 2–4-spored, unitunicate, ellipsoid to oval, evanescent. Ascospores 2-seriate, hyaline when immature and brown to dark brown at maturity, cylindrical to oblong, 4-septate, constricted and dark at septa, smooth-walled. Asexual morph: Phialides rarely seen, ampliform, interspersed between hyphal setae on hyphae (Mibey and Hawksworth 1997). Conidia unicellular, hyaline.

Notes: Cryptomeliola was raised to genus by Hughes & Pirozynski in Mibey and Hawksworth (1997). The genus contains three species (Index Fungorum 2015). There is no sequence data for any species in Cryptomeliola.

Type species: Cryptomeliola orbicularis (Berk. & M.A. Curtis) S. Hughes & Piroz., in Mibey & Hawksworth, Mycol. Pap. 174: 15 (1997)

= Meliola orbicularis Berk. & M.A. Curtis, in Berkeley, J. Linn. Soc., Bot. 10(no. 46): 392 (1868) [1869]

= Englerulaster orbicularis (Berk. & M.A. Curtis) Höhn., Sber. Akad. Wiss. Wien, Math.-naturw. Kl., Abt. 1 119: 454 [62 repr.] (1910)

= Meliolina orbicularis (Berk. & M.A. Curtis) F. Stevens, Annls mycol. 25(5/6): 417 (1927)

Facesoffunginumber: FoF00726

Epiphytes on the surface of host. Superficial hyphae branched, brown or dark brown, with or without hyphopodia, hyphal setae present. Hyphopodia not seen. Hyphal setae 92–320 long × 4–5 μm wide ($\overline{x}=265 \times 5$, n = 10), dense, septate, dark brown, curved and rounded, or hooked at the apex. Sexual morph: Ascomata 280–295 μm diam. ($\overline{x}=288\ \mu m$, n = 5), superficial on upper surface of host, solitary or gregarious, hidden amongst hyphae, subglobose to globose, carbonaceous, verrucose, ascomatal setae and appendages absent. Peridium 51–65 μm ($\overline{x}=56\mu m$, n = 5), comprising dark brown cells of textura angularis when viewed in squash mounts, with two strata, outer stratum with thick-walled, dark brown to reddish cells of textura globulosa to angularis, and inner stratum with hyaline, flattened cells. Hamathecium with evanescent paraphyses. Asci 60–71 × 37–49 μm ($\overline{x}=67\hbox{--} 43\mu m$, n = 10), 2 − 4-spored, unitunicate, ellipsoid to oval, evanescent. Ascospores 57–63 × 17–20 μm ($\overline{x}=58 \times 19\ \mu m$ , n = 10), 2-seriate, hyaline when immature and brown to dark brown at maturity, cylindrical to oblong, 4-septate, constricted and dark at the septa, rounded ends, smooth-walled. Asexual morph: Undetermined.

Notes: Cryptomeliola orbicularis, the type species of Cryptomeliola, was introduced based on Meliola orbicularis Berk. & Curtis (Mibey and Hawksworth 1997). Cryptomeliola orbicularis differs from Meliola species as the hyphae are immersed in the outer layers of host tissue or sometimes superficial, and hyphal setae are crowded, curly, and apically rounded. We re-examined the holotype specimen and could not find ascomatal setae, although the original description illustrated the species with ascomatal setae (Fig. 15). However, other species in the genus Cryptomeliola do not have an ascomatal setae, thus we illustrate Cryptomeliola orbicularis as lacking ascomatal setae base on the holotype specimen.

Material examined

On bark of indet. host, C. Wright 557 comm, Curtis; ex herb. Berkeley (IMI 193125, holotype).

Fig. 15

Cryptomeliola orbicularis (holotype). a, b Herbarium packet, specimen and drawing. c Colony on host surface. d Ascomata on substrate. e Section through ascoma. f Projecting cells on outside of peridium. g, i Setae arising on hyphae. h Upper wall of ascoma in squash mount. j Immature ascus. k, l Mature ascospores. Scale bars: e, f = 50 μm, h, k, l = 20 μm, g, i, j = 10 μm

Endomeliola S. Hughes & Piroz., N.Z. Jl Bot. 32(1): 53 (1994)

Facesoffunginumber: FoF00729

Epiphytes or pathogens on surface of leaves, developing on reddish-brown necrotic areas. Intercellular hyphae branched, septate, cylindrical, smooth, brown to pale brown, irregular, extending into the mesophyll, also penetrating between cells of the 2-layered palisade, with a single terminal hyphopodium. Hyphopodia irregular, mostly ellipsoidal to subglobose to ovoid to obovoid, occasionally angular or lobed in the mesophyll, generally with a short stalk cell, with hyaline pore at the centre of the head cell. Sexual morph: Stromata superficial on surface of host, solitary or gregarious, often discrete and then subglobose, somewhat flattened with a constricted base seated on the stromatic crust, black, comprising dark brown cells of textura angularis in transverse section, surface verrucose, with raised conical protuberant or ampuliform cells. Peridium comprising dark brown cells of textura angularis in transverse section, two strata, outer stratum of thick-walled, dark brown to reddish cells of textura angularis, and inner stratum of hyaline flattened cells. Ascomatal locules 1–4-locules in a pulvinate stroma, with central ostioles lined with periphyses. Hamathecium comprising cylindrical, hyaline, aseptate paraphyses. Asci 4-spored, unitunicate, ellipsoidal to clavate. Ascospores 2–3-seriate, hyaline when immature and brown to dark brown at maturity, broadly ellipsoidal to subcylindrical, 4-septate, slightly constricted and darker at the septa, the central cell sometimes longer than the others, smooth-walled. Asexual morph: Phialides solitary or gregarious on superficial ascostromata, occasionally on ascomatal walls, ampuliform or flask-shaped, brown to dark brown. Conidia ellipsoidal, hyaline (asexual morph redescribed from Hughes and Pirozynski 1994).

Notes: Endomeliola was introduced by Hughes and Pirozynski (1994), with type species E. dingleyae. The genus is distinguished by intercellular hyphae formed in the mesophyll layer of the plant. Endomeliola is a monotypic genus, lacking molecular data. However, the morphology is distinct from other genera in Meliolaceae (Fig. 16) and thus, we accept the genus in Meliolaceae.

Type species: Endomeliola dingleyae S. Hughes & Piroz., N.Z. Jl Bot. 32(1): 54 (1994)

Facesoffunginumber: FoF00728

Epiphytes, or pathogens on surface of leaves, developing on reddish brown necrotic areas. Intercellular hyphae 5–6 μm diam. ($\overline{x}=5\ \mu m$, n = 10), branched, septate, cylindrical, smooth, brown to pale brown, irregular, extending into the mesophyll, also penetrating between cells of the 2-layered palisade, with a single terminal hyphopodium. Hyphopodia 6–8 μm diam. ($\overline{x}=7\ \mu m$, n = 10), irregular, mostly ellipsoidal, subglobose, ovoid or obovoid, occasionally angular or lobed in the mesophyll, generally with a short stalk cell, with hyaline pore around the centre of the head cell. Sexual morph: Stromata 490–785 diam. × 255–575 μm high ($\overline{x}=552 \times 345\ \mu m$, n = 10), superficial on surface of host, solitary or gregarious, often discrete and then subglobose, somewhat flattened with a constricted base seated on the stromatic crust, black, surface verrucose, with raised conical protuberant or ampuliform cells. Peridium 45–63 μm ($\overline{x}=48\ \mu m$, n = 5), comprising dark brown cells of textura angularis in transverse section, with two strata, outer stratum of thick-walled, dark brown to reddish cells of textura angularis, and inner stratum of hyaline flattened cells. Ascomatal locules 1–4-locules in a pulvinate stroma, with central ostioles lined with periphyses. Hamathecium comprising cylindrical, hyaline, aseptate paraphyses. Asci 74–94 × 38–44 μm ($\overline{x}=92 \times 41\ \mu m$, n = 10), 4-spored, unitunicate, ellipsoidal to clavate. Ascospores 65–67 × 29–32 μm ($\overline{x}=66 \times 30\ \mu m$, n = 10), 2–3-seriate, hyaline when immature and brown to dark brown at maturity, broadly ellipsoidal to subcylindrical, 4-septate, slightly constricted and darker at the septa, the central cell sometimes longer than the others, smooth-walled. Asexual morph: Undetermined.

Notes: Molecular data indicate that Endomeliola dingleyae is a member of family Meliolaceae. The species is closely related to Asteridiella intidae with high bootstrap support (Justavino et al. 2015), but differs considerably in morphology and it is possible that the data is incorrect. Therefore, further gene and sequence data are needed to clarity the placement of the genus and species.

Material examined

NEW ZEALAND, Auckland Province, Thames County, Kauaeranga Valley, on living leaves of Coprosma robusta Raoul. (Rubiaceae), 16 November 1965, J.M. Dingley (PDD 24806, isotype).

