Dothideales

Abstract

The type specimens or representative specimens of the potentially dothidealean genera Bagnisiella, Botryochora, Coccostromella, Columnosphaeria, Delphinella, Dictyodothis, Discosphaerina, Dothidea, Dothiora, Endodothiora, Jaffuela, Mycoporis, Omphalospora, Pachysacca, Plowrightia, Saccothecium, Stylodothis, Sydowia and Yoshinagaia were examined while, fresh specimens of Aureobasidium pullulans, Dothidea insculpta, Plowrightia ribesia and Saccothecium sepincola were made from Italy and Thailand. An introduction and the history of these genera, their family placement, morphology, and molecular phylogeny are provided. Morphology plus GenBank data are used to provide a systematic treatment of Dothideales. Phylogenetic analysis of LSU, SSU and ITS gene regions was carried out and in the resulting phylogenetic tree the taxa cluster in two clades with high bootstrap support. Clade A comprises Dothideaceae, the family type of Dothideales. The family Dothioraceae is not recognized as a distinct family and is synonymized under Dothideaceae. Neocylindroseptoria is introduced to accommodate Cylindroseptoria pistaciae as it forms a well-supported distinct clade in Dothideaceae. Clade B comprises Aureobasidium, Kabatiella, Pseudoseptoria, Saccothecium and Selenophoma species and Columnosphaeria fagi, for which we propose a new family, Aureobasidiaceae. The recently introduced Sydowia eucalypti also clustered within Clade B and therefore based on morphology and molecular phylogeny a new genus Pseudosydowia is introduced for Sydowia eucalypti. Celosporium laricicola is separated in a distinct clade, and therefore it is placed in Dothideales, genera, incertae sedis. The genera Bagnisiella, Botryochora, Coccostromella, Jaffuela, Lucidascocarpa, Mycoporis, Omphalospora, Pachysacca and Yoshinagaia are excluded from Dothideales and their placements are discussed.

Introduction

The order Dothideales was introduced by Lindau (1897) to accommodate a single family Dothideaceae. During the next 100 years many families were included or removed from this order depending on the importance authors placed on different taxonomic features. This was a difficult time for researchers who attempted higher fungal classifications. Most fungi had few distinctive characters and therefore classifications became rather subjective as certain characters were chosen as important over others, without any real evidence for their importance. The researches such as M.E Barr, D.L Hawksworth, E.S. Luttrell, J. A. Von Arx and E. Müller however, produced significant classification schemes for their time, considering the vacuum they were working in (Luttrell 1973; Von Arx and Müller 1975; Barr 1987a; Hawksworth et al. 1995).

Order-level classification based on morphology

Following the introduction of Dothideales with a single family, the order was first revised by Theissen and Sydow (1915) who included four heterogeneous families (Dothideaceae, Montagnellaceae, Phyllachoraceae, and Polystomellaceae) comprising unitunicate and bitunicate ascomycetes. Luttrell (1955, 1973) accepted the order for only bitunicate ascomycetes with loculi containing several asci and lacking paraphyses. Luttrell (1951) categorized five families in Dothideales (Table 1), while Luttrell (1955) synonymized Pseudosphaeriales, Capnodiales and Dothiorales under Dothideales and accepted four families in Dothideales in the subclass Loculoascomycetes. Luttrell (1973) included eight families in Dothideales, while Von Arx and Müller (1975) placed 34 families and synonymized Dothioraceae under Dothideaceae. (Barr 1979) treated Dothideales with five families and considered Dothideaceae and Dothioraceae as distinct families. Barr (1987a) accepted six families including a new family Kriegeriellaceae (Barr 1987b). Hawksworth et al. (1995) considered Dothideales to be the largest and most varied group of the Ascomycota including most ascolocular ascomycetes with bitunicate asci and assigned 58 families to Dothideales. They synonymized Asterinales, Capnodiales, Chaetothyriales, Dothiorales, Hysteriales, Melanommatales, Myriangiales, Perisporiales, Pleosporales and Pseudosphaeriales under Dothideales. Barr (1996) introduced a new family Planistromellaceae in Dothideales including six genera. Kirk et al. (2001) included Dothideales with other bitunicate orders, Capnodiales and Myriangiales, in the subclass Dothideomycetidae which was characterized by lack of paraphyses, pseudoparaphyses and paraphysoids. Kirk et al. (2008) treated Dothideales with four familes (Dothideaceae, Dothioraceae, Coccoideaceae and Planistromellaceae).

Table 1 Arrangement of families in Dothideales from 1955 to 2010

Arrangement of dothideales with molecular data

Species identification based on morphology is not always adequate in classification schemes as it may be subjective or even wrong. Therefore, researchers have begun to rely on phylogenetic based identification (Liu et al. 2012; Chomnunti et al. 2014; Ariyawansa et al. 2014; Hongsanan et al. 2014; Hyde et al. 2014; Nilsson et al. 2014; Phookamsak et al. 2014; Wijayawardene et al. 2014). Phylogenetic analyses by Schoch et al. (2006, 2009) showed the Dothideales to be a well-supported order with nine other orders in the class Dothideomycetes and the order Myriangiales to be closely related to the Dothideales. They also validated the concept of Dothideomycetidae sensu Kirk et al. (2001) with several amendments. Schoch et al. (2006, 2009) used phylogenetic analysis to clearly show that the family Botryosphaeriaceae is distinct from the Dothideales and that was confirmed by Liu et al. (2012). Lumbsch and Huhndorf (2010) accepted the families, Dothideaceae, Dothioraceae and Teratosphaeriaceae in Dothideales, while Crous et al. (2007) and Hyde et al. (2013) moved Teratosphaeriaceae to Capnodiales.

In this study, we present new taxonomic and systematic treatment for Dothideales based on morphology and multigene phylogenetic analyses.

History of Dothideaceae

The family Dothideaceae was introduced by Chevallier (1826) as ‘Dothideae’, and later Fuckel (1869) established this family with Dothidea as the type genus and D. gibberulosa as the type species, which were characterized by locules embedded in ascostromata without definite perithecia. This family however, has a rather varied past as can be seen from inclusion of genera by various authors (Table 2) and the follow up account.

Table 2 Accepted genera according to various treatments of Dothideaceae

From its introduction in 1896 to early 2000, the family Dothideaceae underwent numerous changes in concept and inclusion of genera and was largely based on the understanding of morphological characters. Winter (1887) separated Dothideaceae from Hypocreaceae and Sphaeriaceae based on fleshy, black or blackish brown ascostromata with lack of perithecia and included nine genera with 38 species (Orton 1924). Theissen and Sydow (1915) placed this family in Dothideales and divided it into three subfamilies (Coccoideae, Dothideae and Leveillelleae). Gäumann (1952) included Dothideaceae in Pseudosphaeriales along with Dothioraceae and four other families. Luttrell (1973) categorized 22 genera under Dothideaceae, which is the highest number of genera included in the family. Arx and Müller (1975) synonymized Dothioraceae under Dothideaceae and included eleven genera. They accepted Dothideaceae for members having the following characters; ascomata developing as non-ostiolate loculi in stromata, opening by an apical fissure or dehiscence, eight- or many-spored asci borne at the base of the locules and one or many-septate, hyaline or brown, often guttulate ascospores. Sivanesan (1984) treated Dothideaceae with 14 genera and synonymized Botryosphaeriaceae, Dothioraceae and Mycosphaerellaceae under this family. Barr (1987a treated Dothideaceae and Dothioraceae as two separate families and listed 16 genera under Dothideaceae. Hawksworth et al. (1995) removed 13 genera from those of Barr (1987a and added ten other genera. Hawksworth et al. (1995) accepted 13 genera in Dothideaceae characterized by multiloculate ascostromata, saccate or clavate asci and transversely septate ascospores. In a first multigene molecular study, Schoch et al. (2009) confirmed the familial placement of Dothideaceae in the order Dothideales based on Dothidea species including the type D. sambuci and Stylodothis puccinioides (DC.) Arx & E. Müll. Lumbsch and Huhndorf (2010) listed 13 genera including Dictyodothis and Lucidascocarpa to the genera list of Hawksworth et al. (1995), while removing Hyalocrea and Planistroma. Planistroma was later included in Planistromellaceae in Botryosphaeriales based on molecular data (Hyde et al. 2013; Monkai et al. 2013). Wijayawardene et al. (2012) treated the family Dothideaceae with two asexual genera Lecanosticta and Podoplaconema based on the asexual states of Scirrhia and Omphalospora respectively, while Hyde et al. (2013) included Endoconidioma, Kabatina and Podoplaconema.

History of Dothioraceae

The family Dothioraceae was introduced by Theissen and Sydow (1917) in the order Myriangiales along with five other families (Elsinoaceae, Plectodiscelleae, Myxomyriangiaceae, Myriangiaceae and Saccardiaceae) and included five genera Bagnisiella, Dothiora, Pseudosphaeria, Wettsteinina, and Yoshinagaia. This family has been referred to the Dothiorales by Müller and von Arx (1950). Dothiorales was introduced for species having broad, nearly sphaerical or clavate asci, borne in non-ostiolate ascomata, opening at maturity by dehiscence or rupture, or in which the asci develop a naked hymenium (Müller and von Arx 1950). Luttrell (1951b) and Gäumann (1952) included Dothioraceae in Pseudosphaeriales and Gäumann (1952) mentioned that Dothioraceae included the more primitive representatives of the Pseudosphaeriales. Luttrell (1955) synonymized Dothiorales under Dothideales which was followed by some authors (Von Arx and Müller 1975; Sivanesan 1984), but not others (Froidevaux 1972; Luttrell 1973; Barr 1979a, b, 1987a; Hawksworth et al. 1995). Froidevaux (1972) accepted four genera in Dothioraceae, while Barr (1972) included eight. Luttrell (1973) listed seven genera in Dothioraceae. Von Arx and Müller (1975) and Sivanesan (1984) referred Dothideaceae and Dothioraceae as a single family in Dothideales, while (Barr 1979, 1987a) treated Dothideaceae and Dothioraceae as separate families. (Barr 1979) listed 13 genera in Dothioraceae, while Barr (1987a) included five. Hawksworth et al. (1995) listed eight genera characterized by uniloculate ascostromata, clavate asci with septate or muriform ascospores. Lumbsch and Huhndorf (2010) extended the genera in Dothioraceae to ten by adding Phaeocrypotus and Yoshinagaia to those of Hawksworth et al. (1995). Wijayawardene et al. (2012) treated the family Dothioraceae with inclusion of the asexual genera Aureobasidium, Dothichiza, Hormonema, Japonia, Kabatina, Rhizosphaera and Sclerophoma.

Materials and methods

Examination of herbarium material

The type specimens or representative specimens of Bagnisiella, Botryochora, Coccostromella, Columnosphaeria, Delphinella, Dictyodothis, Discosphaerina, Dothidea, Dothiora, Endodothiora, Jaffuela, Mycoporis, Omphalospora, Pachysacca, Phaeocryptopus, Plowrightia, Saccothecium, Stylodothis, Sydowia and Yoshinagaia were obtained from BPI, C, K, S, URM and W. Examination of the type specimens follow Chomnunti et al. (2011). Ascomata were rehydrated in 5 % KOH prior to examination and sectioning. Specimens were examined under a stereo microscope (Motic SMZ 168) and fine forceps were used to remove one or two ascomata and mounted in water. Hand sections were cut with a sharp razor blade. Observations and photomicrographs were made from material mounted in water or lactophenol with cotton blue dye using Nikon ECLIPSE 80i light microscope fitted with a Cannon 450D digital camera. India ink was added to water mounts to detect the presence of gelatinous sheaths or ascospore appendages. Measurements were made with Tarosoft (R) Image Frame Work.

Sample collection, specimen examination and isolation

Fresh specimens were collected in Italy and Thailand. The specimens were observed and examined under Motic SMZ 168 stereomicroscope. Micromorphological characters of the fungus were examined using a Nikon ECLIPSE 80i compound microscope and images captured using a Nikon ECLIPSE 80i compound microscope with a Canon EOS 550D digital camera. Measurements were made with the Tarosoft (R) Image Frame Work and images used for figures processed with Adobe Photoshop CS3 Extended version 10.0 software. Following the method of Chomnunti et al. (2014), a culture was derived from single spore isolation. Germinating spores were transferred to Potato Dextrose Agar (PAD) medium or Malt Extract Agar (MEA) and incubated at 25 °C in the dark. The cultural characteristics such as colour of the mycelium, and shape, texture and growth rate of colonies were recorded after 14 days.

DNA extraction, PCR amplification and sequencing

Fungal isolates were grown on PDA or MEA (Malt Dextrose Agar) at 25 °C for 2–4 weeks. Genomic DNA from mycelia was extracted as in Udayanga et al. (2012) while, Genomic DNA from fruiting bodies was extracted using Biospin Fungus Genomic DNA Extraction Kit (BioFlux®) following the instructions of the manufacturer. Polymerase chain reaction (PCR) was carried out using four partial gene portions in this study. NS1 and NS4 primers were used to amplify a region spanning the small subunit rDNA (White et al. 1990). LROR and LR5 primer pairs were used to amplify a segment of the large subunit rDNA (Vilgalys and Hester 1990) and internal transcribed spacers was amplified by primer pairs ITS1 and ITS4 (White et al. 1990). The amplifications were performed in 25 μL of PCR mixtures containing 9.5 μL ddH2O, 12.5 μL 2 × PCR Master Mix (TIANGEN Co., China), 1 μL of DNA template, 1 μL of each primer (10 μM). The amplification conditions for SSU, LSU and ITS consisted of initial denaturation at 94 °C for 4 mins; followed by 35 cycles of 45 s at 94 °C, 45 s at 56 °C and 1 min at 72 °C, and a final extension period of 10 mins at 72 °C. The PCR products were observed on 1 % agarose electrophoresis gels stained with Ethidium bromide. Purification and sequencing of PCR products were carried at using the above-mentioned PCR primer at Invitrogen Biotechnology Co., Ltd, China.

ATCC American Type Culture Collection, Virginia, USA; CBS Centraalbureau voor Schimmelcultures, Utrecht, The Netherlands; CPC Collection of Pedro Crous housed at CBS; DAOM Plant Research Institute, Department of Agriculture (Mycology), Ottawa, Canada; MFLU Mae Fah Luang University Herbarium Collection; MFLUCC Mae Fah Luang University Culture Collection, ChiangRai, Thailand; The University of Alberta Microfungus Collection and Herbarium.

Phylogenetic analysis

The phylogenetic analysis follows the methods used by Boonmee et al. (2014) with modifications as needed. The generated LSU, SSU and ITS sequences were analyzed using the BLAST search engine of the National Center for Biotechnology Information (NCBI) for the rough identification of fresh isolates used in the analyses. Sequences of the available ex-type cultures and other taxa were obtained from GenBank are provided (Table 4). In addition, several other fungal taxa of the close families (Capnodiaceae, Elsinoaceae, Mycosphaerellaceae and Myriangiaceae) were also included in the analyses. The consensus sequences for each gene were initially aligned by ClustalX v. 1.83 and in Bioedit (Thompson et al. 1997). LSU, SSU and EF datasets were first analyzed separately and then the individual datasets were concatenated into a combined dataset. The model of evolution was performed by using MrModeltest 2.2 (Nylander 2004). A phylogenetic analysis of the concatenated alignment was performed on CIPRES webportal (Miller et al. 2010) using RAxML v. 7.2.7 (Stamatakis 2006; Stamatakis et al. 2008). The best scoring tree was selected with a final likelihood value of −21859.18898. One thousand non parametric bootstrap iterations were run with the GTR model and a discrete gamma distribution. The resulting replicates were plotted on to the best scoring tree obtained previously. Maximum Likelihood bootstrap values (MLBP) equal or greater than 50 % are given above each node (Fig. 1). Posterior probabilities (PP) (Rannala and Yang 1996; Zhaxybayeva and Gogarten 2002) were determined by Markov Chain Monte Carlo sampling (BMCMC) in MrBayes v. 3.0b4 (Huelsenbeck and Ronquist 2001). Six simultaneous Markov chains were run for 1000000 generations and trees were sampled every 100th generation and 10,000 trees were obtained. The first 2,000 trees, representing the burn-in phase of the analyses, were discarded while remaining 8,000 trees used for calculating posterior probabilities in the majority rule consensus tree (Cai et al. 2006). Bayesian Posterior Probabilities (BYPP) with those equal or greater than 0.80 given below each node (Fig. 1). General time reversible model (GTR) was applied with a discrete gamma distribution and four rate classes. Phylogenetic trees were drawn using Treeview v. 1.6.6 (Page 2001). The sequences of novel species and other sequenced taxa in this study are deposited in GenBank.

Fig. 1

RAxML tree based on a combined dataset of ITS, SSU and LSU gene regions. The first set of numbers above or below the nodes are bootstraps from 1000 repetitions, with values above 50 % shown. The second represent Bayesian posterior probabilities expressed as percentages, with values above 80 % shown. Pleospora herbarum is the out-group taxon. The original isolate numbers are noted after the species names and ex-types are in bold

Results

Phylogenetic analysis

The combined 28S (LSU), 18S (SSU) and ITS gene data set consists of 55 taxa of which five are newly generated and 45 are from GenBank, including 45 dothidealean taxa and Pleospora herbarum as the outgroup taxon. All trees were similar in topology and not significantly different (data not shown). Fourty-four taxa in Dothideales separated into two major clades with 65/80 % (RAxML / Bayesian posterior probabilities) bootstrap support, while Celosporium laricicola formed a separate clade but in the Dothideales clade. Coleophoma, Cylindroseptoria, Delphinella, Dothidea, Dothiora, Endoconidioma, Phaeocryptopus, Plowrightia, Pringsheimia, Stylodothis and Rhizosphaera species grouped in one major clade and we name this as Dothideaceae clade as it includes the type of Dothidea (D. sambuci). Other taxa belonging to Aureobasidium, Discosphaerina, Kabatiella, Pseudoseptoria, Saccothecium and Selenophoma grouped in the second major clade. We called that clade Aureobasidiaceae as the type of Aureobasidium A. pullulans (neotypified by Hermanides-Nijhof 1977) clustered here.

Clade A comprises six sub clades with the epitype of Dothidea sambuci (Pers.) Fr. which represents the family Dothideaceae and order Dothideales. In the Dothideaceae clade taxa analyzed cluster in twelve resolved clades. Based on phylogeny, we refer the clade which comprises Dothidea sambuci and D. insculpta Wallr. as Dothidea sensu stricto. The upper resolved clade comprises D. muelleri Loeffler, D. hippophaës (Pass.) Fuckel and D. berberidis De Not. are not Dothidea species in Dothidea sensu stricto and may later require a new genus. The type of Plowrightia, P. ribesia (Pers.) Sacc. clustered in another clade which refers as Plowrightia sensu stricto. Stylodothis puccinioides (the type of Stylodothis) grouped between Dothidea sensu stricto and Plowrightia sensu stricto as a distinct genus. Since the morphology indicates that Dothiora and Sydowia as a distinct genera and the tree is populated with few strains we prefer to maintain Dothiora and Sydowia as distinct genera, although there are no sequences of type strains are available for those genera. The clades Coleophoma, Cylindroseptoria and Endoconidioma, also separate as distinct genera in Dothideaceae with their type strains. Cylindroseptoria pistaciae Quaedvlieg, Verkley & Crous (CBS 471.69) clustered separately in a subclade with 98/100 % bootstrap supports and a new genus is required for this species. Pringsheimia smilacis E. Müll. nested separately with 85/100 % bootstrap supports and it may represent Pringsheimia. However, the type of Pringsheimia should be recollected and epitypified in order to resolve the phylogenetic placement of Pringsheimia in Dothideales. The species grouped in Phaeocryptopus clade (Rhizosphaera kalkhoffii Bubák, R. oudemansii Maubl. and Plowrightia abietis M.E. Barr) along with Phaeocryptopus nudus might be considered to belong in the single genus Phaeocryptopus or could be two distinct genera. Rhizosphaera pini (Corda) Maubl. and Sydowia polyspora (Bref. & Tavel) E. Müll. clustered in the Delphinella clade along with the type of Delphinella, D. strobiligena (Desm.) Sacc. ex E. Müll. & Arx. The tree should be populated with more Sydowia species including the type in order to resolve the placement of Sydowia.

