Families of Dothideomycetes

1. 1.

   Acrospermaceae (DQ Dai, KD Hyde, EBG Jones, J Taylor, NN Wijayawardene)*

2. 2.

   Aigialaceae (EBG Jones, KD Hyde, S Suetrong)

3. 3.

   Aliquandostipitaceae (EBG Jones, S Suetrong, KD Hyde, H Raja, CA Shearer, NN Wijayawardene)

4. 4.

   Anteagloniaceae (A. Mapook, KD Hyde, JK Liu)

5. 5.

   Amniculicolaceae (H Ariyawansa, KD Hyde, EBG Jones, NN Wijayawardene)

6. 6.

   Antennulariellaceae (P Chomnunti, KD Hyde, EBG Jones)

7. 7.

   Argynnaceae (DQ Dai, KD Hyde, NN Wijayawardene)

8. 8.

   Arthopyreniaceae (R Lücking, MP Nelsen)

9. 9.

   Ascoporiaceae (S Boonmee, KD Hyde, EBG Jones)

10. 10.

    Asterinaceae (S Hongsanan, KD Hyde, YM Li, NN Wijayawardene)

11. 11.

    Aulographaceae (S Boonmee, KD Hyde, EBG Jones)

12. 12.

    Bambusicolaceae (DQ Dai, KD Hyde, NN Wijayawardene)

13. 13.

    Biatriosporaceae (KD Hyde, EBG Jones, S Suetrong)

14. 14.

    Botryosphaericeae (EBG Jones, JK Liu, A Dissanayake, KD Hyde, NN Wijayawardene, JY Yan)

15. 15.

    Brefeldiellaceae (KD Hyde, H Ariyawansa, NN Wijayawardene)

16. 16.

    Capnodiaceae (P Chomnunti, KD Hyde, EBG Jones)

17. 17.

    Cladosporiaceae (P Crous, NN Wijayawardene)

18. 18.

    Coccoideaceae (S Boonmee, KD Hyde, EBG Jones)

19. 19.

    Coniothyriaceae (P Crous, NN Wijayawardene)

20. 20.

    Cookellaceae (S Boonmee, KD Hyde, EBG Jones)

21. 21.

    Corynesporascaceae (P Chomnunti, KD Hyde, EBG Jones, NN Wijayawardene)

22. 22.

    Cucurbitariaceae (M Doilom, H Ariyawansa, KD Hyde, NN Wijayawardene)

23. 23.

    Dacampiaceae (P Diederich, R Phookamsak, KD Hyde, JD Lawrey, NN Wijayawardene)

24. 24.

    Delitschiaceae (IC Senanayake, F Doveri, KD Hyde, EBG Jones)

25. 25.

    Diademaceae (H Ariyawansa, KD Hyde, EBG Jones, NN Wijayawardene)

26. 26.

    Didymellaceae (H Ariyawansa, PW Crous, M Doilom, KD Hyde, NN Wijayawardene)

27. 27.

    Didymosphaeriaceae (H Ariyawansa, EBG Jones, KD Hyde, NN Wijayawardene)

28. 28.

    Dissoconiaceae (PW Crous, NN Wijayawardene)

29. 29.

    Dothideaceae (KM Thambugala, H Ariyawansa, KD Hyde, NN Wijayawardene)

30. 30.

    Dothidotthiaceae (H Ariyawansa, KD Hyde, EBG Jones)

31. 31.

    Dycfrolomycetaceae (KL Pang, SA Alias, KD Hyde, S Suetrong)

32. 32.

    Elsinoaceae (R Jayawardena, KD Hyde, NN Wijayawardene)

33. 33.

    Englerulaceae (DQ Dai, KD Hyde, EBG Jones, NN Wijayawardene)

34. 34.

    Eremomycetaceae (DQ Dai, KD Hyde, EBG Jones, NN Wijayawardene)

35. 35.

    Euantennariaceae (DQ Dai, S Wikee, KD Hyde, EBG Jones, NN Wijayawardene)

36. 36.

    Fenestellaceae (R Phookamsak, KD Hyde, EBG Jones)

37. 37.

    Gloniaceae (E Boehm, M Doilom, KD Hyde)

38. 38.

    Halojulellaceae (H Ariyawansa, KD Hyde, EBG Jones, KL Pang, S Suetrong)

39. 39.

    Halotthiaceae (EBG Jones, KD Hyde, S Suetrong)

40. 40.

    Hypsostromataceae (P Chomnunti, KD Hyde, EBG Jones)

41. 41.

    Hysteriaceae (E Boehm, M Doilom, KD Hyde, NN Wijayawardene)

42. 42.

    Kirschsteiniotheliaceae (S Boonmee, KD Hyde, EBG Jones)

43. 43.

    Lentitheciaceae (K Tanaka, K. Hirayama, KD Hyde)

44. 44.

    Leptopeltidaceae (R Phookamsak, KD Hyde, EBG Jones)

45. 45.

    Leptosphaeriaceae (H Ariyawansa, KD Hyde, EBG Jones, NN Wijayawardene)

46. 46.

    Lichenoconiaceae (P Diederich, JD Lawrey)

47. 47.

    Lichenotheliaceae (L Muggia, C Gueidan, K Knudsen)

48. 48.

    Lindgomycetaceae (CA Shearer, K Hirayama, AN Miller, H Raja, K Tanaka)

49. 49.

    Lophiostomataceae (H Ariyawansa, Q Tian, KD Hyde, EBG Jones, NN Wijayawardene)

50. 50.

    Lophiotremataceae (Y Zhang, H Ariyawansa, K Hirayama, K Tanaka)

51. 51.

    Manglicolaceae (EBG Jones, KD Hyde, S Suetrong)

52. 52.

    Massariaceae (KD Hyde, EBG Jones)

53. 53.

    Massarinaceae (Y Zhang, KD Hyde, EBG Jones, WJ Li, K Tanaka)

54. 54.

    Melanommataceae (H Ariyawansa, Q Tian, KD Hyde, EBG Jones, NN Wijayawardene)

55. 55.

    Mesnieraceae (P Chomnunti, KD Hyde, EBG Jones)

56. 56.

    Metacapnodiaceae (P Chomnunti, KD Hyde, EBG Jones, NN Wijayawardene)

57. 57.

    Micropeltidaceae (HX Wu, KD Hyde, EBG Jones, NN Wijayawardene)

58. 58.

    Microthyriaceae (HX Wu, KD Hyde, EBG Jones, NN Wijayawardene)

59. 59.

    Monoblastiaceae (R Lücking, MP Nelsen, A Aptroot, F Schumm)

60. 60.

    Montagnulaceae (H Ariyawansa, KD Hyde, EBG Jones, NN Wijayawardene)

61. 61.

    Morosphaeriaceae (EBG Jones, H Ariyawansa, KD Hyde, S Suetrong, NN Wijayawardene)

62. 62.

    Muyocopronaceae (KD Hyde, S Boonmee)

63. 63.

    Mycoporaceae (S Boonmee, KD Hyde, H Sipman, EBG Jones)

64. 64.

    Mycosphaerellaceae (R Phookamsak, U Braun, PW Crous, KD Hyde, EBG Jones, NN Wijayawardene)

65. 65.

    Myriangiaceae (A Dissanayake, R Jayawardena, KD Hyde, EBG Jones)

66. 66.

    Mytilinidiaceae (E Boehm, M Doilom, KD Hyde, NN Wijayawardene)

67. 67.

    Naetrocymbaceae (DQ Dai, MP Nelsen, R Lücking, P Diederich)

68. 68.

    Natipusillaceae (H Raja, AN Miller, CA Shearer)

69. 69.

    Paranectriellaceae (S Boonmee, KD Hyde)

70. 70.

    Parmulariaceae (DQ Dai, Q Tian, KD Hyde, EBG Jones, NN Wijayawardene)

71. 71.

    Parodiellaceae (P Chomnunti, KD Hyde)

72. 72.

    Patellariaceae (S Yacharoen, DJ Bhat, KD Hyde, EBG Jones, NN Wijayawardene)

73. 73.

    Perisporiopsidaceae (S Boonmee, KD Hyde, EBG Jones)

74. 74.

    Phaeosphaeriaceae (R Phookamsak, KD Hyde, EBG Jones, NN Wijayawardene)

75. 75.

    Phaeotrichaceae (JK Liu, KD Hyde, EBG Jones)

76. 76.

    Phyllostictaceae (S Wikee, P Crous, JK Liu, KD Hyde, EBG Jones)

77. 77.

    Piedraiaceae (P Chomnunti, KD Hyde, EBG Jones)

78. 78.

    Planistromellaceae (J Monkai, JK Liu, KD Hyde, EBG Jones)

79. 79.

    Platystomaceae (KL Pang, EBG Jones)

80. 80.

    Pleomassariaceae (H Ariyawansa, KD Hyde, EBG Jones, NN Wijayawardene)

81. 81.

    Pleosporaceae (H Ariyawansa, KD Hyde, EBG Jones, NN Wijayawardene)

82. 82.

    Polystomellaceae (JK Liu, KD Hyde, EBG Jones)

83. 83.

    Protoscyphaceae (S Boonmee, KD Hyde, EBG Jones)

84. 84.

    Pseudoperisporiaceae (S Boonmee, KD Hyde, EBG Jones)

85. 85.

    Roussoellaceae (JK Liu, R Phookamsak, KD Hyde)

86. 86.

    Salsuginaceae (KD Hyde, S Tibpromma, S Boonmee)

87. 87.

    Schizothyriaceae (R Phookamsak, KD Hyde, EBG Jones)

88. 88.

    Seynesiopeltidaceae (KD Hyde, EBG Jones)

89. 89.

    Shiraiaceae (Liu YX, KD Hyde, WJ Li, Q Tian)

90. 90.

    Sporormiaceae (IC Senanayake, F Doveri, KD Hyde, EBG Jones)

91. 91.

    Strigulaceae (R Lücking, MP Nelsen)

92. 92.

    Sympoventuriaceae (S Hongsanan, PW Crous)

93. 93.

    Teichosporaceae(H Ariyawansa, KD Hyde, EBG Jones, NN Wijayawardene)

94. 94.

    Teratosphaeriaceae (P Chomnunti, P Crous, KD Hyde, NN Wijayawardene)

95. 95.

    Testudinaceae (H Ariyawansa, KD Hyde, EBG Jones)

96. 96.

    Tetraplosphaeriaceae (K Tanaka, K Hirayama)

97. 97.

    Thyridariaceae (Q Tian, KD Hyde)

98. 98.

    Trematosphaeriaceae (Y Zhang, H Ariyawansa, KD Hyde, S Suetrong, EBG Jones)

99. 99.

    Trichothyriaceae (HX Wu, KD Hyde)

100. 100.

     Triposporiopsidaceae (P Chomnunti, KD Hyde, EBG Jones)

101. 101.

     Trypetheliaceae (R Lücking, A Aptroot, MP Nelsen)

102. 102.

     Tubeufiaceae (S Boonmee, KD Hyde, EBG Jones)

103. 103.

     Venturiaceae (S Hongsanan, KD Hyde, EBG Jones, NN Wijayawardene)

104. 104.

     Vizellaceae (JK Liu, R Phookamsak, KD Hyde, EBG Jones, NN Wijayawardene)

105. 105.

     Zopfiaceae (S Hongsanan, KD Hyde, EBG Jones, NN Wijayawardene)

1. 1.

   Acrocordiaceae (KD Hyde)

2. 2.

   Aspidotheliaceae (M Thambugala, KD Hyde, EBG Jones)

3. 3.

   Coccodiniaceae (P Chomnunti, KD Hyde, EBG Jones)

4. 4.

   Diporothecaceae (KM Thambugala, KD Hyde, EBG Jones)

5. 5.

   Eoterfeziaceae (KM Thambugala, KD Hyde, EBG Jones)

6. 6.

   Epigloeaceae (KM Thambugala, KD Hyde, EBG Jones)

7. 7.

   Lautosporaceae (EBG Jones, KD Hyde, S Suetrong)

8. 8.

   Mastodiaceae (KM Thambugala, KD Hyde, EBG Jones)

9. 9.

   Meliolaceae (S Hongsanan, KD Hyde)

10. 10.

    Meliolinaceae (HX Wu, P Chomnunti, YM Li, KD Hyde)

11. 11.

    Microtheliopsidaceae (KM Thambugala, DQ Dai, KD Hyde, EBG Jones)

12. 12.

    Moriolaceae (KD Hyde)

13. 13.

    Myxotrichaceae (KM Thambugala, KD Hyde, EBG Jones)

14. 14.

    Phaneromycetaceae (KM Thambugala, KD Hyde, EBG Jones)

15. 15.

    Phillipsiellaceae (KM Thambugala, KD Hyde, EBG Jones)

16. 16.

    Phyllobatheliaceae (KM Thambugala, KD Hyde, EBG Jones)

17. 17.

    Polystomellopsidaceae (KD Hyde)

18. 18.

    Pseudeurotiaceae (KM Thambugala, KD Hyde, EBG Jones)

19. 19.

    Pyrenotrichaceae (JK Liu, KD Hyde, EBG Jones)

20. 20.

    Roesleriaceae (KM Thambugala, KD Hyde, EBG Jones)

21. 21.

    Seuratiaceae (KM Thambugala, KD Hyde, EBG Jones)

22. 22.

    Thryptosporaceae (KD Hyde)

23. 23.

    Triposporiopsidaceae (P Chomnunti, KD Hyde, EBG Jones)

* = Contributors to each family, underlined names are main authors

Dothideomycetes is the largest class of Ascomycota, with an estimated number of 19,000 species (Kirk et al. 2008). This class largely corresponds to what has historically been called the Loculoascomycetes (Nannfeldt 1932; Luttrell 1955; Janex-Favre 1971; Barr 1979a, b, 1983a, b, 2001; Eriksson 1981; Reynolds 1971; Tehler 1990, 1995; Letrouit-Galinou et al. 1994; Barr and Huhndorf 2001; Liu and Hall 2004), although several lineages previously assigned to that group, such as the Pyrenulales and Verrucariales, are now classified in subclass Chaetothyriomycetidae, class Eurotiomycetes (Lutzoni et al. 2004; Schmitt et al. 2005; Hibbett et al. 2007; Schoch et al. 2009a, b). On the other hand, Arthoniomycetes, which were usually considered to feature an intermediate type of ascoma development between ascolocular ascohymenial fungi (Choisy 1955; Henssen and Jahns 1973; Eriksson 1994; Henssen and Thor 1994; Tehler 1990, 1995), have been shown to be a sister group to Dothideomycetes and the two classes are united in superclass Dothideomyceta (Schoch et al. 2009a).

Dothideomycetes are mostly characterized by ascolocular ascoma development and bitunicate (fissitunicate) asci (Nannfeldt 1932; Luttrell 1955; Eriksson 1981; Barr and Huhndorf 2001). However these characters are partially shared by other classes, such as Arthoniomycetes and Eurotiomycetes and ascoma development has not been thoroughly studied for the large majority of Ascomycota, including Dothideomycetes. Therefore, it is unclear whether Dothideomycetes feature distinct phenotypic apomorphies based on which the class could be characterized. Also, while the class itself receives good support and hence lineages can be assigned to it with confidence once molecular data is available, the subdivision of Dothideomycetes remains unclear, since backbone support is low (Schoch et al. 2009b). An example for the problem of internal classification are the lichenized families Monoblastiaceae, Strigulaceae and Trypetheliaceae, which all have certain morphological and anatomical features in common but are apparently unrelated and have, with few exceptions, no close non-lichenized relative in the class (Nelsen et al. 2009, 2011a). Therefore, all could be treated as separate orders, as already suggested for Trypetheliaceae (Aptroot et al. 2008).

There has been considerable research interest in the phylogeny of the Dothideomycetes in the last 3 years (Schoch et al. 2009b; Nelsen et al. 2009, 2011a). Much of this research has concentrated on the phylogenetic placement of genera based on single and multigene analysis of rDNA sequence data, however, there has been less attention paid to morphology. In particular there have been few attempts to link morphological characters of the morphs into a taxonomic framework, so much so that it is often not clear which characters should be used to define taxa at the order, family, and genus level. This problem has stemmed from the fact the arrangements are based mainly on results of molecular phylogenetic analysis combined with historical placements based on morphological arrangements. Because neither provides a complete assessment of taxa at various levels, the situation for the classification of Dothideomycetes remains confused. In this paper we deal with the families of Dothideomycetes as currently recognized and bring together recent published data and attempt to link morphological data to the findings resulting from molecular analysis. We presently accept 105 families and suggest which genera should be placed in these families, which genera can questionably be placed in these families, and which families and genera should be excluded. We also illustrate 138 dothideomycete genera, with full descriptions and notes and discuss their taxonomic placement.

The recent Outline of the Ascomycota (Lumbsch and Huhndorf 2010) provided the most recent arrangement of Dothideomycetes which included 41 families comprising 249 genera (13 familes and 74 in subclass Dothideomycetidae, 28 and 175 in Pleosporomycetidae) and a further 116 genera which were placed in Dothideomycetes, genera incertae sedis. In the Pleosporales there were 28 families and 175 genera, with 12 genera listed under Pleosporales, genera incertae sedis. Recent research, particularly dealing with the Pleosporales, has provided further data on the class. The study by Zhang et al. (2012a) in particular dealt with 105 genera of Pleosporales, in 28 families and emended the suborder Pleosporineae (Zhang et al. 2009a, b). These studies were mostly based on molecular phylogenetic analysis and did not correlate morphology at the family level with the new phylogenetic arrangements. In this paper we therefore attempt to add some structure to the arrangement of the Pleosporales by discussing the characters of familial arrangements determined at the phylogenetic level.

The Dothideomycetes comprise 22 accepted orders based on molecular phylogenetic studies (Lumbsch and Huhndorf 2010). Each order is briefly discussed. Six families Ascoporiaceae, Coccoideaceae, Cookellaceae, Perisporiopsidaceae, Protoscyphaceae and Pseudoperisporiaceae are included under families incertae sedis because they lack molecular data that can resolve their position in Dothideomycetes. The Meliolales was recently included in Dothideomycetes but produce unitunicate asci (Hosagoudar 2003, 2006, 2013). In addition, molecular analyses revealed that isolates of Meliola (Meliolaceae, Meliolales) clustered with members of Sordariomycetes with strong support, and were unrelated to Dothideomycetes (Saenz and Taylor 1999; Zhuang and Liu 2012).

Acrospermales Minter et al., Boln Soc. argent. Bot. 42(1–2): 112 (2007)

The family Acrospermaceae has been placed in various orders at various times (Brandriff 1936; Eriksson 1982; Barr 1990a, b, c, d; Sherwood 1977; Eriksson and Hawksworth 1991; Hawksworth et al. 1995; Kirk et al. 2001). Minter et al. (2007) introduced the order Acrospermales to accommodate the family Acrospermaceae, which is typified by Acrospermum compressum Tode. Lumbsch and Huhndorf (2010) placed the Acrospermales in Dothideomycetes, order incertae sedis. Stenroos et al. (2010) showed that Acrospermales should be placed in Dothideomycetes, although they clustered with strains of Arthoniomycetes, whereas Pang et al. (2013) clearly show the order belongs in Dothideomycetes.