Fig. 16

Endomeliola dingleyae (isotype). a − c Herbarium packet and specimen. d–e Ascostromata on substrate. f Section through ascostromata. g Peridium between locules. h Periphyses. i Wall cells. j Paraphyses. k Phialides on surface of ascomata. l, m Intercellular hyphae. n Hyphopodia. o, p Asci. q Immature ascospore. r, s Mature ascospores. Scale bars: e, f = 50 μm, h, l–s = 20 μm, g, i, j = 10 μm

Irenopsis F. Stevens, Annls mycol. 25(5/6): 411 (1927)

≡ Meliola subgen. Irenopsis (F. Stevens) Cif., Annls mycol. 36(2/3): 203 (1938)

Facesoffunginumber: FoF00734

Epiphytes on the surface of leaves and stems. Superficial hyphae branched, septate, darker at the septa, brown, with hyphopodia, hyphal setae lacking. Hyphopodia capitate, angular or lobate, alternate or opposite on hyphae, near to hyphal septa, 2-celled, brown. Sexual morph: Ascomata superficial on the surface of hosts, solitary or gregarious on superficial hyphae, globose to subglobose, thick-walled, with raised long setae on the surface. Ascomatal setae curved or hooked at the apex, developing on the ascomata. Peridium comprising dark brown cells of textura angularis when viewed in squash mounts, with two strata, outer stratum thick-walled, dark brown cells of irregular textura angularis, and inner stratum of flattened, hyaline cells. Hamathecium with evanescent paraphyses. Asci 2–4-spored, unitunicate, obovoid to ovoid, evanescent. Ascospores 2-seriate, hyaline to brown, oblong to ovoid, 4-septate, constricted and darker at the septa, rounded ends, smooth-walled. Asexual morph: Phialides ampuliform, mixed with capitate hyphopodia, opposite or alternate, pale brown to brown. Conidia rarely observed, hyaline.

Notes: The genus Irenopsis was established by Stevens (1927). This genus is distinguished by having true setae on ascomata and lacking hyphal setae. Irenopsis comprises 180 species epithets in Index Fungorum (2015). Species were mainly introduced based on host association. Sequence data for I. vincensii places Irenopsis in Meliolaceae (Justavino et al. 2015).

Type species: Irenopsis tortuosa (G. Winter) F. Stevens, Annls mycol. 25(5/6): 439 (1927)

≡ Meliola tortuosa G. Winter, in Gaillard, Le Genre Meliola: 67 (1892)

≡ Irenopsis tortuosa (G. Winter) F. Stevens, Annls mycol. 25(5/6): 439 (1927) var. tortuosa

Possible synonym (from Index Fungorum 2015)

= Meliola tonkinensis var. potomorphes Cif., Annls mycol. 31(3): 149 (1933)

Facesoffunginumber: FoF00732

Epiphytes on the surface of leaves and stems. Superficial hyphae branched, septate, darker at the septa, brown, with hyphopodia, hyphal setae lacking. Hyphopodia capitate, curved, alternate or opposite on hyphae, near to hyphal septa, 2-celled, brown. Sexual morph: Ascomata 175–215 μm diam. × 95–120 μm high ($\overline{x}=208 \times 113\ \mu m$, n = 5), superficial on surface of hosts, solitary or gregarious on superficial hyphae, globose to subglobose, thick-walled, with raised long setae on the surface. Ascomatal setae 8–10 × 92–107 μm ($\overline{x}=9 \times 102\ \mu m$, n = 5), curved or hooked at the apices, developing on the ascomata. Peridium 20–26 μm ($\overline{x}=24\ \mu m$, n = 5), comprising dark brown cells of textura angularis when viewed in squash mounts, with two strata, outer stratum thick-walled, dark brown cells of irregular textura angularis, and inner stratum of flattened, hyaline cells. Hamathecium with evanescent paraphyses. Asci 54–58 × 22–25 μm ($\overline{x}=55 \times 23\ \mu m$, n = 5), 2–4-spored, unitunicate, obovoid to ovoid, evanescent. Ascospores 37–40 × 14–15 μm ($\overline{x}=37 \times 15\ \mu m$, n = 10), 2-seriate, hyaline to brown, oblong to ovoid, 4-septate, or with 4 transverse septa and 2 longitudinal septa, constricted and darker at the septa, rounded ends, smooth-walled. Asexual morph: Phialides 16–17 × 7–8 μm ($\overline{x}=16 \times 7\ \mu m$, n = 10), ampliform, mixed with capitate hyphopodia, opposite or alternate, pale brown to brown. Conidia not seen.

Material examined

DOMINICAN REPUBLIC, on leaves of Potomorpha umbellata Mich. (Piperaceae), August 1929, R. Ciferri 2426 (SF5890, holotype); VENEZUELA, El Limon bei Puerto La Cruz, on leaves of Piperis marginati Jacq. (Piperaceae), 16 January 1928, H. Sydow (SF77640).

Notes: The type specimen was collected in Brazil (Stevens 1927), but we could not locate and examine the type. Therefore a collection of Meliola tonkinensis var. potomorphes from S was studied (Figs. 17 and 18). One ascospore was found with 4 transverse septa and 2 longitudinal septa in Irenopsis tortuosa (S-F5890).

Fig. 17

Irenopsis tortuosa (holotype). a–c Herbarium packet and specimen. d Ascomata on substrate. e, f Ascomata when viewed in squash mounts. g Section through ascoma. h Upper wall of ascomata. i Hyphae with phialides. j Hyphae with capitate hyphopodia. k Seta on ascoma. l Hamathecium. m, n Immature asci. o, p Ascospores with 4 septa. q Ascospore with 4 transverse septa and 2 longitudinal septa. Scale bars: e = 200 μm, f, g = 100 μm, h, m–q = 20 μm, i, j = 5 μm, k, l = 10 μm

Fig. 18

Irenopsis tortuosa (S- F77640). a, b Herbarium packet and specimen. c Ascomata on host surface. d Ascoma when viewed in squash mounts. e Section through ascoma. f Peridium. g Upper wall of ascoma. h Phialides on hyphae. i Hyphae with capitate hyphopodia. j Young ascoma. k Seta on ascoma. l, m Immature asci. n–p Ascospores with 4 septa. Scale bars: d, e = 50 μm, f, g, j–p = 20 μm, h, i = 10 μm

Irenopsis walsurae X.Z. Zeng & K.D. Hyde, sp. nov.

Facesoffunginumber: FoF00733

Index Fungorum: IF551220

Etymology:— walsurae referring to the host on which the taxon was found.

Holotype: MFLU13-0621

Epiphytes on the surface of living leaves. Superficial hyphae 7 μm diam., radiating outwardly, branched, septate, darker at the septa, brown, with hyphopodia, hyphal setae absent. Hyphopodia 14–19 × 9–13 μm ($\overline{x}=16 \times 11\ \mu m$, n = 20), capitate, alternate on hyphae, near to hyphal septa, 2-celled, brown. Sexual morph: Ascomata up to 160 μm, superficial on surface of hosts, scattered, globose to subglobose, thick-walled, with long ascomatal setae. Ascomatal setae up to 170 μm long, raised on ascomata, and rounded at the apex. Peridium comprising dark brown cells of textura angularis when viewed in squash mounts, with two strata, outer stratum a single layer of large, thick-walled, dark brown cells of irregular textura angularis, and inner stratum of flattened, hyaline cells. Hamathecium with evanescent paraphyses. Asci 2–3-spored, unitunicate, obovoid to ovoid, with short pedicel or apedicel, evanescent. Ascospores 33–39 × 12–18 μm ($\overline{x}=36 \times 14\ \mu m$, n = 20), 2–3-seriate, hyaline to brown, oblong to ovoid, 3–4-septate, slightly constricted and darker at the septa, rounded ends, apical cell sometimes slightly longer, smooth-walled. Asexual morph: Phialides 16–21 × 8–10 μm ($\overline{x}=18 \times 9\ \mu m$, n = 10), ampuliform, alternate to opposite, formed on separate hyphae, rarely mixed with capitate hyphopodia. Conidia undetermined.

Material examined

THAILAND, Chiang Mai, Mae Taeng, Pa Pae, Bahn Pa Dheng, 128 Moo 3, Mushroom Research Centre, on the living leaves of Walsura tubulata Hiern. (Meliaceae), 22 November 2013, Xiangyu Zeng (MFLU 13-0621, holotype; isotype, KUN).

Notes: Irenopsis walsurae was found on living leaves of Walsura tubulata (Meliaceae). Irenopsis species known from this host family are I. trichiliae Hosag. & Riju, I. chukrasiae Hosag., I. dysoxyli Jana et al. and I. indica (Anahosur) Hosag. Irenopsis walsurae is most similar to I. dysoxyli, but differs in having longer ascospores and ascomatal setae, with smaller ascomata in Irenopsis walsurae. The new species is also similar to I. trichiliae, but differs in having opposite to unilateral phialides, separated from the hyphopodia in Irenopsis walsurae, while alternate to unilateral phialides, mixed with hyphopodia in I. trichiliae. There are no previous records of Irenopsis species reported on Walsura. Other genera in Meliolaceae known from Walsura are Ectendomeliola walsurae Hosag. & D.K. Agarwal, Meliola walsurae Hansf. and M. walsuricola Bagool & H. Biju, however, ascomatal setae are absent in these three species, while ascomatal setae are present in Irenopsis walsurae. Ectendomeliola is excluded from Meliolaceae in this study (Fig. 19).