Clade B comprises five subclades with the neotype of Aureobasidium pullulans (de Bary) G. Arnaud which represents the new family Aureobasidiaceae. Sydowia polyspora (Bref. & Tavel) E. Müll. clustered in the Dothideaceae clade, while S. eucalypti (Verwoerd & du Plessis) Crous nested in the Aureobasidiaceae clade. Therefore, a new genus is required for S. eucalypti as it is distinct from the Dothideaceae clade. Columnosphaeria fagi (H.J. Huds.) M.E. Barr, Kabatiella lini and Selenophoma mahoniae grouped in the Aureobasidium clade and they might be considered to belong in Aureobasidium. However, the type sequences of Columnosphaeria and Selenophoma are needed to in order to clarify their placement in Aureobasidiaceae. The Kabatiella clade represents Kabatiella microsticta Bubák, the type of Kabatiella and K. caulivora (Kirchn.) Karak. Although the sequences of type species are not available for Pseudoseptoria and Selenophoma they are considered as distinct genera in Aureobasidiaceae based on available sequence data and morphology.

Taxonomic treatment

Dothideaceae Chevall. [as ‘Dothideae’], Fl. gén. env. Paris (Paris) 1: 446 (1826), Facesoffungi number: FoF00066

Biotrophic, necrotrophic or saprobic on twigs and other parts of plants, rarely on leaves. Sexual state: Ascostromata dark brown to black, immersed to erumpent or superficial, solitary or scattered, usually pulvinate or crustose, globose to subglobose, uni or multiloculate, locules up to 13, non-ostiolate, opening by an apical, usually lysigenous pore or by dehiscence, interascal tissue lacking, ascostromata composed of several layers of light brown to dark brown cells of textura angularis. Peridium of locules generally of several layers of lightly pigmented, dark brown, thick-walled cells of textura angularis. Hamathecium generally lacking pseudoparaphyses, hyaline, when present septate. Asci eight- or poly-spored, bitunicate, fissitunicate, saccate or clavate, short-pedicellate, inner membrane apically thickened, apically rounded with an ocular chamber, asci borne at the base of the locules. Ascospores uni-seriate or bi-seriate, partially overlapping, hyaline or brown, transversely septate, constricted at the primary septum, sometimes muriform, small, wall smooth to verrucose, with or without a sheath. Asexual states: mostly coelomycetous or hyphomycetous species of Coleophoma, Cylindroseptoria, Hormonema, Endoconidioma and Kabatina. Conidiomata pycnidial, stromatic, epidermal to subepidermal, solitary or aggregated, immersed to erumpent, globose to subglobose or flask-shaped or irregular, dark brown to black, uni or multi loculate or convoluted with or without central ostioles. Conidiomata wall composed of several layers of hyaline, brown to dark brown cells of textura angularis. Conidiophores present or reduced to conidiogenous cells, hyaline to pale brown at the base, branched, septate, smooth when present. Conidiogenous cells enteroblastic, phialidic, integrated or discrete, determinate, cylindrical to doliiform, hyaline to brown, smooth-walled, lining the inner cavity. Conidia hyaline, aseptate or one-septate, cylindrical or ellipsoid, smooth-walled, granular or not, guttulate.

Type: Dothidea Fr.

Notes: In this study, we synonymize Dothioraceae under Dothideaceae and accept fifteen genera in Dothideaceae. Recollection, epitypification and multigene molecular analyses are needed for the type and other species of Dictyodothis, Dothiora, Endodothiora, Kabatina, Phaeocryptopus and Pringsheimia in order to confirm the familial status in Dothideaceae as no type species sequences are available.

Key to sexual genera of Dothideaceae

1. Ascostromata superficial, gregarious, globose to globose-depressed, black, developing on conifer needles.................................................. Phaeocryptopus

1. Ascostromata immersed, becoming erumpent, clustered, gregarious, or scattered, pulvinate or crustose, locules subglobose to globose, dark brown to black, developing on twigs, woody branches or leaves............................. 2

2. Ascostromata uni-loculate, loculi usually broad............ 3

2. Ascostromata usually multiloculate, loculi often sphaerical...................................................................... 4

3. Asci with many ascospores............................... Sydowia

3. Asci with 8, multiseptate ascospores, constricted at the primary septum......................................... Pringsheimia

4. Asci with many, 1-septate, hyaline ascospores constricted at the septum........................................... Delphinella

4. Asci with 8 or many, 1-septate or multiseptate or muriform, hyaline or pigmented ascospores constricted at the primary septum................................................... 5

5. Ascospores hyaline to brown, 1-many septate or muriform....................................................................... 6

5. Ascospores hyaline, 1-septate, strongly constricted at the septum.......................................................... Plowrightia

6. Asci with 4–8, ellipsoid to fusiform, brown, 1-septate ascospores..................................................... Stylodothis

6. Asci with 8 or more, brown or hyaline, 1-many septate or muriform ascospores...................................................... 7

7. Asci with 8, 1-many septate or muriform ascospores.................................................................... 8

7. Asci with 8 or more, 1- septate or muriform ascospores.................................................................... 9

8. Ascospores brown, muriform, with longitudinal and transverse septa.......................................... Dictyodothis

8. Ascospores hyaline to brown, 1-septate constricted at the septum.............................................................. Dothidea

9. Asci with many, hyaline, ascospores with 5–7 transverse septa.......................................................... Endodothiora

9. Asci usually with 8, hyaline, one to many septate or muriform, ascospores....................................... Dothiora

Key to asexual genera of Dothideaceae

1. Two types of conidia, endoconidia hyaline, unicellular, blastic conidia mostly two-celled, light to dark brown.................................................... Endoconidioma

1. Only one type of conidia, conidia hyaline, smooth-walled, aseptate............................................................. 2

2. Conidiophores absent, conidiophores reduced to conidiogenous cells...................................................... 3

2. Conidiophores present................................................... 4

3. Conidia solitary, hyaline, smooth-walled, granular or not, cylindrical with obtuse apex, tapering at base to truncate scar......................................... Cylindroseptoria

3. Conidia mostly straight, rarely slightly curved, apex subobtuse, base truncate, guttulate

Neocylindroseptoria

4. Conidiogenous cells enteroblastic, phialidic or percurrent, determinate, brown to pale brown, channel and collarette, periclinal thickening present or absent.................................................................. Kabatina

4. Conidiogenous cells discrete or integrated, determinate, hyaline, phialidic........................................ Coleophoma

Dothidea Fr., Observ. mycol. (Havniae) 2: 347 (1818), Facesoffungi number: FoF00066

Synonyms

Phragmodothis Theiss. & Syd., Annls mycol. 12(2): 179 (1914)

Systremma Theiss. & Syd., Annls mycol. 13(3/4): 330 (1915)

Saprobic on dead wood, stems and twigs. Sexual state: Ascostromata dark brown to black, erumpent through the outer layer of the host tissue, to superficial, solitary or scattered, pulvinate, globose to subglobose, coriaceous, multiloculate, with 3–15 locules, cells of ascostromata composed of several layers of dark brown cells of textura angularis. Locules globose to subglobose, broadly or narrowly conical, thick-walled, with or without ostioles. Peridium of locules comprising 1–2 layers of thick-walled, lightly pigmented, dark brown to black cells of textura angularis. Hamathecium lacking pseudoparaphyses. Asci eight-spored, bitunicate, fissitunicate, clavate to sub-cylindrical, with a short broad pedicel, thickened and rounded at apex, with a clear ocular chamber. Ascospores uniseriate to biseriate, partially overlapping, hyaline, sometimes brown, 1-septate, constricted at the septum, ellipsoidal or oblong to obovoid with broadly rounded ends, smooth-walled, thick-walled, with or without a sheath. Asexual state: See notes.

Notes: Dothidea was introduced by Fries (1818) and later Fuckel (1869) assigned Dothidea as the type genus of Dothideaceae with D. gibberulosa (Ach.) Fr. as the type species. Theissen and Sydow (1915) introduced Systremma to accommodate Dothidea typified by D. sambuci Pers., while D. ribesia and D. berberidis were transferred to Dothidella (Shear 1936) which was in the subfamily Dothideae. Theissen and Sydow (1915) chose the oldest species, Sphaeria natans Tode, as the basionym and cited the later Dothidea sambuci as a synonym (Shoemaker and Hambleton 2005). Clements and Shear (1931) changed the type genus and assigned D. sambuci as the type species, in accordance with the recommendation of the Cambridge revision of the International Code (Blain 1927; Orton 1924; Shear 1936). Shoemaker et al. (2003) proposed that the type for Dothidea be formally conserved as Sphaeria sambuci Pers. (Dothidea sambuci (Pers.) Fr.:Fr.) and an epitype specimen was established by Shoemaker and Hambleton (2005). D. sambuci, D. hippophaës and D. insculpta Wallr., form a highly supported monophyletic group, and cluster with high support with other genera classified in Dothideaceae and Dothioraceae in one of the three lineages of dothideomycetous taxa (Hambleton et al. 2003; Lutzoni et al. 2004; Shoemaker and Hambleton 2005). Dothidea presently comprises 499 epithets in Index Fungorum (2014).

In our phylogenetic tree, the epitype of Dothidea sambuci (Pers.) Fr. clustered in the Dothideaceae clade with 99/100 % bootstrap support along with D. insculpta and we refer this clade as Dothidea sensu stricto. D. muelleri, D. hippophaës and D. berberidis are not Dothidea species in Dothidea sensu stricto and may later require a new genus.

Type species: Dothidea sambuci (Pers.) Fr., Syst. mycol. (Lundae) 2(2): 551 (1823), (Fig. 2), Facesoffungi number: FoF00067

Fig. 2

Dothidea sambuci (GZU 78–2002, epitype). a, b Ascostromata on the host substrate. c, d Section through ascostroma showing the arrangement of locules. e Close up of the locules. f, g Asci in cotton blue reagent. Bearing eight ascospores. h–k Ascospores in cotton blue reagent. Scale bars: b = 1000 μm, c, d = 500 μm, e = 100 μm, f–k = 10 μm

≡ Sphaeria sambuci Pers., Syn. meth. fung. (Göttingen) 1: 14 (1801)

For other synonyms see Index Fungorum

Saprobic on dead stems. Sexual state: Ascostromata 700–1000 × 200–310 μm $\left(\overline{x}=900\times 260\mu m,n=5\right)$, black, erumpent through the outer layer of the host tissue, to near superficial, solitary or scattered, globose to subglobose, coriaceous, multiloculate, with 8–15 locules, cells of ascostromata composed of several layers of dark brown cells of textura angularis. Locules 80–120 × 65–90 μm $\left(\overline{x}=95\times 70\mu m,n=20\right)$, globose to subglobose, broadly or narrowly conical, with individual ostioles. Ostiole usually widely porate, with well-developed neck, ostiolar canal filled with a tissue of hyaline cells. Peridium of locules 28–41 μm $\left(\overline{x}=35\mu m,n=15\right)$ comprising 1–2 layers of thick-walled, lightly pigmented, small cells of textura angularis. Hamathecium lacking pseudoparaphyses. Asci 70–80 × 12–15 μm $\left(\overline{x}=73.5\times 13.5\mu m,n=10\right)$, eight-spored, bitunicate, fissitunicate, clavate to sub-cylindrical, thickened with a short broad pedicel and rounded at apex, with a clear ocular chamber. Ascospores 17–20 × 5–6.5 μm $\left(\overline{x}=17.5\times 5.5\mu m,n=20\right)$ uniseriate, partially overlapping, hyaline to light brown when immature, becoming brown to chestnut brown when mature, 1-septate, constricted at the septum, ellipsoidal with broadly rounded ends, smooth-walled, thick-walled, lacking a sheath. Asexual state: Unknown.

Material examined: AUSTRIA, Steiermark (Styria) Grazer Bergland, on Sambucus nigra L. (Adoxaceae), leg D. Baloch 4 October 2002 det. C. Scheuer (GZU 78–2002, epitype).

Notes: Shoemaker and Hambleton (2005) established an epitype specimen from Austria on Sambucus nigra (Adoxaceae). There is no evidence of formation of an asexual state in the culture of the epitype.

Dothidea insculpta Wallr., Fl. crypt. Germ. (Norimbergae) 2: 864 (1833), (Fig. 3), MycoBank: MB 173197; Facesoffungi number: FoF00068

Fig. 3

Dothidea insculpta (MFLU 14–0156) a Appearance of ascostromata on host. b Vertical section through ascostroma with asci. c Section of peridium. d Immature ascus. e–h Mature asci. i Immature ascospore. j–n Mature ascospores. o Colony on PDA medium. (upper) q Colony on PDA medium. (lower) r Germinating ascospore. Scale bars: a, b = 200 μm, c–h = 20 μm. i–n = 10 μm

≡ Plowrightia insculpta (Wallr.) Sacc., Syll. fung. (Abellini) 2: 636 (1883)

≡ Plowrightia insculpta (Wallr.) Sacc., Syll. fung. (Abellini) 2: 636 (1883)

≡ Scirrhia insculpta (Wallr.) M.E. Barr, Contr. Univ. Mich. Herb. 9(8): 565 (1972)

Saprobic on a dead branch of Clematis vitalba. Sexual state: Ascostromata 92–192 μm high × 63–198 μm diam $\left(\overline{x}=144\times 134\mu m,n=7\right)$, dark brown to black, superficial on host tissue, solitary, scattered, globose to subglobose, coriaceous, multiloculate, with 2–4 locules, cells of ascostromata composed of dark brown-walled cells of textura angularis. Locules 91–125 × 109–144 μm $\left(\overline{x}=87\times 114\mu m,n=10\right)$, globose to subglobose, non-ostiolate. Peridium of locules 27–63 μm, comprising few layers of brown to dark brown cells of textura angularis. Hamathecium lacking pseudoparaphyses. Asci 53–79 × 8–14 μm $\left(\ \overline{x} = 65\times 11\mu m,n=10\right)$, eight-spored, bitunicate, fissitunicate, cylindrical to oblong, with a short pedicel, rounded at the apex with an ocular chamber. Ascospores 11–23 × 5–7 μm $\left(\overline{x}=16\times 6\mu m,n=20\right)$, overlapping uni to biseriate, broadly ovoid or ellipsoid, hyaline, 1-septate, constricted at the septum, upper cell broader than the lower cell, smooth. Asexual state: See notes.

Cultural characteristics: Ascospores germinating on MEA or PDA within 36–48 h. Colonies growing on MEA or PAD, reaching 3–5 cm in 5 days at 18 − 20 °C, rhizoid colonies, flat, rough surface, rhizoid edge, opaque opacity, dull green mycelium, dark green around the colonies, produce some odor and clearing effect on media.

Material examined: ITALY, Poggio alla Portacce-Pratomagno (Province of Arezzo [AR]), on a dead branch of Clematis vitalba L. (Ranunculaceae), 30 June 2013, Erio Camporesi (MFLU 14–0156), living culture MFUCC 13–0686.

Notes: We could not locate the type specimen of Dothidea insculpta and our specimen can be considered as an authentic specimen. Hess and Müller (1951) introduced a monotypic genus Asteromellopsis and described Asteromellopsis insculpta H.E. Hess & E. Müll. as the asexual state found in young ascomata of Dothidea insculpta found in nature. Luttrell (1951a) found a microconidial state in young stromata of Dothidea collecta (Eriksson 1981; Shoemaker and Hambleton 2005). We did not observe formation of an asexual state in the culture.

Coleophoma Höhn., Sber. Akad. Wiss. Wien, Math.-naturw. Kl., Abt. 1 116: 637 (1907), Facesoffungi number: FoF00069

Type species: Coleophoma crateriformis (Durieu & Mont.) Höhn., Mitt. bot. Inst. tech. Hochsch. Wien 2(3): 77 (1925)

≡ Ascospora crateriformis Durieu & Mont., Flora Algéricae 1: 590 (1849) [1846–49]

≡ Macrophoma crateriformis (Durieu & Mont.) Berl. & Voglino, Atti Soc. Veneto-Trent. Sci. Nat. 10(1): 194 (1886)

≡ Macrophoma crateriformis (Durieu & Mont.) Berl. & Voglino, Atti Soc. Veneto-Trent. Sci. Nat. 10(1): 194 (1886) f. crateriformis

≡ Macrophoma crateriformis f. macrospora D. Sacc.

≡ Phoma crateriformis (Durieu & Mont.) Sacc., Michelia 2(no. 6): 90 (1880)

≡ Septoria crateriformis (Durieu & Mont.) Sacc., Syll. fung. (Abellini) 3: 496 (1884)

Notes: Coleophoma was introduced by Höhnel (1907) and species of Coleophoma are parasitic, saprobic or endophytic on plants in terrestrial habitats. This genus is characterized by pycnidial conidiomata with well-developed lower and usually poorly developed upper walls; hyaline, septate, collapsed paraphyses among the conidiophores; discrete or integrated, determinate, phialidic conidiogenous cells and hyaline, aseptate, smooth-walled, cylindrical or ellipsoid conidia (Wu et al. 1996; Sutton 1980; Duan et al. 2007). Wijayawardene et al. (2012) placed Coleophoma in Ascomycota, genera incertae sedis. In the phylogenetic analysis of Gruyter et al. (2009), a putative strain of Coleophoma crateriformis clustered in Dothideales, while C. maculans grouped in Helotiales. C. crateriformis is closely related with C. oleae and Duan et al. (2007) reassigned C. oleae to the genus Coleonaema based on conidiomatal development. In our phylogenetic study, C. crateriformis the type of Coleophoma clustered with Coleophoma oleae in one of the subclades in the Dothideaceae clade. Therefore, we accept Coleophoma as an asexual genus in Dothideaceae.

Cylindroseptoria Quaedvlieg et al., Stud. Mycol. 75: 358 (2013)

Type species: Cylindroseptoria ceratoniae Quaedvlieg et al., Stud. Mycol. 75: 358 (2013)

Notes: Cylindroseptoria was introduced by Quaedvlieg et al. (2013) and typified with C. ceratoniae. Cylindroseptoria ceratoniae is characterized by separate, brown, cupulate, short-stipitate conidiomata; a rim with elongated brown, thick-walled cells with obtuse ends, 3–4 wall layers of medium brown cells of textura angularis, becoming hyaline towards inner region, hyaline, smooth, ampulliform conidiogenous cells with prominent periclinal thickening and solitary, hyaline, smooth, granular or not, aseptate, cylindrical conidia with obtuse apex. Quaedvlieg et al. (2013) tentatively placed a new species C. pistaciae in Cylindroseptoria, as it has pycnidial rather than cupulate conidiomata. Multigene phylogenetic analysis of Quaedvlieg et al. (2013) showed that Cylindroseptoria belongs to Dothideaceae. However, the two species clustered in separate clades in the family Dothideaceae in our phylogenetic study. Cylindroseptoria ceratoniae grouped as sister clade to the Coleophoma while, C. pistaciae clustered separately with 98/ 100 % bootstrap support in Dothideaceae. Therefore, we introduce a new genus, Neocylindroseptoria below, for C. pistaciae.

Delphinella (Sacc.) Kuntze, Revis. gen. pl. (Leipzig) 3(2): 74 (1898), Facesoffungi number: FoF00074

Synonyms

Diplosphaerella Grove, J. Bot., Lond. 50: 91 (1912)

Glonium subgen. Delphinella Sacc., Syll. fung. (Abellini) 9: 1103 (1891)

Hariotia P. Karst., J. Bot., Paris 3: 206 (1889)

Pleoglonis Clem., Gen. fung. (Minneapolis): 56, 173 (1909)

Rehmiellopsis Bubák & Kabát, in Bubák, Naturwiss. Z. Forst − Landw. 8: 320 (1910)

Saprobic or parasitic on twigs, wood, leaves and cone scales of gymnosperms and woody dicotyledons (Barr 1972). Sexual state: Ascostromata dark brown to black, immersed and becoming erumpent, solitary or gregarious, globose to subglobose, multiloculate, thick-walled. Locules globose to subglobose, lacking ostioles. Peridium of locules relatively thick, comprising 1–2-layers, lightly pigmented of cells of textura angularis. Hamathecium lacking pseudoparaphyses. Asci polysporous, bitunicate, fissitunicate, oblong, clavate or cylindrical, with a short pedicel, apically rounded, ocular chamber absent, asci borne at the base of the loculus. Ascospores overlapping 2 − 3-seriate to crowded, transversely 1-septate, hyaline or yellowish, with rounded apex, obtuse or pointed at the base, constricted at the septum, smooth-walled. Asexual state: See notes.