Asterinales M.E. Barr ex D. Hawksw. & O.E. Erikss., Syst. Ascom. 5(1): 177 (1986)

The order Asterinales was invalidly introduced by Barr (1976) because of lack of Latin diagnosis, but subsequently validated in Hawksworth and Eriksson (1986). The Asterinales is based on the type family Asterinaceae and is characterized by superficial, thyriothecial, dimidiate, black ascomata, with radial mycelium and often hyphopodiate, appressed hyphae, globose-subglobose asci and 1-septate, brown ascospores (Barr 1987a; Hosagoudar 2012). Molecular analyses confirmed that Asterinales can be included in Dothideomycetes with good support (Hofmann et al. 2010; Wu et al. 2011a). Wu et al. (2011a) also suggested the families Aulographaceae and Parmulariaceae should be included in this order. Therefore, the Asterinales presently include three families Asterinaceae, Aulographaceae and Parmulariaceae.

Botryosphaeriales C.L. Schoch et al., in Schoch et al., Mycologia 98(6): 1050 (2007) [2006]

Synonym: Phyllostictales Seaver, North American Flora 6(1): 1(1922)

The order Botryosphaeriales was introduced by Schoch et al. (2006) based on phylogenetic data and this has been verified in several recent papers (Schoch et al. 2006, 2009a). The order comprises three families: Botryosphaeriaceae, Planistromellaceae and Phyllostictaceae (Liu et al. 2012a; Monkai et al. 2013; Wikee et al. 2013). The order is defined by relatively thick-walled, dark-celled ascostromata, with one to several locules, bitunicate and fissitunicate asci and 1–2-celled ascospores, which are hyaline when immature and become brown in some genera at maturity. Cut ascostromata of dried specimens usually reveal distinct white contents. The asexual states are coelomycetous and produce hyaline to brown conidia from phialidic conidiogenous cells in conidiomata that usually form initially in the ascostromata.

Capnodiales Woron., Annls mycol. 23(1/2): 177 (1925)

The order Capnodiales was introduced by Woronichin (1925), and accepted in Dothideomycetes based on phylogenetic studies (Crous et al. 2009a; Schoch et al. 2006, 2009a; Chomnunti et al. 2011). Lumbsch and Huhndorf (2010) list the order as comprising eight families: Antennulariellaceae, Capnodiaceae, Coccodiniaceae, Davidiellaceae, Dissoconiaceae, Metacapnodiaceae, Mycosphaerellaceae and Piedraiaceae. The Capnodiales are mainly defined by their shared ecological niche as leaf epiphytes associated with the honey dew produced by insects, and saprobes or leaf pathogens and are an ascostromatal order without Pseudoparaphyses (Hughes 1976; Crous et al. 2009a; Chomnunti et al. 2011).

Dothideales Lindau, in Engler & Prantl, Nat. Pflanzenfam., Teil. I (Leipzig) 1: 373 (1897)

The order Dothideales was introduced by Lindau (1897) and typified by Dothidea sambuci (Pers.) Fr. Molecular phylogenetic studies indicate that the order Dothideales belongs in Dothideomycetes with high bootstrap support (Boehm et al. 2009a; Schoch et al. 2009a, b; Shearer et al. 2009; Suetrong et al. 2009). Lumbsch and Huhndorf (2010) list the order as comprising three families: Dothideaceae, Dothioraceae and Teratosphaeriaceae. However, Dothioraceae is shown to be a synonym of Dothideaceae in this study, while Crous et al. (2007a) placed Teratosphaeriaceae in Capnodiales. The order is characterised by immersed, erumpent or superficial, uniloculate or multiloculate ascostromata, with or without a papillate apex, bitunicate asci and usually septate, symmetrical or longitudinally asymmetrical, sometimes muriform, hyaline or pigmented ascospores. The asexual states are coelomycetous and hyphomycetous.

Dyfrolomycetales K.L. Pang, K.D. Hyde & E.B.G. Jones, ordo novus , Mycobank: MB 805281

An order of marine fungi within the class Dothideomycetes. Characterized by perithicoid, coriaceous, immersed ascomata, clypeate, ostiole, papillate; bitunicate, fissitunicate, cylindrical asci, with a short pedicel and distinct ocular chamber with ring-like subapical apparatus; hyphae-like pseudoparaphyses in a gelatinous matrix and overlapping uniseriate, broadly fusiform, symmetrical, hyaline, multiseptate with wide septa, and smooth-walled ascospores. Differing from Monoblastiales in its nonlichenized ascomata, hyphae-like pseudoparaphyses, and hyaline, multiseptate ascospores. Differing from Kirschsteiniotheliaceae in lacking brown, 1-septate ascospores, and an asexual state in Dendryophiopsis.

Typus: Dyfrolomyces K.D. Hyde et al.

Hysteriales Lindau, in Engler & Prantl, Nat. Pflanzenfam., Teil. I (Leipzig) 1: 265 (1896)

The order Hysteriales was introduced by Lindau (1896) and has been placed among the pyrenomycetes and the discomycetes at different times (Rehm 1896). Molecular data now places Hysteriales in the Dothideomycetes (Boehm et al. 2009a, b; Shearer et al. 2009; Suetrong et al. 2009). This order consists of a single family, Hysteriaceae, characterized by a hysterothecium (a carbonaceous navicular ascoma with a longitudinal dehiscence slit). Ascospores are typically dark and vary in septation. The family has been extensively revised based on DNA and amino acid sequence analysis (Boehm et al. 2009a, b).

Jahnulales K.L. Pang et al., in Pang et al., Mycol. Res. 106(9): 1033 (2002)

Pang et al. (2002) introduced the order Jahnulales for species with ascomata with well-developed long hyphal stalks or pedicels. The placement of Jahnulales in Dothideomycetes was confirmed by molecular analysis (Pang et al. 2002; Campbell et al. 2007; Schoch et al. 2009b; Shearer et al. 2009; Suetrong et al. 2009, 2011a). Jahnulales comprises two families: Aliquandostipitaceae and Manglicolaceae (Suetrong et al. 2011a). Members of Jahnulales are aquatic species which are lignicolous, saprobic, and occur on rotting or submerged woody debris and palms (Inderbitzin et al. 2001; Pang et al. 2002; Raja and Shearer 2006; Campbell et al. 2007; Suetrong et al. 2011a). The order is characterized by ca. 40 μm wide, septate, brown, thick-walled mycelial hyphae, that often bear ascomata, hypha-like or trabeculate, septate, unbranched pseudoparaphyses up to 4 μm wide; bitunicate asci with an ocular chamber; and ascospores that are 1-septate, pale to dark brown, with bipolar apical appendages or surrounded by a thin or elaborate wide gelatinous sheath (Suetrong et al. 2011a).

Lichenoconiales Diederich, Lawrey & K.D. Hyde, ordo novus, MycoBank: MB 805297

An order of lichenicolous fungi within class Dothideomycetes. Characterized by a lichenicolous habit, commensalistic to parasitic on crustose, foliose or fruticose lichens; sexual state unknown; conidiomata pycnidial, black, with an irregular pore-like opening; pseudoparenchymatic pycnidial wall composed of 2–4 layers of rounded to polyhedral, flattened cells, dark brown in outer and paler in inner layers, K+ olivaceous; conidiophores lacking; conidiogenous cells covering the entire wall of the inner pycnidial cavity, phialidic, often percurrently proliferating, hyaline and smooth-walled, rarely apically brown and verrucose, elongate ampulliform to subcylindrical; conidia arising singly or indistinctly catenate, subspherical, ellipsoid, obpyriform or elongate-clavate, basally usually truncate, aseptate, not visibly guttulate, surface verruculose to verrucose or echinulate, medium to dark brown, becoming more olivaceous in 10 % KOH. Differing from most pycnidial fungi by the combination of more or less hyaline, elongate ampulliform to subcylindrical, phialides, often percurrently proliferating conidiogenous cells, and ellipsoid, basally truncate, brown conidia with an ornamented wall.

Typus: Lichenoconium Petr. & Syd.

Lichenotheliales K. Knudsen, Muggia & K.D. Hyde, ordo novus, MycoBank: MB 805298

An order of non-lichenized fungi within the class Dothideomycetes. Characterized by a saprobic or lichenicolous lifestyle, epiphytic or endolithic on rock, endokapylic or episubstratic on lichens; thallus black of scattered stroma, connected or not by superficial hyphae, or of areoles scattered to continuous; either with ascomata, perithecioid and ostiolate, with interascal filaments, or stromata non-ostiolate with asci forming in pseudoparenchymatous locules; asci bitunicate, globose to clavate, sometimes with outer layers or apex of ascus K/I+ blue, ascospores hyaline to brown, 1-septate to muriform; macroconidia present or not, black, originating from thallus surface; conidiomata immersed in stromata, present or not, conidia rod-shaped or unknown. Differring from the lichenicolous genus Lichenostigma in Lichenostigmales in the Arthoniomycetes (Ertz et al. 2013) in their ascus type and the presence of a hamathecium, as well as in comprising many non-lichenicolous species occurring on rock.

Typus: Lichenothelia D. Hawksw.

Microthyriales G. Arnaud, Les Astérinées: 85 (1918)

The order Microthyriales was introduced by Arnaud (1918) with the family type Microthyriaceae. Schoch et al. (2009b) provided molecular evidence based on a strain of Microthyrium microscopicum Desm. (the type species of Microthyriaceae) and Stomiopeltis betulae J.P. Ellis (Micropeltidaceae) to represent the Microthyriales and showed that the order can be placed in Dothideomycetes. However, the molecular data based on these limited sequences did not provide enough clarity to support the order. Lumbsch and Huhndorf (2010) placed Microthyriaceae and Micropeltidaceae in Dothideomycetes as family incertae sedis, and Microthyriales was not listed. In this study we accept the Microthyriales based on morphological data and include the families Microthyriaceae and Micropeltidaceae. The Microthyriales are characterized by dark, circular, thyriothecial ascomata with poorly developed bases and central rounded ostioles (Wu et al. 2011a). The bitunicate asci are inclined from the base and outer rim towards the central ostiole and pseudoparaphyses may be present or lacking. Ascospores are hyaline or brown, mostly 2-celled or with a few to several transverse septa. Species are foliar epiphytes, biotrophs, saprobes or epiphytes on dead or living leaves and stems of plants. This group of fungi are relatively poorly studied (Wu et al. 2011a), and thus molecular data is needed to confirm the relationships of these families and their genera, which are generally poorly known. Asexual states are coelomycetous (Asterostomula, ?Cyclopeltella, Leptothyrium, ?Sirothyriella) or hyphomycetous (Hansfordiella, Holubovaniella, Isthmospora, Zalerion) (Kirk et al. 2008; Seifert et al. 2011; Wu et al. 2011a; Wijayawardene et al. 2012).

Monoblastiales Lücking, M.P. Nelsen & K.D. Hyde, ordo novus, MycoBank: MB 805299

An order of lichenized fungi within class Dothideomycetes. Characterized by crustose, mostly epiphytic habit on tree bark; endo- to epiperidermal, ecorticate to corticate thallus; trentepohlioid photobiont; perithecioid ascomata with distinct or reduced involucrellum and psoroplectenchymatous excipulum; branched and anastomosing, thin, net-like physes; fissitunicate asci with ocular chamber and fluorescent ring- or cap-structures; mostly ellipsoid-fusiform, 1-septate (rarely non-septate or 3-septate), euseptate, hyaline to rarely brownish ascospores; conidia produced in conspicuous conidiomata, with the macroconidia typically clustered in groups or masses in a distinct gelatinous matrix; and generally lack secondary substances except lichexanthone and anthraquinones in a few taxa. Differing from Strigulales in the net-like physes and from most Trypetheliales in the euseptate ascospores with few septa and the poor secondary chemistry, and from both in the macroconidia arranged in a gelatinous matrix.

Typus: Monoblastia Riddle

Myriangiales Starbäck, Bih. K. svenska Vetensk. Akad. Handl., Afd. 3 25(no. 1): 37 (1899)

Starbäck (1899) introduced the order Myriangiales for the species characterised by crustose ascostromata and muriform ascospores, similar to the type species, Myriangium duriaei Mont. & Berk. (Berkeley 1845; Miller 1938). Phylogenetic studies based on a multi-gene analysis have indicated that Myriangiales always cluster with Dothideomycetes (Boehm et al. 2009b; Phillips et al. 2008; Schoch et al. 2009b; Zhang et al. 2011a). Kirk et al. (2008) included three families, Cookellaceae, Elsinoaceae and Myriangiaceae in Myriangiales, while Lumbsch and Huhndorf (2010) accepted only Elsinoaceae and Myriangiaceae in the order based on phylogenetic results, while Cookellaceae was treated in Dothideomycetes as family incertae sedis. The order is characterized by pulvinate, irregular ascostromata in which the asci are irregularly arranged in one or more layers in locules. Locules contain single or multiple asci. Asci have minute pedicels and indistinct ocular chambers. Ascospores which are irregularly arranged are liberated only by the breakup of the stromatal layers above them. Asexual states are coelomycetous.

Mytilinidiales Boehm et al., Mycol. Res. 113(4): 468 (2009)

The family Mytilinidiaceae was separated from the order Pleosporales and elevated to a higher taxonomic rank, Mytilinidiales by Boehm et al. (2009a) supported by molecular data. The order comprises Gloniaceae and Mytilinidiaceae (Boehm et al. 2009b; Lumbsch and Huhndorf 2010). Morphologically, the order is characterized by rigid, brittle, hysterothecial, carbonaceous ascomata with the appearance of bivalve shells or are wedge-shaped, and ascospores are of various septate types: didymosporous, phragmosporous and/or dictyosporous (Sivanesan 1984; Alexopoulos et al. 1996; Boehm et al. 2009a, b). Most are pantropical species, saprobic on wood, bark, stem, twigs, gymnosperms and some species are parasitic on leaves (Barr 1975, 1987a, b; Pereira and Barreto 2003; Boehm et al. 2009a).

Natipusillales Raja, Shearer, A.N. Mill. & K.D. Hyde, ordo novus, MycoBank: MB 805300

An order of freshwater fungi within the class Dothideomycetes characterized by small, globose to subglobose Ascomata on submerged wood, erumpent to superficial, hyaline to light brown. Peridial wall membranous, composed of pseudoparenchyma cells forming a textura angularis in surface view. Pseudoparaphyses septate, sparse. Asci 8-spored, bitunicate, fissitunicate, globose, subglobose or obclavate. Ascospores fusiform to cylindrical, one to several septate, multiguttulate or not, hyaline becoming brown with age, with or without a gelatinous sheath and/or appendages. Included family: Natipusillaceae Raja, Shearer & A.N. Mill. Members of Natipusillales differ from other aquatic orders in the Dothideomycetes in having very small, globose, hyaline ascomata, globose asci and appendaged, hyaline ascospores. The establishment of Natipusillales is supported by both morphological characters as well as combined 18S and 28S nrDNA sequence data (Raja et al. 2013a, b, this study).

Typus: Natipusilla A. Ferrer et al.

Patellariales D. Hawksw. & O.E. Erikss., Syst. Ascom. 5(1): 181 (1986)

The order Patellariales was introduced by Hawksworth and Eriksson (1986) based on morphological data to accommodate Patellariaceae, Arthrorhaphidaceae and Phillipsiellaceae, while Barr (1987a) only accepted Patellariaceae in Patellariales. Luttrell (1951) had considered Hysteriaceae and Patellariaceae to be Discomycetes with bitunicate asci and placed them in the bitunicate group. Kutorga and Hawksworth (1997) later monographed several genera of Patellariaceae and 17 genera were accepted. Morphological features of the Patellariales are apothecial or hysterothecial, cup-shaped ascomata, bitunicate asci and hyaline, septate ascospores (Kutorga and Hawksworth 1997). Boehm et al. (2009a) and Schoch et al. (2009b) showed the Patellariales to belong in Dothideomycetes based on molecular data.

Phaeotrichales Ariyawansa, J.K. Liu & K.D. Hyde, ordo novus, MycoBank: MB 805301

An order of coprophilous fungi within the class Dothideomycetes. Characterized by a terrestrial or saprobic habitat. Ascomata solitary, or in small groups, superficial, unilocular, stromatic, globose, black, perithecial or cleistothecial setose. Peridium thin, carbonaceous, membraneous. Hamathecium tissue absent or of evanescent cellular pseudoparaphyses Asci 8-spored, bitunicate, fissitunicate, in irregular or arranged in little groups or bundles and evanescent at maturity and sometimes with an apical ring. Ascospores uniseriate or partially overlapping, dark brown to reddish brown, sometimes fragmenting, composed of terminal germ pores, with or without a sheath. Asexual state: Unknown.

Typus: Phaeotrichum Cain & M.E. Barr

Pleosporales Luttr. ex M.E. Barr, Prodr. Cl. Loculoasc. (Amherst): 67 (1987)

The order Pleosporales was invalidly introduced by Luttrell (1955), and later validated by Barr (1987a), and is based on the family Pleosporaceae with the type species Pleospora herbarum (Pers.) Rabenh. (Barr 1987b). It is the largest order of Dothideomycetes (Kirk et al. 2008). Molecular studies indicated that Pleosporales should comprise 20 accepted families (Boehm et al. 2009a, b; Mugambi and Huhndorf 2009a; Schoch et al. 2009b; Shearer et al. 2009; Suetrong et al. 2009; Tanaka et al. 2009; Zhang et al. 2009a, b), however, more recently Zhang et al. (2012a) included 25 families. The order Pleosporales presently comprises 28 families (Boonmee et al. 2012; Zhang et al. 2012a), while Venturiaceae was excluded from the Pleosporales by Zhang et al. (2011a) and placed in Venturiales. In this study we place 41 families in Pleosporales. Members of Pleosporales are saprobic on dead plant material in fresh water, marine, or terrestrial environments, and a large number, especially asexual states are pathogens on living plants (Zhang et al. 2009c; Boonmee et al. 2012). Pleosporales comprises the suborders Pleosporineae and Massarineae. Zhang et al. (2009a) emended the Pleosporineae, which is a phylogenetically well-supported suborder of Pleosporales and includes seven families: Cucurbitariaceae, Didymellaceae, Didymosphaeriaceae, Dothidotthiaceae, Leptosphaeriaceae, Phaeosphaeriaceae and Pleosporaceae, while Ariyawansa et al. (2013a) and Liu et al. (2013) included two new families, Halojulellaceae and Shiraiaceae. Pleosporineae contains economically important plant pathogens (de Gruyter et al. 2010; Zhang et al. 2009a) and is characterized by broadly to narrowly oblong ascomata, downwardly growing pseudoparaphyses, with 1-multiseptate ascospores. Recent studies using multigene phylogeny of Phoma and its closely related genera, indicated that their representative species cluster in different subclades of Pleosporineae (de Gruyter et al. 2009, 2010). Massarineae is characterized by immersed or superficial ascomata, cylindrical asci with a short pedicel and 1-multiseptate ascospores. Zhang et al. (2012a) included five families in Massarineae viz. Lentitheciaceae, Massarinaceae, Montagnulaceae, Morosphaeriaceae and Trematosphaeriaceae, with most species saprobic in terrestrial or aquatic environments, and we introduce a new family Bambusicolaceae to this suborder.