Fig. 19

Irenopsis walsuri (holotype). a Leaf specimen. b Colony on surface of leaves. c Ascoma on host substrate. d Hyphae with capitate hyphopodia. e Phialides on hyphae. f Section through ascoma. g Long setae forming on ascoma. h Immature ascus. i–j Mature asci. k–m Ascospores with 4 septa. Scale bars: b = 500 μm, c = 100 μm, f = 50 μm, d, e, g = 20 μm, h–m = 10 μm

Meliola Fr., Syst. orb. veg. (Lundae) 1: 111 (1825)

Possible synonyms (from Index Fungorum 2015):

Myxothecium Kunze, Syst. mycol. (Lundae) 3(1): 231 (1829)

Asterina subgen. Asteridium Sacc., Syll. fung. (Abellini) 1: 49 (1882)

Asteridium (Sacc.) Speg. ex Sacc., Syll. fung. (Abellini) 9: 435 (1891)

Facesoffunginumber: FoF00740

Epiphytes on the surface of leaves, stems or branches. Superficial hyphae branched, septate, darker at the septa, brown, with hyphopodia, hyphal setae present. Hyphopodia capitate, alternate or opposite on hyphae, near to hyphal septa, 2-celled, brown. Hyphal setae arising from hyphae, forming around the base of the ascomata in some species, straight or curly, rounded or acute at the apex, or branches, septate or aseptate, brown to dark brown, smooth-walled. Sexual morph: Ascomata superficial on the surface of hosts, solitary to gregarious on superficial hyphae, globose to subglobose, thick-walled, ascomatal setae and appendages absent, surface of ascomata verrucose. Peridium comprising dark brown cells of textura angularis when viewed in squash mounts, with two strata, outer stratum thick-walled, dark brown cells of irregular textura angularis, and inner stratum of flattened, hyaline cells. Hamathecium with evanescent paraphyses. Asci 2–4-spored, unitunicate, broadly clavate to oblong, evanescent. Ascospores 2–4-seriate, hyaline to brown, oblong to broadly cylindrical, 3–4-septate, constricted and darker at the septa, rounded ends, smooth-walled, verrucose when immature. Asexual morph: Phialides ampuliform, alternate or opposite on hyphae, sometimes curved, pale brown to brown. Conidia hyaline.

Notes: The genus Meliola was introduced by Fries (1825), and is the largest genus in the family Meliolaceae. Meliola contains over 1200 species (Kirk et al. 2008). Most species were introduced based on host association. Molecular analysis of Meliola members were provided by Gregory and John (1999), Justavino et al. (2015) and Pinho et al. (2012a, 2014) and show their placement in Meliolaceae.

Type species: Meliola nidulans (Schwein.) Cooke, Grevillea 11(no. 57): 37 (1882)

≡ Sphaeria nidulans Schwein., Schr. naturf. Ges. Leipzig 1: 45 (1822)

≡ Chaetosphaeria nidulans (Schwein.) Rehm, Ascomyceten, fasc.: no. 287 (1875)

≡ Meliola nidulans (Schwein.) Cooke, Grevillea 11(no. 57): 37 (1882) var. nidulans

Facesoffunginumber: FoF00738

Epiphytes on the surface of leaves, stems or branches. Superficial hyphae branched, septate, darker at the septa, brown, with hyphopodia, hyphal setae present. Hyphopodia not seen. Hyphal setae up to 350 μm, forming around the base of the ascomata in some species, straight or curly, rounded or acute at the apex or branches, septate or aseptate, brown to dark brown, smooth-walled. Sexual morph: Ascomata 235–345 μm diam. ($\overline{x}=312\ \mu m$, n = 5), superficial on the surface of hosts, solitary to gregarious on superficial hyphae, globose to subglobose, thick-walled, ascomatal setae and appendages absent, surface of ascomata verrucose. Peridium 53–62 μm diam. ($\overline{x}=56\ \mu m$, n = 5), comprising dark brown cells of textura angularis when viewed in squash mounts, with two strata, outer stratum a single layer or multi-layered when immature, of large, thick-walled, dark brown cells of irregular textura angularis, and inner stratum of flattened, hyaline cells. Hamathecium with evanescent paraphyses. Asci 58–63 × 32–36 μm ($\overline{x}=60 \times 31\ \mu m$, n = 5), 2–4-spored, unitunicate, broadly clavate to oblong, evanescent. Ascospores 45–54 × 15–19 μm ($\overline{x}=51 \times 18\ \mu m$, n = 5), 2–4-seriate, hyaline to brown, oblong to broadly cylindrical, 3–4-septate, constricted and darker at the septa, rounded ends, smooth-walled, verrucose when immature. Asexual morph: Undetermined.

Notes: The species Meliola nidulans was established by Cooke (1882), and was based on Sphaeria nidulans Schwein. We examined the type specimen of M. nidulans and could not obtain good photographs of the superficial hyphae, hyphopodia, and phialides because the specimen is not in good condition (Fig. 20).

Material examined

USA, Pennsylvania, New Garden, on branches of Cornus sp. (Cornaceae), Michener Collection 711 (BPI 800359, holotype of Sphaeria nidulans).

Fig. 20

Meliola nidulans (holotype) a, b Ascomata on host substrate c Long setae developed from hyphae. d Section through ascoma. e Peridium. f Paraphyses. g, h Immature asci in cotton blue reagent. i Immature ascus in Melzer’s reagent. j Mature ascospore with 4 septa and rounded ends. k Immature ascospores in Melzer’s reagent with verrucose surface. Scale bars: a = 1000 μm, b = 200 μm, e = 50 μm, c, f–k = 20 μm, d = 100 μm

Meliola clerodendricola Henn., Hedwigia 37: 288 (1898)

Facesoffunginumber: FoF00736

Epiphytes on surface of living leaves. Superficial hyphae 7 μm diam. radiating outwardly, straight to substraight, branched, septate, darker at the septa, brown, with hyphopodia, hyphal setae present. Hyphopodia 13–17 × 9–13 μm ($\overline{x}=15 \times 11\ \mu m$, n = 20), capitate, unilateral on hyphae, near to hyphal septa, 2-celled, brown. Hyphal setae long, straight, acute at the apex, septate, brown, and smooth-walled. Sexual morph: Ascomata up to 160 μm diam., superficial on surface of host, dense, globose to subglobose, thick-walled, ascomatal setae and appendages absent. Peridium comprising dark brown cells of textura angularis when viewed in squash mounts, with two strata, outer stratum a single layer of large, thick-walled, dark brown cells of irregular textura angularis, and inner stratum of flattened, hyaline cells. Hamathecium with evanescent paraphyses. Asci 2–4-spored, unitunicate, obovoid to ovoid, with short pedicel, evanescent. Ascospores 31–35 × 10–13 μm, ($\overline{x}=33 \times 12\ \mu m$, n = 20), 2–4-seriate, hyaline to light brown, oblong to ellipsoid, 3–4-septate, constricted and darker at the septa, rounded ends, apical cell sometimes slightly larger, smooth-walled. Asexual morph: Undetermined.

Material examined

THAILAND, Chiang Mai, Mushroom Research Centre, Mae Taeng, Pa Pae, Bahn Pa Dheng, 128 Moo 3, on the living leaves of Clerodendrum sp. (Lamiaceae), 22 November 2013, Xiangyu Zeng (MFLU 13-0620; KIB, reference specimen designated here).

Notes: Meliola clerodendricola (MFLU 13-0620) was found on living leaves of Clerodendrum sp. (Lamiaceae). The morphology of MFLU13-0620 is most typical of Meliola clerodendricola based on host association and size of hyphal setae, ascomata, and ascospores. We were unable to locate the holotype material of Meliola clerodendricola, but an isotype is preserved in S (F8150) (Fig. 21). However, DNA cannot be extracted from the type specimen, consequently, we designate this as a reference specimen (Ariyawansa et al. 2014). Meliolaceae species known from Clerodendrum sp. are Asteridiella clerodendricola Hosag. and A. vivekananthanii Hosag., however, hyphal setae are absent in these species, but are found in M. clerodendricola. In addition, Asteridiella clerodendricola can produce a pathogenic effect on the host, while there is no pathogenic effect with Meliola clerodendricola. Molecular analyses place M. clerodendricola in the Meliolaceae clade, with a putatively named specimen of Endomeliola dingleyae (97 % ML support and 1.0 PP support). This result is strange as Meliola clerodendricola has superficial hyphae and hyphal setae, while Endomeliola dingleyae has intercellular hyphae without hyphal setae. Meliola clerodendricola was found on a host in the order Lamiales and Endomeliola dingleyae was found on host in the order Gentianales; their hosts belong in closely related orders in the euasterids (Jansen et al. 2007; Justavino et al. 2015). The identification of Endomeliola dingleyae may be incorrect and although the sequence has been placed in GenBank it does not appear to be linked to a voucher specimen.