Notes: Delphinella was introduced by Kuntze (1898) based on Sphaeria strobiligena. Müller and von Arx (1962) assigned Delphinella strobiligena as the type species, and transferred D. abietis (O. Rostr.) E. Müll., D. balsameae (Waterman) E. Müll., D. cookie (Linds.) E. Müll., D. deviata (Petr.) E. Müll. and D. polyspora (Johanson) E. Müll. Barr (1972) introduced D. tsugae (House) M.E. Barr, while Barr et al. (1986) added D. peckii (Lindau) M.E. Barr which had been previously been referred to Sphaerella and Mycosphaerella. The locules of D. polyspora are smaller and more conic than the other species of the genus (Barr 1972). Von Arx and Müller (1975) included the genus Delphinella under Dothideaceae. Hawksworth (1979) synonymized D. cookei under Muellerella lichenicola (Sommerf.) D. Hawksw. and placed it in Verrucariaceae based on its polysporous asci. However, the genus Delphinella should be placed under Dothideaceae, Dothideomycetes (Barr 2001; Hyde et al. 2013).

Barr (1972) described the asexual state of Delphinella abietis as Dothiorella (Phoma bohemica Bubák and Kabát) which is characterized by thick-walled pycnidia with aspects similar to that of ascostromata and hyaline, fusoid, one-celled conidia. However, modern taxonomic and molecular data has shown the Dothiorella belongs in Botryosphaeriaceae (Liu et al. 2012).

In the phylogeny (Fig. 1) a putative strain of Delphinella strobiligena (CBS 735.71) clustered in Dothideaceae clade close to a putative strain of Sydowia polyspora (CBS 116.29). Considering the close relationship of the two strains it may be that one of these two strains is wrongly identified or these two species should be in one genus. However, we named this clade as Delphinella which comprises D. strobiligena, Rhizosphaera pini and S. polyspora pending on more fresh collections of Delphinella and Sydowia species.

Type species: Delphinella strobiligena (Desm.) Sacc. ex E. Müll. & Arx, in Müller & von Arx, Beitr. Kryptfl. Schweiz 11(no. 2): 25 (1962), (Fig. 4), Facesoffungi number: FoF00075

Fig. 4

Delphinella strobiligena (PC 0084688, holotype). a–c Ascostromata on host surface. d Section through ascostromata showing locules. e Ascus arrangement in locules. f Cells of ascostroma and peridium of locule. c–h Asci. i Ascus stained in cotton blue reagent. j–k Ascospores. l Ascospores stained in cotton blue reagent. Scale bars: d = 100 μm, e, g–i = 50 μm, f = 20 μm, j–l = 10 μm

≡ Didymella strobiligena (Desm.) Sacc., Syll. fung. (Abellini) 1: 552 (1882)

≡ Glonium strobiligenum (Desm.) Mouton, Bull. Soc. R. Bot. Belg. 28(C.R.): 79 (1889)

≡ Hariotia strobiligena (Desm.) P. Karst., J. Bot., Paris 3: 206 (1889)

≡ Pleoglonis strobiligena (Desm.) Clem., Gen. fung. (Minneapolis): 1–227 (1909)

≡ Sphaeria strobiligena Desm., Annls Sci. Nat., Bot., sér. 3 6: 75 (1846)

Parasitic on twigs, wood, needles and cone scales of conifer plants. Sexual state: Ascostromata 220–310 μm high × 290–330 μm diam $\left(\overline{x}=295\times 320\mu m,n=5\right)$, black, initially immersed and becoming erumpent at maturity on host surface, scattered, subglobose, carbonaceous, 2 − 3-loculate, cells of ascostromata of brown-walled textura angularis. Locules 140–209 μm high × 162–191 μm diam $\left(\overline{x}=180\times 170\mu m,n=5\right)$, globose to subglobose, lacking ostioles. Peridium of locules 22–80 μm $\left(\overline{x}=46\mu m,n=10\right)$, thin-walled, composed of a single layer of light brown to hyaline cells of textura angularis. Hamathecium lacking pseudoparaphyses. Asci 79–87 × 9–11 μm $\left(\overline{x}=82\times 10\mu m,n=7\right)$, poly-sporous, bitunicate, fissitunicate, oblong, clavate or cylindric-clavate with a short pedicel, rounded at the apex. Ascospores 8–10 × 3–5 μm $\left(\overline{x}=9\times 4\mu m,n=15\right)$, overlapping 1 − 3-seriate, hyaline, 1-septate, ellipsoid to fusiform, upper cell rounded and wider than lower cell, slightly constricted at the septum, smooth-walled, surrounded by a thin sheath. Asexual state: Unknown.

Material examined: FRANCE, on strobili of Pinus sp., Desmaziere (PC 0084688, holotype).

Dictyodothis Theiss. & Syd., Annls mycol. 13(3/4): 346 (1915), Facesoffungi number: FoF00076

Saprobic on dead twigs, stems and branches of land plants. Sexual state: Ascostromata black, immersed, erumpent at maturity, aggregated or in clusters, scattered, discoid to pulvinate, globose to subglobose, coriaceous, multiloculate, with 8–10 locules, cells of ascostromata composed of several layers of dark brown to black cells of textura prismatica and textura angularis. Locules globose to subglobose, ostiolate. Hamathecium lacking pseudoparaphyses. Peridium of locules comprising light brown to brown cells of textura angularis. Hamathecium lacking pseudoparaphyses. Asci eight-spored, bitunicate, clavate to sub-clavate or cylindrical, pedicellate, thickened with a short broad pedicel and rounded at apex, with a clear ocular chamber. Ascospores overlapping uniseriate to biseriate, yellowish brown to dark brown, muriform, with 3–6 transverse septa and 1–5 longitudinal septa, constricted at the primary septum, oblong, ovoid to fusoid, part above the central septum wider. Asexual state: Unknown.

Notes: Dictyodothis was introduced by Theissen and Sydow (1915) to accommodate Dictyodothis berberidis (Rehm) Theiss. & Syd. and Dictyodothis excavata (Cooke & Ellis) Theiss. & Syd. under the family Dothideaceae. As the type genus they assigned Dictyodothis berberidis which has been referred to Curreya. Tilak and Kale (1969) introduced two species with paraphyses, Dictyodothis acaciae Tilak, S.B. Kale & S.V.S. Kale and D. grewiae Tilak, S.B. Kale & S.V.S. Kale,. Later Von Arx and Müller (1975) included this genus in the family Pleosporaceae based on “paraphysoids” in the locules. Barr (1981) showed that “paraphysoids” are the walls and strands of cytoplasmic remnants of discharged asci and ascospores are similar with those of Dothidea sambuci. Therefore, Barr (1981) included Dictyodothis again in the Dothideaceae. Barr (1987a), Lumbsch and Huhndorf (2010) also listed Dictyodothis under the Dothideaceae. No asexual states have been reported for this genus. Currently there are around eight species epithets listed in Index Fungorum (2014).

Because of the erumpent, multiloculate ascostromata this genus should be included in Dothideaceae. However, Dictyodothis is distinct in having yellowish-brown, muriform ascospores. No sequence data is available for this species.

Type species: Dictyodothis berberidis (Rehm) Theiss. & Syd., Annls mycol. 13(3/4): 346 (1915), (Fig. 5), Facesoffungi number: FoF00077

Fig. 5

Dictyodothis berberidis (holotype). a. Ascostromata on host surface. b. Close up of ascostroma c. Section through ascostroma. d. Close up of the cells of ascostroma. e–g. Asci with short, broad pedicel bearing eight ascospores. h–k Reddish brown to dark yellowish brown, muriform ascospores. Scale bars: c = 500 μm, d = 100 μm, e–g = 20 μm, h–k = 5 μm

≡ Curreya berberidis Rehm, Bih. K. svenska Vetensk Akad. Handl., Afd. 3 25(no. 6): 4 (1899)

Saprobic on branches of Berberis buxifolia. Sexual state: Ascostromata 1000–1500 × 150–180 μm $\left(\overline{x}=1100\times 150\mu m,n=20\right)$, black, superficial, semi-immersed to erumpent, solitary or scattered, subglobose to broadly ellipsoid, coriaceous, multiloculate, with 8–10 locules, cells of ascostromata dark brown to black cells of textura angularis, sometimes ostiolate. Peridium of locules 20–40 $\left(\overline{x}=35\mu m,n=20\right)$ comprising lightly pigmented to light brown cells of textura angularis. Hamathecium lacking pseudoparaphyses. Asci 50–60 × 14–20 μm $\left(\overline{x}=55\times 17\mu m,n=20\right)$ eight-spored, bitunicate, fissitunicate, clavate to broadly-clavate, with a short, broad pedicel, thickened and rounded at apex with an ocular chamber. Ascospores 15–20 × 6–10 μm $\left(\overline{x}=16\times 8\mu m,n=40\right)$, uniseriate or discontinuously arranged, partially overlapping, reddish brown to dark yellowish brown, muriform with three transverse septa and 1–2 vertical septum in the central cells when mature, constricted at the septa, oblong, smooth to verruculose, without a sheath. Asexual state: Unknown.

Material examined: SOUTH AMERICA, Pantagonia, Rio Argopardo, on dead wood of Berberis buxifolia Lam. (Berberidaceae), 4 March 1896, Dunsen (W, holotype).

Dothiora Fr., Summa veg. Scand., Section Post. (Stockholm): 418 (1849), Facesoffungi number: FoF00078

Synonyms

Dothiora subgen. Metadothis Sacc., Syll. fung. (Abellini) 8: 766 (1889)

Jaapia Kirschst., Krypt. − Fl. Brandenburg (Leipzig) 7(3): 444 (1938)

Keisslerina Petr., Annls mycol. 17(2/6): 74 (1920) [1919]

Leptodothiora Höhn., Ber. dt. bot. Ges. 36: 311 (1918)

Metadothis (Sacc.) Sacc., Syll. fung. (Abellini) 10: 857 (1892)

Stigmea Bonord., Abh. naturforsch. Ges. Halle 8: 79 (1864)

Saprobic or parasitic on leaves, branches or twigs in terrestrial habitatas. Sexual state: Ascostromata black, immersed to erumpent, pulvinate to depressed globose, multiloculate, thick-walled, cells of ascostromata composed of lightly pigmented or dark brown cells of textura angularis. Locules globose to subglobose, broadly rounded to short papillate, apex opening by an irregular, small pore. Peridium of locules composed of several layers of thick-walled dark brown cells of textura angularis. Hamathecium lacking pseudoparaphyses. Asci 8 or more spored, bitunicate, fissitunicate, oblong to clavate, pedicellate, with a small ocular chamber. Ascospores overlapping biseriate to crowded, one to many septate, usually constricted at the primary median septum, sometimes with a vertical septum in one or several of the central cells and rarely in the end cells, hyaline, rarely yellow to pale brown, obovate to elliptic or fusoid, often inequilateral or slightly curved, smooth, occasionally surrounded by a thin mucilaginous sheath. Asexual states: Dothichiza sp.: Pycnidia frequently found on host and produced in culture conidial stroma similar to ascostromata. Conidiogenous cells lining the cavity of the pycnidium. Conidia avoid to oblong, hyaline, one celled formed singly on phialidic, obpyriform, simple, smooth-walled. (asexual morph description follows D. europaea) (Eriksson 1981; Sivanesan 1984).

Notes: Dothiora was introduced by Fries (1849) with D. pyrenophora (Fr.) Fr. as the type species. Saccardo (1889) and Lindau (1897) included the genus in the Discomycetes, while Theissen and Sydow (1915) placed it in Dothideales. Theissen and Sydow (1917) moved the genus to the new family Dothioraceae under the order Myringiales. Clements and Shear (1931) placed it both in the Phacidiacede and Myriangiaceae (Miller and Burton 1943). Froidevaux (1972) placed Dothiora in Dothioraceae along with four other genera (Table 3) and assigned D. sorbi (Dothiora pyrenophora synonymized with D. sorbi) as the type species. Barr (1972) listed eleven species with, D. pyrenophora as type species, while Froidevaux (1972) accepted 14 species. Von Arx and Müller (1975) and Sivanesan (1984) treated Dothiora under the family Dothideaceae as they synonymized Dothideaceae and Dothioraceae. Barr (1987a), Hawksworth et al. (1995) and Lumbsch and Huhndorf (2010) however, categorized Dothiora under Dothioraceae as they treated Dothideaceae and Dothioraceae as separate two families in Dothideales.

Table 3 Accepted genera according to various treatments of Dothioraceae

Table 4 Taxa used in the phylogenetic analysis and their corresponding GenBank accession numbers. Newly deposited sequences are shown in bold

The asexual state of Dothiora pyrenophora has been reported as Dothichiza sorbi Lib. by Sivanesan (1984) as pycnidia formed in culture and abundantly on the host. The type species of Dothichiza is D. populea Sacc. & Briard, which causes cankers of Populus sp. (Hedgcock and Hunt 1916; Waterman 1957). Dothiora populea forms multiloculate pustules on the surface of Populus sp., but have not been linked to a sexual state. Although some species of Dothichiza have been linked to Dothideales via molecular data (Bills et al. 2004; Zalar et al. 2008), it has not been established that Dothiora pyrenophora and Dothichiza populea are related.

Type species: Dothiora pyrenophora (Fr.) Fr., Summa veg. Scand., Section Post. (Stockholm): 418 (1849), (Fig. 6), Facesoffungi number: FoF00079

Fig. 6

Dothiora pyrenophora (BPI 674269). a Material label. b Ascostromata on host surface. c Cross section through ascostroma and peridium. d Peridial wall. e, f Immature and mature asci. g–i Muriform ascospores. j Conidia associated with base of ascostromata. Scale bars: b, c = 200 μm, d, g–j = 20 μm, e, f = 50 μm

Saprobic on branches of Sorbus sp. Sexual state: Ascostromata (639−) 650 − 756.5 μm diam, black, immersed to erumpent, breaking through the host surface through angular splits, multiloculate, cells of ascostromata composed dark brown to black cells of textura angularis. Locules 140–172 μm high × 152–180 μm diam., subglobose or obpyriform, arranged at periphery of stroma in a single layer, widest at the base, short papillate, with small ostiole pores. Peridium of locules 78–88 μm thick, composed cell layers of textura prismatica and textura angularis, thick-walled and black at the outer wall. Hamathecium lacking pseudoparaphyses. Asci (85−)93 − 108(−112) × (13−)16 − 20(−23) μm $\left(\overline{x}=99\times 18\mu m,n=15\right)$, eight-spored, bitunicate, fissitunicate, oblong to subclavate, saccate, short-pedicellate, with a distinct ocular chamber, ca. 2(−3) μm wide. Ascospores (28−)29 − 36(−40) × 8 − 10(−11) μm $\left(\overline{x}=32\times 10\mu m,n=10\right)$, 2 − 3-seritate overlapping in the ascus, sometime irregularly arranged, hyaline or pale grayish, muriform, 7 − transverse septate, with 1 − longitudinal septum in upper part, deeply constricted at middle septum, cylindrical to fusiform, pointed at both ends, smooth. Asexual states: Unknown.

Material examined: SWITZERLAND, Graubuenden, Davos, Dischmatal, on Sorbus aucuparia L. (Rosaceae), May 26 1964, E. Müller (BPI 674269).

Notes: Conidiomata with numerous, cylindrical, hyaline conidia were found at the base of the ascostromata (Fig. 6). It is not clear if these are the asexual state, another associated taxon or a fungicolous taxon. We could not loan the type so illustrate a specimen from BPI.

Endoconidioma Tsuneda et al., Mycologia 96(5): 1129 (2004), Facesoffungi number: FoF00080

Type species: Endoconidioma populi Tsuneda et al., Mycologia 96(5): 1129 (2004)

Notes: Endoconidioma was introduced by Tsuneda et al. (2004) as a monotypic genus in order to accommodate E. populi in Dothideaceae. Endoconidiogenesis in E. populi is similar to the endoconidial hyphomycete Phaeotheca and no coelomycetous taxa have been reported to produce endoconidia. Morphological characters and DNA sequence data showed that Endoconidioma is distinct from the previously established endoconidial genera (Tsuneda et al. 2004). This genus is characterized by subglobose to flask-shaped, entirely closed conidiomata, forming on a black subiculum, a darkly pigmented peridium and locules filled with conidiogenous cells. Endoconidia are formed endogenously and hyaline, unicellular and released by dissolution of the conidiogenous and the peridial cells of the conidioma. Blastic conidia, light to dark brown, mostly two-celled, produced holoblastically from pigmented, undifferentiated hyphae (Tsuneda et al. 2004). In our phylogenetic analysis (Fig. 1) E. populi is clustered in Dothideaceae with 65/100 % bootstrap support.

Endodothiora Petr., Annls mycol. 27(5/6): 345 (1929), Facesoffungi number: FoF00082

Parasitic on Dothidea puccinioides. Sexual state: Ascostromata black, immersed, becoming erumpent, solitary or gregarious, subglobose to broadly ellipsoid, coriaceous, multiloculate, with 2–5 locules, cells of ascostromata composed of dark brown to black cells of textura angularis. Locules globose to subglobose, without ostioles. Peridium of locules thin-walled, composed of light brown to hyaline cells of textura angularis. Asci 20 − 26-spored, bitunicate, cylindrical to broadly cylindrical with a short pedicel, rounded at the apex. Ascospores overlapping, crowded, hyaline, oblong, 5–7 septate, constricted at the primary septum, smooth-walled. Asexual state: Unknown.

Notes: Endodothiora was introduced by Petrak (1929) to accommodate a single species E. sydowiana Petr., which is immersed in ascostromata of Dothidea puccinioides (DC.) Fr., Syst. (Barr 1972). Polyspored asci, multiseptate hyaline ascospores of Endodothiora share similar characteristics with those of Sydowia, but immersed parasitic habitat excludes it from Sydowia (Barr 1972, 2001). Luttrell (1973) placed this genus in Dothioraceae based on it being parasitic on Dothidea collecta (Schwein.) Ellis & Everh., and immersed in the stroma of the host fungus. We re-examined the type and we found different asci with brown ascospores (Dothidea puccinioides) which share the same ascomata that is good proof for E. sydowiana being parasitic on the Dothidea puccinioides. No sequence data is available for this monotypic genus.

Type species: Endodothiora sydowiana Petr., Annls mycol. 27(5/6): 345 (1929), (Fig. 7), Facesoffungi number: FoF00083

Fig. 7

Endodothiora sydowiana (W 12058, holotype). a, b Ascostromata of E. sydowiana. c, d Vertical section through ascostromata illustrating the structure. e Vertical section of ascostromata showing the loculi bearing the asci inside. f Asci. g Asci in cotton blue reagent. h, i Ascospores in cotton blue reagent. Scale bar: b = 500 μm. c–e = 100 μm. f–i = 10 μm

Parasitic on Dothidea puccinioides. Sexual state: Ascostromata 250–310 μm high × 200–600 μm diam., gregarious, black, immersed, becoming erumpent solitary or scattered, subglobose to broadly ellipsoid, coriaceous, multiloculate, with 2–5 locules, cells of ascostromata dark brown to black cells of textura angularis. Locules 120–140 μm × 110–120 μm $\left(\overline{x}=137\times 114\mu m,n=5\right)$ globose to subglobose, without ostioles. Peridium of locules 20–28 μm $\left(\overline{x}=23\mu m,n=5\right)$, thin-walled, composed of light brown to hyaline cells of textura angularis. Asci 90–115 μm × 21–35 μm $\left(\overline{x}=96\times 32\mu m,n=10\right)$, 20 − 26 − spored, bitunicate, cylindrical to broadly cylindrical with a short pedicel, rounded at the apex. Ascospores 16–30 × 6–9 μm $\left(\overline{x}=19\times 7\mu m,n=20\right)$, overlapping, crowded, hyaline, oblong, hyaline, 5–7 septate, constricted at the primary septum, smooth-walled. Asexual state: Unknown.

Material examined: ABKHAZIA, on dead stem of Buxus sempervirens L. (Buxaceae), 27 March 1912, G. Woronow (W 12058, holotype).