Strigulales Lücking, M.P. Nelsen & K.D. Hyde, ordo novus, MycoBank: MB 805302

An order of lichenized fungi within class Dothideomycetes. Characterized by crustose, mostly epiphytic habit on leaves, tree bark, and rocks; sub or supracucicular (leaves) or endo to epiperidermal (bark), ecorticate to corticate thallus; trentepohlioid photobiont; perithecioid ascomata with distinct or rarely reduced involucrellum and psoroplectenchymatous excipulum; unbranched to more rarely branched and sometimes anastomosing, but not net-like physes; fissitunicate asci with ocular chamber; mostly fusiform to oblong, 1-septate (rarely 3–7-septate to muriform), euseptate, hyaline ascospores; conidia produced in pycnidia, with the macroconidia typically being septate to muriform and having terminal, gelatinous appendages; and general lack of secondary substances. Differing from most Trypetheliales in the euseptate ascospores with few septa and the lack of secondary chemistry, and from both Monoblastiales and Trypetheliales in the mostly unbranched physes and the macroconidia producing gelatinous appendages.

Typus: Strigula Fr.

Trypetheliales Lücking et al., in Aptroot et al., BibliothecaLichenol. 97: 13 (2008)

The order Trypetheliales was introduced in Aptroot et al. (2008) to accommodate a single family Trypetheliaceae of lichenized fungi. This order was accepted in the Dothideomycetes based on molecular phylogenetic results (Del Prado et al. 2006; Schoch et al. 2009a, b; Nelsen et al. 2009, 2011b). The order includes mostly lichenized species characterized by a branched and anastomosing hamathecium embedded in a distinct gelatinous matrix, and by usually hyaline ascospores that feature a combination of eusepta and angular distosepta, giving the lumina a diamond-shaped appearance. These characters are not constant, however, as some lineages produce ascospores with eusepta only and in some lineages they are dark brown. Also the degree of anastomosing of the hamathecium can vary, especially in basal lineages.

Tubeufiales S. Boonmee et al., Fungal Divers. (in prep.)

The order Tubeufiales will be introduced by Boonmee et al. (2013, in prep) based mainly on a distinct habit, most being saprobes on decaying lignocellulosic material, sexual characters, helicosporous asexual morphs, and strongly supported by molecular data. The ascomata are characterized by superficial, light brown, dark brown to black, uniloculate pseudothecia, and asexual states are hyphomycetous helicospores or chlamydospores (Barr 1979a, 1980; Sivanesan 1984; Kodsueb et al. 2006a; Tsui and Berbee 2006; Tsui et al. 2006; Boonmee et al. 2011).

Typus: Tubeufia Penz. & Sacc.

VenturialesYing Zhang & K.D. Hyde, Fungal Divers. 51(1): 252 (2011)

The order Venturialeswas introduced by Zhang et al. (2011a), based on morphological and ecological characteristics, as well as strong support from multigene phylogenetic analyses. This order presently includes Sympoventuriaceae and Venturiaceae (Zhang et al. 2011a). Some species are plant pathogens and others are saprobic, mostly on leaves or stems of dicotyledons. Important characters are small to medium-sized ascomata, with or without setae, a mostly evanescent hamathecium, 8-spored, broadly or usually obclavate asci, usually lacking a pedicel, hyaline, light greenish olivaceous to brown, 1-septate, symmetrical, asymmetrical or apiosporous ascospores and hyphomycetous asexual states.

Layout of the paper

Each family is treated with a family description, notes on its history and presence of any asexual morphs, an account of the genera and including phylogenetic data where applicable. The type is listed along with a description of the type genus except in cases where there is only a single genus in the family. Notes on the type genus are provided along with full citations for the type species. This is followed by accepted genera and their type species; important synonyms and occasional notes are given. Synonymies follow the basis of one name for a biological genus or species (see Hawksworth et al. 2011; Hawksworth 2012) and follow published papers (e.g. Crous et al. 2006; Wingfield et al. 2012) or Wijayawardene et al. (in prep.) or Index Fungorum (2013). A key to the genera included in the family is provided unless the number of genera is small. The type genus is illustrated with a representative plate of its key morphological features. For consistency, authorities and their abbreviations for all taxa and citations of place of publication and abbreviations follow Index Fungorum (2013), even those these are not always entirely consistent.

Arrangement of dothideomycetes

The circumscription of the Dothideomycetes is based on the previous arrangements given in Lumbsch and Huhndorf (2010) which is then updated based on consideration of recent publications and interpretation of genera from the literature. We consider this a working circumscription which will be further refined as more genera are studied both at the morphological and molecular levels. Basionyms and synonyms are listed for the type genus and species of each family, unless there are more than three. Other synonyms are only listed where we have either made changes or examined material. All other possible synonyms can be found in Index Fungorum.

Molecular phylogeny

Sequences were obtained from GenBank following mostly previous publications (e.g. Schoch et al. 2006, 2009a; Suetrong et al. 2009; Zhang et al. 2012a; Li et al. 2012; Liu et al. 2012a) and are listed in Supplementary Table 1.

Sequences for each DNA region were initially aligned using Bioedit (Hall 2004) and ClustalW v. 1.6 (Thompson et al. 1997). Alignments were checked and manually optimized. The dataset was refined visually in BioEdit v. 7.0.1 (Hall 2004). By doing this we were able to make sure the data included in our tree were reliable. Following the recommendation of Wiens (2006), we included taxa in our multi-locus matrix even if they did not have all genes present. All absent genes were coded as missing data. Schismatomma decolorans (Turner & Borrer ex Sm.) Clauzade & Vezda (Arthoniomycetes) was chosen as the outgroup sequence based on its placement as sister clade to the Dothideomycetes (Schoch et al. 2006, 2009b). Four genes small subunits ribosomal RNA (SSU), large subunits ribosomal RNA (LSU), the translation elongation factor-1 alpha (TEF1) and the second largest subunit of RNA polymerase II (RPB2) were applied in this study.

Phylogenetic trees based on individual LSU, individual SSU, combined LSU and SSU and combined LSU, SSU and TEF1 datasets (data not shown) were congruent with the combined LSU, SSU, TEF1 and RPB2 data sets. However, the position of the families Kirschsteiniotheliaceae, Lichenoconiaceae and Monoblastiaceae were not constant. The phylogenetic analyses of the combined LSU, SSU, TEF1 and RPB2 data were performed using maximum parsimony, Bayesian and maximum likelihood algorithms.

Maximum-parsimony analyses were performed by PAUP v. 4.0b10 (Swofford 2002) using the heuristic search option with 1,000 random taxa addition and tree bisection and reconnection (TBR) as the branch swapping algorithm. All characters were unordered and of equal weight and gaps were treated as missing data. The Tree Length (TL), Consistency Indices (CI), Retention Indices (RI), Rescaled Consistency Indices (RC) and Homoplasy Index (HI) were calculated for each tree generated. Maxtrees were unlimited, branches of zero length were collapsed and all multiple, equally parsimonious trees were saved. Clade stability was assessed using a bootstrap (BT) analysis with 1,000 replicates, each with 10 replicates of random stepwise addition of taxa (Hillis and Bull 1993). Maximum parsimony bootstrap values (MPBP) equal or greater than 50 % are given above each node (Fig. 2).

Maximum likelihood (ML) analysis was performed at the CIPRES webportal (Miller et al. 2010) using RAxML v. 7.2.8 as part of the “RAxML-HPC2 on TG” tool (Stamatakis 2006; Stamatakis et al. 2008). A general time reversible model (GTR) was applied with a discrete gamma distribution and four rate classes. Fifty thorough maximum likelihood (ML) tree searches were done in RAxML v. 7.2.7 under the same model, with each one starting from a sepa-rate randomised tree and the best scoring tree selected with a final ln value of −13,974.356237. One thousand non para-metric 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. 2).

Bayesian analyses (Larget and Simon 1999): The model of evolution was estimated by using MrModeltest 2.2 (Nylander 2004). Independent Bayesian phylogenetic analyses were performed in MrBayes v. 3.1.2 (Huelsenbeck and Ronquist 2001) using a uniform [GTR+I+G] model, lsetnst = 6 rates = invgamma; prsetstatefreqpr = dirichlet (1,1,1,1). 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 5,000,000 generations and trees were sampled every 100th generation (resulting in 10,000 total trees). The first 5,000 trees were discarded as burn-in prior to convergence of the four chains. The remaining trees were used to construct a 50 % majority rule consensus tree and to calculate Bayesian Posterior Probabilities (BYPP) with those equal or greater than 0.90 given below each node (Fig. 2).

Phylogenetic trees were drawn using Treeview v. 1.6.6 (Page 1996) and MEGA 5 (Tamura et al. 2011). The alignments were deposited in TreeBASE (www.treebase.org).

Definitions of previously inconsistently defined terms used in the study

We tried to be consistent in the terminology we have used. However, it was not always possible, especially because some terms are not commonly accepted and some have been interpreted differently by different authors. This applies especially to the denominations of the different types of interascal hyphae, for which we have defined the terminology used in this work as follows (Eriksson 1981; Ryan et al. 2005; Kirk et al. 2008).

Paraphyses: originating from only the bottom of the ascomatal cavity.

Paraphysoids: originating from both the bottom and top of the ascomatal cavity.

Pseudoparaphyses: originating from only the top but eventually attaching to the bottom and becoming free apically, in mature stages often not distinguishable from paraphysoids or paraphyses.

If the ontogeny and hence the type was unknown or unclear, we used as far as possible the neutral terms physes or interascal hyphae. Also, instead of distinguishing between ascolocular pseudothecia and ascohymenial perithecia, we used the neutral terms perithecioid ascomata or else the term perithecia in a broad sense, since by definition all ascomata in the higher Ascomycota (Pezizomycotina) are homologous and thus a differential terminology using the term pseudothecia for ascolocular perithecioid ascomata does not reflect phylogenetic relationships.

The numbers cited for taxonomic entries up to 489999 had originally been assigned by Index Fungorum and have been adopted by MycoBank, and we cite them in this paper as MycoBank numbers.

The combined LSU, SSU, TEF1 and RPB2 gene data set consists of 414 taxa, with Schismatomma decolorans as the outgroup taxon. The dataset consists of 4,584 characters after alignment, 1,615 characters were constant, and 2,303 characters were parsimony informative, while 666 variable characters are parsimony-uninformative. A heuristic search with random addition of taxa (1,000 replicates) and treating gaps as missing characters generated six equally parsimonious trees. The best scoring RAxML trees are shown in Figs. 1 and 2. Bootstrap support (BS) values of MP and ML (equal to or above 50 %) are shown on the upper branches. Values of the Bayesian posterior probabilities (PP) (equal to or above 90 %) from MCMC analyses are shown under the branches (TL = 39,381, CI = 0.140, RI = 0.617, RC = 0.086, HI = 0.860).

The best scoring RAxML Dothideomycetes tree (compress overview tree) from 414 taxa based on a combined dataset of LSU, SSU, TEF1 and RPB2 sequences with all lineages collapsed to family level where possible. The tree was rooted with Schismatomma decolorans

Full size image

The best scoring RAxML Dothideomycetes tree from 414 taxa based on a combined dataset of LSU, SSU, TEF1 and RPB2 sequences with all lineages collapsed to family level where possible. The tree is the same as Fig. 1. Bootstrap support values for maximum likelihood (ML, green) and maximum parsimony (MP, red) greater than 50 % are given above the nodes; Bayesian posterior probabil-ities (BYPP, blue) above 0.90 are given below the nodes. The original isolate numbers are given after the species names. The tree was rooted with Schismatomma decolorans

Full size image

In the phylogenetic trees (Figs. 1 and 2), the 414 strains of Dothideomycetes included in the analysis cluster into the two Dothideomycete subclasses as previously described based on presence or absence of pseudoparaphyses (Schoch et al. 2006). Subclass Pleosporomycetidae includes Pleosporales, Mytilinidiales, and Hysteriales and is the same as in previous publications (Schoch et al. 2009b; Boehm et al. 2009b; Shearer et al. 2009; Suetrong et al. 2011a). However, the position of Jahnulales was not consist and clustered together with two newly introduced orders Dyfrolomycetales and Strigulales; and Acrospermales in this study. The second subclass Dothideomycetidae representing Capnodiales, Dothideales and Myriangiales is confirmed as well-supported.

Twenty-two orders (Pleosporales, Mytilinidiales, Hysteriales, Acrospermales, Dyfrolomycetales, Strigulales, Jahnulales, Tubeufiales, Patellariales, Botryosphaeriales, Dothideales, Myriangiales, Capnodiales, Lichenoconiales, Phaeotrichales, Microthyriales, Natipusillales, Venturiales, Asterinales, Lichenotheliales, Monoblastiales and Trypetheliales) within the class Dothideomycetes are supported in Figs. 1 and 2. The order Venturiales was recently proposed (Zhang et al. 2011a) with two families Venturiaceae and Sympoventuriaceae to accommodate the groups which have a parasitic or saprobic lifestyle, occurring on leaves or stems of dicotyledons; small to medium-sized ascomata, often with setae; deliquescing pseudoparaphyses; 8-spored, broadly cylindrical to obclavate asci; 1-septate, yellowish, greenish or pale brown to brown ascospores; and hyphomycetous asexual morphs (Zhang et al. 2011a). The family Tubeufiaceae which was not placed in an order by Schoch et al. (2006), is now accommodated in Tubeufiales (Boonmee et al. 2013, in press). The order Tubeufiales includes taxa with uni-loculate, superficial and pigmented (e.g. pale brown, brown, and dark brown to black) ascomata, and mostly multi-celled and hyaline relatively long ascospores and mostly produce helicosporous asexual states. Both Venturiaceae and Tubeufiaceae were traditionally assigned to Pleosporales, although they have diagnostic characters which readily distinguish them from other pleosporalean families. Phylogenetically, the core genera of Venturiaceae and Tubeufiaceae form monophyletic clades within Dothideomycetes, and represent separate sister lineages from the current orders. The nearest relatives of the Tubeufiales are Botryosphaeriales. The species-rich order Botryosphaeriales included the single family Botryosphaeriaceae in Schoch et al. (2006). With more extensive taxa sampling of diversity in this order, the Planistromellaceae was shown to cluster adjacent to the Botryosphaeriaceae (Minnis et al. 2012; Monkai et al. 2013), and Phyllostictaceae reinstated (Wikee et al. 2013). Three well-supported clades are representative of these families in Botryosphaeriales (Figs. 1 and 2). Saccharata protea Saccharata proteae (Wakef.) Denman & Crous does not reside in any of the above clades, and is placed on early diverging branches of the order, representing an underscribed family (Saccharataceae; Slippers et al. 2013, in press). The monotypic genus Melanops (type = Melanops tulasnei Fuckel) clustered together with Planistromellaceae, but without strong support, and is placed in the Melanopsaceae by Slippers et al. (2013, in press). Their somewhat similar morphological characters however, may show that Melanops belongs in Planistromellaceae. In addition, the two asexual genera Aplosporella and Pseudofusicoccum did not cluster in any clade in this analysis. Aplosporella is placed in the Aplosporellaceae by Slippers et al. (2013, in press).

Families with good support but not clustering in any known orders include Kirschsteiniotheliaceae, Lichenoconiaceae, Lichenotheliaceae, Monoblastiaceae, Natipusillaceae, Phaeotrichaceae, and Strigulaceae (Figs. 1 and 2). Dyfrolomycetaceae clustered together with Acrospermaceae, Kirschsteiniotheliaceae and Strigulaceae and formed a sister group with Jahnulales. Monoblastiaceae (lichenized fungi) formed a separate lineage comprising two species, Megalotremis verrucosa and Anisomeridium ubianum (Vain.) R.C. Harris with 100 % bootstrap support and is closest to the lichen family Trypetheliaceae Zenker (1827). The recently introduced family Kirschsteiniotheliaceae (Boonmee et al. 2012) is also related to Dyfrolomycesillaceae and Acrospermaceae, however its placement is not stable in all phylogenetic analyses (Schoch et al. 2009b; Suetrong et al. 2009; Boonmee et al. 2012) and this will require further study. Lichenoconiaceae, a lichenicolous group of coelomycetous asexual morphs, is introduced as a new family in this paper. Natipusillaceae, introduced by Raja et al. (2011) comprises freshwater species with unique characters (e.g. thin-walled, light-coloured, ascomata, subglobose asci and two-celled appendaged ascospores), and forms a distinct lineage (100 % and 100 % BS and 94 % PP) in a clade with Phaeotrichaceae, Microthyriaceae and Venturiales. The poorly studied Phaeotrichaceae is a distinct lineage (100 % and 100 % BS and 100 % PP) that clusters in a clade with Asterinales, Microthyriales and Venturiales (Figs. 1 and 2). The Lichenotheliaceae are lichenicolous or loosely associated with algae and form a separate clade without any close relatives. Based on their distinct morphological characters and habitat, and phylogenetic analysis (of available sequence data) we therefore introduce seven orders, namely Dyfrolomycetales K.L. Pang, K.D. Hyde & E.B.G. Jones, ordo novus, Lichenoconiales Diederich, Lawrey & K.D. Hyde, ordo novus, Lichenotheliales K. Knudsen, Muggia & K.D. Hyde, ordo novus, Monoblastiales Lücking, M.P. Nelsen & K.D. Hyde, ordo novus, Natipusillales, Raja, Shearer, A.N. Mill. & K.D. Hyde, ordo novus, Phaeotrichales Ariyawansa, J.K. Liu & K.D. Hyde, ordo novus and Strigulales Lücking, M.P. Nelsen & K.D. Hyde, ordo novus (Table 1).

Phylogeny of orders of dothideomycetes

Pleosporomycetidae C.L Schoch et al., Mycologia 98(6): 1048 (2007) [2006]

Pleosporomycetidae was introduced as a subclass by Schoch et al. (2006) and included the single order Pleosporales. Subsequently, Schoch et al. (2009a) included the two other orders Mytilinidiales and Hysteriales within this subclass, and suggested that the order Jahnulales can be tentatively included within Pleosporomycetidae. Our phylogenetic analysis showed similar results, however the newly introduced order Dyfrolomycetales and Strigulales, and Acrospermales clustered together with Jahnulales. Because the lack of strong phylogenetic evidence, we only accept Pleosporales, Mytilinidiales and Hysteriales representing the subclass Pleosporomycetidae.