Fig. 21

Meliola clerodendricola (MFLU13-0620). a Leaf specimen. b Colony on surface of leaves. c Ascomata on host substrate. d, e Section through ascomata. f Hyphae with capitate hyphopodia. g, h Immature asci. i, j Ascospores with 4 septa. Scale bars: b, c = 100 μm, d, e = 50 μm, f–l = 10 μm

Meliola thailandicum Hongsanan & K.D. Hyde, sp. nov.

Facesoffunginumber: FoF00739

Index Fungorum: IF551218

Etymology:—from Latin thailandicum meaning Thailand, referring to the location where the fungus was found.

Holotype: MFLU15-0044

Epiphytes on the surface of living leaves. Superficial hyphae 6 μm diam., radiating outwardly, branched, septate, darker at the septa, brown to dark brown, with hyphopodia. Hyphal setae up to 300 μm long, septate, darker at the septa, 1–3 forked at the apex. Hyphopodia 7 μm diam., capitate, mostly alternate or sometimes opposite on hyphae, near to hyphal septum, 2-celled, brown. Sexual morph: Ascomata 185–200 μm diam. × 120–180 μm high ($\overline{x}=190 \times 134\ \mu m$, n = 10), superficial on surface of host, mostly scattered or solitary, globose to subglobose, thick-walled, ascomatal setae and appendages absent. Peridium 18–20 μm ($\overline{x}=20\ \mu m$, n = 10), comprising dark brown cells of textura angularis when viewed in squash mounts, with two strata, outer stratum a single layer of large, thick-walled, dark brown cells of irregular textura angularis, and inner stratum of flattened, hyaline cells. Hamathecium with evanescent paraphyses. Asci 59–60 × 26–28 μm ($\overline{x}=60 \times 27\ \mu m$, n = 10), 2–4-spored, unitunicate, obovoid to ovoid, or broadly clavate, with short pedicel, sessile at maturity, evanescent. Ascospores 39–41 × 14–16 μm ($\overline{x}=40 \times 15\ \mu m$, n = 20), 2–4-seriate, hyaline to brown, oblong to cylindrical, 4-septate, constricted and darker at the septa, rounded ends, with thin sheath when immature, smooth-walled. Asexual morph: Phialides 7–8 μm ($\overline{x}=7\ \mu m$, n = 10), ampuliform, alternate to opposite, formed on separate hyphae, mixed with capitate hyphopodia. Conidia undetermined.

Material examined

THAILAND, Chiang Rai, Amphoe Thoeng, on the living leaves of Dimocarpus longan Lour. (Sapindaceae), 18 January 2015, S. Hongsanan (MFLU15-0044 holotype; KIB, isotype); Chiang Rai, Mueang, Rai Chun Tawan Meditation Centre, on the living leaves of Dimocarpus longan (Sapindaceae), 3 January 2015, S. Hongsanan (MFLU15-0047); Chiang Rai, Mueang, Tasud, on the living leaves of Dimocarpus longan (Sapindaceae), 11 February 2015, S. Hongsanan (MFLU15-0377); Chiang Rai, Mueang, Agricultural research center, on living leaves of Acacia auriculiformis A.Cunn. ex Benth. (Fabaceae), 23 January 2015, S. Hongsanan (MFLU15-0378); Chiang Rai, Mueang, Mae Fah Luang University, on the living leaves of Dimocarpus longan (Sapindaceae), 12 February 2015, S. Hongsanan (MFLU15-0379).

Notes: Meliola thailandicum is common on living leaves of Dimocarpus longan in northern Thailand. Meliola capensis (Kalchbr. & Cooke) Theiss. and M. dimocapi Hosag. & T.K. Abraham are also known from Dimocarpus longan, however, they differ in having apically acute hyphal setae, and ascospores without a thin-sheath, while M. thailandicum has longer hyphal setae with 1 to many branches at the apex, and ascospores with thin-sheath when immature and that may be present at maturity (Figs. 22 and 23). Meliola thailandicum was also collected on Acacia auriculiformis. Meliola species known from this host genus are M. acaciarum Speg., M. acaciae-binervatae Hansf., M. acaciae-confusae Sawada, M. acaciicola Hansf. and M. acaciorum Speg., however, all species differ from M. thailandicum in having shorter hyphal setae, acute or rounded at the apex, and ascospores lacking a sheath. In M. thailandicum hyphal setae are longer and branched at the apex, and ascospores have thin sheaths when immature and sometimes when mature. Molecular analyses indicate that M. thailandicum from D. longan and A. auriculiformis are the same species which belong in Meliolaceae within the genus Meliola (100 % ML support and 1.0 PP support). Hence, we conclude that the species is not host-specific. This may have important implications on the numbers of Meliolaceae species, as most taxa were introduced based on host occurrence.

Fig. 22

Meliola thailandicum (holotype). a Leaf specimens. b Ascomata on host substrate. c Section through ascoma. d Hyphal seta. e Hyphal setae with forked apices. f Peridium. g Hyphae with capitate hyphopodia. h Terminal hyphae. i Upper wall of ascoma. j Phialides. k Immature ascospore in Melzer’s reagent. l, m Mature ascospores. n, o, q Immature asci. p Immature ascus in Melzer’s reagent. Scale bars: c, d = 100 μm, e, f = 20 μm, g–j = 10 μm, k–q = 20 μm

Fig. 23

Meliola thailandicum (from Acacia auriculiformis, MFLU 15-0378). a Leaf specimens. b Ascoma on host substrate. c Section through ascoma. d Hyphal seta. e Young ascoma. f Phialides on hyphae. g Capitate hyphopodia on hyphae. h Germinated ascospore. i, j Hyphal setae with branches at apex. k Mature ascospore with 4 septa. Scale bars: c = 100 μm d = 200 μm e = 50 μm, f–k = 20 μm

Meliola mucunicola Hongsanan & K.D. Hyde, sp. nov.

Facesoffunginumber: FoF00737

Index Fungorum: IF551219

Etymology:— mucunicola referring to the host on which the taxon was found.

Holotype: MFLU15-0386

Epiphytes on the surface of Mucuna living leaves. Superficial hyphae 5–8 μm ($\overline{x}=6\ \mu m$, n = 10), radiating outwardly, branched, septate, darker at the septa, brown to dark brown, with hyphopodia. Hyphal setae 167–195 × 8–10 μm ($\overline{x}=185 \times 9\ \mu m$, n = 10), rounded to acute at the apex. Hyphopodia 10–13 μm ($\overline{x}=12\ \mu m$, n = 10), capitate, mostly alternate or sometimes opposite on hyphae, near to hyphal septa, 2-celled, brown. Sexual morph: Ascomata 120–145 μm diam. × 90–105 μm high ($\overline{x}=122 \times 96\ \mu m$, n = 10), superficial on surface of hosts, mostly scattered or solitary, globose to subglobose, thick-walled, ascomatal hyphae or appendages lacking. Peridium 15–19 μm ($\overline{x}=17\ \mu m$, n = 10), comprising dark brown cells of textura angularis when viewed in squash mounts, with two strata, outer stratum a single layer of large, thick-walled, dark brown cells of irregular textura angularis, and inner stratum of flattened, hyaline cells. Hamathecium with evanescent paraphyses. Asci 44–53 × 22–23 μm ($\overline{x}=48 \times 23\ \mu m$, n = 10), 2-spored, unitunicate, oblong to cylindrical, with short pedicel, sessile at maturity, evanescent. Ascospores 37–41 × 12–15 μm ($\overline{x}=39 \times 13\ \mu m$, n = 20), 2-seriate, hyaline to brown, oblong to cylindrical, 4-septate, constricted and darker at the septa, rounded ends, with thin sheath when immature, smooth-walled. Asexual morph: Phialides 7–9 μm ($\overline{x}=8\ \mu m$, n = 10), ampuliform, alternate to opposite on hyphae, mixed with capitate hyphopodia. Conidia undetermined.

Material examined

THAILAND, Chiang Rai, Mueang, Agricultural Research Center, on the living leaves of Mucuna pruriens (L.) DC. (Fabaceae), 23 March 2015, S. Hongsanan (MFLU15-0386, holotype; KIB, isotype).

Notes: Meliola mucunicola was found on living leaves of Mucuna pruriens. Meliola species known from this host genus are M. mucunae Hansf. and M. mucunae-acuminatae Hansf. The new species is most typical of M. mucunae-acuminatae in terms of morphology and host association, as M. mucunae-acuminatae can be found on the same host species. Meliola mucunicola differs from M. mucunae-acuminatae in having ascomata surrounded by many short setae (Fig. 24), while in M. mucunae-acuminatae ascomata are surrounded by a few long setae. Moreover, phialides of Meliola mucunicola are larger than those of M. mucunae-acuminatae. Molecular analyses place Meliola mucunicola in the Meliolaceae clade close to Meliola centellae Pinho & O.L. Pereira with 55 % ML support and 0.8 PP support.