Kabatina R. Schneid. & Arx, Phytopath. Z. 57: 179 (1966), Facesoffungi number: FoF00084

Type species: Kabatina thujae R. Schneid. & Arx, Phytopath. Z. 57: 180 (1966)

Parasitic on branches of Abies, Juniperus, Mahonia, Populus and Thuja. Sexual state: Unknown. Asexual state: Mycelium immersed, branched, septate, hyaline to pale brown or black. Conidiomata dark brown to black, sporodochial or acervular, epidermal to subepidermal, pulvinate, pale to dark brown, partially immersed inside the host, composed of branched, septate hyphae, amphigenous or hypophyllous, thick-walled textura angularis. Dehiscence by irregular rupture of the cuticle and epidermis. Conidiophores stromatic or branched, clustered together resembling a synnema, septate, hyaline to pale brown, smooth. Conidiogenous cells enteroblastic, phialidic or percurrent, determinate, cylindrical to doliiform or subclavate, brown to pale brown, channel and collarette minute, periclinal thickening present or absent, borne terminally and intercalary. Conidia hyaline, aseptate, cylindrical to ellipsoid, terminal in basipetal chains or singly, conidial secession schizolytic, smooth (Sutton 1980; Butin and Pehl 1993; Seifert et al. 2011).

Notes: Kabatina was introduced by Schneider and von Arx (1966) to accommodate K. juniperi R. Schneid. & Arx, and K. thujae R. Schneid. & Arx,. Butin and Schneider (1976), Ramaley (1992) and Butin and Pehl (1993) added K. populi Butin & R. Schneid., K. mahoniae A.W. Ramaley and K. abietis Butin & Pehl, respectively. This genus was shown to group as a sister clade with Dothidea hippophaës and D. insculpta Wallr. by Tsuneda et al. (2004). Bills et al. (2004) also showed that the phylogenetic placement of K. thujae and K. juniperi (CBS 239.66, CBS 466.66) in Dothideales with Hormonea species based on the phylogenetic analysis of ITS1-5.8S-ITS2 rDNA (ITS) data. Wijayawardene et al. (2012) placed this genus under Dothioraceae, while Hyde et al. (2013) categorized Kabatina as an asexual genus in the family Dothideaceae. We also accept Kabatina as an asexual genus in Dothideaceae considering above facts. Although, Kabatina presently comprises nine epithets (Index Fungorum 2014), only ITS sequence data are available in GenBank therefore, we did not include the Kabatina species in to our phylogenetic tree. Species of Kabatina need to be sequenced for protein-coding genes and nuclear ribosomal genes in order to obtain a better resolution.

Kabatina species cause several desease known as “Evergreen Disease, Needle Cast of Firs” and economic losses in Abies, Juniperus, Mahonia and Populus and is associated with needle diebacks in conifers (Sutton 1980; Tisserat and Pair 1997; Bills et al. 2004; Cech et al. 2009).

Neocylindroseptoria K. M. Thambugala & K. D. Hyde, gen. nov., Index Fungorum number: IF  550730

Etymology: The generic epithet, neo (Lat., new), refers to the similarity to Cylindroseptoria.

Type species: Neocylindroseptoria pistaciae (Quaedvlieg, Verkley & Crous) K. M. Thambugala & K. D. Hyde, com. nov., Index Fungorum number: IF 550731≡ Cylindroseptoria pistaciae Quaedvlieg, Verkley & Crous, Stud. Mycol. 75: 359 (2013) 

Generic description

Asexual state: Conidiomata pycnidial, globose, black, erumpent, separate, with black crusty outer layer of cells, with central ostiole; wall of 3–6 layers of brown cells of textura angularis. Conidiophores reduced to conidiogenous cells. Conidiogenous cells phialidic (mostly monophialidic, but a few observed to also be polyphialidic), lining the inner cavity, hyaline, smooth, ampulliform, proliferating percurrently (inconspicuous) or with periclinal thickening at apex (also occurring as solitary loci on superficial hyphae surrounding pycnidia). Conidia cylindrical, mostly straight, rarely slightly curved, hyaline, smooth-walled, apex subobtuse, base truncate, guttulate, aseptate (Quaedvlieg et al. 2013). Sexual state: Unknown.

Notes: Cylindroseptoria pistaciae was introduced by Quaedvlieg et al. (2013) and tentatively placed in Cylindroseptoria as it has pycnidial rather than cupulate conidiomata. Cylindroseptoria pistaciae separates from C. ceratoniae and formed a strong clade in Dothideaceae with 98/100 % bootstrap support. Therefore, we introduce Neocylindroseptoria to accommodate C. pistaciae.

Phaeocryptopus Naumov, Bull. Soc. mycol. Fr. 30(1): 424 (1915), Facesoffungi number: FoF00086

Synonyms

Adelopus Theiss., in Theissen & Sydow, Annls mycol. 15(6): 482 (1918) [1917]

Cryptopus Theiss., Annls mycol. 12(1): 72 (1914)

Growing on conifer needles. Sexual state: Ascostromata black, globose to globose-depressed, erumpent or superficial, uniloculate, irregularly ostiolate, base embedded in the matrix, fused to erumpent hypostroma. Hamathecium lacking pseudoparaphyses. Asci eight-spored, cylindrical to cylindrical clavate, sessile and rounded at apex with a small ocular chamber. Ascospores uniseriate to triseriate, partially overlapping, hyaline, 1-septate, constricted at the septum, clavate or oblong with rounded ends, smooth-walled (Saccardo 1925; Von Arx and Müller 1954). Asexual state: See notes.

Notes: Phaeocryptopus was introduced by Naumov (1914) as a monotypic genus and Phaeocryptopus abietis Naumov. was assigned as the type. Petra (1938) included another three species Phaeocryptopus gaeumannii (T. Rohde), Phaeocryptopus nudus (Peck) Petr. and Phaeocryptopus pinastri (Sacc. & Ellis) Petr. while Petrak (1962) added P. podocarpi (Syd. & P. Syd.) Petr.,. Butin (1970) included P. araucariae Butin, and P. australis Butin, while Farr (1984) added P. saxegotheae (Henn.) M.L. Farr, which has been referred to Dimerosporium. Müller and Von Arx (1950) included Phaeocryptopus in Venturiaceae. Barr (1987a) also classified Phaeocryptopus under Venturiaceae. Rhizosphaera species are generally accepted as asexual states of Phaeocryptopus, however the relationship has never been conclusively established (Winton et al. 2007). Presently P. nudus (Asterina nuda Peck), is considered as the type of Phaeocryptopus as the earliest introduced one. Phylogenetic analysis of Winton et al. (2007) and Schoch et al. (2009) showed that Phaeocryptopus gaeumannii clustered in Mycosphaerellaceae, Capnodiales while P. nudus nested in Dothioraceae, Dothideales. In our phylogenetic analysis Rhizosphaera kalkhoffii, R. oudemansii and Plowrightia abietis along with Phaeocryptopus nudus clustured in a subclade which might be considered to belong in a single genus Phaeocryptopus or could be two distinct genera. We name this subclade as Phaeocryptopus since Rhizosphaera abietis L. Mangin & Har. the type of Rhizosphaera, has no molecular data.

Type species: Phaeocryptopus nudus (Peck) Petr., Annls mycol. 36(1): 15 (1938), (Fig. 8), Facesoffungi number: FoF00087

Fig. 8

Phaeocryptopus nudus (F67760, holotype). a, b Herbarium material. c, d Ascomata on the host surface. e, f Asci. g Ascospores. Scale bars: d = 1000 μm, e, f = 20 μm, g = 10 μm

Parasitic on conifer needles. Sexual state: Mycelium superficial, extensive, brown. Ascostromata 13–21 μm diam, black, superficial, gregarious, solitary or scattered, globose to subglobose, coriaceous, forming linear spots parallel along the middle vein. Hamathecium lacking pseudoparaphyses. Asci 41–48 × 11–19 μm $\left(\overline{x}=45\times 17\mu m,n=5\right)$, eight-spored, bitunicate, fissitunicate, clavate, widely oblong, lacking a pedicel, ocular chamber not observed. Ascospores 11–18 × 3–9 μm $\left(\overline{x}=14\times 6\mu m,n=5\right)$, uniseriate to triseriate, partially overlapping, hyaline,1-septate, stongly constricted at the septum, upper cell slightly larger than lower cell, clavate, widely oblong, guttulate. Asexual state: Unknown.

Material examined: USA, Adirondack mountains, on dead needles of Abies balsamea (S- F67760 holotype?).

Plowrightia Sacc., Syll. fung. (Abellini) 2: 635 (1883), Facesoffungi number: FoF00088

Synonyms

Elmerococcum Theiss. & Syd., Annls mycol. 13(3/4): 282 (1915)

Parasitic or saprobic on leaves, twigs and wood in terrestrial habitats. Sexual state: Ascostromata dark brown to black, immersed, becoming erumpent through the epidermis, solitary or scattered, pulvinate, subglobose to globose, coriaceous, multiloculate, with 2 to many locules, cells of ascostromata composed of several layers of dark brown cells of textura angularis. Locules subglobose to globose, thick-walled. Peridium of locules comprising several layers of dark brown cells of textura angularis or prismatica, vertical to the host surface. Hamathecium lacking pseudoparaphyses. Asci eight-spored, bitunicate, fissitunicate, cylindrical, elongate to ellipsoid, pedicellate, rounded at the apex. Ascospores overlapping, biseriate, hyaline, 1-septate, slightly constricted at septum, fusiform, subglobose to globose, tapering towards both ends, smooth-walled. Asexual state: See notes.

Notes: Plowrightia was introduced by Saccardo (1883) in the family Dothideaceae to accommodate P. berberidis (De Not.) Sacc., P. bullata Sacc., P. hippophaës, P. insculpta (Wallr.) Sacc., P. martianoffiana (Niessl & Thüm.) Sacc., P. mezerei (Schleich. ex Fr.) Sacc., P. morbosa Sacc., P. periclymeni (Fuckel) Sacc., P. ribesia, P. tuberculiformis (Ellis) Sacc., and P. virgultorum (Fr.) Sacc., and P. ribesia assigned as the type species. Barr (1972) synonimysed Plowrightia with Dothiora, but Von Arx and Müller (1975) and Barr (1987a) reinstated it in Dothideaceae and separated it from Dothiora based on the pulvinate, erumpent ascostromata of Plowrightia with small locules often higher than broad. Barr (1987a) transferred Plowrightia to Dothideaceae, while Hawksworth et al. (1995) and Lumbsch and Huhndorf (2010) included it in the family Dothioraceae. Recent molecular and phylogenetic studies carried out by Winton et al. (2007) also confirmed the position of Plowrightia in the family Dothioraceae. Based on their phylogenetic analysis, Winton et al. (2007) showed that Plowrightia abietis is similar to Phaeocryptopus nudus and identical to Rhizosphaera oudemansii Maubl. However, in this study we observed the type specimen of P. nudus and P. abietis and it is clear that they are not similar. In our phylogenetic tree, Phaeocryptopus nudus and R. oudemansii are clustered separately in two sister clades in Dothideaceae while, Plowrightia abietis clustered with R. oudemansii in the same clade. Therefore, we can conclude that Plowrightia abietis and R. oudemansii are identical with the latter as the asexual state of P. abietis. However, Rhizosphaera abietis L. Mangin & Har. the type of Rhizosphaera, has no molecular data and needs to be re-collected and sequenced in order to resolve the affinities of Rhizosphaera with Plowrightia in Dothideaceae. Plowrightia shares a common morphology with Dothidea. We collected P. ribesia, the type species of Plowrightia and it clustered in Dothideaceae along with other Dothidea species. Other Plowrightia species P. abietis and P. periclymeni grouped with P. nudus and Rhizosphaera species. Rhizosphaera species are also known as the asexual states of Phaeocryptopus. Conidia are unicellular but occasionally two-celled (Orton 1915). Plowrightia also produces Hormonema asexual states in culture and classified in Venturiaceae, but grouping with Capnodiales (Winton et al. 2007). Petrak (1923b) proposed a new genus Systremmopsis with Systremmopsis ribesia as the asexual morph of Dothidea (Plowrightia) ribesia found in stromata in nature with hyaline conidia without conidiophores. Systremmopsis ribesia needs recollecting and sequencing to establish the relationship between Plowrightia ribesia and Systremmopsis ribesia.

Type species: Plowrightia ribesia (Pers.) Sacc., Syll. fung. (Abellini) 2: 635 (1883), (Figs. 9, 10), Facesoffungi number: FoF00089

Fig. 9

Plowrightia ribesia (BPI 642930). a Herbarium material. b Ascostromata on the host surface. c Partial section through ascostromata. d Close up of the locules. e, f Asci. g, h Ascospores. Scale bars: b = 1000 μm, c, d = 100 μm, e, f = 50 μm, g, h = 20 μm

Fig. 10

Plowrightia ribesia (MFLU 14–0040). a, b. Ascostromata on the host surface. c–e. Vertical section through ascostromata. f. Close up of the peridium of locus. g–j. Asci with ascospores. k. Germination of ascospore. l–p. Ascospores. q. Colonies on PDA from above and r blow. Scale bars: b = 1000 μm, c = 100 μm, d = 50 μm, e, f = 20 μm, g–k = 10 μm, l–p = 5 μm

≡ Sphaeria ribesia Pers., Ann. Bot. (Usteri) 11: 24 (1794)

Parasitic or saprobic on wood and twigs in terrestrial habitats. Sexual state: Ascostromata (683−)955 × 1257(−1297) μm, black, immersed, becoming erumpent thorough bark at maturity, solitary or scattered, globose to subglobose, coriaceous, multiloculate, with many locules, cells of ascostromata composed of several layers of dark brown to black cells of textura angularis. Locules 408–413 μm high × (76−)84 − 115(−122) μm diam. $\left(\overline{x}=410.5\times 93\mu m,n=10\right)$, subglobose to obpyriform, arranged at the periphery 1 to 2 layers of locules, evidently ostioles at the surface. Peridium of locules thick, cell layers with ca. 11 μm thick, composed of dark brown to lightly pigmented cells of textura prismatica and angularis. Hamathecium lacking pseudoparaphyses. Asci (108.5−) 111–123 (−125) × 13–16 (−18) μm $\left(\overline{x}=116\times 15\mu m,n=15\right)$, eight-spored, bitunicate, oblong, cylindrical to subclavate, pedicellate, and rounded at the apex with a flattened ocular chamber ca. 2.5 μm wide, stalks ca. 16 μm long. Ascospores (20−)23 − 30.5(−32) × 7–9.5(−10) μm $\left(\overline{x}=26\times 8.5\mu m,n=25\right)$, overlapping, uni to biseriate, hyaline, broadly fusiform, rounded at both ends, with upper broad cell, 1-septate, with a median septum, constricted at the septum, smooth. Asexual state: Unknown.

Material examined: CZECH REPUBLIC, Böhmen, Tabor, on Ribes rubrum L. (Grossulariaceae), April 1904, Fr. Bubák (BPI 642930).

Description of Plowrightia ribesia (MFLU 14–0040)

Saprobic on woody plants. Sexual state: Ascostromata 1–1.3 mm high × 0.45 − 0.53 mm diam $\left(\overline{x}=1.23\times 0.48 mm,n=10\right)$, black, superficial or semi-immersed to erumpent, solitary, scattered, or sometimes gregarious, globose to subglobose, coriaceous, multiloculate, with 10–13 locules, cells of ascostromata composed of dark brown-walled of textura angularis. Locules $\left(\overline{x}=131\times 98\mu m,n=10\right)$, globose to subglobose, non-ostiolate. Peridium of locules 15–31 μm $\left(\overline{x}=21\mu m,n=7\right)$, comprising few layers of lightly pigmented to brown cells of textura angularis. Hamathecium lacking pseudoparaphyses. Asci 81–122 × 11–17 μm $\left(\overline{x}=98\times 14\mu m,n=10\right)$, eight-spored, bitunicate, fissitunicate, clavate, with a short pedicel, thick-walled, rounded at the apex. Ascospores 18–28 × 5–11 μm $\left(\overline{x}=23\times 8\mu m,n=10\right)$, uniseriate, partially overlapping, hyaline, 1-septate, constricted at the septum, upper cell often broader than the lower cell, fusiform to ellipsoid, gradually tapering towards the apex, smooth-walled. Asexual state: Unknown.

Culture characteristics: Ascospores germinating on PDA within 12 h and germ tubes arise from both end cells. Colonies growing fast on PDA, reaching a diam. of 3 cm after 5 days at 29 °C, velvety, radiating towards the edge. Mycelium initially hyaline and light pink at the margin.

Material examined: ITALY, Fonte al Fringuello - Pratomagno (Province of Arezzo [AR]), on dead and not land branch of Daphne sp. (Thymelaeaceae), 10 May 2013, Camporesi Erio (MFLU14 − 0040), living culture MFLUCC 13–0670.

Notes: Plowrightia ribesia (Sphaeria ribesia Pers.) was introduced by Saccardo (1883) as the type specimen of Plowrightia. We could not loan the holotype of Plowrightia ribesia (Pers.) Sacc., but observed a specimen from BPI (BPI 642930) from the Czech Republic. This was compared with our recent collection from Italy from dead branches of Daphne sp. and is indistinguishable. Plowrightia ribesia is widely distributed in Europe and is parasitic on Ribes rubrum (Grossulariaceae) (Saccardo 1883; Hoggan 1927). Since the host of the MFLU specimen from Italy is from Daphne sp. (Thymelaeaceae) and not Ribes we have not designated this as an epitype. However, this strain can be considered as an authentic until proven otherwise.

Plowrightia abietis (M.E. Barr) M.E. Barr [as ‘Plowrighthia’], Sydowia 41: 32 (1989), (Fig. 11), Facesoffungi number: FoF00090

Fig. 11

Plowrightia abietis (NY00914455, holotype) a Herbarium material b, c Ascostromata on the host surface. d Section through ascostroma. e Close up of locules. f–h Bitunicate asci. i–l Ascospores. Scale bars: d = 200 μm, e = 50 μm, f–h = 20 μm, i–l = 5 μm

≡ Xenomeris abietis M.E. Barr, Can. J. Bot. 46: 842 (1968)

Saprobic on twigs in terrestrial habitatas. Sexual state: Ascostromata 400–800 μm wide black, superficial, solitary, globose to subglobose, multiloculate, cells of ascostromata composed of several layers of dark brown cells of textura angularis. Locules 40–50 μm × 32–40 μm $\left(\overline{x}=44\times 37\mu m,n=5\right)$, globose to subglobose, non-ostiolate. Peridium of locules 8–12 μm wide, comprising dark brown to lightly pigmented cells of textura angularis. Pseudoparaphyses not observed. Asci 35–45 μm × 11.7 − 13 μm $\left(\overline{x}=39\times 12.4\mu m,n=15\right)$, eight-spored, bitunicate, broadly cylindrical, obovoid, short pedicellate and rounded at the apex with an ocular chamber. Ascospores 10–13.5 μm × 3.5 − 5.5 μm $\left(\overline{x}=11.8\times 4.3\mu m,n=20\right)$,uni to biseriate, partially overlapping, hyaline, 1-septate, with the upper cell slightly broader than the lower one, rounded end at both apices, constricted at the septa, oblong to obovoid, smooth-walled. Asexual state: Unknown.

Material examined: USA, Idaho, Ida Creek Spur Road, on twigs of Abies grandis (Douglas ex D. Don) Lindley (Pinaceae), 5 October 1939, A.W. Slipp (NY00914455, holotype).

Notes: Sequence data of a putative strain of Plowrightia abietis (ATCC 24339) clustered in the Phaeocryptopus clade with Rhizosphaera species (R. kalkhoffii and R. oudemansii). This species should therefore probably be synonymised under Rhizosphaera or Phaeocryptopus, but we refrain from doing so until a sequence can be linked with morphology.

Stylodothis Arx & E. Müll., Stud. Mycol. 9: 11 (1975), Facesoffungi number: FoF00091

Saprobic on bark and stems. Sexual state: Ascostromata black, immersed, erumpent to superficial, solitary or scattered, gregarious, coriaceous, multiloculate, with four to numerous locules, cells of ascostromata composed of several layers of dark brown cells of textura angularis. Locules globose to subglobose, non-ostiolate. Peridium of locules a single layer heavily or lightly pigmented cells of textura angularis. Hamathecium comprising hyaline, septate, anastomosing pseudoparaphyses, embedded in a gelatinous matrix. Asci 4–8-spored, bitunicate, clavate to cylindrical, pedicellate, rounded at the apex. Ascospores uni or biseriate, sometimes partially overlapping, yellowish-brown to dark brown, 1-septate, rarely 2 or 3-septate, constricted at the primary septum, upper cell wider, ellipsoid to fusiform, lower cell narrow and tapering to rounded base, smooth-walled, sometimes guttulate. Asexual state: Unknown.