Pleosporales

The 23 or 27 families included in Pleosporales by Kirk et al. (2008) and Lumbsch and Huhndorf (2010) respectively, were previously groupedbased on morphological characters. These families were recently resolved at molecular level, for example Schoch et al. (2006) found reasonable support for 7 to 23 families in a phylogeny generated from four genes. Subsequently, Zhang et al. (2012a) accepted 28 families in this order based on both morphology and phylogeny (four genes). Our analysis in Figs. 1 and 2 provide support for 33 family clades, which correspond to the suborders Pleosporineae with seven families and Massarianeae with six families. The other seven newly introduced families are Anteagloniaceae, Bambusicolaceae, Biatriosporaceae, Halojulellacaeae, Roussoellaceae, Salsugineaceae and Thyridariaceae, belong in Pleosporales, while the newly introduced family Bambusicolaceae forms a clade within the suborder Massarianeae. Lophiostomataceae comprise two distinct clades Lophiostoma-clade (100 %, 100 % BS and 0.97 PP) and Misturarosphaeria-clade (100 %, 95 % BS and 0.99 PP) which may represent distinct families, however, we refrain from introducing a new family until more species are sequenced. Roussoellaceae (100 %, 100 % BS and 1.00 PP) and Biatriosporaceae are newly introduced families in this paper. They form distinct clades (94 % and 79 % BS) which comprise morphologically and phylogenetically distinct families. The Anteagloniaceae (100 %, 100 % BS and 0.95 PP) is also introduced as a new family and is related to Lophiotremataceae. Salsuginaceae (54 %, 52 % BS and 0.99 PP) is also a distinct family introduced in this paper, close to Aigialaceae and Lindgomycetaceae.

Mytilinidiales

No new data are available for this order which appears to be well-supported (Boehm et al. 2009a, b, c; Schoch et al. 2009b).

Hysteriales

Phylogenetic analyses indicate that the Hysteriales are closely related to the Pleosporales (Schoch et al. 2006, 2009a, b, c; Boehm et al. 2009a). In addition, the shared morphological characters, such as the similar centrum confirms their close relationship. However, there is still strong branch support for its separation from Pleosporales (Boehm et al. 2009a, b). Luttrell (1955) retained the Hysteriales based on the elongated hysteriaceouslocule, capable of relatively indeterminate linear growth which is distinct from the strict Pleospora-type centrum, defined as it is by constrained concentric growth. In contrast to the close association between the Hysteriales and the Pleosporales, the Mytilinidiales is a distant clade which is close to Pleosporales within the Pleosporomycetidae (Boehm et al. 2009a, b).

Acrospermales

The order Acrospermales was established by Minter et al. (2007) to accommodate Acrospermaceae which the asci and ascospores more than 1,000 μm. The phylogenetic studies of Stenroos et al. (2010) used five-gene analysis to show that this order belongs in Dothideomycetes. In our phylogenetic study, it is close to the other two newly introduced order Dyfrolomycetales and Strigulales. However, whether a subclass is required to accommodate this order requires further research.

Dyfrolomycetales

The order Dyfrolomycetales is introduced in this study.

Strigulales

The order Strigulales is introduced in this study.

Jahnulales

The order Jahnulales was introduced in Dothideomycetes based on molecular analysis of SSU nrDNA sequence data from Aliquandostipite, Jahnula and Patescospora (Pang et al. 2002). Most taxa of Aliquandostipitaceae are freshwater fungi occurring on woody substrata. Suetrong et al. (2011a) introduced a new family Manglicolaceae for the marine ascomycete Manglicola guatemalensis Kohlm. & E. Kohlm. In this study, taxa of Aliquandostipitaceae and Manglicolaceae from Jahnulales formed a well-supported clade related to the newly introduced orders Dyfrolomycetales and Strigulales, and Acrospermales, and is close to Hysteriales, as was indicated in earlier studies (Schoch et al. 2009a, b; Boehm et al. 2009a, Shearer et al. 2009). Although in our trees (Figs. 1 and 2) these orders form a distinct clade, the results are not stable, and therefore we do not treat these orders as members of Pleosporomycetidae, nor do we introduce a new subclass.

Tubeufiales

This strongly supported order, with a distinct habitat, being mostly saprobes on decaying lignocellulosic material with helicosporous asexual morphs is introduced in Boonmee et al. (2013) and is not discussed further here.

Patellariales

The Patellarialesis a poorly known order with discomycetous ascomata. A more extensive taxon sampling is required to address the diversity in this order and determine its phylogenetic relationships.

Botryosphaeriales

The order Botryosphaeriales was introduced by Schoch et al. (2006), following molecular analysis, and comprised a single family Botryosphaeriaceae. Subsequent studies suggested that Botryosphaeriales most likely should accommodate additional families (Schoch et al. 2009a; Liu et al. 2012a, b; Minnis et al. 2012; Monkai et al. 2013). Based on morphology and phylogeny (four genes), Liu et al. (2012a) presented a classification of Botryosphaeriales, and two major clades representing Botryosphaeriaceae and Phyllostictaceae resulted from their analysis. Minnis et al. (2012) also included Planistromellaceae as a distinct family, in an unresolved position relative to other genera within Botryosphaeriales. In our phylogenetic analysis, these three families formed well-supported clades (Figs. 1 and 2). The placement of the monotypic genera Saccharata and Melanops were unresolved, Saccharata proteae being basal in the order and Melanops tulasnei clustering together with Planistromellaceae with low bootstrap support. Slippers et al. (2013, in press) introduce the Saccharataceae and Melanopsaceae to accommodate these taxa. A strongly supported clade which included Aplosporella prunicola Damm & Crous and a putative strain of Bagnisiella examinans was formed between Planistromellaceae and Phyllostictaceae, for which Slippers et al. (2013, in press) introduce the Aplosporellaceae. The asexual genus Pseudofusicoccum clustered together with Phyllostictaceae with good support, however, the position of the strain was not stable; we therefore do not include this genus in Phyllostictaceae in our study. The family Botryosphaeriaceae, typified by Botryosphaeria dothidea (Moug. ex Fr.) Ces. & De Not and related genera formed a well-supported clade and includes most sexual and asexual genera presently placed in the family. An extensive taxon sampling is required to address the diversity in this order and there is still much research to be carried out with resolution of families and genera, linkage of sexual and asexual morphs and differentiation of cryptic species (Liu et al. 2012a, b).

Dothideomycetidae P.M. Kirk et al., ex C.L. Schoch et al., Mycologia 98(6): 1048 (2007) [2006]

This subclass always comprises Capnodiales, Dothideales and Myriangiales (Schoch et al. 2006, 2009b; Zhang et al. 2009a, b, c; Boehm et al. 2009a, b, c; Figs. 1 and 2)

Dothideales

The order Dothideales was introduced by Lindau (1897) to accommodate a single family Dothideaceae and the families of the Dothideales varied from author to author. The Dothideales previously comprised Dothideaceae, Dothioraceae and Teratosphaeriaceae (Lumbsch and Huhndorf 2010). Morphological examination and multi-gene phylogenetic analysis indicate that the Dothioraceae should be combined under Dothideaceae, while Aureobasidium, Selenophoma, Kabatiella and Columnosphaeria may need a new family (Thambugala et al. in prep., Figs. 1 and 2).

Myriangiales

This order comprises Myriangiaceae and Elsinoaceae which are separated by 100 % BS. Further collections and sequence data of species in this order are needed to establish if these families can be maintained.

Capnodiales

The order Capnodiales comprises six clades (Figs. 1 and 2) representing Capnodiaceae, Cladosporiaceae, Dissoconiaceae, Mycosphaerellaceae and Teratosphaeriaceae plus one unnamed clade comprising mostly rock inhabiting species. The Teratosphaeriaceae is recently included (Crous et al. 2009c), while the Capnodiaceae has been redefined (Chomnunti et al. 2011). The family Dissoconiaceae comprises several asexual genera of plant pathogens and has been studied by Li et al. (2012).

Lichenoconiales

The order Lichenoconiales is introduced in this study.

Phaeotrichales

The order Phaeotrichales is introduced in this study.

Microthyriales

The Microthyriales is a poorly known order with thyrothecial ascomata and it is represented by only a single taxon, Microthyrium microscopicum (type species) in our phylogenetic analysis. This result requires additional molecular data.

Natipusillales

The order Natipusillales is introduced in this study.

Venturiales

The order Venturiales has been detailed by Zhang et al. (2011a) and no new data are available.

Asterinales

The order Asterinales is represented by Asterinaceae in our phylogenetic analysis. Few molecular studies have been carried out on this group, which mostly focused on morphology. Hofmann et al. (2010) presented the first molecular hypothesis of the Asterinaceae within Dothideomycetes based on the LSU and SSU sequences. In our phylogenetic analysis, it is close to Venturiales and the newly introduced orders Natipusillales and Phaeotrichales.

Lichenotheliales

The order Lichenotheliales is introduced in this study.

Monoblastiales

The order Monoblastiales is introduced in this study.

Trypetheliales

The order Trypetheliales was recently introduced by Aptroot et al. (2008) and represents the largest lichen forming clade in Dothideomycetes.

Acrospermaceae Fuckel, Jb. nassau. Ver. Naturk. 23–24: 92 (1870) [1869–70], MycoBank: MB 80430

Saprobic, epiphytic or symbiotic on herbaceous plants. Sexual state: Ascomata solitary or in groups, superficial or immersed in stromata, often long, yellowish white to brown, black when dry, club-shaped to conoid, with a short stipe, flattened when dry, swelling when moist, ostiole large. Peridium in horizontal section comprising three layers, an outer layer composed of dark brown cells of textura angularis, a central layer, composed of pale brown tissue of elongated cells, and an inner layer composed of dense tissue of small, light brown cells. Hamathecium of narrow long, pseudoparaphyses or pseudoparaphyses lacking. Asci 8-spored, bitunicate, narrowly cylindrical, pedicellate, apically rounded with an ocular chamber. Ascospores fasciculate, filiform, hyaline, multi-septate, almost as long as the asci, smooth-walled. Asexual state: hyphomycetous. Conidiophores micronematous, pale brown, septate, branched or unbranched. Conidiogenous cells holoblastic, sympodial with denticles, pale brown, smooth-walled. Conidia cylindrical, long ellipsoid, pale yellow, 1–3-septate, smooth-walled.

Notes: The family Acrospermaceae presently includes two genera Acrospermum and Oomyces (Lumbsch and Huhndorf 2010). Acrospermum has club-shaped, brown, uni-locular ascomata and narrow, long paraphyses which resemble ascospores. However, Oomyces has conoid, yellowish white, multi-locular stromata and lacks pseudoparaphyses (Eriksson 1981). The asexual states of this family include members of Dactylaria and Gonatophragmium which are linked to asexual Acrospermum (Wijayawardene et al. 2012). However, Dactylaria was shown to be polyphyletic by Bussaban et al. (2005) and Seifert et al. (2011) also stated that Dactylaria is heterogenous, hence the phylogeny needs clarification. Kirk et al. (2008) mentioned Gonatophragmium is an asexual state of Acrospermum, but Seifert et al. (2011) did not assign any taxonomic affiliation. Whether Gonatophragmium is polyphyletic is not clear. No asexual state has been reported for Oomyces. Minter et al. (2007) established the order Acrospermales to accommodate this family, and Stenroos et al. (2010) confirmed this unique order belonging to Dothideomycetes by using five-gene.

Type: Acrospermum Tode, Fung. mecklenb. sel. (Lüneburg) 1: 8 (1790), MycoBank: MB 54 Fig. 3

Acrospermum cf. compressum (Material examined: VANUATU, Santo Island, South Santo, on decaying leaves of Pandanus sp., 31 October 1996, E.H.C. McKenzie HKU (M) 5005, IFRD 210–003). a, b Dry black ascomata on host surface. c Ascomata with short stipe. d Ascomata with large ostiole. e Ascoma in horizontal section. f, g Three layers of peridium. h-j Narrowly cylindrical asci. k, l Filiform ascospores. Scale bars: a–d = 200 μm, e–j = 50 μm, h–l = 10 μm

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Possible synonyms:

Scleroglossum Pers., J. Colorado-Wyoming Acad. Sci.: no. 671 (1820)

Xyloglossum Pers., Traité sur les Champignons Comestibles (Paris): 140,144 (1818)

Saprobic or epiphytic on herbaceous plants. Sexual state: Ascomata solitary or in groups, superficial, club-shaped to conoid, erect, uni-locular, brown to blackish when dry, with a short stipe or sessile, flattened when dry, swelling when moist, ostiole large, apex rounded. Peridium in horizontal section comprising three layers, an outer layer comprising dark brown cells of textura angularis, a central thick layer, comprising pale brown tissue of gelatinized hyphae with elongated cells, and an inner layer comprising dense tissue of small, light brown cells. Hamathecium of narrow, long, pseudoparaphyses. Asci 8-spored, bitunicate, narrowly cylindrical, pedicellate, with an ocular chamber. Ascospores fasciculate, filiform, hyaline, multi-septate, almost as long as the asci, smooth-walled. Asexual state: Hyphomycetous. Conidiophores micronematous, pale brown, septate, branched or un-branched. Conidiogenous cells holoblastic, sympodial with denticles, long cylindrical, pale brown, smooth-walled. Conidia cylindrical, long ellipsoid, pale yellow, 1–3-septate, apically rounded, smooth-walled (Webster 1956).

Notes: Acrospermum includes 66 species epithets (Index Fungorum 2013). The genus was established by Tode (1790) and has superficial, club-like ascomata, narrowly cylindrical, bitunicate asci and filiform, multi-septate, hyaline ascospores (Riddle 1920). Eriksson (1981) redescribed the type species A. compressum and another similar species A. graminum. Molecular data showed that these two species can be clearly separated (Stenroos 2010). Webster (1956) also concluded after studying the asexual morphs in culture.

Type species: Acrospermum compressum Tode, Fung. mecklenb. sel. (Lüneburg) 1: 8 (1790), MycoBank: MB 140891

Other genera included

Gonatophragmium Deighton, in Cejp & Deighton, Mycol. Pap. 117: 13 (1969)

Type species: Gonatophragmium mori (Sawada) Deighton, in Cejp & Deighton, Mycol. Pap. 117: 13 (1969)

Oomyces Berk. & Broome, Ann. Mag. nat. Hist., Ser. 2 7: 185 (1851)

Type species: Oomyces carneoalbus (Lib.) Berk. & Broome, Ann. Mag. Nat. Hist., Ser. 2 7: 185 (1851)

Key to sexual genera of Acrospermaceae

1. Fruiting bodies ascomata club-shaped, brown, paraphyses long ...................................................................Acrospermum

1. Fruiting bodies stomata conoid, yellowish white, paraphyses lacking .................................................................Oomyces

Aigialaceae Suetrong et al., in Suetrong et al., Stud. Mycol. 64: 166 (2009), MycoBank: MB 515957

Saprobic on submerged wood in mangroves and terrestrial habitats. Sexual state: Ascomata scattered or rarely clustered, immersed, erumpent to superficial, black, globose or conical, coriaceous to carbonaceous, ostiolate, with rounded or slit-like ostiole, apapillate. Peridium 50–130 μm wide, composed of dark brown thick-walled cells. Hamathecium comprising 1.5–2 μm wide, hyaline, anastomosing, trabeculate pseudoparaphyses, embedded in a gelatinous matrix. Asci 8-spored, bitunicate, fissitunicate, obclavate to cylindrical, long-pedicellate, with a non-amyloid ocular chamber. Ascospores bi-seriate, fusiform, hyaline to brown, septate to muriform, smooth-walled, slightly constricted at the septa, surrounded by an evanescent mucilaginous sheath or with caps at the ends. Asexual state: Unknown.

Notes: The family Aigialaceae was introduced by Suetrong et al. (2009) for taxa with carbonaceous ascomata without papilla, trabeculate pseudoparaphyses, cylindrical asci and ascospores with a sheath or gelatinous appendages around the apical cells. Three genera, Aigialus, Ascocratera and Rimora, all marine taxa, were included. These genera were previously referred to Massarinaceae (Lumbsch and Huhndorf 2010). Zhang et al. (2012a) commented on the wide range of morphological variation between members of Aigialaceae, with taxa sharing very few features in common. Aigialus grandis, the type species of Aigialaceae has flattened, brown, muriform ascospores, with mucilage surrounding their apices (Jones et al. 2009a). The asci are cylindrical and the hamathecium comprises narrow anastomosing pseudoparaphyses (trabeculae), while the ascomata are mostly immersed in wood and asymmetrical, forming in the same plane as the wood fibres and have an apical slit-like opening. This is quite different from the second genus presently included in Aigialaceae, Ascocratera (Suetrong et al. 2009). This monotypic genus represented by Ascocratera manglicola Kohlm., has raised superficial carbonaceous ascomata, lacking a papilla, and 1(−3)-septate hyaline ascospores surrounded by a mucilaginous sheath (Kohlmeyer 1986). The third genus, shown by molecular data to belong in the family, Rimora (=Lophiostoma mangrovei Kohlm. & Vittal), is similar to Ascocratera, with raised apapillate ascomata and septate hyaline ascospores. Despite these morphological similarities Ascocratera and Rimora are not congeneric as shown by the sequence data (Fig. 1) (Suetrong et al. 2009; Zhang et al. 2012a).

The introduction of the family Aigialaceae is strongly supported (100/100 support) by phylogenetic analyses of five nuclear genes (LSU, SSU, rRNA, RPB2 and TEF1) in Pleosporales as shown in Fig. 1. Subsequently two additional genera, from terrestrial habitats, have been included in the family: Fissuroma and Neoastrosphaeriella (Liu et al. 2011). These two genera form a well-supported subclade in the Aigialaceae, and share morphological features with Ascocratera and Rimora, i.e. carbonaceous apapillate ascomata, trabeculate pseudoparaphyses, cylindrical asci and septate ascospores with a sheath. The genus Fissuroma was introduced to accommodate species previously assigned to Astrosphaeriella with slit-like ostioles. Further Astrosphaeriella, Lophiostoma and Massarina species should be sequenced to determine their phylogenetic relationships.

The family includes five genera and eleven species, of which three genera with marine/mangrove affinities and two only from terrestrial habitats. The family type, Aigialus, and Fissuroma are illustrated below.