Fig. 24

Meliola mucunicola (holotype). a Leaf specimen. b Ascomata and hyphal setae on host substrate. c Section through ascoma. d Hyphal setae. e Peridium. f Phialides on hyphae. g Hyphae with capitate hyphopodia. h Rounded apex of hyphal seta. i Upper wall of young ascoma. j Immature ascus. k Immature ascus in Melzer’s reagent. l Upper wall of ascoma when viewed in squash mounts. m, n Mature ascospores with 4 septa. Scale bars: c, d = 50 μm, e, i–k, m, n = 20 μm, f–h, l = 10 μm

Genera / species synonymized under Meliola

Laeviomeliola Bat., Atas Inst. Micol. Univ. Recife 1: 224 (1960)

≡ Laeviomeliola psidii Bat., Atas Inst. Micol. Univ. Recife 1: 225 (1960)

Meliola psidicola Hongsanan & K.D. Hyde, nom. nov.

Facesoffunginumber: FoF00847

Index Fungorum: IF551222

Epiphytes on the surface of leaves. Superficial hyphae 6–7 μm diam. ($\overline{x}=6\ \mu m$, n = 10), branched, septate, darker at the septa, brown, with hyphopodia. Hyphal setae 260–284 × 5–7 μm ($\overline{x}=277 \times 7\ \mu m$, n = 10), rounded to acute at the apex. Hyphopodia 8–10 μm diam. ($\overline{x}=9\ \mu m$, n = 10), capitate, cylindrical to subcylindrical, alternate on hyphae, near to hyphal septa, brown, 2-celled. Sexual morph: Ascomata 158–182 μm diam. ($\overline{x}=163$, n = 5), superficial on upper surface of leaves, solitary or gregarious on superficial hyphae, globose to subglobose, thick-walled, lacking ascomata setae and appendages. Peridium 22–25 μm ($\overline{x}=23\ \mu m$, n = 5), comprising dark brown cells of textura angularis when viewed in squash mounts, with two strata, outer stratum a single layer of large, thick-walled, dark brown cells of irregular textura angularis, and inner stratum of flattened, hyaline cells. Hamathecium with evanescent paraphyses (Batista and da Silva 1960). Asci 37–43 × 16–22 μm ($\overline{x}=41 \times 19\ \mu m$, n = 3), 2-spored, unitunicate, clavate to obovoid, evanescent. Ascospores 44–52 × 15–16 μm ($\overline{x}=46 \times 15\ \mu m$, n = 10), 2-seriate, hyaline when immature, brown at maturity, ellipsoid to fusiform, 4-septate, not constricted when immature, slightly constricted and darker at the septa at maturity, acute ends, smooth-walled. Asexual morph: Undetermined.

Material examined

BRAZIL, Pernambuco, Caruaru, on leaves of Psidium guajavae Linnaeus (Myrtaceae), 9 March 1958, Dr. Epaminondas de Barros Correia (URM 12749, holotype).

Notes: The genus Laeviomeliola is monotypic and was introduced by Batista and da Silva (1960), with the type species L. psidii. We re-examined the type specimen and concluded that the morphology of Laeviomeliola is quite similar to Meliola based on its hyphal setae. Laeviomeliola psidii differs in having fusiform ascospores with tapering ends, while in Meliola the ends are rounded (Fig. 25). Thus, Laeviomeliola should be synonymized under Meliola. As Meliola psidii is already in use, we provide a new name Meliola psidiicola. Fresh collection and molecular data are needed to confirm the position of the genus.

Fig. 25

Meliola psidiicola (holotype). a Ascoma on substrate. b Section through ascoma. c Section through peridium. d Hyphal seta. e Wall of ascoma in a squash mount. f Hyphae with capitate hyphopodia. g Hyphal setae on superficial hyphae. h Ascus when immature. i–l Ascospores with 4 septa. Scale bars: b, d, e, g–l = 20 μm, c, f = 10 μm

Doubtful genera within Meliolales

Ceratospermopsis Bat., Mycopath. Mycol. appl. 5: 165 (1951)

Notes: Ceratospermopsis was established by Batista (1951), and comprises two species, C. cupaniae Bat. and C. xylopiae Bat. The type species was not mentioned (Batista 1951; Index Fungorum 2015). We were unable to examine either species which are kept in URM, Brazil despite several requests. Ceratospermopsis is characterized by hyphal setae, lack of hyphopodia on the hyphae, globose to subglobose, membranous ascomata, 8-spored, subglobose asci, and muriform, slightly olivaceous, cylindrical to fusoid ascospores, with hyaline appendages (Batista 1951). We suggest that C. xylopiae should be the type species of the genus Ceratospermopsis based on its morphology. There is no sequence data to clarify the placement of the genus, which is poorly known and appears to lack herbarium material, thus we treat it as doubtful.

Type species: Ceratospermopsis xylopiae Bat., Mycopath. Mycol. appl. 5: 166 (1951)

Ectendomeliola Hosag. & D.K. Agarwal, Indian Phytopath. 59(1): 99 (2006)

≡ Ectendomeliola walsurae Hosag. & D.K. Agarwal, Indian Phytopath. 59(1): 99 (2006)

Notes: Ectendomeliola was established by Hosagoudar and Agarwal (2006) with the type species E. walsurae. This genus is characterized by 2-celled, ovate to subcylindrical hyphopodia on the hyphae, straight to curly hyphal setae, acute or rounded at the apex, and oblong to cylindrical ascospores, with 4 septa, and constricted at the septa (Figs. 26 and 27). Ectendomeliola is most typical of Meliola based on its hyphal setae, but differs in having hyphopodia immersed in host epidermal cells. This genus is also similar to Endomeliola, however the hyphopodia form in the epidermal layer, while hyphopodia extend into the mesophyll layer in Endomeliola. There are two species placed in the genus, E. walsurae from Walsura trifolia (A. Juss. ) Harms. and E. otonephelii from Otonephelium stipulaceum (Bedd.) Radlk. (the latter species has longer hyphal setae than E. walsurae).

Fig. 26

Ectendomeliola walsurae (holotype, redrawn from Hosagoudar and Agarwal 2006). a Ascoma on the surface of host (notes: not sectioned and only showing ascoma shape and asci arrangement). b Septate hyphae. c Capitate hyphopodia on hyphae. d Phialides on hyphae. e Apex of the hyphal setae. f Ascospores with 4 septa

Fig. 27

Ectendomeliola otonephelii (holotype, redrawn from Hosagoudar and Archana 2013). a Capitate hyphopodia on hyphae. b Phialides on hyphae. c Ascospores with 4 septa. d Hyphopodia immersed in host epidermal cells. e Apex of the hyphal setae

Haraea Sacc. & P. Syd., in Saccardo, Annls mycol. 11(3): 312 (1913)

Facesoffunginumber: FoF00731

Epiphytes on the surface of leaves, stems, and straw. Superficial hyphae branched, septate, aggregated, hyphopodia and hyphal setae lacking. Sexual morph: Ascomata superficial on surface of hosts, gregarious or solitary, globose to subglobose, flattened when immature, membranous to subcarbonaceous, brown to dark brown, and with aggregated hyphae at the margin, ascomatal setae and appendages absent. Peridium comprising dark brown of radially arranged cells when immature, with two strata, outer stratum of dark brown cells of textura angularis, and inner stratum of hyaline, flattened cells. Hamathecium with paraphyses. Asci 4–8-spored, rounded at the apex, with thick pedicel (Saccardo 1913). Ascospores 2–4-seriate, pale brown to brown, or reddish, oblong to fusoid, 3-septate, slightly constricted at the septa, smooth-walled. Asexual morph: Undetermined.

Notes: The genus Haraea was established by Saccardo (1913) with type species H. japonica Sacc. & P. Syd. It is a good genus but its placement is doubtful because hyphopodia are lacking, and asci were not illustrated. Hence, Haraea should be place as a doubtful genus until molecular data are available to verify the placement of this genus. Five species are listed in Index Fungorum (2015).

Type species: Haraea japonica Sacc. & P. Syd., Annls mycol. 11(3): 312 (1913)

Faces of fungi number: FoF00730

Epiphytes on the surface of leaves, stems, and straw. Superficial hyphae branched, septate, aggregate, hyphopodia and hyphal setae lacking. Sexual morph: Ascomata 100–140 μm diam. × 85–100 μm high ($\overline{x}=135 \times 95\ \mu m$, n = 5), superficial on surface of hosts, gregarious or solitary, globose to subglobose, flattened when immature, membranous to subcarbonaceous, brown to dark brown, and aggregated hyphae at the margin, ascomatal setae and appendages absent. Peridium 25–34 μm diam. ($\overline{x}=31\ \mu m$, n = 5), comprising dark brown radially arranged cells when immature, with two strata, outer stratum of thick-walled, dark brown cells of textura angularis, and inner stratum of hyaline, flattened cells. Hamathecium with paraphyses. Asci 4–8-spored, rounded at the apex, with thick pedicel (Saccardo 1913). Ascospores 39–40 × 12–14 μm ($\overline{x}=39 \times 14\ \mu m$, n = 5), 2–4-seriate, pale brown to brown, somewhat reddish, oblong to fusoid, 3-septate, slightly constricted at the septa, smooth-walled. Asexual morph: Undetermined.