Notes: Stylodothis was introduced by Von Arx and Müller (1975) to accommodate Stylodothis puccinioides and Stylodothis indica (Loeffler & S.K. Bose) Arx & E. Müll. Morphologicaly these two species are similar to Dothidea. S. puccinioides has four ascospores in ascus and develops on Buxus sempervirens while S. indica bear eight ascospores sometimes with 2 or 3-septate and grows on Berberidis lycii. Recent phylogenetic analyses by Schoch et al. (2006) show that S. puccinoides is closely related to Dothidea. In the present study Stylodothis is well-supported genus in Dothideaceae with 81/100 % bootstrap support.

Type species: Stylodothis puccinioides (DC.) Arx & E. Müll., Stud. Mycol. 9: 11 (1975), (Fig. 12), Facesoffungi number: FoF00092

Fig. 12

Stylodothis puccinioides (PC 0084648, holotype). a Herbarium material. b, c Ascostromata on host substrate. d Section through ascostroma. e Cells of ascostroma. f Peridium of locules. g Ascus. h Ascus stained in Melzer’s reagent. i Ascus stained in cotton blue reagent. j Pseudoparaphyses. k Ascospore in 70 % lactic acid. l Ascospore stained in Melzer’s reagent. m Ascospore stained in cotton blue reagent. Scale Bar: d = 100 μm, e–i, k–m = 20 μm, j = 10 μm

≡ Sphaeria puccinioides DC., in de Candolle & Lamarck, Fl. franç., Edn 3 (Paris) 5/6: 118 (1815)

Saprobic on twigs or wood. Sexual state: Ascostromata 197–374 μm high × 495–713 μm diam $\left(\overline{x}=345\times 570\mu m,n=5\right)$, black, formed on erumpent basal stroma, gregarious, thick at the basal of ascostromata, coriaceous, multiloculate, with 4–5 locules, cells of ascostromata composed of several layers of dark brown cells of textura angularis. Locules 69–134 μm high × 83–116 μm diam $\left(\overline{x}=88\times 110\mu m,n=10\right)$, globose to sub-globose, without an ostiole. Peridium of locules 31–40 μm $\left(\overline{x}=34\mu m,n=10\right)$, comprising 1-layer, with heavily pigmented, brown to dark brown cells of textura angulari. Hamathecium of 5 μm wide, septate, anastomosing, pseudoparaphyses embedded in a gelatinous matrix. Asci 135–147 × 21–24 μm $\left(\overline{x}=138\times 23\mu m,n=10\right)$, 2 − 4-spored, bitunicate, clavate to cylindrical, short and rounded pedicellate, rounded at the apex without an ocular chamber. Ascospores 36–40 × 17–20 μm $\left(\overline{x}=39\times 19\mu m,n=10\right)$, uniseriate, yellowish when immature, brown to dark brown at maturity, 1-septate, constricted and dark at septum, upper cell wider than lower cell, ellipsoid to fusiform, smooth-walled, sometimes guttulate. Asexual state: Unknown.

Material examined: GERMANY, near Aix, on bark and stem of Buxus sempervirens L. (Buxaceae), Mont du Chat (PC 0084648, holotype)

Sydowia Bres., Hedwigia 34(Beibl.): 66 (1895), Facesoffungi number: FoF00093

Synonyms

Pleodothis Clem., Gen. fung. (Minneapolis): 49, 173 (1909)

Plowrightia sect. Plowrightiella Sacc., Syll. fung. (Abellini) 11: 376 (1895)

Plowrightiella (Sacc.) Trotter, in Saccardo, Syll. fung. (Abellini) 24(1): 542 (1926)

Parasitic or saprobic on conifer plants or branches and wood. Sexual state: Ascostromata black, immersed to erumpent, solitary or gregarious, globose to subglobose, coriaceous, uniloculate, ostiolate. Peridium several layers thick, composed of dark brown cells of textura angularis or sometimes prismatica, more darkened on the outside, hyaline to lightly pigmented towards the interior. Hamathecium lacking pseudoparaphyses. Asci 8-poly-spored, bitunicate, fissitunicate, clavate to oblong with a short pedicel, apically rounded. Ascospores crowded to overlapping, hyaline, transversely multiseptate, constricted at the primary septum, sometimes with a vertical septum in the mid cells or rarely in the end cells, straight to inequilateral, guttulate, upper part usually wider and shorter than the lower part, elliptic, obovate, smooth. Asexual state: Hormonema and Sclerophoma (Wijayawardene et al. 2012; Hyde et al. 2013). Pycnidia stromatic, immersed at first, later erumpent, globose to irregular, solitary or aggregated, uni or multi locular or convoluted. The wall is thick and dehisces by breaking down the overlying walltissue. Conidiogenous cells phialidic, enteroblastic, determinate, discrete, hyaline to pale brown, doliiform to ampulliform, with a wide channel and minute collarette, formed from the inner cells of the wall lining the locule. Conidia hyaline, aseptate, ellipsoid, and rarely tapered at the base smooth-walled, sometimes guttulate (Sivanesan 1984).

Notes: Sydowia was formally established by Bresadola (1895) based on its gregarious ascomata in the host surface and remained monotypic until Saccardo (1899) added S. carestiae. Later, Luttrell (1973) transferred this genus into Dothioraceae based on its phragmosprous ascospores. Arx and Müller (1975) re-evaluated the bitunicate ascomycetes and placed this genus in Dothideacea based on its broad unilocular stromata and ascospores with only transverse septa. Barr (1972) and Sivanesan (1984) accepted only polysporous species in this genus. Barr (1972) accepted five species in Sydowia while Sivanesan (1984) included two. Barr (2001) accepted phragmosporous species in both eight-spored and poly-spored asci. She introduced three new phragmosporous species (S. slippii, S. taxicola and S. wolfii) which have been referred to Dothiora and S. ceanothi. Recent phylogenetic studies by Schoch et al. (2006) also proved that the placement of Sydowia in the Dothioraceae. The type species of Hormonema, Hormonema dematioides Lundberg and Melin has been suggested as the asexual state of Sydowia polyspora (Butin 1964; Cheewangkoon et al. 2009). In phylogeny analysis in Bills et al. (2004) Sydowia polyspora and H. dematioides clustered in the same clade. H. dematioides do not have the sequences of ex-type strain and also no authentic specimen has been described. We did not include this H. dematioides in our phylogenetic analysis as lack of LSU sequence data. Sequences dat of LSU and other protein coding genes and culture based analysis are required in order to confirm this relationship. Hence, we do not synonymize Hormonema under Sydowia and further investigations are suggested. Crous (in Crous et al. 2003) introduced Sydowia eucalypti (Sphaerulina eucalypti) and accommodated it in this genus based on having Sclerophoma and Hormonema synanamorphs in culture and the thick-walled ascostromata. In present phylogenetic study Sydowia eucalypti separates from the Dothideaceae clade and clusters in the Aureobasidiaceae clade with other Selenophoma species. Therefore, we exclude S. eucalypti from Sydowia and accommodate it in a new genus, Pseudosydowia. In our phylogeny analysis (Fig. 1) a putative strain of Sydowia polyspora (CBS 116.29) clustered in Dothideaceae close to a putative strain of Delphinella strobiligena (CBS 735.71). Considering the close relationship of the two strains it may be that one of these two strains is wrongly identified or these two species should be in one genus as both of them are polysporous. However, we named this clade as Delphinella which comprises D. strobiligena, Rhizosphaera pini and S. polyspora pending fresh collections of Delphinella and Sydowia species. Sydowia differs from other genera of Dothideaceae in having uniloculate ascostromata. However, we accommodate Sydowia in Dothideaceae pending molecular data of the type species.

Type species: Sydowia gregaria Bres., Hedwigia 34(Beibl.): 66 (1895), (Fig. 13), Facesoffungi number: FoF00094

Fig. 13

Sydowia gregaria (S-F6473, holotype). a Herbarium material. b, c Ascostromata on host surface. d, e Section of ascostromata. f Close up of the peridium. g–i Bitunicate asci. j–m Ascospores. Scale bars: d, e = 100 μm, f = 50 μm, g–i = 50 μm, j–i = 5 μm

Parasitic or saprobic on branches of conifer trees. Sexual state: Ascostromata 190–255 μm wide, black, immersed, becoming erumpent, gregarious, scattered, globose to subglobose, coriaceous, uniloculate, with a papillate to rounded short ostiole. Peridium up to 32–65 μm thick several layers thick, composed of dark brown cells of textura angularis, more darkened on the outside. Hamathecium lacking pseudoparaphyses. Asci 78–105 × 12–17 μm $\left(\overline{x}=95.5\times 13.3\mu m,n=15\right)$, polyspored, bitunicate, clavate to oblong, with a short pedicel, apically rounded. Ascospores 10–14 × 3–4.2 μm $\left(\overline{x}=12.3\times 3.6\mu m,n=30\right)$, crowded to overlapping, hyaline, transversely 2–3-septate, constricted at the primary septum, elliptic, obovate, smooth-walled. Asexual state: Unknown.

Material examined: GERMANY, Berlin, Steglitz bei Berlin, on branches of Picea abies (L.) H. Karst (Pinaceae), October 1894, P. Sydow (S-F6473, holotype).

Genera not studied

Pringsheimia Schulzer, Verh. zool.-bot. Ges. Wien 16: 57 (1866)

Type species: Pringsheimia rosarum Schulzer, Verh. zool.-bot. Ges. Wien 16: 57 (1866)

Notes: Pringsheimia was introduced by Schulzer (in Schulzer et al. 1866) to accommodate P. rosarum and currently comprises 17 epithets (Index Fungorum 2014). We could not locate the type of this genus. A sequence from a putative strain of Pringsheimia smilacis is available in GenBank and clustered in Dothideaceae as a separate clade in our phylogenetic tree. The sequences of the type and other species are not available and they need recollecting, molecular analysis and epitypifying in order to resolve the family placement of this genus.

Aureobasidiaceae K. M. Thambugala & K. D. Hyde, fam. nov., Index Fungorum number: IF 550732; Facesoffungi number: FoF00097

Parasitic or saprobic on twigs, wood and leaves or human skin. Sexual state: Ascomata black, subglobose, immersed to erumpent, uniloculate, sometimes ostiolate. Peridium composed of several layers of brown to dark brown cells of textura angularis. Hamathecium lacking pseudoparaphyses. Asci eight-spored, bitunicate, saccate to broadly clavate with a short bifurcate pedicel or apedicellate, apex broadly rounded with a distinct ocular chamber. Ascospores biseriate to triseriate, partially overlapping, hyaline, 3 to many septate or aseptate, muriform to phragmosporous, obovoid or elliptic with broad to narrow rounded ends. Asexual state: Coelomycetous or hyphomycetous. Hyphomycetous asexual state: Stromata present or absent, visible in substomatal cavity, hyaline or lightly pigmented, dark brown with globose to broadly ellipsoidal, round or elongated pseudo-parenchymatous cells when present. Colonies spreading, smooth, often covered with slimy masses of conidia, usually with sparse aerial mycelium; light brown, yellow, pink or black. Hyphae with cells commonly wider than long, hyaline, frequently soon becoming brown and thick-walled sometimes thin, smooth, transversely septate. Conidiogenous cells on hyaline hyphae, lateral, terminal or intercalary, cylindrical, clavate or globose, integrated, terminal, with holoblastic, polyblastic conidiogenesis, with numerous synchronously produced conidia. Conidia blastic, hyaline to dark brown, smooth, aseptate, straight, ellipsoidal to spherical, reniform to sickle-shaped, oblong to cylindrical, sometimes cylindrical with obtuse ends and occasionally with a slightly truncate base. Secondary conidia common; endoconidia often present. Coelomycetous asexual state: Conidiomata dark brown, pycnidial or acervular, separate or aggregated, scattered, immersed or superficial, globose, unilocular, thin-walled; walls 2–3 layers thick, composed of lightly pigmented to brown, thick-walled cells of textura angularis, with or without an ostiole. Conidiophores reduced to conidiogenous cells or hyaline, irregularly branched at the base and above, 1 to 3-septate, smooth when present. Conidiogenous cells enteroblastic, phialidic, discrete, determinate or ampulliform, hyaline to pale brown, smooth, subglobose, obpyriform or obovoid, collarette and channel minute, with apical periclinal thickening, guttulate, formed from the inner cell of the pycnidial wall, slightly tapered toward the apex. Conidia holoblastic, aseptate, falcate, fusiform, ellipsoidal to obovoid, eguttulate or irregularly guttulate, smooth, at first thin-walled, hyaline or pale brown, later pale brown with thicker walls, smooth-walled or verruculose.

Type: Aureobasidium Viala & G. Boyer

Notes: The Aureobasidiaceae clade comprises Aureobasidium, Kabatiella, Pseudoseptoria, Saccothecium and Selenophoma species and Columnosphaeria fagi, for which we propose a new family, Aureobasidiaceae. The CBS 584.75 ex-neotype strain of Aureobasidium pullulans clustered here. Aureobasidiaceae separates from Dothideaceae with 65/80 % bootstrap support. Morphologically, Aureobasidiaceae can be distinguished from Dothideaceae usually by having only immersed to erumpent, uniloculate ascostromata and aseptate to many-septate, hyaline ascospores. The recently introduced Sydowia eucalypti also clustered within Aureobasidiaceae and we propose a new genus, Pseudosydowia for Sydowia eucalypti. The asexual states of Aureobasidiaceae Aureobasidium, Kabatiella, Pseudoseptoria and Selenophoma species cluster here, while Columnosphaeria, Pseudosydowia and Saccothecium nested as sexual genera. Columnosphaeria fagi, Pseudosydowia eucalypti and Saccothecium sepincola are the only sexual species cluster in the Aureobasidiaceae clade. The sexual species Discosphaerina (Columnosphaeria) fagi is closely related with Aureobasidium pullulans (Schoch et al. 2006; Zalar et al. 2008). Our phylogenetic tree also shows the close relationship of these two species which form a single clade. We accept three sexual genera and four asexual genera in Aureobasidiaceae.

Key to sexual genera of Aureobasidiaceae

1. Ascospores septate, obovoid to ellipsoid...................... 2

1. Ascospores aseptate, ellipsoidal-fusiform, or fusiform............................................ Columnosphaeria

2. Asci broadly ellipsoid to clavate, straight to slightly curved, subsessile, ascospores bi- to multiseriate, overlapping, hyaline, (1–)3(−4)-septate at maturity, constricted at the median septum, obovoid to ellipsoid with obtuse ends, tapering towards both ends, but more prominently towards the lower end................................................... Pseudosydowia

2. Asci saccate to broad-clavate with a short bifurcate pedicel or apedicellate, ascospores biseriate to triseriate, partially overlapping, hyaline, 3 to many septate, muriform to phragmosporous, obovoid or elliptic with broad to narrowly rounded ends............. Saccothecium

Key to asexual genera of Aureobasidiaceae

1. Coelomycetous asexual states....................................... 2

1. Hyphomycetous asexual states...................................... 3

2. Conidia falcate, fusiform, ellipsoidal to obovoid, aseptate, eguttulate or irregularly guttulate, smooth, at first thin-walled, hyaline or pale brown, later pale brown with thicker walls and verruculose.......... Selenophoma

2. Conidia fusiform, lunate or irregular, curved, aseptate, colourless, smooth-walled with or without an excentric basal appendage, continuous with conidium body, plectronoid to podiform, or with a blunt or spathulate distal end................................................ Pseudoseptoria

3. Conidia blastic, produced simultaneously in dense groups, hyaline to dark brown, smooth, aseptate, straight, ellipsoidal to spherical, reniform to sickle-shaped................................................... Aureobasidium

3. Conidia clustered on swollen apex, borne on tiny sterigmata, aseptate, hyaline, oblong to cylindrical.............................................................. Kabatiella

Aureobasidium Viala & G. Boyer, Rev. gén. Bot. 3: 371 (1891), Facesoffungi number: FoF00098

Synonyms:

Aureobasis Clem. & Shear, Gen. fung., Edn 2 (Minneapolis): 343, 381 (1931)

Dematoidium Stautz, Phytopath. Z. 3: 204 (1931)

Pachybasidiella Bubák & Syd., Annls mycol. 13(1): 9 (1915)

Polyspora Laff., Scientific Proc. R. Dublin Soc., N.S. 16: 258 (1921) [1920–22]

Protocoronis Clem. & Shear, Gen. fung., Edn 2 (Minneapolis): 197, 344 (1931)

Protocoronospora G.F. Atk. & Edgerton, J. Mycol. 13(5): 185 (1907)

Pullularia Berkhout, De Schimmelgeslachten Monilia: 55, 64 (1923)

Parasitic or saprobic on plants, fruits and causes human skin diseases. Some plant pathogens form Kabatiella synanamorphs which cause leaf spots. They form sub circular irregular, amphigenous, necrotic, sunken, pale to medium brown leaf spots with a raised, dark brown margin. Stromata present or absent, visible in sub-stomatal cavity, hyaline or lightly pigmented, dark brown with globose to broadly ellipsoidal, round or elongated pseudo-parenchymatous cells when present. Colonies spreading, smooth, often covered with slimy masses of conidia, usually with sparse aerial mycelium; light brown, yellow, pink or black. Hyphae with cells commonly wider than long, hyaline, frequently soon becoming brown and thick-walled sometimes thin, smooth, transversely septate. Asexual state: Conidiomata acervular to sporodochial, amphigenous, substomatal, subepidermal, pulvinate, dry or crystaline in appearance, pale brown, discrete. Conidiogenous cells on hyaline hyphae, lateral, terminal or intercalary, cylindrical, clavate or globose, integrated, terminal, with holoblastic, polyblastic conidiogenesis, with numerous synchronously produced conidia. Conidia blastic, hyaline to dark brown, smooth-walled, aseptate, straight, ellipsoidal to spherical, reniform to sickle-shaped, sometimes cylindrical with obtuse ends and occasionally with a slightly truncate base, rather variable in shape and size. Secondary conidia common; endoconidia often present. Occasionally dark, one or two-celled arthroconidia are formed. Hyaline conidia one-celled, smooth, ellipsoidal, very variable in shape and size often with an indistinct hilum. Dark brown conidia 1–2 celled, two celled slightly constricted at septum. (Hermanides-Nijhof 1977; Zalar et al. 2008). Sexual state: See notes.

Notes: Aureobasidium was introduced by Viala and Boyer (1891) and Aureobasidium vitis assigned as the type specimen. Thereafter a number of species were introduced by various authors and several Kabatiella species were also transferred to Aureobasidium considering morphological characters and ability to form leaf spots on a restricted host range (Hermanides-Nijhof 1977). The type specimen Aureobasidium vitis has not been preserved. De Bary (1884) introduced Dematium pullulans and Arnaud (1910) suggested that Dematium pullulans and Aureobasidium vitis are the same. Then Hermanides-Nijhof (1977) described A. pullulans as the oldest name for the type species of Aureobasidium designating CBS 584.75 as a ex-neotype strain (Zalar et al. 2008). In this paper, we introduce a new family Aureobasidiaceae in order to accommodate species in the second major clade (Aureobasidiaceae) of our phylogenetic tree. The Aureobasidium clade consist of two putative strains of Discosphaerina (Columnosphaeria) fagi (CBS 171.93), Selenophoma mahoniae (CBS 388.92) and the ex-type strain of Kabatiella lini (CBS 125.21) in addition to Aureobasidium species. They might be considered to belong in Aureobasidium, but we retain Columnosphaeria fagi and Selenophoma mahoniae in related genera until the type specimens of Columnosphaeria and Selenophoma are collect and sequenced.

Type species: Aureobasidium pullulans (de Bary) G. Arnaud var. pullulans – Annales École Nat. Agric. Montpellier 16: 39, 1918, (Fig. 14), Facesoffungi number: FoF00099

Fig. 14

Aureobasidium pullulans (MFLUCC 14–0288). a Culture on PDA incubated for a. 2 weeks b. 7 weeks at 25 °C. c–g Conidiophores and conidiogenesis. h Conidia. Scale bars: c–h = 10 μm

For other synonyms see Index Fungorum

Cultural characteristics: Colonies on PDA at 25 °C attaining about 70–80 mm diam after 14 days, appearing smooth and slimy due to abundant sporulation, pinkish white. Within first 6 weeks colonies filamentous and there after develop white, setae-like mycelia then turning to brown first and then black at the irregular margin.

Vegetative hyphae hyaline, transversely septate, smooth, sometimes becoming brown to dark brown when getting older, smooth, thin-walled, 2–9 μm wide, transversely septate. Conidiogenous cells undifferentiated, intercalary or terminal on hyaline hyphae or arising as short lateral branches. Conidia 5.5 − 16 × 3–6 μm $\left(\overline{x}=9.2\times 4.2\mu m,n=60\right)$, produced synchronously in dense groups from small denticles, and also formed percurrently on short lateral denticles, hyaline sometimes becoming brown, ellipsoidal, aseptate, smooth. Endoconidia are produced by an intercalary cells.