Type: Aigialus Kohlm. & S. Schatz, Trans. Br. mycol. Soc. 85(4): 699 (1985), MycoBank: MB 6002 Fig. 4

Aigialus grandis (Material examined: THAILAND, Chanthaburi Province, Tha Mai, Khung Kraben Bay Royal Development Study Center, on mangrove wood, 28 June 2008, S Suetrong et al. (BBH 23439 with isolates BCC 32284, 32285, 32286, 32287). a Surface view of mature ascomata of A. grandis on mangrove wood. b Released ascospores from the ostiole. c Surface of wood cut to show immersed ascomata with thick-walled peridium. d Longitudinal section through mature ascoma. e Sagittal section of ascoma. f Trabeculate pseudoparaphyses. g Cylindrical ascus. h-j Ascus with refractive apical ring. Apex of endoascus with some ascospores already released. k-l Ascospores in asci. m-s Ascospores. Scale bars: a = 1 mm, b–d = 500 μm, e = 100 μm, f, k–s = 25 μm, g = 50 μm, h–j = 15 μm

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Saprobic on mangrove wood, marine and with a tropical distribution. Sexual state: Ascomata gregarious, completely immersed, subglobose, laterally compressed, black, ostiolate, with a slit-like opening, apapillate. Peridium coriaceous to carbonaceous, two-layered, outer layer 50–100 μm, composed of elongate cells, inner layer 30–50 μm, of smaller hyaline cells. Hamathecium comprising 1.5–2 μm wide, trabeculate pseudoparaphyses, unbranched at the base, anastomosing above the asci, embedded in a gelatinous matrix. Asci 8-spored, bitunicate, fissitunicate, cylindrical, long-pedicellate, with a refractive non-amyloid, apical ring. Ascospores bi-seriate, ellipsoidal to broadly fusiform, muriform, dark brown except for hyaline to pale brown apical cells, glabrous, with a gelatinous cap or a mucilaginous sheath. Asexual state: Unknown.

Notes: A tropical to subtropical marine Ascomycete genus found on mangrove wood, frequent in occurrence. The genus Aigialus was initially classified in the Melanommatales (Kohlmeyer and Schatz 1985) sensu Barr (1979a) because of the trabeculate pseudoparaphyses, but it was not assigned to any family in the Dothideomycetes. Hawksworth et al. (1995) assigned the genus to Massariaceae in the order Pyrenulales, based on the possession of a refractive apical ring in the ascus that is J–. Subsequently, Tam et al. (2003), based on a single gene sequence analysis, referred it to the Pleosporales, a proposal accepted by Kirk et al. (2008). Aigialus can now be classified with confidence in the Pleosporales, family Aigialaceae based on a wider taxon sampling (five species) and four gene sequence analysis. Thus the classification of this genus is resolved within a wider study of taxa in the Dothideomycetes (Schoch et al. 2009b; Suetrong et al. 2009; Zhang et al. 2012a). Five species are accepted in the genus: A. grandis, A. mangrovei Borse, A. parvus S. Schatz & Kohlm., A. rhizophorae Borse and A. striatispora K.D. Hyde (Jones et al. 2009a).

Type species: Aigialus grandis Kohlm. & S. Schat., Trans. Br. mycol. Soc. 85(4): 699 (1985), MycoBank: MB 103973

Other genera included

Ascocratera Kohlm., Can. J. Bot. 64: 3036 (1986)

Type species: Ascocratera manglicola Kohlm., Can. J. Bot. 64: 3036 (1986)

Fissuroma J.K. Liu et al., Fungal Divers. 51: 145 (2011)

Type species: Fissuroma maculans (Rehm) J.K. Liu et al., Fungal Divers. 51: 145 (2011)

Neoastrosphaeriella J.K. Liu et al., Fungal Divers. 51: 148 (2011) MycoBank: MB 563462

Figure 5

Neoastrosphaeriella krabiensis (THAILAND, Krabi Prov., Krabi College of Agriculture and Technology, on petiole of Metroxylon sagu, 26 September 2010, J.K. Liu, JKA0050, MFLU11-1151, holotype). a Appearance of ascomata on host surface. b Section of an ascoma. c Structure of peridium. d Pseudoparaphyses. e-h Asci. i-n Ascospores. Note the sheath in young ascospores. Scale bars: a = 1 mm, b = 100 μm, c, d = 10 μm, e–h = 30 μm, i–n = 10 μm

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Saprobic on woody palm tissue, tropical in distribution. Sexual state: Ascomata scattered, usually immersed to semi-immersed beneath host tissue, or erumpent, black, carbonaceous, with a slit-like ostiole. Peridium composed of dark brown thick-walled cells. Hamathecium comprising 1–1.5 μm wide, anastomosing, trabeculate pseudoparaphyses, embedded in a gelatinous matrix. Asci 8-spored, bitunicate, fissitunicate, obclavate to cylindrical, short-pedicellate, with a small ocular chamber. Ascospores 2–3-seriate, fusiform, hyaline when young, becoming brown to dark brown, 1-septate, constricted at the septum, spore wall verrucose when mature, surrounded by a mucilaginous sheath. Asexual state: Unknown.

Notes: A monotypic tropical terrestrial ascomycete genus found on palm petioles and forming a sister group (Fig. 1) with four other genera in the Aigialaceae (Liu et al. 2011). The genus is characterized by its semi-immersed ascomata with slit-like ostioles, obclavate asci and brown, verrucose ascospores. The genus differs from Fissuroma in having smaller obclavate asci and brown verrucose ascospores, while Fissuroma has cylindro-clavate asci and hyaline ascospores. Molecular analyses based on a combined dataset of 18S and 28S nrDNA sequences place Neoastrosphaeriella in subclade to the genera Fissuroma, Aigialus, Rimora and Ascocratera. Neoastrosphaeriella shares features with Astrosphaeriella, a genus which is basal to Aigialaceae in the Pleosporales (Schoch et al. 2009b).

Type species: Neoastrosphaeriella krabiensis Jian K. Liu et al., in Liu et al., Fungal Divers. 51(1): 148 (2011), MycoBank: MB 563463

Rimora Kohlm. et al., in Suetrong et al., Stud. Mycol. 64: 166 (2009)

Type species: Rimora mangrovei (Kohlm. & Vittal) Kohlm et al., in Suetrong et al., Stud. Mycol. 64: 166 (2009)

Key to genera of Aigialaceae

1. Ascomata immersed or conical and crater-like, with rounded often papillate ostioles ......................................................2

1. Ascomata elongate with flat tops, opening with a slit-like ostiole .....................................................................................3

2. Ascomata immersed in a stroma, asci with a refractive apical plate in the ectoascus and a refractive ring in endoascus, ascospores dark brown, muriform with hyaline to light brown apical cells ..................................................................Aigialus

2. Ascomata conical and crater-like, on a black stroma, asci with refractive apical apparatus, ascospores hyaline, 1-septate ...............................................................Ascocratera

3. Ascomata forming on black stromatic crust, superficial, asci without apical apparatus, ascospores fusiform, 3-septate ......................................................................................Rimora

3. Ascomata immersed to semi-immersed, ascospores 1-septate ..................................................................................4

4. Asci cylindro-clavate, ascospores elongate-fusiform, hyaline, smooth ......................................................Fissuroma

4. Asci obclavate, ascospore fusiform, brown to dark brown, verrucose when mature ...........................Neoastrosphaeriella

Aliquandostipitaceae Inderb., in Inderbitzin et al., Am. J. Bot. 88(1): 54 (2001), MycoBank: MB 82130

Saprobic on submerged or trapped decorticated wood in freshwater streams, peat swamps and marine habitats. Sexual state: Ascomata scattered or rarely in small groups, immersed-erumpent or superficial, if superficial often attached to substrate by repent or pendant thick brown hyphal strands, hyaline, pale brown to black, coriaceous, ostiolate, short papillate. Peridium variable, 3–7 cell layers thick, comprising textura globulosa to angularis of very large, light brown to brown, thin-walled, cells. Hamathecium comprising 2–7 μm wide, hyaline, septate, sparsely branched, filamentous pseudoparaphyses, anastomosing above the asci. Asci 8-spored, bitunicate, fissitunicate, clavate to cylindrical, pedicel absent or very short, with or without an ocular chamber. Ascospores bi-seriate to multi-seriate, 1-septate, ellipsoid to fusiform, hyaline to pale brown, guttulate, with or without various types of appendages and/or sheaths. Asexual states: hyphomycetous Brachiosphaera, Xylomyces. Conidiophores reduced or un-branched or sparingly branched, hyaline. Conidiogenous cells monoblastic, thallic or sympodial or percurrent, monilioid, hyaline or brown. Conidia phragmosporous or staurosporous, with globose central cell, with 4-several, radiating septate arms, hyaline or brown, single or branched or un-branched chains (Seifert et al. 2011) or tetraradiate with 4–8 appendages.

Notes: The family Aliquandostipitaceae (Jahnulales) was introduced by Inderbitzin et al. (2001) for taxa with mycelium five times wider than those of normal ascomycetes. Ascomata, borne either on these thick hyphae or on hyphal stalks, are membranous, have a hamathecium of persistent pseudoparaphyses, bitunicate asci and hyaline to pale brown, 1-septate ascospores with or without a sheath or with appendages. Two new species were described, Aliquandostipite khaoyaiensis Inderb. and A. sunyatsenii Inderb. from wood collected in Thailand and China, respectively, and assigned to the family Aliquandostipitaceae based on 18S rDNA data (Inderbitzin et al. 2001). Inderbitzin et al. (2001) were unable to assign the family to any order, e.g., Dothideales or Pleosporales. Based primarily on analyses of 18S rDNA sequence data, Pang et al. (2002) subsequently established Jahnulales Pang et al. (Dothideomycetes, Ascomycota) for Aliquandostipitaceae that was monophyletic and distinct from other clades of bitunicate fungi, such as the Pleosporales and Dothideales. More recently, based on a review of the Jahnulales, Suetrong et al. (2011a) accepted the family Aliquandostipitaceae and included Aliquandostipite, Jahnula and Megalohypha and the asexual genera Brachiosphaera, Speiropsis and Xylomyces (Fig. 1). Molecular data also supported the inclusion of Manglicola in Jahnulales, but its familial placement was distinct from Aliquandostipitaceae (Suetrong et al. 2011a). Within the Jahnulales, generic limits are not well-delineated, and Jahnula was shown to be polyphyletic in a multi-gene study by Suetrong et al. (2011a). Jahnula species grouped in three clades, Clade 1, Jahnula sensu stricto, included J. aquatica (Plöttner & Kirschst.) Kirschst., (type species), J. granulosa K.D. Hyde & S W Wong, J. rostrata Raja & Shearer, J. potamophila K.D.Hyde & S W Wong and Megalohypha aqua-dulces A. Ferrer & hearer. Clade II comprised seven Jahnula species, while J. australiensis K.D.Hyde, Brachiosphaera tropicalis Nawawi & Descals and Speiropsis pedatospora Tubaki group in Clade III. Species grouping in Clade II (Jahnula sensu lato) may comprise more than one genus, for example, J. bipolaris (K.D. Hyde) K.D. Hyde, J. seychellensis K.D. Hyde & S.W. Wong and J. sunyatsensii (Inderb.) L.L. Pang & E.B.G. Jones all have ascospores with apical gelatinous pads and could be referred to the genus Ascagilis (Hyde 1992b). However, no taxonomical changes were proposed for Jahnula species not grouping in the Jahnula sensu stricto clade, until further species are collected and sequenced (Suetrong et al. 2011a). All Aliquandostipite species have well developed ascospore sheaths, with the exception of A. siamensis. (Sivichai & E.B.G. Joes J.) Campbell et al. 2007).

Type: Aliquandostipite Inderb., Am. J. Bot. 88(1): 54 (2001), MycoBank: MB 483979

Figure 6

Aliquandostipite khaoyaiensis (Material examined: a, c, d, g, j = ILLS A441-1; A441-2; b, e, f, h, i, k, l = F89-1). a Ascoma with a stalk. b Sessile ascoma on wood. c Longitudinal section of ascoma. d Peridial wall. e Longitudinal section of sessile ascoma. f Pseudoparaphyses. g Asci. h Dehiscing asci. i-l Ascospores with gelatinous sheath; stained with Indian ink in figure j and in aqueous nigrosin in figure l. Scale bars: a = 40 μm; b = 500 μm; c–l = 20 μm

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Saprobic on woody material in freshwater habitats. Sexual state: Ascomata immersed to superficial, globose to subglobose, hyaline, becoming brown to dark brown, coriaceous, papillate, developing on hyphal strands, or long or short hyphal stalks, often dimorphic. Peridium membranous, 1–4-layered, of cells of textura angularis to globosa. Hamathecium consisting of septate, hyaline, sparsely branched, hypha-like pseudoparaphyses. Asci 8-spored, bitunicate, fissitunicate, clavate to cymbiform, short-pedicellate, with thickened apical region, and ocular chamber. Ascospores bi-seriate to multi-seriate, oval to fusiform, 1-septate, constricted at the septum, hyaline becoming pale-brown with well-developed sheath, smooth-walled. Asexual state: Unknown.

Type species: Aliquandostipite khaoyaiensis Inderb., Am. J. Bot. 88(1): 54 (2001)

Notes: The genus was introduced by Inderbitzin et al. (2001) for two freshwater lignicolous ascomycetes collected in Thailand and China: A. khaoyaiensis Inderb., and A. sunyatsenii Inderb. (= Jahnula sunyatsenii). The genus is characterized by sessile and/or stalked ascomata (dimorphic), wide mycelium, bitunicate asci, the presence of pseudoparaphyses and 1-septate ascospores with or without a sheath (Inderbitzin et al. 2001). Currently, five species are accepted in the genus: A. crystallinus Raja et al., A. khaoyaiensis, A. minuta Raja & Shearer, A. separans (Abdel-Wahab & El Sharouney) J. Campb. et al., and A. siamensis (Suetrong et al. 2011a). Shearer et al. (2009) showed that four Aliquandostipite species form a well-supported monophyletic group. Strains of A. khaoyaiensis and A. siamensiae group together, with ascospore measurements in the same range. They differ in that the sheath in A. khaoyaiensis balloons out, while in A. siamensis it is narrow and ascospores in the latter are dimorphic: small and brown (33–45 × 10–13 μm) or large (58–73 × 15–25 μm) and hyaline to pale brown (Pang et al. 2002). Both types of ascospores, small and brown and large and hyaline to pale brown were observed in the same ascus in A. siamensis (Campbell et al. 2007). A combined analysis (SSU rDNA, LSU rDNA) of taxa in the Jahnulales showed that Aliquandostipite species formed a well-supported clade (Campbell et al. 2007), which is also supported in a larger study by Suetrong et al. (2011a).

Other genera included

Brachiosphaera Nawawi, in Descals et al., Trans. Br. mycol. Soc. 67: 213 (1976)

Type species: Brachiosphaera jamaicensis (J.L. Crane & Dumont) Nawawi, in Descals et al., Trans. Br. mycol. Soc. 67: 216 (1976)

Jahnula Kirschst., Ann. mycol. 34(3): 196 (1936), MycoBank MB 2526 Fig. 7

Jahnula seychellensis (Material examined: THAILAND, Nakorn Ratchassima Province, Tad Tha Phu, KhaoYaiNational Park, on Sindora siamensisin freshwater, 2 May 2001, S. Sivichai, BCC SS1536.2). a Ascomata with short stalk (arrowed). b Section of ascoma. c Ascus apical pore (arrowed). d Asci and pseudoparaphyses. e, f Asci. g-m Ascospores with bipolar pads (arrowed). Scale bars: a, b = 100 μm, c, g = 10 μm, d = 40 μm, e–f = 20 μm, h–m = 10 μm

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Saprobic on soft, decorticated woody material, predominately a freshwater genus, but also marine. Sexual state: Ascomata immersed, semi-immersed or erumpent, sessile or attached to a long stalk, globose to subglobose, black, coriaceous, ostiolate, short-papillate, easily detached from the substratum, solitary or gregarious. Peridium comprising 3–5 layers of relatively large, thin-walled, brown or hyaline, angular cells, arranged in a textura angularis. Hamathecium comprising 2–4 μm wide, hypha-like, filamentous, septate pseudoparaphyses, unbranched between the asci, branching and anastomosing above the asci. Asci 8-spored, bitunicate, fissitunicate, cylindrical or obclavate, pedicellate, thick-walled, with an ocular chamber and faint ring. Ascospores 1–3-seriate, ellipsoid-fusiform, pale brown to dark brown, 1-septate, constricted at the septum, guttulate, smooth or verruculose, some with mucilaginous appendages, pads or sheaths. Asexual state: Unknown.

Notes: Jahnula comprises saprobic species on woody material (generally decorticated wood) in streams, with the exception of J. appendiculata Pinruan et al. found on the trunk of the palm Licuala longecalycata in a peat swamp (Pinruan et al. 2002) and Xylomyces chlamydosporus Goos et al. and X. rhizophorae Kohlm. & Volkm.-Kohlm. which occur in marine habitats (Kohlmeyer and Volkmann-Kohlmeyer 1998a, b). In a molecular study by Campbell et al. (2007) the genus would appear to be paraphyletic, while in a the combined analysis of SSU and LSU rDNA, Jahnula species grouped into three clades, hence Suetrong et al. (2011a) concluded that it is polyphyletic. Suetrong et al. (2011a) refrained from introducing new genera for species assigned to Clades II and III, and suggested further taxon sampling in order to resolve the phylogenetic relationships and delineation of Jahnula species. Xylomyces is therefore regarded as a synonym of Jahnula based on the one name rule (McNeill et al. 2011). Likewise, Brachiosphaera tropicalis and Speiropsis pedatospora group with Jahnula species in Clade III, but these cannot be synonymized until the phylogenetic relationships of Jahnula species are resolved further. Moreover, the genera Brachiosphaera, Speiropsis and Xylomyces have been shown to be polyphyletic and not all belong in the Jahnulales.

Type species: Jahnula aquatica (Plöttner & Kirschst.) Kirschst., Ann. Mycol. 34: 196 (1936), MycoBank: MB 257114

The species illustrated in Fig. 2 is Jahnula seychellensis from material collected in Thailand, S. Sivichai (SS1536.2) on submerged wood.