Material examined

JAPAN, Prov. Mino, Kawauye-mura, on straw of Sasa paniculata (Poaceae), January 1913, K. Hara 97 (F10662, holotype).

Note: The holotype specimen of Haraea japonica is not in good condition. In this study, we could not find the characters to illustrate the asci and hamathecium (Fig. 28). This genus is also typical of Iodosphaeria Samuels et al. based on dark brown ascomatal setae, arising from cells at the ascomata surface with a stellate arrangement (Senanayake et al. 2015). Thus, H. japonica may be an over mature collection of Iodosphaeria, however, many characters were not seen in the type specimen. Hence, a new collection with sequence data are needed to illustrate its morphology and to clarify its placement in phylogenetic tree.

Fig. 28

Haraea japonica (holotype). a, b Herbarium packet and specimen. c Ascomata on host substrate. d, e Section through ascomata. f Hyphae at the base of ascomata. g Upper wall of ascoma. h Hyphae with septa. i–k Conidia. Scale bars: e = 500 μm, f–g = 100 μm, h–m = 10 μm

Hypasteridium Speg., Boln Acad. nac. Cienc. Córdoba 26(2-4): 349 (1921)

Notes: The genus was introduced with Metasteridium Speg. and Parasteridium Speg by Spegazzini (1921). However, no species name was mentioned, and the publication was noted as uncertain by Eriksson and Hawksworth (1988). Thus, we treat the three genera as doubtful until the original publication or herbarium specimens can be examined and emended.

Leptascospora Speg., Physis, Rev. Soc. Arg. Cienc. Nat. 4(no. 17): 284 (1918)

Facesoffunginumber: FoF00735

Epiphytes on the surface of leaves. Superficial hyphae branched at the base of ascomata. Sexual morph: Ascomata superficial, globose to subglobose, flattened above, with colorless appendages. Hamathecium with paraphyses. Asci 8-spored, cylindrical, rounded at the apex, narrowed at the base. Ascospores linear, multiseptate, rounded ends, hyaline, smooth-walled (Raciborski 1909). Asexual morph: Undetermined.

Notes: The monotypic genus Leptascospora was introduced by Spegazzini (1918), with the type species L. uredinis.

Type species: Leptascospora uredinis (Racib.) Speg., Physis, Rev. Soc. Arg. Cienc. Nat. 4(no. 17): 284 (1918)

Material examined

INDONESIA, Java Bogor, on the lower surface of leaves of Mucuna sp. (Fabaceae), M. Raciborski (ZT Myc 24176, holotype).

Notes: Leptascospora uredinis was predated as Hyaloderma uredinis Racib. We re-examined the holotype material and it is in poor condition, thus, we treat Leptascospora as doubtful until it can be recollected and sequenced.

Metasteridium Speg.

Notes: see under Hypasteridium Speg.

Ophiociliomyces Bat. & I.H. Lima, Anais Soc. Biol. Pernambuco 13(2): 29 (1955)

Facesoffunginumber: FoF00743

Epiphytes or saprobes on leaves of Bauhinia. Superficial hyphae and hyphopodia lacking. Sexual morph: Ascomata superficial on the surface of leaves, solitary, globose, collapsing when dry. Peridium dark brown, comprising interwoven cells when viewed in squash mounts, darker at rim, with two strata, outer stratum of thick-walled, dark brown cells, and inner stratum of hyaline to pale brown cells. Asci 8-spored, bitunicate, clavate to ovoid, or subglobose, apedicellate, apically rounded, without ocular chamber. Ascospores fasciculate, hyaline to greyish, fusiform-clavate, 10–13-septate, not constricted at septa, straight to slightly curved, smooth-walled. Asexual morph: Undetermined.

Notes: Ophiociliomyces bauhiniae is the type species of Ophiociliomyces and has different morphology from the members of Meliolaceae. Thus, this genus is placed as doubtful. Four species are listed in Index Fungorum.

Type species: Ophiociliomyces bauhiniae Bat. & I.H. Lima, Anais Soc. Biol. Pernambuco 13(2): 30 (1955)

Facesoffunginumber: FoF00742

Epiphytes or saprobic on leaves of Bauhinia raddiana. Superficial hyphae and hyphopodia lacking. Sexual morph: Ascomata 185–240 μm diam. ($\overline{x}=211\ \mu m$, n = 5), superficial on the surface of leaves, solitary, globose, collapsing when dry. Peridium 18–27 μm, dark brown, comprising interwoven cells when viewed in squash mounts, darker at rim, with two strata, outer stratum of thick-walled, dark brown cells, and inner stratum of hyaline to pale brown cells. Hamathecium lacking pseudoparaphyses. Asci 46–67.5 × 13–32 μm ($\overline{x}=55.5 \times 21\ \mu m$ n = 10), 8-spored, bitunicate, clavate to ovoid, or subglobose, apedicellate, apically rounded, without ocular chamber. Ascospores 52–77 × 5–9.5 μm ($\overline{x}=63.5 \times 7\ \mu m$, n = 20), fasciculate, hyaline to greyish, fusiform-clavate, 10–13-septate, not constricted at septa, straight to slightly curved, smooth-walled. Asexual morph: Undetermined.

Material examined

BRAZIL, Pernambuco; Recife, Dois Irmãos., on leaves of Bauhinia raddiana Bong. (Fabaceae), 5 February 1955, Osvaldo Soares da Silva, (URM 1235, holotype).

Notes: The type specimen was re-examined and lacked superficial hyphae, had bitunicate asci and multi-septate, hyaline ascospores (Fig. 29). We therefore treat this genus as doubtful within Meliolales.

Fig. 29

Ophiociliomyces bauhiniae (holotype). a Herbarium material and specimen. b Ascoma on substrate. c Squash mount of ascoma. d, e Peridium and ascoma wall. f, g Asci at young and mature stages (g mounted in cotton blue reagent). h, i Ascospores. Scale bars: b, c =100 μm, d = 20 μm, e = 10 μm, f–i = 40 μm

Ophioirenina Sawada & W. Yamam., in Imazeki et al., Special Publication College of Agriculture, National Taiwan University 8: 35 (1959)

Facesoffunginumber: FoF00745

Epiphytes on the surface of leaves. Superficial hyphae branched, septate, with 1-celled hyphopodia. Sexual morph: Ascomata superficial on surface of leaves, solitary or gregarious, globose to subglobose, glabrous, dark brown to black, central ostiole at maturity. Peridium thick, comprising two strata, outer stratum of thick-walled, brown to dark brown cells of textura angularis to globulosa, inner stratum of hyaline to pale brown, scleroparenchymatous cells of textura angularis to globulosa. Hamathecium aparaphysate. Asci 8-spored, unitunicate, clavate, short pedicellate or sometimes apedicellate, ocular chamber absent. Ascospores fasciculate, hyaline to brown, clavate to narrowly fusiform, 4–6-septate, not constricted at septa, slightly darker at septa at maturity, smooth-walled. Asexual morph: Undetermined.

Notes: Ophioirenina was introduced and placed in Meliolaceae by Sawada and Yamamoto (in Sawada 1959) based on its superficial hyphae with hyphopodia, and globose ascomata developing in black colonies (Lumbsch and Huhndorf 2010; Patil and Mahamulkar 1999). Ophioirenina differs from Meliolales species in having 1-celled hyphopodia on the hyphae, papilla with a narrow pore or opening via a rupture, a peridial wall comprising two strata of scleroparenchymatous cells of textura angularis and hyaline ascospores. The genus is most typical of Leptosphaeriaceae species because of its scleroplectenchyma cells. We treat Ophioirenina as Dothideomycetes genera, incertae sedis until molecular data available.

Type species: Ophioirenina theae Sawada & W. Yamam., Special Publication College of Agriculture, National Taiwan University 8: 36 (1959)

Facesoffunginumber: FoF00744

Epiphytes on the surface of leaves. Superficial hyphae branched, septate, with 1-celled hyphopodia. Sexual morph: Ascomata 170–220 μm diam. ($\overline{x}=197\ \mu m$, n = 5), superficial on surface of leaves, solitary or gregarious, globose to subglobose, glabrous, dark brown to black, central ostiole at maturity. Peridium 43–52 μm ($\overline{x}=46\ \mu m$, n = 5), thick, comprising two strata, outer stratum of thick-walled, brown to dark brown cells of textura angularis to globulosa, inner stratum of hyaline to pale brown, scleroparenchymatous cells of textura angularis to globulosa. Asci 78–81 × 18–20 μm ($\overline{x}=80 \times 18\ \mu m$, n = 10), 8-spored, unitunicate, clavate, short pedicellate or sometimes apedicellate, ocular chamber absent. Ascospores 50–62 × 5–6 μm ($\overline{x}=58 \times 5\ \mu m$, n = 10), fasciculate, hyaline to brown, clavate to narrowly fusiform, 4–6-septate, not constricted at the septa, slightly darker at each septum at maturity, smooth-walled. Asexual morph: Undetermined.