Material examined: THAILAND, Chiang Rai, on PDA, 12 Aprial 2013 (MFLUCC 14–0288, ICMP 20350).

Columnosphaeria Munk, Dansk bot. Ark. 15(no. 2): 103 (1953), Facesoffungi number: FoF00100

For other synonyms see Index Fungorum

Saprobic or parasitic on dead wood or leaves in terestial habitats Sexual state: Ascostromata black to dark brown, immersed under the host tissue or erumpent, solitary or gregarious or scattered, globose to subglobose, unilocular, ostiolate at maturity usually by dehiscence of the thin wall apex. Peridium of several layers, dark brown wall cells of textura angularis. Hamathecium lacking pseudoparaphyses. Asci eight-spored, bitunicate, clavate to cylindro-clavate, short pedicellate, apically rounded. Ascospores uni-seriate or irregularly bi-seriate, partially overlapping, hyaline to brown, aseptate, ellipsoidal-fusiform, or fusiform, smooth-walled, sometimes with a thin mucilaginous sheath. Asexual state: See under C. fagi.

Notes: Columnosphaeria was introduced by Munk (1953) as a monotypic genus and typified by Columnosphaeria sarothamni. Von Arx and Müller (1954, 1975) and Munk (1957) considered Columnosphaeria as a synonym of Guignardia (Luttrell 1973) while Sivanesan (1984) synonymized the genus under Discosphaerina and C. sarothamni under D. cytisi. Luttrell (1973) accommodated this genus in Dothideaceae. Barr (2001) included new five species which have been referred to Guignardia and Physalospora and placed Columnosphaeria in Dothioraceae. However, in our phylogenetic tree Columnosphaeria (Discosphaerina) fagi nests in the family Dothideaceae. Barr (2001) transfered Discosphaerina fagi (H.J. Huds.) M.E. Barr, to Columnosphaeria fagi based on its morphology. Phyllosticta Pers., is considered as the current name of Columnosphaeria (Index Fungorum 2014). However, Columnosphaeria sarothamni, the type of Columnosphaeria does not share morphological features with Phyllosticta (Wikee et al. 2013). Hormonema, Sarcophoma and Aureobasidium are reported as the asexual morphs in culture (Barr 2001) and these asexual morphs are usually reported in Dothideales. We examined the type material of Columnosphaeria (C. sarothamni) and C. fagi, which grouped in Aureobasidiaceae. Based on the dothidealen asexual morphs of Columnosphaeria species and the similarity between C. sarothamni and C. fagi tentatively we accept Columnosphaeria in Aureobasidiaceae, Dothideales although the type has morphological characters like Botryosphaeriaceae.

Type species: Columnosphaeria sarothamni Munk, Dansk bot. Ark. 15(no. 2): 103 (1953), (Fig. 15), Facesoffungi number: FoF00101

Fig. 15

Columnosphaeria sarothamni (C-F70846, holotype). a Herbarium material. b Ascostromata on the host surface. c Section through ascostroma. d Peridium. e–g Bitunicate asci. h–j Ascospores. Scale bars: c = 100 μm, d–g = 50 μm, h–j = 15 μm

Fig. 16

Columnosphaeria fagi (IMI 110819, holotype). a Herbarium material b, c Ascostromata on the host surface. d Section through ascostroma. e Peridium. f–h Bitunicate asci i–l Ascospores. Scale bars: d = 25 μm, f–h = 20 μm, i–l = 5 μm

Saprobic on twigs of Sarothamnus sp. Sexual state: Ascostromata 235–300 μm high, 260–295 μm wide $\left(\overline{x}=267\times 282\mu m,n=10\right)$ solitary or scattered, black, immersed under the host tissue, globose to subglobose, uniloculate, ostiolate at maturity usually by dehiscence of the thin wall at the apex. Peridium 30–50 μm $\left(\overline{x}=42\mu m,n=15\right)$ wide, comprising several layers of dark brown cells of textura angularis. Hamathecium lacking pseudoparaphyses. Asci 80–90 × 15–18 μm $\left(\overline{x}=86\times 16.5\mu m,n=20\right)$ eight-spored, bitunicate, clavate to cylindro-clavate, short pedicellate, apically rounded. Ascospores 22 − 31× 5.5 − 9 μm $\left(\overline{x}=27\times 7\mu m,n=20\right)$ uni-seriate or irregularly bi-seriate, partially overlapping, hyaline when young, brown at maturity, aseptate, ellipsoidal-fusiform, or fusiform, smooth-walled. Asexual state: Unknown.

Material examined: DENMARK, on Cytisus scoparius (L.) Link (Fabaceae), 12 May 1948, Anders Munk (C-F70846, holotype)

Columnosphaeria fagi (H.J. Huds.) M.E. Barr, Harvard Pap. Bot. 6(1): 28 (2001), (Fig. 16), Facesoffungi number: FoF00102

≡ Guignardia fagi H.J. Huds., Nova Hedwigia 10(3/4): 323 (1966) [1965]

Parasitic on leaves of a deciduous tree. Sexual state: Ascomata 65–100 μm high, 70–115 μm wide $\left(\overline{x}=76\times 93\mu m,n=5\right)$, epiphyllous, subcuticular, black, immersed to erumpent, scattered, uniloculate without a distinct ostiole. Peridium 9–20 μm $\left(\overline{x}=14\mu m,n=15\right)$ wide, comprising several layers, dark brown to hyaline cells of textura angularis. Hamathecium lacking pseudoparaphyses. Asci 35–45 × 9–14 μm $\left(\overline{x}=40.5\times 11.5\mu m,n=20\right)$, eight-spored, sometimes less than eight, bitunicate, broadly cylindrical, ellipsoidal, short pedicellate or sessile, apically rounded with an ocular chamber. Ascospores 11–19 × 3–6 μm $\left(\overline{x}=13.6\times 4.4\mu m,n=20\right)$, overlapping bi-triseriate, hyaline, aseptate, ellipsoidal to fusiform, sometimes with a thin mucilaginous sheath. Asexual state: aureobasidium-like; Colonies white to black. Conidiogenous cells intercalary or terminal, forming blastic conodia, denticulate or with minute scar. Conidia aseptate, rarely 1-septate, hyaline, smooth, elliptic-oblong. Secondary conidia and endoconidia are sometimes produced (asexual morph description follows Sivanesan 1984).

Material examined: UK, Cambridgeshire: Cambridge, Gog magog Hills, on Fagus sylvatica (Fagaceae), 18 May 1964, H.J. Hudson (IMI 110819, holotype).

Kabatiella Bubák, Hedwigia 46: 297 (1907), Facesoffungi number: FoF00103

Synonymy:

Exobasidiopsis Karak., Notul. syst. Inst. cryptog. Horti bot. petropol. 1: 81 (1922)

Subepidermal on leaves. Sexual state: Unknown. Asexual state: Conidiomata sporodochia or acervuli, stromata, pale or pale brown. Conidiophores stromatic, unbranched, or branched, hyaline. Conidiogenous cells polyblastic synchronous, inconspicuous scars orndenticles, vesiculate, hyaline. Conidia clustered on swollen apex, borne on tiny sterigmata, aseptate, hyaline, oblong to cylindrical, conidial secession schizolytic (Bubák and Kabát 1907; Seifert et al. 2011).

Type species: Kabatiella microsticta Bubák, Hedwigia 46: 297 (1907)

Notes: Kabatiella was introduced by Bubák (in Bubák and Kabát 1907) in order to accommodate Kabatiella microsticta. Several authors have been synonymized Kabatiella under Aureobasidium (Hermanides-Nijhof 1977; Sivanesan 1984) but Seifert et al. 2011 treated these two genera as separate. In our phylogenetic analysis, Kabatiella forms a separate clade with 90/ 95 % bootstrap support. Kabatiella microsticta, the type species of Kabatiella and K. caulivora clustered in the Kabatiella clade. However, the ex-type stain of K. lini clustered with Aureobasidium species in our phylogenetic tree. Therefore, K. lini may belong to Aureobasidium.

Pseudoseptoria Speg., Anal. Mus. nac. B. Aires, Ser. 3 13: 388 (1910), Facesoffungi number: FoF00134

≡ Lunospora Frandsen, Meddr Plantepatol. Afd. Kgl. Veterin. Landb., (Københaven) 26: 70 (1943)

≡ Aphanofalx B. Sutton, Trans. Br. mycol. Soc. 86(1): 21 (1986)

Type species: Pseudoseptoria donacicola Speg., Anal. Mus. nac. B. Aires, Ser. 3 13: 388 (1911), Facesoffungi number: FoF00135

Notes: Pseudoseptoria was introduced by Spegazzini (1910) as an asexual genus in order to accommodate Pseudoseptoria donacicola (P. donacis). This genus is characterized by immersed, branched, septate, pale brown mycelium, pycnidial, solitary or linearly aggregated, immersed, brown, globose, unilocular, thin-walled conidiomata of pale brown cells of textura angularis with a distinct, central, circular ostiole. Conidiogenous cells are discrete, determinate or indeterminate, hyaline, smooth, ampulliform with a prominent cylindrical papilla and falcate. Conidia are fusoid, hyaline, aseptate, guttulate, smooth, thin-walled, and acutely rounded at each end (Sutton 1980; Quaedvlieg et al. 2013). Wijayawardene et al. (2012) placed this genus in Ascomycota, genera incertae sedis. Quaedvlieg et al. (2013) introduced two new species, P. collariana and P. obscura and placed them in Dothioraceae, based on molecular phylogeny and stated that P. donacis (CBS 291.69, 313.68 and 417.51), the type species of Pseudoseptoria formed a monophyletic lineage with P. collariana and P. obscura. In this study P. collariana and P. obscura clustered in Aureobasidiaceae as a sister clade to Selenophoma linicola (CBS 468.48). Considering these facts we accept Pseudoseptoria in Aureobasidiaceae as a distinct genus.

Pseudosydowia K. M. Thambugala & K. D. Hyde gen. nov., Index Fungorum number: IF 550733, Facesoffungi number: FoF00105 Etymology:  Referring to its similarity with Sydowia 

Type species: Pseudosydowia eucalypti (Verwoerd and du Plessis) K. M. Thambugala & K. D. Hyde sp. comb. Nov., Index Fungorum number: IF 550734; Facesoffungi number: FoF00106

≡ Sydowia eucalypti (Verwoerd & du Plessis) Crous, in Crous et al., Sydowia 55(2): 143 (2003)

≡ Selenophoma eucalypti Crous, C.L. Lennox & B. Sutton, Mycol. Res. 99(6): 648 (1995)

≡ Sphaerulina eucalypti Verwoerd & du Plessis, S. Afr. J. Sci. 28: 296 (1931)

Leaf spots amphigenous, sub circular, becoming confluent, covering large areas of the leaf, pale brown, surrounded by a narrow, raised, dark brown margin. Mycelium internal, medium brown, consisting of septate, branched, smooth hyphae. Sexual state: Ascostromata amphigenous, black, subepidermal, becoming erumpent, separate or aggregated in clusters of up to 8, globose, apical ostiole wall consisting of several layers of dark brown textura angularis. Asci eight-spored, bitunicate, fasciculate, broadly ellipsoid to clavate, straight to slightly curved, sub sessile with a well-developed ocular chamber. Ascospores overlapping, bi- to multiseriate, hyaline, (1-)3(−4)-septate at maturity, constricted at the median septum, aguttulate, thin-walled, straight to slightly curved, obovoid to ellipsoid with obtuse ends, tapering towards both ends, but more prominently towards the lower end (Crous et al. 2003). Asexual state: Conidiomata pycnidial to avervular, amphigenous, subepidermal, dark brown, dehiscence by irregular rupture of the upper wall. Conidiomata wall comprising 3 to 6 layers of dark brown to black cells of textura angularis. Conidiophores reduced to conidiogenous or hyaline, smooth, branched, 1 to 7-septate, constricted at the septa after incubation in moist chambers. Conidiogenous cells annellidic, integrated, indeterminate, formed from the inner cells of the conidiomatal wall, hyaline to pale brown, smooth-walled, slightly thick-walled, ampulliform to cylindrical, slightly tapered toward the apex. Conidia aseptate, medium brown to olivaceous-brown, ellipsoid to ovoid, aguttulate, thin-walled (Crous et al. 2003; Cheewangkoon et al. 2009).

Notes: Sydowia eucalypti was originally described by Verwoerd and du Plessis (1931) as Sphaerulina eucalypti. Crous et al. (2003) accommodated this species in Sydowia based on its Sclerophoma and Hormonema asexual states and the thick-walled ascostromata. His phylogenetic analysis also showed that of Sydowia eucalypti clustered in Dothioraceae, Dothideales. Sydowia eucalypti is linked to the sexual morph of Selenophoma eucalypti, which clustered with other Aureobasidium and Hormonema species (Crous et al. 2003; Cheewangkoon et al. 2009). In our study Sydowia eucalypti clustered in Aureobasidiaceae with species of Aureobasidium, Selenophoma and Kabatiella. Therefore, we propose a new monotypic genus Pseudosydowia for Sydowia eucalypti.

Saccothecium Fr., Fl. Scan.: 349 (1836), Facesoffungi number: FoF00107

Synonyms

Metasphaeria Sacc., Syll. fung. (Abellini) 2: 156 (1883)

Phaeodothiora Petr., Sydowia 2(1–6): 82 (1948)

Pleosphaerulina Pass., Atti R. Acad. Lincei, Rendiconti Cl. Sci. Fis., sér. 5 7(2): 46 (1891)

Schizostege Theiss., Annls mycol. 14(6): 415 (1917) [1916]

Parasitic or saprobic on woody branches and rarely on leaves in terrestrial habitats. Sexual state: Ascomata black, immersed to erumpent, solitary or gregarious, globose to subglobose, usually uniloculate, thick-walled, sometimes ostiolate. Peridium composed of several layers of brown to dark brown pseudoparenchymatous cells of textura angularis. Hamathecium lacking pseudoparaphyses. Asci eight-spored, bitunicate, fissitunicate, saccate to broad-clavate, becoming cylindric-clavate just before rupture, with a short bifurcate pedicel or apedicellate, apex broadly rounded with a distinct ocular chamber. Ascospores biseriate to triseriate, partially overlapping, hyaline, 3 to many septate, muriform to phragmosporous, obovoid or elliptic with broadly to narrowly rounded ends. Asexual state: aureobasidium-like; conidia hyaline or brownish, ovate, aseptate (Barr 1972).

Notes: Saccothecium was introduced by Fries (1836) and Kirschstein (1938, 1939) introduced several species which had been assigned to Metasphaeria, Sphaerulina, Leptosphaeria and Sphaeria. Kirschstein (1939, 1941) added two new species to Saccothecium, while Vasilyeva and Mel’nik (2006) introduced Saccothecium cornicola Lar. N. Vassiljeva,. Pande (2008) placed several species in Saccothecium, which had been classified under Pringsheimia or Metasphaeria, but most are invalid (MycoBank 2014). Wehmeyer (1957) and Holm (1957) proposed to lectotypify the genus with Saccothecium sepincola. Froidevaux (1972) and Sivanesan (1984) synonymized Saccothecium sepincola under Pringsheimia sepincola and they considered P. sepincola as the type species of Pringsheimia. Saccothecium has been assigned to Dothioraceae, Dothideales (Barr 1972; 1987a and 2001; Kirk et al. 2008; Lumbsch and Huhndorf 2010) based on similar morphological characters of peridium, asci and ascospores (Barr 1972), while Pringsheimia is considered to be distinct from Saccothecium (Barr 1987a; 2001). Molecular phylogenetic analysis, Schoch et al. (2009) shows that Saccothecium sepincola (CBS 278.32) clustered within Didymellaceae and Zhang et al. (2012) included Saccothecium in Pleosporales genera incertae sedis. As there are no morphology-molecular link between the strain CBS 278.32 and S. sepincola it is difficult to show the affinities of Saccothecium in Pleosporales. However, we collected S. sepincola from Italy and directly isolated DNA from ascostromata. Our collection of S. sepincola from Italy clustered in Aureobasidiaceae in our phylogenetic analysis. Therefore, we assign Saccothecium in Aureobasidiaceae.

Type species: Saccothecium sepincola (Fr.) Fr. [as ‘saepincola’], Summa veg. Scand., Section Post. (Stockholm): 398 (1849), (Fig. 17), Facesoffungi number: FoF00108

Fig. 17

Saccothecium sepincola (MFLU 14–0276). a, b Appearance of immersed ascostromata on the host surface. c, d Section through ascostromata. f–i Bitunicate asci. j–m. Released ascospores. Scale bars: c–e = 50 μm, f–i = 25 μm, j–m = 10 μm

≡ Sphaeria sepincola Fr. [as ‘saepincola’], Observ. mycol. (Havniae) 1: 181 (1815)

Saprobic on twigs, and branches of Rosa and Crataegus sp. Sexual state: Ascomata 105–135 μm high, 80 − 105 μm wide $\left(\overline{x}=121\times 95\mu m,n=10\right)$, black, immersed to erumpent, solitary or scattered, globose to subglobose, usually uniloculate, rarely biloculate without a distinct ostiole. Peridium 14–36 μm $\left(\overline{x}=22\mu m,n=15\right)$ wide, a single layer, composed of brown to lightly pigmented cells of textura angularis, near the base connected to the host tissue. Hamathecium lacking pseudoparaphyses. Asci 38–64 × 11–16 μm $\left(\overline{x}=50\times 14\mu m,n=20\right)$, eight-spored, bitunicate, saccate to broadly clavate or cylindric-, with a short bifurcate pedicel and a distinct ocular chamber. Ascospores 14–20 × 4–6 μm $\left(\overline{x}=16.9\times 5\mu m,n=25\right)$, overlapping biseriate, hyaline, 3–6-septate, sometimes 1 vertical septate, asymmetric, obovoid, fusiform to clavate, with broadly to narrowly rounded ends, with broad upper cells, smooth-walled. Asexual state: Unknown.

Material examined: ITALY, Collinaccia - Castrocaro e Terra del Sole (province of Forlì-Cesena [FC]), on the twigs of Cornus sanguinea (Cornaceae), 3 March 2013, Erio Camporesi (MFLU 14–0276).

Selenophoma Maire, Bull. Soc. bot. Fr. 53: clxxxvii (1907), Facesoffungi number: FoF00109

Synonyms

Falcispora Bubák & Serebrian., Hedwigia 52: 269 (1912)

Ludwigiella Petr., Annls mycol. 20(5/6): 319 (1922)

Neopatella Sacc., in Sydow & Sydow, Annls mycol. 6(6): 530 (1908)

Pseudosarcophoma Urries, An. Jard. bot. Madr. 10: 227 (1952)

Selenophomopsis Petr., Annls mycol. 22(1/2): 182 (1924)

Mycelium immersed, branched, septate, pale brown. Sexual state: Unknown. Asexual state: Conidiomata pycnidial or avervular, separate or aggregated, scattered, immersed or superficial, globose, dark brown, uniloculate, thin-walled; walls 2 to 3 layers thick, composed of brown, thick-walled, large-celled, textura angularis, lacking ostioles, dehiscence by circular or irregular rupture of upper wall. Conidiophores reduced to conidiogenous cells or hyaline, irregularly branched at the base and above, 1 to 3-septate, smooth. Conidiogenous cells enteroblastic, phialidic, discrete, determinate, hyaline to pale brown, smooth, subglobose, obpyriform or obovoid, collarette and channel minute, with apical periclinal thickening, guttulate, formed from the inner cell of the pycnidial wall, slightly tapered toward the apex. Conidia holoblastic, aseptate, falcate, fusiform, ellipsoidal to obovoid, eguttulate or irregularly guttulate, smooth-walled, at first thin-walled, hyaline or pale brown, later pale brown with thicker walls and verruculose.