Megalohypha A. Ferrer & Shearer, in Ferrer et al., Mycologia 99: 458 (2007)

Type species: Megalohypha aqua-dulces A. Ferrer & Shearer, in Ferrer et al., Mycologia 99(3): 456 (2007)

Speiropsis Tubaki, J. Hattori bot Lab. 20: 171 (1958)

Type species: Speiropsis pedatospora Tubaki, J. Hattori Bot Lab. 20: 171 (1958)

Xylomyces Goos et al., Mycologia 69(2): 282 (1977)

Type species: Xylomyces chlamydosporus Goos et al. [as ‘chlamydosporis’], Mycologia 69(2): 282 (1977)

Key to sexual genera of Aliquandostipitaceae

3. Ascospores with a thin or broad gelatinous sheath, lacking appendages ...................................................Aliquandostipite

3. Ascospores with apical caps, appendages, or lacking such structures ................................................................................3

4. Ascospores with longitudinal sulcate striations, lacking appendages or a gelatinous sheath ...................Megalohypha

4. Ascospores lacking longitudinal sulcate striations, with or without apical pads .....................................................Jahnula

Key to asexual genera of Aliquandostipitaceae

1. Asexual morph hypha-like, filamentous, septate, conidia and conidiophores absent .......................................................2

1. Sexual morphs ....................................................................3

2. Conidiophores and conidia lacking, chlamydospores intercalary, multi-septate ................................................Xylomyces

2. Conidia tetraradiate with 4–8 appendages (arms) ........................................................................Brachiosphaera

Amniculicolaceae Y. Zhang et al., in Zhang et al., Stud. Mycol. 64: 95 (2009), MycoBank: MB 515469

Saprobic in freshwater habitats. Sexual state: Ascomata solitary, scattered or in small groups, erumpent, immersed or nearly superficial, globose, subglobose to lenticular, with rough black surface, usually staining the woody substrate shades of purple, ostiolate. Ostiole with elongate apex and ostiolar canal filled with hyaline cells. Peridium two-layered, outer layer of small heavily pigmented thick-walled cells of textura angularis, inner layer of hyaline thin-walled cells textura angularis. Hamathecium comprising numerous, hyaline, septate, narrow, trabeculate, pseudoparaphyses, embedded in a gel matrix. Asci 8-spored, bitunicate, fissitunicate, long cylindrical to clavate, short-pedicellate, apially truncate, with an ocular chamber. Ascospores 1–2-seriate, fusiform or narrowly fusiform, hyaline, reddish-brown or pale, one to multi-septate, or muriform, constricted at the median septum, generally surrounded by an irregular, hyaline, gelatinous sheath. Asexual states: closely related to Anguillospora longissima (Sacc. & P. Syd.) Ingold, Spirosphaera cupreorufescens Voglmayr and Repetophragma ontariense (Matsush.) W.P. Wu (Zhang et al. 2008b, 2012a; Shearer et al. 2009).

Notes: The family Amniculicolaceae was formally introduced by Zhang et al. (2009b) to accommodate various fresh water fungi collected from Europe in the order Pleosporales and accepted by Shearer et al. (2009). The family is characterized by ascomata with a rough black surface, usually staining the woody substrate purple, trabeculate pseudoparaphyses and short-pedicellate asci bearing hyaline, reddish-brown or pale, 1-multi-septate or muriform ascospores, generally with hyaline gelatinous sheath (Zhang et al. 2009b). Currently the family comprises three genera, Amniculicola (type), Murispora and Neomassariosphaeria that form a well supported clade in the Pleosporales (Zhang et al. 2009a). Seifert et al. (2011) also confirmed the placement of Anguillospora in Amniculicolaceae but the placement of Repetophragma and Spirosphaera is confused as it has some lineages in Sordariomycetes. Hence we conclude that Amniculicolaceae have Anguillospora, “Spirosphaera”-like and “Repetophragma”-like hyphomycetous asexual morphs.

Type: Amniculicola Y. Zhang & K.D. Hyde, Mycol. Res. 112(10): 1189 (2008), MycoBank: MB 511328 Fig. 8

Amniculicola lignicola (Material examined: FRANCE, Rimont, Ruisseau de Peyrau on driftwood of Alnus glutinosa, 23 July 2006, J. Fournier, HKU (M) 17515, holotype). a, b Placement of the ascomata on host surface. c Close up of ascomata with slit like opening. d, e Section of ascoma (TS) with purple basal consisting of remnants of the host. f Close up of the peridium. g Slot-like ostiole, apex well-differentiated into two tuberculate flared lips. h Septate, long, hyaline, anastomosing and branched pseudoparaphyses placing between and above the asci. i-l Eight-spored asci with short, furcate pedicel. m-o Hyaline ascospores, with deeply constricted median septa. p Ascospores surrounded by an irregular hyaline gelatinous sheath in Indian ink. Scale bars: d–e = 150 μm, f = 25 μm, g = 40 μm, h = 5 μm, i–l = 50 μm, m–p = 10 μm

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Saprobic on wood in freshwater habitats. Sexual state: Ascomata scattered, solitary, or in small groups, initially immersed, becoming erumpent, to nearly superficial, with basal wall remaining immersed in host tissue, laterally flattened and base not easy to remove from the host, with black wall, rough and often consisting of remnants of the host, usually staining the woody substrate purple, globose to subglobose, broadly or narrowly conical, ostiolate. Ostiole slit-like, with apex well-differentiated into two tuberculate flared lips, long, ostiolar canal filled with periphyses. Peridium thick laterally and at the apex, thinner at the base, coriaceous, two-layered, outer layer composed of small heavily pigmented thick-walled cells of textura angularis, inner layer composed of hyaline thin-walled cells of textura angularis. Hamathecium comprising numerous, hyaline, septate, narrow, trabeculate, branched pseudoparaphyses, anastomosing between and above the asci, and embedded in gel matrix. Asci 8-spored, bitunicate, fissitunicate, long cylindrical to narrowly fusiform, with a short, twisted, narrow, furcate pedicel, truncate to rounded at the apex with a minute ocular chamber, inconspicuous when observed in water. Ascospores obliquely uni-seriate, partially overlapping, or bi-seriate, broadly fusiform to fusiform with broadly to narrowly rounded ends, hyaline, 1-septate, deeply constricted at the median septum, smooth-walled, containing four refractive oil globules, surrounded by an irregular, hyaline, gelatinous sheath, observed only when mounted in Indian ink. Asexual state: Unknown.

Notes: Based on ascomata with slit-like ostioles, thin, branching and anastomosing pseudoparaphyses and 1-septate, deeply constricted, fusiform ascospores, with a wide gelatinous sheath, Zhang et al. (2008b) introduced this new freshwater genus and referred it to the Pleosporales (Zhang et al. 2009a, 2012a). Initially treated as monotypic, the genus Amniculicola is represented by A. lignicola Y. Zhang ter & K.D. Hyde, but subsequently two other species were described, A. immersa Y. Zhang ter. et al. and A. parva Y. Zhang ter. et al. (Zhang et al. 2009b). All three Amniculicola species were collected from submerged wood in freshwater (Zhang et al. 2008b, 2009b). Shearer et al. (2009) showed that Amniculicola parva, Anguillospora longissima and Amniculicola immersa formed well-supported sister group to Amniculicola lignicola, with Massariosphaeria typhicola (P. Karst.) Leuchtm. (mistakenly designated as Semimassariosphaeria typhicola in Shearer et al. 2009) in a sister clade. In a four gene analysis Repetophragma ontariense, A. immersa, A. parva formed a sister group to Massariosphaeria (current name Neomassariosphaeria) typhicola and M. grandispora (Sacc.) Leuchtm. in a well-supported clade (Zhang et al. 2012a). However, the type species A. lignicola was not included in their analysis. The sexual state of Anguillospora longissima, has been reported as an undescribed species of ‘Massarina’ (Willoughby and Archer 1973; Webster 1992), which possesses superficial ascomata with a dark peridium, cylindrical, 8-spored asci and fusiform 1-septate ascospores, characters compatible with the morphological characters similar to Amniculicola lignicola. Therefore the sexual state of Anguillospora longissima may be Amniculicola parva, or a new species of Amniculicola, however this cannot currently be resolved as no asexual state has been reported for Amniculicola lignicola.

Type species: Amniculicola lignicola Y. Zhang ter & K.D. Hyde, Mycol. Res. 112(10): 1189 (2008), MycoBank MB 511330

Other genera included

Murispora Y. Zhang bis et al., in Zhang et al., Stud. Mycol. 64: 95 (2009)

Type species: Murispora rubicunda (Niessl) Y. Zhang ter et al., in Zhang et al., Stud. Mycol. 64: 96 (2009)

Neomassariosphaeria Y. Zhang bis et al., in Zhang et al., Stud. Mycol. 64: 96 (2009)

Type species:Neomassariosphaeria typhicola (P. Karst.) Y. Zhang ter et al., in Zhang, et al., Stud. Mycol. 64: 96 (2009)

Key to genera of Amniculicolaceae

1. Ascospores two-celled ...................................Amniculicola

1. Ascosporeswith more than two septa ................................2

2. Ascospores muriform, pale or reddish-brown .................................................................................Murispora

2. Ascospores with trans-septa only, hyaline to reddish-brown ..............................................................Neomassariosphaeria

Anteagloniaceae K.D. Hyde, J.K. Liu & A. Mapook, fam. nov. MycoBank: MB 804658

Saprobic on dead wood. Sexual state: Ascomata hysterothecial, superficial or sunken in substrate, oval to elongate, or globose to subglobose, black, carbonaceous, straight or curved, rarely branched, without subiculum. Ostiole central, slit-like. Peridium thick, dark brown, comprising a single stratum of dark brown cells of textura epidermoidea. Hamathecium comprising 1.4–1.5 μm wide, cylindrical to filiform pseudoparaphyses in a gelatinous matrix. Asci 8-spored, bitunicate, fissitunicate, elongate cylindric-clavate, straight or slightly curved, short- pedicellate, apically rounded, with an ocular chamber. Ascospores uniseriate, 2-celled, very small, ellipsoidal, hyaline, 1-septate, constricted at septa, widest at the middle and tapering towards the narrow ends,straight, smooth-walled. Asexual state: Unknown.

Typus: Anteaglonium Mugambi & Huhndorf

Notes: Although the ascomata of Anteaglonium are hysteriothecial and characteristic of Hysteriaceae, the genus was shown to cluster as a distinct lineage in Pleosporales (Mugambi and Huhndorf 2009a, b) showing a parallel evolution of hysterothecial ascomata in Dothideomycetes. A similar topology was shown in the study of Schoch et al. (2009a), Zhang et al. (2012a), Mugambi and Huhndorf (2009b) and is confirmed in our analysis (Figs. 1 and 2). We therefore introduce a new family to accomodate this distinct lineage. Anteaglonium comprises four species in Index Fungorum (2013).

Type: Anteaglonium Mugambi & Huhndorf, Syst. Biodiv. 7(4): 460 (2009) Fig. 9

Anteaglonium abbreviatum (Material examined: USA, Tennessee, Sevier County, Great Smoky Mountains National Park, Cherokee Orchard, vic. of Gatlinburg, Trillium tral, 35° 40′ 30.88″ W, 780 m elev.,on wood, branch, on ground, 1″ diam., A.N. Miller et al., 12 July 2004, ANM 37). a Herbarium specimen. b Ascomata on host surface. c Section through ascoma. d Peridium. e Pseudoparaphyses. f-h Immature and mature asci. Note fissitunicate ascus and strained by lactophenol cotton blue (see in h). i–l Ascospores. Scale bars: b = 500 μm, c = 50 μm, f–h = 20 μm, d = 10, e, i–l = 5 μm

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Type species: Anteaglonium abbreviatum (Schwein.) Mugambi & Huhndorf, Syst. Biodiv. 7(4): 460 (2009)

≡ Hysterium abbreviatum Schwein., Trans. Am. phil. Soc., Ser. 2 4(2): no. 2094 (1832)

Antennulariellaceae Woron., Annls mycol. 23 (1/2): 178 (1925), MycoBank: MB 80461

Foliar epiphytes as sooty moulds living on insect exudates. Subiculum dark brown to black, effuse, densely velutinous, somewhat dense or velvety, forming on the surface of leaves and twigs. Hyphae deeply pigmented at the margin, cylindrical to moniliform, or regular, septate hyphae with smooth or roughened walls. Erect hyphae dark, broad and irregularly anastomosing in a network, forming a brown to black, dense or scant covering. Sexual state: Ascomata developing from repeated divisions of hyphae, brown to dark brown, subglobose, obovoid to broadly ellipsoidal, or ovoid, sessile, or with a robust stalk, with or without appendages, ostiolate at maturity, arising from terminal or intercalary cells on aerial hyphae. Peridium thin-walled, pseudoparenchymatous. Asci 8-spored, bitunicate, fissitunicate, small, pyriform to ellipsoidal, or clavate. Ascospores minute, not numerous, irregularly arranged, ovoid, more or less oblong, hyaline, 1–3-septate, slightly constricted at the septum, with the upper half slightly shorter and broader than the lower one, rounded at both ends. Asexual state: coelomycetous or hyphomycetous (see under notes).

Notes: Antennulariellaceae, a family of sooty moulds, was described by Woronichin (1925) in Capnodiales, including six genera with 27 species (Kirk et al. 2008), while Lumbsch and Huhndorf (2010) listed only two genera. Species in the family have a widespread distribution, and are found in warm temperate to tropical locations, where they grow as black sooty molds on plants (Cannon and Kirk 2007). The asexual states of Antennulariaceae have been recorded as pycnidial (Antennariella, Batista and Ciferri 1963) and hyphomycetous (Capnodendron, Hughes 1976, 2000). In Antennariella pycnidia are reported to be small and dark brown, with a smooth or roughened pseudoparenchymatous wall, subglobose to obovoid, terminal on a short stalk or intercalary, somewhat lateral on conidiophores, with a short neck and ostioles at maturity. Conidiogenous cells are rare and not reported by Hughes (2000). Conidia are minute, hyaline, more or less ellipsoidal and 1-celled (Hughes 1976). In Capnodendron hyphae are dark brown, cylindrical or irregular, septate, smooth or roughened. Conidiophores are scattered, or gregarious, and velutinous, straight or irregularly bent, barely different from aerial hyphae, arising as upright branches of repent hyphae or as modified upturned ends of hyphae and very variable in length, brown to dark brown, more or less cylindrical, smooth to coarsely roughened, with lateral branches terminating in a characteristic conidiogenous cell (Hughes 1976, 2000). The terminal conidiogenous cell is very characteristic and more or less ovoid with a flat terminal scar left by the fallen conidium, sessile. Conidia are brown to dark brown, somewhat pale brown to brown, holoblastic, narrowly clavate to ellipsoidal or fusiform, straight or curved or irregularly curved, slightly constricted at the septa, smooth or coarsely roughened, sometimes with longitudinal striations, gently or abruptly tapered at the base to a flattened or denticulate scar, rounded at the apex or scarred at both ends of conidia in chains. Conidia and conidiophores can anastomose with each other; the asexual state has been found attached to ascostromata, with flexuous hyphal appendages in Antennulariella (Hughes 1976, 2000).

Woronichin (1925) referred this family along with Capnodiaceae and Coccodiniaceae to Capnodiales. Antennulariellaceae may differ from these two families in having completely closed ascomata with irregular hyphae and in its conidial states. Woronichin (1925) mentions that the Antennulariellaceae were the most representative family in the order, the two families (Capnodiaceae and Coccodiniaceae) with ostiolate ascomata, being the most advanced. Hughes (1976) however, reported that the ascomata of Antennulariella are ostiolate at maturity. Woronichin (1926) added Achaetobotrys as a second genus in Antennulariellaceae with two species, both referred to Limacinia by Barr (1961) and to Euantennaria by Hughes (1974). Hughes (2003) introduced the hyphomycetous sooty mould genus, Capnofrasera from New Zealand, Chile, Venezuela, Brazil, Canada and USA in the Antennulariellaceae, and compared the genus with Tomenticola, Paratomenticola, Sporidesmiopsis and Capnondendron. The family needs a detailed study of both sexual and asexual states to establish the relationships among the various genera. We tentatively include all names mentioned in Antennulariellaceae pending further studies.

Type: Antennulariella Woron., Trudy Byuro Prikl. Bot. 8(6): 771 (1915), MycoBank: MB 221 Fig. 10

Antennulariella fuliginosa (Material examined: RUSSIA, Sochi, Kraevskij Mountain, on living leaf of Ilex, 14 May 1913, Woronichin, holotype). a Herbarium specimen. b–d, h Hypha with ascomata. e, f Ascomata with long cylindrical appendage. g Hyphae. i Vertical section through ascoma. j Ascoma with ostiole. k Ascoma with a robust stalk. l Asci and ascospores (redrawn from Woronichin, 1915). Scale bars: b–d = 50 μm, e, f, h, i, k = 20 μm, g, j = 10 μm

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Foliar epiphytes as sooty moulds living on insect exudates. Mycelium brown to dark brown, thin to dense, cylindrical, irregular, septate, slightly constricted at the septa, smooth-walled. Sexual state: Ascomata developing from repeated divisions of hyphae, dark brown, sessile, with a pseudoparenchymatous wall, subglobose to globose, with a robust stalk, with long, cylindrical, hyphal appendages, wide at the base, tapering to the apex, ostiolate at maturity, terminally or laterally on hyphae. Peridium thin-walled, cells brown to pale brown, of textura angularis. Hamathecium aparaphysate. Asci 8-spored, bitunicate, fissitunicate, cylindrical, clavate to ellipsoidal. Ascospores overlapping, hyaline, 1-septate, with the upper cell slightly shorter and broader than the lower one, rounded end at both apices. Asexual state: coelomycetous, Antennariella pycnidial, terminal on long stalk, or lateral or intercalary, dark brown with pseudoparenchymatous walls, subglobose to ovoid, with ostioles. Conidia ellipsoidal, hyaline, one-celled; hyphomycetous asexual state, Capnodendron Hughes, mycelium superficial, scattered to dense, pale brown to dark brown, smooth-walled, cylindrical, branched, upright hyphae, straight or branched, septate, long, tapering to the apex. Some upright hyphae and their branches may become conidiogenous cells. Conidia in simple chains, subglobose, or ellipsoidal to cylindrical, brown to dark brown, with or without septa, slightly constricted at the septa, various lengths in multi-septate conidia, with a rounded or slight scar at one or both ends.

Notes: Antennulariella was introduced by Woronichin (1915) and now includes five species (Kirk et al. 2008, Index Fungorum 2013). Hughes (1976) has seen many collections which have characteristics similar to those of the type species as illustrated by Woronichin (1915). Phylogenetic estimates of Capnodiaceae and distribution of characters and classifications using character compatibility analysis show that Antennulariella and Acrogenotheca form one group; this group produces ascospores which have only one septum at maturity in the ascus, the spore dispersal strategy incorporates delayed states of hyphal initial germination and growth so that septation and pigmentation develop in steps, usually outside the ascus (Reynolds 1986).

Antennulariella batistae S Hughes (2000) was described as a new species with a hyphomycetous synanamorph Capnodendron trichomericola, which is the type of the genus Capnodendron. We therefore synonymize Capnodendron under Antennulariella above. Antennulariella batistae is a sooty mold which occurs alongside Metacapnodium moniliforme (L.R. Fraser) S. Hughes (Fraser) Hughes (1981), Euantennaria mucronata (Mont.) S Hughes (1974) and Trichopeltheca asiatica Batista et al. (Hughes 1965) and has been reported on more than 80 hosts, including ferns, conifers, monocotyledons and dicotyledons. Capnodendron trichomericola (Cif., Bat. & Nascim.) S. Hughes is also a synonym of Antennulariella batistae.