Material examined

TAIWAN, Shin-Chu, Formosa, on leave of Camellia sinensis (L.) Kuntze (Theaceae), 11 November 1925, K. Sawada (BPI 698821)

Notes: We found ascomata developing in the black colonies which are quite similar to Meliolales species, but differ based on their 1-celled hyphopodia, ascoma wall, and ascospore septation (Fig. 30).

Fig. 30

Ophioirenina theae (holotype). a, b Herbarium packet and specimen. c, d Ascomata on substrate. e Section through ascomata. f Peridium and ascoma wall. g Ascoma when immature. h Hyphae with 1-celled appressoria. i, j Asci with 8-spores. k, l Ascospores. Scale bars: e = 50 μm, f–l = 20 μm

Ophiomeliola Starbäck, Bih. K. svenska VetenskAkad. Handl., Afd. 3 25(no. 1): 22 (1899)

Facesoffunginumber: FoF00747

Epiphytes or pathogens on surface of living leaves. Superficial hyphae septate, dark brown, without hyphopodia. Sexual morph: Ascomata superficial on surface of leaves, solitary, subglobose to ovoid, covered with aerial hyphae, with apical, papillate ostiole. Peridium comprising two strata, outer stratum of thick-walled, dark brown to black cells of textura globulosa, and inner stratum of hyaline to pale brown cells, scleroparenchymatous cells of textura globulosa to angularis. Hamathecium comprising filiform paraphyses embedded in a gelatinous matrix. Asci 8-spored, unitunicate, clavate to cylindrical, with long pedicel. Ascospores fasciculate, hyaline, cylindrical to narrowly fusiform, 7-septate, not constricted at septa, slightly curved, tapering toward the ends, smooth-walled. Asexual morph: Undetermined.

Notes: Ophiomeliola was introduced by Starbäck (1899), with the type species Ophiomeliola lindmanii. Four species are listed for this genus in Index Fungorum and all lack molecular data. This genus cannot be placed in Meliolaceae based on its fusiform, 7-septate, hyaline ascospores, but it may belong to Lasiosphaeriaceae based on its ascomata wall. However, we treat this as Sordariomycetes genus incertae sedis until molecular data becomes available.

Type species: Ophiomeliola lindmanii Starbäck, Bih. K. svenska VetenskAkad. Handl., Afd. 3 25(no. 1): 22 (1899)

Facesoffunginumber: FoF00746

Epiphytes or pathogens on surface of living leaves. Superficial hyphae septate, dark brown, without hyphopodia. Sexual morph: Ascomata 310–345 μm high × 270–345 μm diam. ($\overline{x}=334 \times 295\ \mu m$, n = 3), superficial on surface of leaves, solitary, subglobose to ovoid, covered with aerial hyphae, with apical, papillate ostiole. Peridium 35–40 μm, comprising two strata, outer stratum of dark brown to black cells of textura globulosa, and inner stratum of hyaline to pale brown, scleroparenchymatous cells of textura globulosa to angularis. Hamathecium comprising filiform paraphyses embedded in a gelatinous matrix. Asci 121–141 × 17.5–20 μm ($\overline{x}=135 \times 19\ \mu m$, n = 15), 8-spored, unitunicate, clavate to cylindrical, with long pedicel. Ascospores 88–125 × 3–6 μm ($\overline{x}=109 \times 5\ \mu m$, n = 20), fasciculate, hyaline, cylindrical to narrowly fusiform, 7-septate, not constricted at septa, slightly curved, tapering toward the ends, smooth-walled. Asexual morph: Undetermined.

Material examined

BRAZIL, Rio Grande do Sul, Hamburgerberg pr Porto Alegre, on leaves of Myrtacea cujusdam (Myrtaceae), September 1892, C.A.M. Lindman No. 37 (SF10033, holotype).

Notes: Ophiomeliola lindmanii is typical of species in Lasiosphaeriaceae in having aerial hyphae, a peridium comprising two strata, with the outer strata of dark brown to black cells of textura globulosa to angularis, unitunicate asci, and hyaline, multi-septate ascospores (Fig. 31). Molecular data is needed to clarify the placement.

Fig. 31

Ophiomeliola lindmanii (holotype). a Herbarium packet and material. b. Ascomata on substrate. c Aerial hyphae. d Vertical section of ascoma. e Peridium. f Paraphyses. g–k Unitunicate asci. i–n Ascospores. Scale bars: b = 1 mm, c, d, g–n = 50 μm, e = 40 μm, f = 20 μm

Parasteridium Speg., Boln Acad. nac. Cienc. Córdoba 26(2-4): 349 (1921)

Notes: see under Hypasteridium Speg.

Pauahia F. Stevens, Bulletin of the Bernice P. Bishop Museum, Honolulu, Hawaii 19: 17 (1925)

Facesoffunginumber: FoF00749

Epiphytes on the surface of Sideroxylon. Superficial hyphae and hyphopodia absent. Sexual morph: Ascostromata superficial or erumpent at the base, pseudostromata, solitary, subglobose or irregular, thick-walled. Locules immersed in ascostromata, 1–multi-loculate. Peridium of locules comprising two strata, outer stratum of dark brown cells of textura angularis, and inner stratum of hyaline to pale brown, flattened cells. Hamathecium with paraphyses. Asci 2-spored, bitunicate, broadly clavate to oval, with short pedicel, evanescent. Ascospores 2-seriate, hyaline to brown, oblong to cylindrical, 3-septate, constricted and darker at the septa, terminal cells markedly smaller and slightly lighter than the others, thin sheath present at maturity, smooth-walled. Asexual morph: Undetermined.

Notes: The genus Pauahia was established within the family Dothideaceae in the section Leveillelleae by Stevens (1925). Pauahia is monotypic with Pauahia sideroxyli as its type species. We re-examined the type specimen of P. sideroxyli and found that the species differs from other genera in Meliolaceae in having pseudostromata, and lacking superficial hyphae.

Type species: Pauahia sideroxyli F. Stevens, Bulletin of the Bernice P. Bishop Museum, Honolulu, Hawaii 19: 17 (1925)

Facesoffunginumber: FoF00748

Epiphytes on the surface of Sideroxylon. Superficial hyphae and hyphopodia absent. Sexual morph: Ascostromata 700–1200 μm diam. × 353–410 μm high ($\overline{x}=1000 \times 365\ \mu m$, n = 2), superficial or erumpent at the base, solitary, pseudostromata, subglobose or irregular, thick-walled. Locules 200–235 × 165–219 μm ($\overline{x}=208 \times 179\ \mu m$, n = 5), immersed in ascostromata, 1–multi-loculate. Peridium of locules comprising two strata, outer stratum of dark brown cells of textura angularis, and inner stratum of hyaline to pale brown, flattened cells. Hamathecium with paraphyses. Asci 46–82 × 25–45 μm ($\overline{x}=70 \times 36\ \mu m$, n = 5), 2-spored, bitunicate, broadly clavate to oval, with short pedicel, evanescent. Ascospores 20–23 × 62–77 μm ($\overline{x}=22 \times 65\ \mu m$, n = 5), 2-seriate, hyaline to brown, oblong to cylindrical, 3-septate, constricted and darker at the septa, terminal cells markedly smaller and slightly lighter than the others, thin sheath present at maturity, smooth-walled. Asexual morph: Undetermined.

Notes: Pauahia sideroxyli is a single species in the genus Pauahia. We re-examined the type specimen which is very small and in poor condition, and found that Pauahia sideroxyli differs from Meliolales species in having a large pseudostromata, erumpent in host epidermal cells, and bitunicate asci (Fig. 32). Molecular data is lacking, so we cannot clarify the placement of this genus. Therefore, Pauahia sideroxyli is placed as Dothideomycetes genera, incertae sedis until it can be recollected and sequenced.

Material examined

USA, Hawaii, Maui, Nahiku, on Planchonella sandwicensis (A.Gray) Pierre (Sapotaceae), January 1909, Lyon H. L. no. 61 (ILL00011484, holotype; BISH 499038, isotype).

Fig. 32

Pauahia sideroxyli (holotype). a Specimen. b Ascostromata on substrate. c Section through ascostromata showing locule. d, e Peridium of ascostromata. f Hamathecium. g, h Asci with 2-spores in cotton blue reagent. i, j Ascospores with 3 septa and lighter end cells. k Ascospores in congo red reagent. Scale bars: c = 100 μm, d, e, g–k = 20 μm, f = 10 μm

Pleomeliola (Sacc.) Sacc., Syll. fung. (Abellini) 14(1): 17 (1899)

≡ Meliola subgen. Pleomeliola Sacc., Syll. fung. (Abellini) 1: 70 (1882)

Notes: Pleomeliola contains three species which are listed in Index Fungorum (2015) with the type species as Pleomeliola fenestrata (Cooke & Ellis) Sacc. The genus is characterized by subglobose, membranacous, reddish brown ascomata, 8-spored, ovoid asci, and cylindrical or clavate, muriform, 4–5-septate, hyaline to brown ascospores. We could not found any ascomata or hyphae on the holotype specimen. Hansford examined this specimen in 1952, and noted on the packet “This specimen is not Meliola” (Fig. 33) . We also exclude this genus from Meliolales. There is no molecular data available for this genus.