Type species: Selenophoma catananches Maire [as ‘catanaches’], Bull. Soc. bot. Fr. 53: clxxxvii (1906) MB 182152; Facesoffungi number: FoF00110

Notes: Selenophoma was introduced by Selenophoma (1907) with Selenophoma catananches Maire as the type species. Sutton (1980) accepted five species in Selenophoma and the graminicolous Selenophoma species have been accommodated in Pseudoseptoria which is the earliest available generic name for them. The type species differs from those subsequently included in the genus Selenophoma inasmuch as that a proportion of the conidia become pale brown with thicker walls and develop verrculose ornamentation (Sutton 1980). Crous et al. (2003) introduced Sydowia eucalypti with its Selenophoma asexual state and coniothyrium-like and hormonema-like synanamorphs. Sutton (1996) introduced S. anniae and recently Cheewangkoon et al. (2009) introduced S. australiensis with Hormonema as a synanamorph. In our phylogenetic analysis three Selenophoma species clustered in three different clades. Selenophoma australiensis has been shown to be a sister group to Pseudosydowia, while S. linicola to Pseudoseptoria.

Dothideales, genera incertae sedis

Celosporium Tsuneda & M.L. Davey, Botany 88: 472 (2010), Facesoffungi number: FoF00111

Type species: Celosporium laricicola Tsuneda & M.L. Davey [as ‘larixicolum’], Botany 88(5): 473 (2010), Facesoffungi number: FoF00112

Notes: Celosporium was introduced by Tsuneda et al. (2010) to accommodate a single species C. laricicola. This is characterized by dematiaceous hyphae forming terminal or intercalary, black, irregular shaped cellular clumps, conidiomata with aggregated cellular clumps and 1 to 3 celled, hyaline endoconidia, variable size and shape released by cell-wall dissolution of the conidiogenous cells. Phylogenetic analyses (LSU and ITS) of Tsuneda et al. (2010) shows that C. laricicola clustered in Dothideales. In their ITS analyses C. laricicola nested with endophytes of spruce and slow growing dematiaceous endolithic fungi by forming a strongly supported clade. Although C. laricicola clustered among endophytes and slow growing dematiaceous endolithic fungi, it separates from the other endoconidial dothidealean taxa. Our phylogenetic analysis also confirmed the placement of C. laricicola in Dothideales but it separates from both clades A and B. Therefore, we remain Celosporium in Dothideales, genera incertae sedis.

Genera excluded from Dothideales

Genus and type species descriptions are given unless the genus is monotypic when only the species description is given.

Aplosporellaceae Slippers, Boissin & Crous, Studies in Mycology 76: 41 (2013)

The family Aplosporellaceae was introduced by Slippers et al. (2013) and includes Aplosporella and Bagnisiella in the order Botryosphaeriales. Aplosporellaceae is characterized by multiloculate ascomata, bitunicate, stalked or sessile asci with a well-developed apical chamber and hyaline to pigmented, septate or aseptate ascospores, without a mucoid appendage or sheath. The asexual states of Aplosporellaceae are characterized by uni-multilocular pycnidial conidiomata, hyaline, phialidic conidiogenous cells. proliferating percurrently or with periclinal thickening at apex, hyaline, smooth-walled, septate, hyphae-like, branched or unbranched paraphyses and ellipsoid to sub cylindrical, aseptate conidia initially hyaline becoming pigmented (Slippers et al. 2013).

Bagnisiella Speg., Anal. Soc. cient. argent. 10(5–6): 146 (1880), (Fig. 18), Facesoffungi number: FoF00114

Fig. 18

Bagnisiella australis (S-F227223) a–c Ascostromata on host substrate, d Section of ascostroma. e Asci arrangment within hamathecium f Close up of the peridium. g Dehiscence of asus h–j Asci with short pedicel bearing discontinuously arranged partially overlapping 6–8 ascospores. k Ascus stained with cotton blue reagent. l–m Smooth, hyaline ascospores. n–o Ascospore stained with cotton blue reagent. Scale bars: d = 20 μm, e, f = 500 μm, g = 15 μm, h–k = 5 μm, l–o = 10 μm

Saprobic on branches, wood. Sexual state: Ascostromata black, immersed to erumpent or superficial, pulvinate, scattered or aggregated, apex plane or depressed, coriaceous, multiloculate, with 4 to numerous locules, opening widely and irregularly at maturity, cells of ascostromata arranged in upright rows, blackened externally, dark brown to hyaline cells of textura angularis. Locules globose to subglobose, thicked-walled. Peridium of locules thin-walled, lightly pigmented or dark brown cells of textura angularis. Hamathecium lacking pseudoparaphyses. Asci eight-spored, bitunicate, clavate to cylindro-clavate, with a short pedicel, apically rounded, with a small ocular chamber. Ascospores overlapping uni-triseriate, hyaline to dark brown, aseptate, tapered to pointed ends, elliptic, oblong to ovate, straight to inequilateral, minutely guttulate, smooth-walled without a sheath. Asexual state: Unknown.

Notes: Bagnisiella was introduced by Spegazzini (1880) and Bagnisiella australis Speg. assigned as the type species. Saccardo (1883) placed this genus in Dothideaceae. Theissen (1916) established family Dothioraceae of the Myriangiales and included Bagnisiella in Dothioraceae. Previously (1915) this genus was included in an “Anhang” to their arrangement of the Dothideales. The placement of Bagnisiella in Dothioraceae was followed by Theissen and Sydow (1917), Petrak (1923a); (Barr 1979, 1987a); Hawksworth et al. (1995) and Kirk et al. (2001, 2008). Luttrell (1955) suggested that Bagnisiella may be representing a link between Myriangiales and Dothioraceae. Lumbsch and Huhndorf (2007, 2010) categorized Bagnisiella in Dothideaceae. We examined B. australis, the type of Bagniesiella and its morphology suggests Bagniesiella should be excluded from Dothideales and placed in Botryosphaeriales. Slippers et al. (2013) showed that there is a consistent connection between Aplosporella and Bagniesiella and placed it in family Aplosporellaceae and Bagnisiella reduced to synonymy with Aplosporella. Therefore, we transfer Bagnisiella from Dothideales to Aplosporellaceae based on molecular phylogeny and morphology.

Type species: Bagnisiella australis Speg., Anal. Soc. cient. argent. 10(1): 22 (1880), Facesoffungi number: FoF00115

Parasitic on woody branches. Sexual state: Ascostromata 900–1200 × 1000–1400 μm $\left(\overline{x}=1100\times 1200\mu m,n=10\right)$, black, outwardly grey, flattened at the upper surface, erumpent, solitary or scattered, globose to sub globose, coriaceous, multiloculate, with 5 to numerous locules, cells of ascostromata composed of several layers of dark brown cells of textura angularis. Locules 12.3 − 19.1 × 2.3 − 4.2 μm $\left(\overline{x}=13\times 2.9\mu m,n=24\right)$ rectangular. Peridium comprising several layers of thick-walled, pale brown cells of textura angularis. Hamathecium lacking pseudoparaphyses. Asci 29–49 × 6–9 μm $\left(\overline{x}=39\times 9\mu m,n=10\right)$ 6–8 spored, bitunicate, fissitunicate, clavate to sub-cylindrical, aggregated, short pedicellate, apically rounded with a small ocular chamber. Ascospores 25–40 × 5–9 μm $\left(\overline{x}=29.5\times 8.23\mu m,n=2\right)$ discontinuously arranged partially overlapping, hyaline, aseptate, oblong or ovate, smooth-walled, without a sheath. Asexual state: Unknown.

Material examined: ARGENTINA, La Plata, Punta Lara, on Acacia bonariensis Hook. & Arn (Fabaceae), August 1951, J.C. Lindquist (S-F227223).

Notes: Liu et al. (2012) has observed and illustrated the holotype of Bagnisiella australis in LPS which is immature, but does not appear to be botryosphaeriaceous based on the characters of the sunken ascostromata and cylindrical asci. Therefore, we examined an authentic specimen from S in order to clarify its familial placement.

Botryosphaeriaceae Theiss. & Syd. [as ‘Botryosphaeriacae’], Annls mycol. 16(1/2): 16 (1918)

Botryosphaeriaceae was introduced by Theissen and Sydow (1918) and previously this family had been referred to various orders including Myrangiales, Dothideales, and Pseudosphaeriales/Pleosporales. Schoch et al. (2006) included Botryosphaeriaceae in Botryosphaeriales and the family is characterized by uni- to multilocular ascostromata, hyphae-like, branched or unbranched, septate, pseudoparaphyses, bitunicate, fissitunicate asci with apically rounded with an ocular chamber and hyaline to brown, aseptate to septate smooth to verrucose ascospores. Asexual morphs of Botryosphaeriaceae with uni to multilocular pycnidial conidiomata, with hyaline, phialidic conidiogenous cells and hyaline to pigmented, thin to thick-walled conidia which sometimes have mucoid appendages or sheaths, striations, verrucose walls and germ slits (Liu et al. 2012; Hyde et al. 2013). Kirk et al. (2008) estimated that there are 26 genera and 1517 species in the family, while Liu et al. (2012) accepted 29 genera and approximately 1485 species.

Coccostromella Petr., Sydowia 21: 267 (1968) [1967], Facesoffungi number: FoF00117

Notes: Coccostromella was introduced by Petrak (1967) to accommodate a single species Coccostromella puttemansii which has been referred to Auerswaldia. Lumbsch and Huhndorf (2010) grouped the genera Auerswaldia and Coccostromella within Dothideaceae, while Liu et al. (2012) included Auerswaldia into Botryosphaeriaceae based on molecular and phylogenetic data. Although Lumbsch and Huhndorf (2010) placed Coccostromella in Dothideaceae, presence of filiform pseudoparaphyses and clavate to cylindro-clavate asci and aseptate ascospores exclude it from Dothideaceae. Therefore, we refer Coccostromella to Botryosphaeriaceae.

Type species: Coccostromella puttemansii (Henn.) Petr., Sydowia 21: 267 (1968) [1967], (Fig. 19), Facesoffungi number: FoF00118

Fig. 19

Coccostromella puttemansii (S-F11552, isotype) a Herbarium packet and material. b, c Ascostromata on leaf surface. d Section of ascostroma showing locules. e Close up of cells of ascostromata f Pseudoparaphyses. g–h Asci with 8 ascospores j–m Ascospores. Scale bars: d = 700 μm, e–f = 100 μm, g–i = 50 μm, j–m = 5 μm

≡ Auerswaldia puttemansii Henn., Hedwigia 41: 111 (1902)

Parasitic on leaves. Sexual state: Ascostromata up to 1000 μm diam, black, gregarious, superficial, epiphyllous, sometimes hypophyllous, globose to subglobose, gregarious, globose to subglobose, coriaceous, multiloculate, with 3–4 locules, cells of ascostromata thick-walled, brown textura angularis. Locules 180–350 × 200–360 μm $\left(\overline{x}=265\times 280\mu m,n=15\right)$, globose to subglobose, without individual ostiole, opening by dehiscence. Peridium of locules composed of thin-walled, light brown to hyaline cells of textura angularis. Hamathecium composed of 2.5 − 4 μm wide, filiform, hyphae − like, aseptate, hyaline pseudoparaphyses. Asci 100–125 × 20–34 μm $\left(\overline{x}=113\times 27.5\mu m,n=15\right)$, eight-spored, bitunicate, fissitunicate, clavate to cylindro-clavate, with a short pedicel, apically rounded. Ascospores 24–32 × 8.5 − 15 μm $\left(\overline{x}=27\times 10.8\mu m,n=30\right)$ overlapping uniseriate to biseriate, hyaline, aseptate, ellipsoidal to obovoid, rough-walled. Asexual state: Unknown.

Material examined: BRAZIL, São Paulo, on leaves of Lauracearum (Lauraceae), 01 April 1901, Puttemans (S-F11552, isotype).

Mycosphaerellaceae Lindau, in Engler and Prantl, Nat. Pflanzenfam., Teil. I (Leipzig) 1(1): 421 (1897)

Synonym:

Sphaerellaceae Nitschke, Verh. naturh. Ver. preuss. Rheinl. 26: 74 (1869)

The family Mycosphaerellaceae was introduced by Lindau (1897) with the family type Mycosphaerella Johanson. Hawksworth et al. (1995) placed Mycosphaerellaceae in Dothideales. Kirk et al. (2001) assigned Mycosphaerellaceae to the new order Mycosphaerellales, while Kirk et al. (2008) placed the family in Capnodiales. Mycosphaerellaceae is characterized by uni to multiloculate ascostromata with a central ostiole, bitunicate cylindrical to cylindrical-clavate asci and hyaline to pale yellowish, aseptate or septate ascospores often constricted at the septum. Asexual morphs of Mycosphaerellaceae include mucedinaceous and dematiaceous hyphomycetes (conidiophores solitary, fasciculate, sporodochial or synnematous), or acervular to pycnidial coelomycetes with hyaline to pigmented conidiophores and conidia. Hyde et al. (2013) accepted Mycosphaerellaceae in Capnodiales with 12 sexual and 32 asexual genera.

Mycoporis Clem., Gen. fung. (Minneapolis): 50, 173 (1909), Facesoffungi number: FoF00120

Type species: Mycoporis perexigua (Müll. Arg.) Clem., Gen. fung. (Minneapolis): 1–227 (1909), (Fig. 20), Facesoffungi number: FoF00121

Fig. 20

Mycoporis perexigua (G 00110864, holotype). a Ascomata on the host surface. b Squash mount of ascomata c, d Sections of the ascomata e Peridium. f Ascus in water g–h Asci in lactophenol cotton blue i–j Ascospores stained in cotton blue reagent. Scale bars: a = 1000 μm, b, c = 100 μm, d, e = 50 μm, f–j = 10 μm

≡ Mycoporellum perexiguum Müll. Arg., Nuovo G. bot. ital. 23: 399 (1891)

Parasitic on leaves. Sexual state: Ascomata 120–180 μm, appearing as black spots on the host surface, gregarious, scattered, superficial, very easily removed from the host surface, globose, uniloculate, ostiolate. Peridium 11–18 μm, one-layered, composed of dark to brown cells of textura angularis. Hamathecium lacking pseudoparaphyses. Asci 48–57 × 11–22 μm $\left(\overline{x}=50\times 20\mu m,n=15\right)$, eight-spored, bitunicate, broadly cylindrical to fusiform, sessile, with a large ocular chamber, up to 5–9 μm. Ascospores 22–27 × 4–8 μm $\left(\overline{x}=24\times 7\mu m,n=15\right)$, overlapping, uniseriate at the apex to tri-seriate near the base, hyaline, 5-septate, strongly constricted at the primary septum, broadly fusiform to cylindrical with broadly rounded ends. Asexual state: Unknown.

Notes: Mycoporis was introduced by Clements (1909) as a monotypic genus in order to accommodate Mycoporellum perexiguum Müller Arg. (Mycoporis perexigua). Zahlbruckner (1922) categorized Mycoporis under Mycoporaceae, while Hawksworth et al. (1995) and Lumbsch and Huhndorf (2010) included this genus in Dothideaceae. We examined M. perexiguum which did not match the taxonomic concepts of Dothideaceae or Dothideales as ascomatal characters mainly differ from other dothidealean species but similar with Mycosphaerellaceae. Therefore, we exclude Mycoporis from Dothideales and transfer it to Mycosphaerellaceae.

Material examined: AUSTRALIA, Brisbane, Bailey (G 00110864, holotype).

Pseudoperisporiaceae Toro, in Seaver & Palacios Chardon, Scient. Surv. P. Rico 8(1): 0 (1926)

Synonym:

Epipolaeaceae Theiss. & P. Syd., Annls mycol. 16(1/2): 7(1918)

The family Pseudoperisporiaceae was introduced by Toro in Seaver and Chardón (1926) and is characterized by superficial ascomata surrounded by brown mycelium at the base, brown to red brown, thin-walled, peridium composed of cells of textura angularis, bitunicate, sessile, or short pedicellate asci with small ocular chamber and 1-septate, hyaline to brown ascospores constricted at the septum (Hyde et al. 2013). Chaetosticta is reported as the asexual state of this family (Hyde et al. 2013). Lumbsch and Huhndorf (2010) and Hyde et al. (2013) listed 22 genera in Pseudoperisporiaceae in Dothideomycetes incertae sedis.

Jaffuela Speg., Boln Acad. nac. Cienc. Córdoba 25: 39 (1921), Facesoffungi number: FoF00123

Type species: Jaffuela chilensis Speg., Boletín de la Academia Nacional de Ciencias de Córdoba 25: 39 (1921), (Fig. 21), Facesoffungi number: FoF00124

Fig. 21

Jaffuela chilensis (LPS 667, holotype). a Appearance of ascomata on the host surface. b Mycelium on the host surface. c Mycelium. d, e Sections through ascomata. f, g Asci. h, i Ascospores. Scale bar s: a, b = 500 μm, c = 10 μm, d = 25 μm, e–i = 10 μm

Growing on leaves of Puya chilensis. Sexual state: Mycelium up to 8–13 μm long spreading below host surface which easily removed from the host tissue, composed of dark-brown moniliform shape cells. Ascostromata 100–130 μm high × 90–130 μm diam., black, immersed to erumpent, gregarious, scattered, globose to subglobose, coriaceous, unilocular, composed of one layer of brown cells of textura angularis fused to the host. Hamathecium lacking pseudoparaphyses Asci 40–60 × 20–30 μm $\left(\overline{x}=48\times 25\mu m,n=15\right)$, eight-spored, bitunicate, elliptic-obovate, sessile or short pedicellate, apically rounded with a small ocular chamber. Ascospores 25–30 × 7–9 μm $\left(\overline{x}=27\times 8.5\mu m,n=15\right)$, irregularly arranged, overlapping, 2 to 3 seriate, brown, 1-septate, constricted at the septum, with broadly to narrowly rounded ends, smooth-walled. Asexual state: Unknown.

Notes: Jaffuela was introduced by Spegazzini (1921) based on J. chilensis and has remained monotypic until now. Luttrell (1973) placed this genus in Parodiopsidaceae which is characterized by intact pseudothecia opening by a broad pore or crumbling at the apex. Arx and Müller (1975) accommodated Jaffuela in Dothideaceae, while Lumbsch and Huhndorf (2010) included it in Dothioraceae.

We examined the type specimen of Jaffuela and it does not share common morphology with those genera of Dothideales. However, Jaffuela shows similarities with Pseudoperisporiaceae, such as superficial ascomata surrounded by brown mycelium, brown to red brown, thin-walled, peridium composed of cells of textura angularis, bitunicate, short pedicellate asci with a small ocular chamber and 1-septate, brown ascospores, constricted at the septum Therefore, we tentatively assign Jaffuela to Pseudoperisporiaceae based on the morphology. The type needs recollecting, sequencing and epitypifying in order to confirm its familial status.

Material examined: CHILE, on Puya chilensis Molina (Bromeliaceae), 1918, Spegazzini (LPS 667, holotype).

Teratosphaeriaceae Crous & U. Braun, Stud. Mycol. 58: 8 (2007)

Teratosphaeriaceae was introduced by Crous et al. (2007) with its phylogenetic position within Capnodiales. This family is characterized by pseudothecial, immersed to superficial ascomata with papillate ostioles, branched, septate, anastomosing pseudoparaphyses, bitunicate, obclavate to globose or saccate asci and 1-septate, hyaline ascospores. Crous et al. (2009) accepted Baudoinia, Capnobotryella, Catenulostroma, Devriesia, Penidiella, Phaeothecoidea, Readeriella, Staninwardia and Teratosphaeria in Teratosphaeriaceae based on DNA sequence data derived from the LSU gene. Hyde et al. (2013) accepted 24 sexual and asexual genera in this family.

Pachysacca Syd., Annls mycol. 28(5/6): 435 (1930), Facesoffungi number: FoF00126

Type species: Pachysacca eucalypti Syd., Annls mycol. 28(5/6): 435 (1930), (Fig. 22), Facesoffungi number: FoF00127

Fig. 22

Pachysacca eucalypti (K(M) 176511, holotype). a, b Appearance of ascostromata on the host surface. c Section through ascostroma. d–f Asci. g Ascospores. Scale bars: b = 800 μm, c = 50 μm, d, e = 20 μm

Saprobic on Eucalyptus leaves. Sexual state: Ascostromata up to 900–1200 μm diam $\left(\overline{x}=1150\mu m\right)$, black, immersed at the base, fused with host surface, nearly flattened at the top, solitary or scattered, multiloculate, with 2–4 locules. Peridium of locules composed of pale brown to brown cells of textura angularis. Hamathecium lacking pseudoparaphyses. Asci 120–150 × 36–43 μm $\left(\overline{x}=135\times 41\mu m,n=5\right)$, eight-spored, bitunicate, fissitunicate, narowly cylindrical at the upper and broadly oblong at the end, with a short pedicel. Ascospores 59–65 × 8–10 μm $\left(\overline{x}=63\times 9\mu m,n=10\right)$, over-lapping, uniseriate to multiseriate, hyaline, 1 − 2-septate, narrowly cylindrical, with broadly rounded ends, smooth-walled. Asexual state: Unknown.