Antennariella unedonis (Maire & Sacc.) Bat. & Cif., is regarded as a synonym of Capnodium = Polychaeton (Capnodiaceae), while Microxiphium footii var. ciliolatum Sacc. is also listed as a synonym of Capnodium (= Polychaeton) (Index Fungorum 2013). Hughes (2000) lists Capnodendron and Antennariella as synanamorphs of Achaetobotrys batistae while Hughes (1976) links Achaetobotrys with Antennariella. Antennulariella concinnata (Fraser) S Hughes (1976) is neotypified, illustrated and described by Hughes (2007) who also mentions that Heteroconium is a synanamorph. It is therefore synonymized above. There is, however, considerable confusion in such linkages, as many different sooty mold species may occur on a single host. Linkage by association alone is not good enough evidence and in future studies individual spores should be isolated and subjected to sequence analysis to establish which sexual and asexual states are the same biological species.

Type species: Antennulariella fuliginosa Woron., Trudy Byuro Prikl. Bot. 8: 769–807 (1915), MycoBank: MB 224166

Other genera included

Achaetobotrys Bat. & Cif., Saccardoa 2: 49 (1963)

Type species: Achaetobotrys affinis (L.R. Fraser) Bat. & Cif., Saccardoa 2: 49 (1963)

Capnofrasera S. Hughes, N.Z. Jl Bot. 41(1): 139 (2003)

Type species: Capnofrasera dendryphioides S. Hughes, N.Z. Jl Bot. 41(1): 141 (2003)

Key to sexual genera of Antennulariellaceae

1. Ascostromata subglobose to globose, with hyphal appendages ................................................................Antennulariella

1. Ascostromata obovoid to broadly ellipsoidal, lacking hyphal appendages ...............................................Achaetobotrys

Argynnaceae Shearer & J.L. Crane, Trans. Br. mycol. Soc. 75(2): 193 (1980), MycoBank: MB 80470

Saprobic on plant stems or wood and in fresh water and terrestrial habitats. Sexual state: Ascomata apothecial or perithecial, solitary, superficial to slightly immersed, discal, dark brown to black, coriaceous, non-ostiolate, lacking a stipe. Excipulum thick, ectal excipulum composed of brown to dark brown tissues mixed with host cells, medullary excipulum composed of hyaline cells of textura porrecta. Hymenium composed of wide, septate, branching, hyaline pseudoparaphyses. Epithecium lacking. Asci 8-spored, bitunicate, fissitunicate, clavate, globose to subglobose, pedicellate, thick-walled, lacking a ocular chamber, exposed or not. Ascospores 2–3-seriate, or multi-seriate, papilionaceous to broadly reniform, 1-septate, light brown, orange to light orange brown, smooth-walled, bi- to multi-guttulate, with a dark band at the septum. Asexual states: Unknown.

Notes: The family Argynnaceae is characterized by taxa with apothecia and perithecia and papilionaceous, 1-septate, ascospores (Shearer and Crane 1980). Argynna has superficial, discal apothecia, with light brown ascospores, with a dark band at the septum (Shearer and Crane 1980). Lepidopterella has superficial to slightly immersed, globose to subglobose ascomata, and papilionaceous to broadly reniform, brown to orange-brown ascospores (Shearer and Crane 1980; Raja and Shearer 2008). Lepidopterella is therefore quite dissimilar to Argynna, however, Lepidopterella is included in this family because of the butterfly-shaped ascospores. Lumbsch and Huhndorf (2010) also include Argynna and Lepidopterella in Argynnaceae and we retain both genera in the family until molecular data become available. Shearer and Crane (1980) misinterpreted the asci of this family as being unitunicate when they introduced this family. However, our studies on herbarium material of Argynna polyhedron (Schwein.) and Morgan also indicate that this species has bitunicate asci.

Type: Argynna Morgan, J. Cincinnati Soc. Nat. Hist. 18: 41 (1895), MycoBank: MB 306

Figure 11

Argynna polyhedron (Material examined: USA, Missouri, Big Spring, on stalks of Zea mays, 15 April 1939, G.W. Martin (3922), BPI 618534). a Herbarium material. b Black ascoma on host surface. c Section through ascoma and peridium. d Septate and branched pseudoparaphyses. e-g Immature and mature asci. h-k Papilionaceous ascospores, with a dark band at the septum. Scale bars: b = 500 μm, c = 200 μm, d = 5 μm, e–g = 10 μm, h–k = 5 μm

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Saprobic on plant stems of maize and in terrestrial habitats. Sexual state: Mycelium branched, septate, finely roughened, yellowish-brown, growing around apothecium. Apothecia thin, solitary, superficial, discal, dark brown to black, non-ostiolate, with flat apex. Excipulum thick, formed on the host sufface, lacking side layers, basal layer thin, ectal excipulum composed of brown to dark brown tissues, mixed with host cells, medullary excipulum composed of hyaline cells of textura porrecta. Hymenium composed of wide, septate, branching, hyaline pseudoparaphyses. Epithecium lacking. Asci 8-spored, bitunicate, clavate, pedicellate, thick-walled, lacking an evident ocular chamber. Ascospores 2–3-seriate, multi-seriate, papilionaceous, 1-septate, light brown, smooth-walled, with a dark band at the septum. Asexual states: Unknown.

Notes: Argynna includes one species according to Shearer and Crane (1980). This genus is characterized by solitary, superficial apothecia and 1-septate papilionaceous, light brown, smooth-walled ascospores with a dark band at the septum. The genus Argynna is similar to members of the discomycetes based on the apothecium-like fruiting bodies.

Type species: Argynna polyhedron (Schwein.) Morgan [as ‘polyedron’], J. Cincinnati Soc. Nat. Hist. 18: 41 (1895), MycoBank: MB 140004

≡ Physarum polyhedron Schwein. [as ‘polyaedron’], Trans. Am. phil. Soc., Ser. 2 4(2): 257 (1832)

Other genus included

Lepidopterella Shearer & J.L. Crane, Trans. Br. mycol. Soc. 75(2): 194 (1980)

Type species: Lepidopterella palustris Shearer & J.L. Crane, Trans. Br. mycol. Soc. 75(2): 194 (1980)

Key to genera of Argynnaceae

1. Asci clavate, ascospores light brown, with a dark band at the septum ..................................................................Argynna

1. Asci globose to subglobose, ascospores brown to orange brown, without a dark band at the septum ..........Lepidopterella

Arthopyreniaceae Walt. Watson, New Phytol. 28: 107 (1929), MycoBank: MB 80473

Non-lichenized on bark in terrestrial, temperate forest habitats. Thallus reduced and ecorticate, whitish or indistinct. Sexual state: Ascomata scattered, prominent, brown-black, hemisphaerical, carbonaceous, ostiolate, ostiole round. Involucrellum not distinctly separated from excipulum, dark brown to carbonized. Excipulum dense, proso- to paraplectenchymatous in thin sections, brown. Hamathecium comprising 0.5–0.7 μm wide, densely packed paraphysoids connected to both the base and the top of the centrum, hyaline, straight to irregularly bent, branched and anastomosing. Asci forming in locules between the paraphysids, 8-spored, bitunicate, fissitunicate, broadly clavate, short-pedicellate, with rather broad, non-amyloid ocular chamber. Ascospores irregularly arranged to bi-seriate, oblong with the proximal end slightly tapering, hyaline, (1–3)-septate, with thin eusepta and rectangular lumina, smooth-walled, not or very slightly constricted at the septa, the upper cells often slightly larger than the lower cells. Asexual state: Pycnidia erumpent to prominent, visible as brown-black dots. Conidia acrogenous, either macro- or microconidia; macroconidia non-septate, bacillar, hyaline, 10–13 × 2–2.5 μm; microconidia non-septate, acicular, hyaline, 7–14 × 0.8–1 μm. Chemistry: Unidentified perithecial wall pigment reacting K+ green.

Notes: The family Arthopyreniaceae was originally established by Watson (1929) to include a large number of presumably lichenized fungi with perithecioid ascomata: Acrocordia, Athrismidium, Arthopyrenia, Bottaria, Celothelium, Laurera, Leptorhaphis, Microthelia, Microtheliopsis, Polyblastiopsis, Pseudosagedia, Raciborskiella, and Tomasellia. Watson (1929) united this amalgam of genera as a result of their shared trentepohlioid photobiont and branched, anastomosing paraphyses; however, Watson’s (1929) concept of Arthopyreniaceae included taxa now assigned to three different classes, at least six different orders, and eight different families (Harris 1975, 1995; Aptroot et al. 2008; Nelsen et al. 2009, 2011b). Thus, Acrocordia forms part of Monoblastiaceae (Dothideomycetes) and Microthelia is a synonym of Anisomeridium in the same family; Athrismidium is a synonym of Tomasellia which is currently classified in Naetrocymbaceae (Dothideomycetes), as well as Leptorhaphis; Bottaria is a synonym of Mycoporum in Mycoporaceae (Dothideomycetes); Celothelium belongs in Celotheliaceae (Chaetothyriomycetidae: Pyrenulales) and Microtheliopsis in Microtheliopsidaceae in the same subclass (Chaetothyriomycetidae: Chaetothyriales); Laurera is a synonym of Astrothelium in Trypetheliaceae (Dothideomycetes: Trypetheliales); Polyblastiopsis is a synonym of Julella (Dothideomycetes); Pseudosagedia belongs in Porinaceae (Lecanoromycetes: Ostropales); and Raciborskiella is a synonym of Strigula in Strigulaceae (Dothideomycetes).

Overall, Watson’s (1929) definition of Arthopyreniaceae encompassed several lichenized and some non-lichenized lineages currently classified in Dothideomycetes but unrelated to each other (Nelsen et al. 2009, 2011b). The family circumscription was eventually refined, but at the same time, some authors subsumed Arthopyreniaceae under Mycoporaceae (Riedl 1961; von Arx and Müller 1975) or Pleosporaceae (Poelt 1974; Harris 1975). The family was later reinstated by Barr (1979a), but Eriksson (1981) considered it a synonym of Xanthopyreniaceae; this stems from the author’s observations on Arthopyrenia rhyponta (Ach.) A. Massal., a species now included in Naetrocymbe (Harris 1995).

Following a broad revision of Arthopyrenia sensu lato (Harris 1975), Arthopyreniaceae was redefined to include only the genera Arthopyrenia, Julella, and Mycomicrothelia, distinguished from each other by their ascospore septation and pigmentation (Harris 1995). According to Harris (1995), the main characters uniting these genera are clypeate ascomata (i.e. with distinct involucrellum), presence of cellular pseudoparaphyses (i.e. developing from the top of the centrum; see Eriksson 1981), and fissitunicate asci with an ocular chamber. While two of these characters are rather universal among many perithecial fungi, the type of interascal hyphae actually varies within Arthopyrenia sensu Harris (1975, 1995), with different types found in the Arthopyrenia cerasi (Schrad.) A. Massal. group, A. cinchonae (Ach.) Müll. Arg. and relatives, and A. salicis A. Massal. (Foucard 1992). Arthopyrenia cerasi and the closely related A. lapponina Anzi, which represent Arthopyrenia sensu stricto (see below), have persistent, very densely arranged paraphysoids that develop from both the top and the base of the centrum and form locules for the asci (see below); these two species also share the K+ sordid-green perithecial wall. In A. cinchonae and relatives, the persistent physes develop from the bottom of the centrum, as correctly stated by Harris (1995), and taper towards the top. These species phylogenetically nest in Trypetheliaceae and are unrelated to Arthopyrenia sensu stricto (Nelsen et al. 2009, 2011a). The hitherto sequenced species of Julella and Mycomicrothelia, with a similar type of paraphyses, also cluster in Trypetheliaceae (Nelsen et al. 2009, 2011a). Finally, Arthopyrenia salicis forms periphysoids, whereas the paraphyses become gelatinous at maturity; it is the only other species of Arthopyrenia sensu lato sequenced besides the tropical group centered around A. cinchonae and occupies a position within Pleosporales (Nelsen et al. 2009, 2011a).

Thus, with Arthopyreniaceae and Arthopyrenia itself being obviously heterogeneous, the exact definition of the family hinges on its typification. Massalongo (1852) included 13 species in its original circumscription of Arthopyrenia, but did not define a holotype. Of these, five belong in Naetrocymbaceae, three in Monoblastiaceae, and one in Porinaceae, while a further one is a non-lichenized member of Dothideomycetes in the genus Cyrtidula. The remaining three species are A. analepta (Ach.) A. Massal., A. cerasi, and A. salicis, all temperate in their distribution. Arthopyrenia was first lectotypified by Fink (1910) with A. analepta; however, this name is illegitimate since Acharius (1798) cited Verrucaria olivacea Pers., a synonym of Porina borreri (Trevis.) D. Hawks. & P. James, as synonym of Lichen analeptus Ach., and hence, as already noted by Riedl (1961), A. analepta is automatically typified by the type of Verrucaria olivacea and thus a synonym of P. borreri. Riedl (1961) selected a different lectotype, A. rhyponta, which, however, is a member of Naetrocymbe (Harris 1995) and would make Arthopyrenia the valid name for that group of species. The problem was eventually solved by Jørgensen’s request (following suggestion by Harris) to reconsider the proposal of Hawksworth and David (1989), which ultimately lead to the conservation of Arthopyrenia with a conserved type, A. cerasi (Gams 1999). This conservation turned out to be quite fortunate, since based on phylogenetic and anatomical evidence, this is the only way to conserve Arthopyrenia and Arthopyreniaceae as a separate entity in a sense closest to what Harris (1995) had envisioned, albeit with only a single genus and few species belonging here. Thus, with Arthopyrenia forming four separate groups based on molecular and anatomical data, one including the largely tropical species now assigned to Trypetheliaceae (Nelsen et al. 2011a), one comprising the species now classified as Naetrocymbe (Harris 1995), and the two remaining groups representing the unusual A. salicis and the anatomically unique A. cerasi and relatives, the selection of the latter as the type defines Arthopyrenia and Arthopyreniaceae through the presence of densely arranged paraphysoids with the asci formed in locules, and by the peculiar, K + sordid-green reaction of the perithecial wall.

Type: Arthopyrenia A. Massal., Ric. auton. lich. crost. (Verona): 165 (1852), MycoBank: MB 325 Fig. 12

Arthopyrenia cerasi (Material examined: Rabenhorst, Lichenes Europaei 145) a-c Surface view of thallus and perithecial ascomata, showing dark brown hyphae. d-f Section through perithecium showing centrum, paraphysoids, and asci. g Branched and anastomosing paraphysoids. h, i Asci with ascospores embedded in hamathecium. j, k Individual asci with ascospores. l Ascus enlarged showing ascospores. Scale bars: a–c = 1 mm, d–l = 10 μm

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Non-lichenized on bark in terrestrial, temperate forest habitats. Thallus reduced and immersed in the substrate, whitish or indistinct, often forming dark brown hyphae over the substrate. Sexual state: Ascomata scattered, prominent, brown-black, hemisphaerical, carbonaceous, ostiolate, ostiole round. Involucrellum not distinctly separated from excipulum, dark brown to carbonized, K+ sordid-green in section. Excipulum dense, proso- to paraplectenchymatous in thin sections, brown. Hamathecium comprising 0.5–0.7 μm wide, densely packed paraphysoids connected to both the base and the top of the centrum, hyaline, straight to irregularly bent, branched and anastomosing. Asci forming in locules between the paraphysids, 8-spored, bitunicate, fissitunicate, broadly clavate, short-pedicellate, with rather broad, non-amyloid ocular chamber. Ascospores irregularly arranged to bi-seriate, oblong with the proximal end slightly tapering, hyaline, (1–)3-septate, with thin eusepta and rectangular lumina, smooth-walled, not or very slightly constricted at the septa, the upper cells often slightly larger than the lower cells. Asexual state: Pycnidia erumpent to prominent, visible as brown-black dots. Conidia acrogenous, either macro- or microconidia; macroconidia non-septate, bacillar, hyaline; microconidia non-septate, acicular, hyaline. Chemistry: Unidentified perithecial wall pigment reacting K+ green.

Notes: Arthopyrenia, once a large collective genus including over hundred accepted species, and with over 600 epithets associated with that name (Harris 1975), is now reduced to less than a handful of species which are essentially northern temperate in their distribution and not lichenized. The unique features of these species are the densely arranged paraphysoids developing from both the top and base of the centrum, and the K+ sordid-green perithecial wall (Foucard 1992; Harris 1995; Smith et al. 2009. While Harris (1975) transferred a large number of species to genera unrelated to Arthopyrenia, he suggested that Arthopyrenia could still be further reduced and might contain groups of unrelated taxa. Harris (1995) continued his refinement of Arthopyrenia by transferring a number of species from the A. rhyponta-A. saxicola group to Naetrocymbe and again noted the genus may contain further aberrant taxa. Not all agreed with Harris’ transfer of these species to Naetrocymbe and have continued to recognize species from this group as Arthopyrenia (Aptroot 1998, 2002a; Coppins 2002).

A number of molecular studies have demonstrated the placement of Arthopyrenia salicis and an unidentified Arthopyrenia species in Pleosporales (Lumbsch et al. 2005; Mugambi and Huhndorf 2009a; Schoch et al. 2009b; Tanaka et al. 2009; Zhang et al. 2009a, 2012a). Arthopyrenia salicis has been considered an unusual Arthopyrenia species, as it forms short periphysoids and produces gelatinizing physes at maturity (Harris 1973; Coppins 1988; Foucard 1992), both of which are aberrant characters for Arthopyrenia sensu lato. Harris (1995) ultimately retained this taxon in Arthopyrenia, but noted his uncertainty about this decision. Molecular data have demonstrated that A. salicis is related to Roussoella and Roussoellopsis (Tanaka et al. 2009; Schoch et al. 2009b; Nelsen et al. 2011a, b), two genera of bambusicolous fungi. Arthopyrenia salicis forms periphysoids produced in a gelatinized matrix, while these two genera, differ from A. salicis by the formation of paraphysoids and oblong to fusiform, hyaline spores lacking striation (Hino and Katumoto 1965; Hyde 1997). In contrast, A. cinchonae and related species were found to cluster in Trypetheliales (Nelsen et al. 2011b). Notably, neither the A. cinchonae nor the A. salicis group have names available at generic level. In contrast, Arthopyrenia sensu stricto has at least two generic synonyms in Ciferriolichen and Mycociferria, both based on A. lapponina (Tomaselli and Ciferri 1952, Ciferri and Tomaselli 1953). Further anatomical and phylogenetic revision of many additional species is required to adequately place all species currently assigned to Arthopyrenia sensu lato in one of these clades.