Type species: Pleomeliola fenestrata (Cooke & Ellis) Sacc., Syll. fung. (Abellini) 14(1): 17 (1899)

Material examined

USA, New Jersey, on scales of pine cones (Pinaceae), MC. Cooke 2465 ex herb. (K(M) 193124, holotype).

Fig. 33

Pleomeliola fenestrata (holotype). a–d Herbarium packet and specimen

Pleomerium Speg., Physis, Rev. Soc. Arg. Cienc. Nat. 4(no. 17): 284 (1918)

Facesoffunginumber: FoF00751

Epiphytes on the lower surface of leaves. Superficial hyphae branched, septate, darker at each septum, brown, lacking hyphopodia and hyphal setae. Sexual morph: Ascomata superficial, solitary on superficial hyphae, easily removed, subglobose to globose, surrounded and covered by hyphae, thin-walled. Peridium thin, membranous, comprising brownish to dark brown, interwoven cells when viewed in squash mounts. Hamathecium lacking pseudoparaphyses. Asci 8-spored, bitunicate, obovoid to subglobose, short pedicel or absent, ocular chamber not well developed. Ascospores 3–4-seriate, hyaline to brown, oblong to fusiform, muriform, 4–8 transverse septa, 1–2 longitudinal septa, not constricted but darker at septa, surrounded and covered by mucilage, smooth-walled. Asexual morph: Conidia oblong, 1-celled (Rehm 1901).

Notes: The monotypic genus Pleomerium was established within the family Englerulaceae by Spegazzini (1918). This genus differs from Meliolales species in having superficial hyphae without hyphopodia, bitunicate asci, and muriform ascospores. Molecular data is not available in GenBank. Hence, we suggest that the genus should be placed as Dothideomycetes genera, incertae sedis, until sequence data can clarify its placement.

Type species: Pleomerium fuscoviridescens (Rehm) Speg., Physis, Rev. Soc. Arg. Cienc. Nat. 4(no. 17): 284 (1918)

≡ Limacinia fuscoviridescens Rehm, Hedwigia 40: 168 (1901)

≡ Naetrocymbe fuscoviridescens (Rehm) Bat. & Cif., Saccardoa 2: 159 (1963)

Facesoffunginumber: FoF00750

Epiphytes on the lower surface of leaves. Superficial hyphae branched, septate, darker at septa, brown, lacking hyphopodia and hyphal setae. Sexual morph: Ascomata 280–350 μm diam. ($\overline{x}=312\ \mu m$, n = 5), superficial, solitary on superficial hyphae, easily removed, subglobose to globose, surrounded and covered by hyphae, thin-walled. Peridium thin, membranous, comprising brownish to dark brown, interwoven cells when viewed in squash mounts. Hamathecium lacking pseudoparaphyses. Asci 84–96 × 58–63 μm ($\overline{x}=87 \times 60\ \mu m$, n = 5), 8-spored, bitunicate, obovoid to subglobose, short pedicel or absent, ocular chamber not well-developed. Ascospores 44–47 × 16–17 μm ($\overline{x}=45 \times 16\ \mu m$, n = 10), 3–4-seriate, hyaline to brown, oblong to fusiform, muriform, 4–8 transverse septa, 1–2 longitudinal septa, not constricted but darker at septa, surrounded and covered by mucilage, smooth-walled. Asexual morph: Undetermined.

Notes: Pleomeliola fuscoviridescens was introduced as Limacinia fuscoviridescens by Rehm (1901). We re-examined the holotype specimen of P. fuscoviridescens and found that it differs from other species in Meliolales (Fig. 34) This genus may belong in Englerulaceae because of its membranous ascomata, containing few asci, lack of ostiole, and dissolving at centre to release bitunicate asci. However, molecular data is needed to clarify its placement.

Material examined

BRAZIL, Santa Catarina, Blumenau, on leaves of undetermined host, November 1887, E. Ule 941 b (SF7084, holotype of Limacinia fuscoviridescens Rehm).

Fig. 34

Pleomerium fuscoviridescens (holotype). a, b Herbarium packet and specimen. c Ascoma on substrate. d Ascoma viewed in squash mount. e Section through ascoma. f Upper wall cells of young ascoma. g Mycelium. h Ascus with 8-spores. i Ascus in 70 % lactic acid. j Ascus in Melzer’s reagent. k Ascus in cotton blue reagent. l–m Ascospores at maturity. Scale bars: d, e = 100 μm, f, h–k = 50 μm, g, l = 20 μm

Prataprajella Hosag., Nova Hedwigia 55(1-2): 224 (1992)

Notes: Prataprajella was introduced by Hosagoudar (1992). The type species was found on leaves of Turpinia malabarica, and was deposited in HCIO, which will not loan specimens outside India. Therefore, a re-drawing from Hosagoudar (2006) is provided here (Fig. 35). Prataprajella is characterized by superficial hyphae with globose to stellate hyphopodia, hyphal setae lacking, globose to subglobose ascomata, with raised ascomatal setae, and conical appendages, brown, 3-septate ascospores (Hosagoudar 2006). We treat Prataprajella as a doubtful genus until the type specimen can be re-examined, or fresh collection can be collected and sequenced.

Type species : Prataprajella turpiniicola (Hosag.) Hosag., Nova Hedwigia 55(1-2): 225(1992)

Fig. 35

Prataprajella sp. (a redrawn from Hosagoudar 2006), Prataprajella turpiniicola (b–e redrawn from Hosagoudar and Riju 2013). a Ascoma with appendages and setae surrounded by septate hyphae with capitate hyphopodia and phialides. b Stellate hyphopodia on hyphae. c Stellate hyphopodia mixed with phialides on hyphae. d Setae with rounded ends. e Ascospores with 3 septa and germinated ascospore

Ticomyces Toro, J. Agric. Univ. Puerto Rico 36: 48 (1952)

Possible synonyms

= Tonduzia F. Stevens, Illinois Biol. Monogr. (Urbana) 11(2): 16 (1927)

= Dontuzia L.D. Gómez, Brenesia 2: 21 (1973)

Notes: Ticomyces was predated as Tonduzia Stevens in the family Perisporiaceae (Stevens 1927), with the type species T. psychotriae Stevens. Toro (1952) introduced Ticomyces in Meliolaceae (Kirk et al. 2008). However, the genus differs from other species in Meliolales in having superficial aseptate hyphae without hyphopodia. Ascospores are hyaline with several septa (Batista 1951). There is no molecular data available for this genus, thus we treat Ticomyces in Sordariomycetes genera, incertae sedis.

Type species: Ticomyces psychotriae (F. Stevens) Toro, J. Agric. Univ. P. Rico 36: 49 (1952)

Tonduzia psychotriae F. Stevens, Illinois Biol. Monogr. (Urbana) 11(2): 168 (1927)

Dontuzia psychotriae (F. Stevens) L.D. Gómez, Brenesia 2: 21 (1973)

Material examined

COSTA RICA, Columbiana, on Psychotria brachiate Sw. (Rubiaceae), 19 July 1923, F.L. Stevens 570 (ILL00011485, glass slide from holotype).

Notes: We re-examined the type specimen of Ticomyces psychotriae but could not see asci and ascospores (Fig. 36). Therefore, new fresh collection and sequence data are needed to clarify placement of the genus.

Fig. 36

Ticomyces psychotriae (holotype). a Herbarium packet. b Glass slide. c, d Ascomata developed on hyphae. e Hyphae without hyphopodia

Urupe Viégas, Bragantia 4(1-6): 125 (1944)

Notes: Urupe was introduced by Viégas (1944) and is a monotypic genus. The genus is characterized by intercellular hyphae, reddish ascomata, clavate asci with 8-spores, and hyaline, 3-septate ascospores (Viégas 1944). We were unable to borrow the type species, therefore, we treat Urupe as Sordariomycetes genera, incertae sedis based on morphology as illustrated in Viégas (1944).

Type species: Urupe guaduae Viégas, Bragantia 4(1-6): 125 (1944)

Xenostigme Syd., Annls mycol. 28(5/6): 434 (1930)

Notes: Xenostigma was established by Sydow (1930) with the type species X. trichophila Syd. The genus is monotypic and differs from Meliolales species in having slimy ascomata at maturity, with light grey cells, asci with 8-spores, and dark brown to black, 1-septate ascospores (Sydow 1930). We were unable to locate type material of Xenostigme trichophila. Thus, we place Xenostigme as Sordariomycetes genera, incertae sedis until molecular data is available.

Type species: Xenostigme trichophila Syd., Annls mycol. 28(5/6): 434 (1930)