Notes: Pachysacca was introduced by Sydow (1930) in order to accommodate P. eucalypti. Later, Swart (1982) introduced another two new species P. pusilla H.J. Swart and P. samuelii (Hansf.) H.J. Swart. Hawksworth et al. (1995) and Lumbsch and Huhndorf (2010) placed this genus in Dothideaceae. We examined and illustrated the type specimen of Pachysacca and it should be excluded from Dothideales as morphology of Pachysacca differs from other members of Dothideales that we accept here. Pachysacca shares morphologically similar characterstics with Teratosphaeriaceae. Therefore, we tentatively refer Pachysacca in Teratosphaeriaceae.

Material examined: AUSTRALIA, Noorlunga, on Eucalyptus rostrata Schlecht. (Myrtaceae), May 1924, G. Samuel (K(M)176511, holotype).

Dothideomycetes, genera incertae sedis

Dothideomycetes is the largest class in the division Ascomycota and is characterized by bitunicate (fissitunicate) asci. Lumbsch and Huhndorf (2010) included 34 families and over 150 genera in Dothideomycetes incertae sedis in the Outline of Ascomycota—2009, while Hyde et al. (2013) included 26 families. We exclude the following genara from Dothideales and include them in Dothideomycetes, genera incertae sedis pending fresh collections and phylogenetic investigations.

Botryochora Torrend, Brotéria, sér. bot. 12: 65 (1914), Facesoffungi number: FoF00128

Type species: Botryochora nigra (Torrend) Torrend, Brotéria, sér. bot. (1914), (Fig. 23),

Fig. 23

Botryochora nigra (BPI 643038). a. Herbarium packet b Conidiomata on the host surface. c Section through conidioma d Conidiophores and developing conidia. e Conidia. Scale bars: b = 1000 μm, c = 200 μm, d–e = 20 μm

Facesoffungi number: FoF00129

Saprobic on bark. Sexual state: not observed. Asexual state: Conidiomata 1272–1704 (−1941) μm diam $\left(\overline{x}=1673\mu m,n=5\right)$, black, solitary or scattered, immersed at base, superficial, globose to subglobose, multiloculate with 3–5 locules. Peridium composed of pale to brown cells of textura angularis. Conidiophores (10−)13 − 16(−20) × 5–5.5 μm long $\left(\overline{x}=16\times 5\mu m,n=20\right)$, hyaline, unbranched. Conidiogenous cells integrated, philaidic. Conidia 15.5 − 19 (−20) × 9.5 − 11 μm $\left(\overline{x}=18\times 10\mu m,n=30\right)$, hyaline, aseptate, oval-ellipsoid, rounded to narrow ends, smooth-walled.

Notes: Botryochora was introduced by Torrend (1914) in order to accommodate Megalonectria nigra Torrend which was classified in Nectriaceae, Sordariomycetes. Hawksworth et al. (1995) and Lumbsch and Huhndorf (2010) placed this genus in Dothioraceae. Although the sexual morph could not be found on this specimen, we observed the asexual state associated with in same herbarium material. The sexual state is characterized by black, erumpent to superficial, multiloculate ascostromata, eight-spored asci and brown ascospores (Saccardo 1925–1928). We could not observe the sexual state in the herbarium material and the morphology of its asexual state does not justify placement in Dothideales. Therefore, we transfer Botryochora to Dothideales, genera incertae sedis. Recollection, epitypification and molecular analysis are required to confirm the placement of Botryochora.

Material examined: MOZAMBIQUE, Zumbo, May 1913, A. Cruz (BPI 643038); CONGO, Kisantu, leg. by H. Vanderyst (URM 9415, 9416).

Lucidascocarpa A. Ferrer, et al., Mycologia 100(4): 642 (2008)

Type species: Lucidascocarpa pulchella A. Ferrer et al. Mycologia 100(5): 644 (2008)

Notes: Lucidascocarpa was introduced by Ferrer et al. (2008) in order to accommodate Lucidascocarpa pulchella. Based on ascostromatic ascomata, fissitunicate asci and lack of pseudoparaphyses, they placed Lucidascocarpa in Dothideaceae. Lucidascocarpa is distinct from other species of Dothideaceae as it does not share common morphologies. In L. pulchella, ascomata are white, with long periphysate necks, peridium comprises hyaline textura angularis cells and ascospores have seven septate and surrounded by a large gelatinous sheath. Therefore, we exclude Lucidascocarpa from Dothideaceae and transfer in to Dothideomycetes genera incertae sedis. Lucidascocarpa shares some morphological affinities with Tubeufiaceae such as superficial, unilocular, globose-subglobose, papillate ascomata, eight-spored, clavate, pedicellte asci and multiseptate, hyaline ascospores. Lucidascocarpa differs from other species of Tubeufiaceae in having hyaline ascomata with a long cylindrical neck, a hamathecium lacking pseudoparaphyses, and verruculose ascospores with a large gelatinous sheath.

Omphalospora Theiss. & Syd., Annls mycol. 13(3/4): 361 (1915), Facesoffungi number: FoF00132

Synonyms

Plectosphaerella Kirschst., Krypt. − Fl. Brandenburg (Leipzig) 7(3): 310 (1938)

Plectosphaerina Kirschst., Annls mycol. 36(5/6): 368 (1938)

On leaves and stalks of dicotyledons. Sexual state: Ascostromata forming shiny black erumpent bands on both leaf surfaces between the cuticle and epidermis, gregarious, superficial to immersed, crustose, multiloculate, locules globose to subglobose, without a distinct ostiole. Peridium consists of two layers of cells; inner layer vertically arranged dark-brown cells of textura angularis, outer layer; consist with cells of textura epidermoidea. Hamathecium lacking pseudoparaphyses. Asci eight-spored, bitunicate, broadly cylindrical to subglobose or ovoid, rounded at the apex, sessile or very short pedicellate. Ascospores overlapping, uniseriate to biseriate, hyaline, 1-septate, septate near the base, not constricted at the septum, ovoid to oblong with broadly rounded ends, tapered to a pointed base, smooth-walled (Theissen and Sydow 1915; Barr 1972). Asexual state: Podoplaconema (Wijayawardene et al. 2012).

Notes: Omphalospora was introduced by Theissen and Sydow (1915) to accommodate Omphalospora ambiens (Lib.) Theiss. & Syd. and O. stellariae (Lib.) Theiss. & Syd. which have been referred to Dothidea. It is a very poorly studied genus. Höhnel (1919) introduced three new species O. himantia (Pers.) Höhn., O. melaena (Fr.) Höhn. and O. silenes (Niessl) Höhn. which were classified in Sphaeria and Asteroma, but later O. himantia (Pers.) Höhn and O. silenes (Niessl) Höhn has been excluded from Omphalospora. Omphalospora stellerae Murashk., O. tragacanthae (Lév.) Petr. and O. acanthaceifolii Bat. et al. have been introduced recently. No molecular data are available for this genus in GenBank. Omphalospora is considerd as a genus of Dothideaceae (Von Arx and Müller 1975; Barr 1987a; Hawksworth et al. 1995; Lumbsch and Huhndorf 2010), but we believe Omphalospora belongs to neither Dothideaceae nor Dothideales and tentatively we refer this genus in to Dothideomycetes, genera incertae sedis. Podoplaconema is considered as the asexual state of Omphalospora (Wijayawardene et al. 2012). More molecular and morphology studies are needed to resolve these affinities.

Type species: Omphalospora stellariae (Lib.) Theiss. & Syd., Annls mycol. 13(3/4): 361 (1915), (Fig. 24), Facesoffungi number: FoF00133

Fig. 24

Omphalospora stellariae (W 421). a Herbarium material. b, c Ascostromata on the host surface. d Section through ascostroma. e, f Asci in lacto-phenol cotton blue reagent. g Ascus in water. h, i Ascospores. Scale bars: b = 100 μm, d = 50 μm, e–g = 10 μm, h–i = 5 μm

≡ Dothidea stellariae Lib., Pl. crypt. Arduenna, fasc. (Liège) 2(nos 101–200): no. 172 (1832).

For other synonyms see Index Fungorum

On leaves and stems of Stellaria nemorum in terrestrial habitats. Sexual state: Ascostromata forming black areas on the host, superficial, gregarious, scattered, elongate, multiloculate, with 5–6 locules, cells of ascostromata composed of few layers of brown cells of textura angularis. Locules globose to subglobose, non-ostiolate. Peridium of locules one-layered, composed of brown to lightly pigmented cells of textura angularis. Hamathecium lacking pseudoparaphyses. Asci 18–25 × 7–12 μm $\left(\overline{x}=21\times 10\mu m,n=15\right)$, eight-spored, bitunicate, broadly cylindrical to clavate, sessile, apex narrowly rounded. Ascospores 6–9 × 4–5.6 μm $\left(\overline{x}=8\times 5\mu m,n=15\right)$, overlapping, uniseriate to triseriate, hyaline, 1-septate, apiosporous, slightly constricted at the septum, ovoid to oblong with broadly rounded ends, smooth-walled . Asexual state: Unknown.

Material examined: GERMANY, Brandenburg Province, on Stellaria nemorum L. (Caryophyllaceae), 18 May 1910, indentified by Otto Jaap. (W421).

Yoshinagaia Henn., Hedwigia 43: 143 (1904), Facesoffungi number: FoF00136

Type species: Yoshinagaia quercus Henn., Hedwigia 43: 143 (1904), (Fig. 25), Facesoffungi number: FoF00137

Fig. 25

Yoshinagaia quercus (IMI 348745, holotype). a, b Appearance of ascostromata on the host surface. c Vertical section through ascostroma. d, e Close up of the peridium. f–i Asci. Scale bars: c = 500 μm. d, e = 100 μm. f–i = 50 μm

≡ Japonia quercus Höhn., Sber. Akad. Wiss. Wien, Math.-naturw. Kl., Abt. 1 118: 880 [69 repr.] (1909)

Parasitic on living leaves of Cyclobalanopsis morii. Sexual state: Ascostromata scattered, erumpent on both sides of the leave surface, with irregular pore at the surface, globose, black. Peridium uniloculate, with a foot-like hypostroma at the base connected to the host surface, surrounded by the brown to black pseudoparenchymatous cells. Hamathecium lacking psudoparaphyses. Asci 190–220 × 19–27 μm $\left(\overline{x}=210\times 23\mu m,n=10\right)$, eight-spored, bitunicate, fissitunicate, cylindrical, rounded at the apex, with a pedicel up to 30 μm. Ascospores 19–25 × 11–15 μm $\left(\overline{x}=22\times 13\mu m,n=10\right)$, uniseriate, sometimes overlapping, hyaline, aseptate oval to oblong, with broadly rounded ends. Asexual state: Japonia quercus, develops in the same stromatic locule; conidiogenous cells are holoblastic; hyaline, conidia are 2-septate, long fusoid to navicular, and bear an apical branched appendage (asexual morph description follows Barr 2001).

Notes: Yoshinagaia was established by Hennings (1904) as a monotypic genus and has remained monotypic until now. Eriksson and Hawksworth (1993) placed Yoshinagaia in Seuratiaceae based on its ascostromatic character. Sivanesan and Hsieh (1995) re-appraised the systematic status of Yoshinagaia, discussed the similarity in the morphology between Bagnisiella and Yoshinagaia and placed this genus under the family Dothioraceae based on ascomatic, hamathecial, ascus and ascospore characters. Lumbsch and Huhndorf (2010) in outline of Ascomycota, also included Yoshinagaia in the family Dothioraceae, but in this study we exclude this genus from Dothideales to Dothideomycetes genera incertae sedis as it is not typical of any existing family of Dothideomycetes.

Material examined: TAIWAN, on the leaves of Cyclobalanopsis morii (Hayata) Schottky (Fagaceae), 29 May 1991, W.H. Hsieh (IMI 348745, holotype).

Sordariomycetes

Hyponectriaceae Petr., Annls mycol. 21(3/4): 305 (1923)

Hyponectriaceae was introduced by Petrak (1923b) which is typified by Hyponectria buxi and accommodates the genera Hyponectria and Anisostomula. Barr (1994) placed Hyponectriaceae in Xylariales, Wang and Hyde (1999) refer eleven genera to the Hyponectriaceae, while Lumbsch and Huhndorf (2010) treated Hyponectriaceae in Xylariales with 16 genera. The family Hyponectriaceae is characterized by immersed, erumpent or nearly superficial ascomata, septate paraphyses, unitunicate, eight-spored asci and one-celled or one to several septate ascospores, surrounded by mucilaginous sheath (Barr 1990; Wang and Hyde 1999).

Discosphaerina Höhn., Sber. Akad. Wiss. Wien, Math.-naturw. Kl., Abt. 1 126(4–5): 353 (1917), (Fig. 26), Facesoffungi number: FoF00138

Fig. 26

Discosphaerina discophora (S-F10728, holotype). a Herbarium material. b, c Ascomata on host surface. d Section through ascoma. e Peridium. f–h Apparetly unitunicate asci. i Ascospores. Scale bars: d = 50 μm, e–g = 25 μm, h = 15 μm, i = 10 μm

Saprobic on leaves and twigs in terrestrial habitats. Sexual state: Ascomata black, depressed, globose to tympaniform, scattered to aggregated, immersed to superficial, visible as minute black dots on host tissue, uniloculate, ostiolate at maturity usually by dehiscence of the thin wall at the apex. Peridium 2-layered: outer layer comprising several layers, dark brown to black cells of textura angularis, inner layer of fattened cells, 2–3 layers of hyaline to light brown cells of textura angularis. Paraphyses not observed. Asci eight-spored, ?unitunicate, oblong to ovoid, sessile to sub-sessile, apically rounded with an ocular chamber. Ascospores overlapping biseriate to triseriate, hyaline, elliptic, obovate, oblong to fusoid, aseptate, smooth-walled. Asexual state: Unknown.

Notes: Discosphaerina was introduced by Höhnel (1917) and typified by D. discophora. Höhn. The placement of this genus is still confused as the unitunicate nature of D. discophora is unclear and some members of this genus have bitunicate asci. Barr (1972) treated Discosphaerina in Dothideaceae, Dothideales and accepted five species in this genus. Sivanesan (1984) also placed this genus in Dothideaceae with nine species. Lumbsch and Huhndorf (2010) placed Discosphaerina in Hyponectriaceae. The phylogenetic placement of Discosphaerina (=Columnosphaeria) fagi was confirmed in Dothideaceae, Dothideales (Schoch et al. 2006; 2009; Zalar et al. 2008) and some Discosphaerina species produce Aureobasidium, Kabatia, Sarcophoma and Selenophoma asexual morphs in culture (Sivanesan 1984; Barr 1972). We examined the type of Discosphaerina and D. fagi in order to obtain a better morphological understanding of this problem. We believe that D. discophora should not be placed in Dothideales because of the apparently unitunicate nature of its asci (Fig. 26 f, h). Our phylogenetic analysis shows a close relationship between Discosphaerina (Columnosphaeria) fagi and the Aureobasidium pullulans group which cluster in the same clade in the family Aureobasidiaceae. Discosphaerina fagi has been classified under Columnosphaeria (Barr 2001) and we discuss this further in Columnosphaeria section. Fresh collections of D. discophora are needed to establish a natural classification.

Type species: Discosphaerina discophora Höhn., Sber. Akad. Wiss. Wien, Math.-naturw. Kl., Abt. 1 126(4–5): 353 (1917), Facesoffungi number: FoF00139

≡ Guignardia discophora (Höhn.) Petr., Annls mycol. 19(1/2): 111 (1921)

Saprobic on leaves and twigs in terrestrial habitats. Sexual state: Ascomata 68–122 μm high 90–130 μm wide, black, depressed, globose to tympaniform, scattered to aggregated, immersed to superficial, visible as minute black dots on host tissue, uniloculate, ostiolate at maturity usually by dehiscence of the thin wall at the apex. Peridium 15–30 μm thick, 2-layered: outer layer thick, comprising several layers of dark brown to black cells of textura angularis, inner layer thin, of fattened cells, 2–3 layers of hyaline to light brown cells of textura angularis. Paraphyses not observed. Asci 28–45 × 6–8.5 μm $\left(\overline{x}=37\times 7.5\mu m,n=15\right)$, eight-spored, ?unitunicate, oblong to ovoid, sessile to sub-sessile, apically rounded with an ocular chamber. Ascospores 8–12 × 2.4 − 3.8 μm $\left(\overline{x}=9.5\times 3\mu m,n=15\right)$, overlapping biseriate to triseriate, hyaline, elliptic, obovate, oblong to fusoid, aseptate, smooth-walled. Asexual state: Unknown.

Material examined: CZECH REPUBLIC, Theusing in Böhmen, Bernklau, on Solidago virgaurea L. (Asteraceae), June 1917, R. Steppan (S-F10728, holotype).

Discussion

In this pape,r we re-examined the types or representative specimens of dothidealean genera including the genera which were previously referred to Dothideales by various authors. Based on the molecular phylogeny and modern taxonomic concepts, we accept two families in Dothideales including the Dothideaceae and new family Aureobasidiaceae. Dothioraceae is not recognized as a distinct family and is synonymized under Dothideaceae. However, several genera Columnosphaeria, Dictyodothis, Dothiora, Endodothiora, Kabatina, Pringsheimia, Selenophoma and Sydowia which do not have ex-type sequences need to be re-collected and sequenced in order to confirm their familial placements. We exclude several genera from Dothideales to appropriate families or Dothideomycetes, genera incertae sedis or Sordariomycetes genera incertae sedis based on morphology and modern taxonomic concepts.

In this study, we re-collected and described P. ribesia and it forms a separate subclade in Dothideaceae while, other Plowrightia species (P. abietis and P. periclymeni) grouped in another separate clade along with Delphinella strobiligena, Sydowia polyspora, Phaeocryptopus nudus and Rhizosphaera species. This clade may represent a new genus, but needs more sequenced strains especially Hormonema and Kabatina in order to resolve their phylogenetic placement in Dothideaceae.

Dothideaceae is characterized by immersed to erumpent or superficial, uni or multiloculate ascostromata, 8- or poly-spored, bitunicate asci and hyaline or brown, transversely septate, sometimes muriform ascospores while Aureobasidiaceae consists of immersed to erumpent, uniloculate ascostromata, eight-spored, bitunicate asci with a short bifurcate pedicel or apedicellate and hyaline, 3 to many septate or aseptate, muriform to phragmosporous, obovoid or elliptic ascospores with broad to narrow rounded ends. The asexual morphs of Dothideales are coelomycetous or hyphomycetous with a wide host range. The asexual genera which we accept in Dothideaceae include only coelomycetous species (Coleophoma, Cylindroseptoria, Endoconidioma, Kabatina, Neocylindroseptoria) while Aureobasidiaceae consist with both coelomycetous and hyphomycetous species (Aureobasidium, Kabatiella, Pseudoseptoria and Selenophoma). However, there are hyphomycetous species that have been reported as the asexual morphs of Dothideaceae. Rhizosphaera species are known as the asexual states of Phaeocryptopus (Orton 1915), while the type of Hormonema, H. dematioides Lundberg and Melin has been suggested as the asexual state of Sydowia polyspora (Butin 1970; Cheewangkoon et al. 2009). We do not refer Hormonema and Rhizosphaera as separate genus in Dothideaceae and further morphology and phylogeny studies are needed. Wijayawardene et al. (2012) and Hyde et al. (2013) listed Podoplaconema under Dothideaceae as an asexual genus. In this study, we transfer Omphalospora to Dothideomycetes, genera incertae sedis. Therefore, the placement of Podoplaconema which is known as the asexual state of Podoplaconema still confused, as no molecular data are available in GenBank.

Aureobasidiaceae has 65/80 % bootstrap support in the phylogenetic tree (Fig. 1) we propose as a new family based on both morphology and phylogeny. Previous phylogenetic studies (Lumbsch and Lindemuth 2001; Schoch et al. 2006, 2009; Boonmee et al. 2012; Hyde et al. 2013) also showed that the separation of A. pullulans and other associated species which we accommodate in Aureobasidiaceae with high bootstrap support in Dothideales. In this study we used only ITS, SSU and LSU in our phylogenetic analysis due to the unavailability of other genes in GenBank. We suggest to include protein-coding genes (EF, RBP2 or BTUB) in future phylogenetic analysis of this order as it is evident that Aureobasidium species form a species complex (Zalar et al. 2008).