The other names tentatively listed as synonyms of Arthopyrenia require further taxonomic studies. Thus, Giacominia was described as the ‘fungal’ element of Arthonia parolinii Beltram. and the status of that species is unclear; it is not included in the current checklist of Italian lichens. The same applies to Arthopyrenia maroccana (M. Choisy) Zahlbr. the type of Mesopyrenia, and Mycarthopyrenia sorbi Keissl., the type of Mycarthopyrenia. Other genera commonly listed as synonyms of Arthopyrenia belong elsewhere: Leiophloea (Ach.) Gray, based on Verrucaria subgen. Leiophloea Ach. [the name Leiophloea Trevis. does not exist and is a lapsus for (Ach.) Gray], was lectotypified with Lichen analeptus Ach. by Riedl (1961) and hence is a synonym of Porina s.lat. (see above). The same applies to Mycoarthopyrenia Cif. & Tomas., which is also based on Lichen analeptus. Magmopsis Nyl. appears to be a synonym of Pyrenopsis in Lichinaceae. The names Jattaeolichen and Jattaeomyces are both based on Verrucaria epidermidis var. pyrenastrella Nyl., which is a synomyn of Naetrocymbe punctiformis, and hence these names are synonyms of Naetrocymbe.

Type species: Arthopyrenia cerasi (Schrad.) A. Massal., Ric. auton. lich. crost. (Verona): 167 (1852), MycoBank: MB 377023

≡ Verrucaria cerasi Schrad., Nov. gen. pl. (Lipsiae) 22: 86 (1797)

Ascoporiaceae Kutorga & D. Hawksw., Syst Ascom 15(1–2): 25 (1997), MycoBank: MB 81923

Saprobic on decorticated wood in terrestrial habitats. Sexual state: Ascostromata relatively large, superficial, circular or cupulate, slightly raised, stipitate, solitary, exterior surrounded by brown mycelium, orange to red-brown at the margin or edge, dark brown to black in the centre, cells of ascostromata comprising a maze-like arrangement of mixed, brown-walled cells of textura angularis to subglobosa or epidermoidea, multi-loculate. Locules in a single layer at the periphery of the ascostromata, subcylindric-elongate, obpyriform to ovoid, crowded, opening by small, narrow, vertical ostiole. Hamathecium comprising numerous filiform, branched, septate pseudoparaphyses embedded in a gelatinous matrix. Asci 8-spored, bitunicate, fissitunicate, narrowly cylindric-clavate, pedicellate, apically rounded with a small ocular chamber, staining pinkish to purplish in 3 % KOH. Ascospores bi-seriate, fusiform, 1-septate, constricted at the septum, hyaline to yellowish when immature, olive-brown to dark brown when mature. Asexual state: see notes under genus.

Notes: Ascoporia was introduced by Samuels and Romero (1991) and placed in Patellariaceae. There is considerable variation in ascostroma structures, ascus shape and ascospore colour of A. lateritia Samuels & A.I. Romero. Typically the ascostromata are darkly pigmented in section, although bright coloured features may be seen on the substrate as well; asci are obviously bitunicate with a small ocular chamber; ascospores are typically fusiform, 1-septate and dark brown at maturity (Samuels and Romero 1991). Kutorga and Hawksworth (1997) investigated the isotype of A. lateritia and found that overall morphologies distinguish it from the type species of Patellariaceae (Patellaria Fr. 1882). Therefore, they separated Ascoporia from Patellariaceae and introduced the family Ascoporiaceae (Kutorga and Hawksworth 1997) comprising a single genus Ascoporia.

An earlier name for Ascoporia was found in Pseudosolidum by Rossman et al. (1999) when they examined a portion of the type specimen of Pseudosolidum at K.

Type: Ascoporia Samuels & A.I. Romero, Bolm Mus. paraense ‘Emílio Goeldi’, sér. bot.7(2): 264 (1993) [1991]

Synonym:

Pseudosolidum Lloyd (as McGinty), Mycol. Writ. 7: 1206 (1923), MycoBank: MB 26458 Fig. 13

Pseudosolidum solidum (Material examined: BRAZIL, Belem, Iha do Combu, Estacao Experimental Combu, Para. 01d30’s, 48d27’w, on decorticated wood, January 1989, G.J. Samuels; K.F. Rodrigues (6207), BPI1109903, isotype of Ascoporia lateritia). a Ascostroma on substrate in surface view. b Crowded brown mycelium surrounding ascostroma. c, d Section through ascostroma showing arrangement of locules. e Cells of ascostroma. f Hamathecium. g–i Immature and mature asci in 3 % KOH. Note colour changes. j–l Ascospores. Note remnants of mucilage in j and k. Scale bars: a = 5 mm, b = 1 mm, c–d = 200 μm, e = 10 μm, f = 5 μm, g–i = 50 μm, j–l = 20 μm

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Notes: Pseudosolidum (= Ascoporia) is a unique genus with locules lying in a peripheral layer around the cushion-like ascostromata, but is typical of Dothideomycetes in its hamathecium characters (Fig. 13). The most similar genus may be Shiraia which also has large orange ascostromata; however the habit (an endophyte on bamboo) and ascospores (muriform) are distinct (Hennings 1900; Liu et al. 2012a). Pseudosolidum contains a single species P. solidum (Berk. & M.A. Curtis) Lloyd (Fig. 13). Rossman et al. (1999) noted that Ascoporia lateritia Samuels & A.I. Romero, which is illustrated here, is a synonym of Pseudosolidum solidum. Kutorga and Hawksworth (1997), in their examination of the isotype specimen of A. lateritia, found a coelomycete similar to Plectophomella, which they considered as a possible asexual state. This coelomycetous asexual morph has cylindrical, hyaline, smooth, phialidic conidiogenous cells and ellipsoid to ovate, 1-celled, hyaline conidia and was described by Kutorga and Hawksworth (1997). There are no molecular data to support this link.

Type species: Ascoporia lateritia Samuels & A.I. Romero, Bolm Mus paraense ‘Emílio Goeldi’, sér bot 7(2): 264 (1993) [1991], MycoBank: MB 360813

= Pseudosolidum solidum (Berk. & M.A. Curtis) Lloyd (as McGinty), Mycol. Writ. 7: 1206 (1923)

≡ Hypoxylon solidum Berk. & M.A. Curtis, J. Acad. nat. Sci. Philad., N.S. 2(6): 286 (1854) [1853]

Asterinaceae Hansf., Mycol. Pap. 15: 188 (1946), MycoBank: MB 80492

Possible synonyms:

Leveillellaceae Theiss. & Syd., Annls mycol. 16(1/2): 33 (1918)

Lembosiaceae Hosag., in Hosagoudar, Abraham & Biju, J. Mycopathol. Res. 39(1): 61 (2001)

Foliar epiphytes obligately biotrophic on the upper or lower surface of leaves, rarely on stalks or shoots of plants, often forming web-like circular structures containing superficial black ascomata. Sexual state: Mycelium superficial, usually hyphopodiate. Ascomata superficial, forming below blackened mycelia, flattened, roundish in outline, elongate or linear, opening by radiating star-like or longitudinal splits, interascal tissue absent. Upper wall comprising rows of cells, radiating from the central ostiole to the periphery, base poorly developed. Asci 8-spored, bitunicate, fissitunicate, globose or broadly saccate, with short indistinct pedicel, lacking a distinct ocular chamber, occasionally with J+ (blue) gel remnants. Ascospores 2–5-seriate or fasciculate or conglobate, ellipsoidal, occasionally cylindrical, hyaline or brown, mostly 2-septate, mostly brown when mature, wall smooth or roughened. Asexual state: coelomycetous states with pycnidia or pycnothyria. Conidiophores branched or unbranched, hyaline or brown. Conidiogenous cell monoblastic or percurrent, hyaline or brown. Conidia brown, phragmosporous, single; hyphomycetous states without conidiomata or sporodochia then gelatinous, pale, mycelium hyaline or darkly pigmented. Conidia forming on stalked conidiophores, non-septate or 1-septate, variable in shape, e.g. globose to ellipsoidal, clavate, mostly brown to dark brown at maturity, mostly with hyaline areas or germ pores.

Notes: Eriksson (1981) suggested Asterinaceae should be restricted to certain tropical Melastomataceae and a remarkable common feature was the J+ granular coating on the surface of the asci. We have studied many genera of Asterinaceae and the J+ coating on the surface of the asci seems rare, while species have been recorded from a wide range of non melastomataceous hosts. Eriksson (1981) also compared the Asterinaceae with Englerulaceae; ascomata forming beneath the superficial mycelium in the former and on the mycelium in the latter.

The important features of Asterinaceae are the superficial, web-like, black colonies that form on the upper and lower surface of leaves and are typical of Asterinaceae (see Fig. 14). The mycelium is usually hyphopodiate. The thyriothecia form below the mycelium and comprise radiating cells and have a concentrically ridged surface – albeit hard to observe under the light microscope. The thyriothecia open by gelatinization of the shield cells and are usually stellate but may be longitudinal. Asci are saccate and ascospores are mostly conglobose and hyaline when young and becoming brown at maturity. The significance of the J+ ascal coating as mentioned by Eriksson (1981) is unclear.

Asterina millettiae Sexual state. (Material examined: PHILIPPINES, Luzon, Laguna Province, Mount Makiling, on living leaves of Cynometra ramiflora, February 2012, KD Hyde, MFLU 13–0087). a Ascomata superficial on host substrate.b, c Star-shaped opening.d, e Superficial mycelium. f, g Ascoma wall cells radially arranged. h Mycelium with hyphopodiate. i Immature ascus. j Immature ascus in Melzer’s reagent. k Ascus at maturity. l Immature ascospore in Melzer’s reagent. m Ascospore at maturity in Melzer’s reagent. n-p Ascospores at maturity. Scale bars: b–e = 100 μm, f = 50 μm, g, h = 20 μm, i–p = 10 μm

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Sivanesan (1984) stated that Asterostomella, Clasterosporium, Mitteriella, Sarcinella, Sporidesmium, Triposporium and Septothyrella are the asexual states of Asterinaceae. In addition to these, Wijayawardene et al. (2012) listed Acarella, Asterostromina, Asterostomula, Clasterosporium, Clasterosporium, Elachopeltis, Eriothyrium, Leprieurina, Mahanteshamomyces, Peltasterella, Pirozynskia, Septoidium, Septothyrella, Shivamyces, Sirothyriella, Thyrinula and Triposporium as also belonging to Asterinaceae. However the placement of Septoidium in Asterinaceae is confused as Ellis (1971) and Sivanesan (1984) stated that Septoidium is the asexual state of Perisporiopsis (Parodiopsidaceae). Chaverri and Gazis (2010) also confirmed this link. Seifert et al. (2011) also concluded that Septoidium belonged to Parodiopsidaceae and Parodiellaceae, hence we exclude Septoidium from Asterinaceae. Hosagoudar (2004) showed that Shivamyces is the asexual state of Alina (Parodiopsidaceae) hence we remove Shivamyces also from Asterinaceae. Most of these placements are based on morphological characters and presence on the same host. Hofmann et al. (2010) showed that this family clustered together with Venturiaceae within Dothideomycetes based on LSU and SSU data. Our phylogenetic results (Figs. 1 and 2) showed that it belongs to Asterinales and is chose to Venturiales.

Type: Asterina Lév., Annls Sci. Nat., Bot., sér. 3 3: 59 (1845), MycoBank: MB 409

Figures 14 and 15

Asterina phlogacanthi Asexual state (Material examined: THAILAND, Chiang Rai, Tasud, on living leaves of Clinacanthus nutans, May 2012, Saowanee Wikee, MFLU 13–0088). a Pycnothyrium superficial on host substrate. b Star-shaped opening. c Conidia arranged in pycnothyrium. d Conidioma wall cells radially arranged. e Conidioma when immature. f Thin basal layer of the pycnothyrium wall. g Mycelium bearing hyphopodia. h-i Conidia. j Conidiogenous cells producing conidia. Scale bars: e, g, h, i, j = 20 μm, b, d, f = 50 μm, c = 100 μm

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Foliar epiphytes obligately biotrophic on leaves, forming blackened, circular, elongate or X-or Y-shaped areas. Mycelium spreading over the leaf surface, with hyphopodia and scattered with superficial ascomata. Hyphopodia circular, lateral,alternating, sparsely septate, penetrating the host cells with haustoria. Thyriothecia solitary, gregarious, superficial, easily removed from the host surface, rounded, black, dehiscing to open by slit-like, star-like, or X or Y-shaped fissures when mature. Upper wall comprising a thin layer of mostly neatly arranged dark cells, which are branched at the outer rim, from which mycelium strands develop, basal wall layer poorly developed. Hamathecium usually lacking pseudoparaphyses. Asci 8-spored, bitunicate, fissitunicate dehiscence not observed, oblong to subglobose, lacking a pedicel, ocular chamber not observed, not staining blue in IKI, asci arranged vertically in the thyriothecium. Ascospores overlapping, conglobate, hyaline when immature, brown when mature, 1-septate, strongly constricted at the septum. Asexual states “Asterostomella”, “Clasterosporium”-like, “Mahanteshamyces”. Mycelium brown, superficial, with appressoria. Conidiomata pycnothyria flattened, dimidiate, radiate, orbicular, stellately dehisced at the centre. Conidia pycnothyriospores, brown, ovate, pyriform, angular, or wall straight to sinuate.

Type species: Asterina melastomatis Lév., Annls Sci. Nat., Bot., sér. 3 3: 59 (1845), MycoBank: MB 218181

Other genera included

Aphanopeltis Syd., Annls mycol. 25(1/2): 82 (1927)

Type species: Aphanopeltis phoebes Syd., Annls mycol. 25(1/2): 82 (1927)

Asterotexis Arx, Fungus, Wageningen 28: 6 (1958)

Type species: Asterotexis cucurbitacearum (Rehm) Arx, Fungus, Wageningen 28: 6 (1958)

Batistinula Arx, Publções Inst. Micol. Recife 287: 4 (1960)

Type species: Batistinula gallesiae Arx, Publções Inst. Micol. Recife 287: 6 (1960)

Cirsosia G. Arnaud, Annals d’École National d’Agric. de Montpellier, Série 2 16(1–4): 127 (1918) [1917]

Type species: Cirsosia manaosensis (Henn.) G. Arnaud [as ‘manaoensis’], Annals d’École National d’Agric. de Montpellier, Série 2 16(1–4): 127 (1918) [1917]

Halbania Racib., Crypt. Par. Java: no. 89 (1889)

Type species: Halbania cyathearum Racib., Crypt. Par. Java: 89 (1889)

Lembosia Lév., Annls Sci. Nat., Bot., sér. 3 3: 58 (1845). MycoBank: MB2724

Type species: Lembosia tenella Lév., Annls Sci. Nat., Bot., sér. 3 3: 58 (1845)

Meliolaster Höhn., Ber. dt. bot. Ges. 35: 701 (1918)

Type species: Meliolaster clavispora (Pat.) Höhn., Ber. dt. bot. Ges. 35: 701 (1918)

Prillieuxina G. Arnaud, Annals d’École National d’Agric. de Montpellier, Série 2 16(1–4): 161 (1918) [1917], MycoBank: MB4365

Type species: Prillieuxina winteriana (Pazschke) G. Arnaud, Annals d’École National d’Agric. de Montpellier, Série 2 16(1–4): 162 (1918) [1917], MycoBank: MB160955

Schenckiella Henn., Bot. Jb. 17: 523 (1893)

Type species: Schenckiella marcgraviae Henn., Bot. Jb. 17: 523 (1893)

Trichasterina G. Arnaud, Arnaud, Annals d’École National d’Agric. de Montpellier, Série 2 16(1–4): 172 (1918) [1917]

Type species: Trichasterina styracis (Theiss.) G. Arnaud, Annales de l’École Nationale d’Agriculture de Montpellier, N.S. 16(1–4): 172 (1918)

Uleothyrium Petr., Annls mycol. 27(5/6): 388 (1929)

Type species: Uleothyrium amazonicum Petr., Annls mycol. 27(5/6): 388 (1929)

Key to sexual genera of Asterinaceae

1. Ascospores with more than two cells ................................2

1. Ascospores 2-celled ...........................................................6

2. Ascospores hyaline .........................................Uleothyrium

2. Ascospores brown ..............................................................3

3. Ascospores cylindrical, 4-septate, light brown .......................................................................Schenckiella

3. Ascospores not cylindrical, with less than 4 septa ...........4

4. Ascospores clavate, reddish brown ...................Meliolaster

4. Ascospores ellipsoidal-fusiform, 3-septate, dark brown, ............................................................................................5

5. On dicotyledon leaves .......................................Batistinula

5. On fern leaves ......................................................Halbania

6. Ascospores clavate, apical cell large and rounded, asci cylindrical to clavate, on Cucurbitaceae ...............Asterotexis

6. Ascospores conglobose, ellipsoidal, mostly on other hosts ............................................................................................7

7. Thyriothecium mostly elongate, opening by a stellate fissure, ascospores oblong to obvoid, hyaline to brown .................................................................................... Lembosia

7. Thyriothecium mostly rounded, opening by a stellate fissure, ascospores conglobate, brown ......................................8

8. Ascospores hyaline at maturity, hyphopodia arrow-like, lateral and alternative, asexual state Elachopeltis-like ............................................................................ Aphanopeltis

8. Ascospores brown at maturity, not with the above combination of characters ................................................................9

9. Superficial mycelia without spines ...............Trichasterina

9. Superficial mycelia with spines .......................................10

10. Ascospores ellipsoidal, lower cell longer and narrower ........................................................................Prillieuxina

10. Ascospores conglobose ..................................................11

11. Hyphopodia lateral and alternating, composed of three cells, with the middle cell swelling, ascospores 2-celled, clearly ornamented ...................................................................Cirsosia

11. Hyphopodia circular, lateral and alternating, sparsely septate, some ascospores ornamented ................Asterina

Aulographaceae Luttrell ex P.M. Kirk et al., in Kirk et al., Ainsworth & Bisby’s Dictionary of the Fungi, Edn 9 (Wallingford): ix (2001), MycoBank: MB 82122

Possible synonyms:

Aulographaceae Luttr., in Ainsworth, Sparrow & Sussman, The Fungi (London) 4A: 154 (1973)

Lembosiaceae Hosag., in Hosagoudar, Abraham & Biju, J. Mycopathol. Res. 39(1): 61 (2001)

Lembosiaceae Höhn., Annls mycol. 16(1/2): 146 (1918)

Saprobic on dead leaves. Sexual state: Ascomata superficial, a thyriothecium, solitary, some fused, scattered, usually elongate, slightly convex or shield-like, mostly linear, some furcate or branched, anastomosed, triangular in shape, dehiscing by a slit, dark brown to black, red-brown at the margin, mycelium appressed towards the base. Peridium composed of very thin, tightly packed textura epidermoidea, somewhat brittle, easily broken. Hamathecium of asci and lacking pseudoparaphyses. Asci 8-spored, bitunicate, fissitunicate, broadly ellipsoid or subclavate-oblong, or globose to subglobose, apex thickened, pedicel stumpy or lacking, ocular chamber not distinct. Ascospores 2–3-seriate, narrowly obovoid to clavate, 1-septate, not constricted at the septum, upper cell wider, hyaline, guttulate, smooth-walled. Asexual state: Unknown.

Notes: The family Aulographaceae was introduced by Luttrell (1973) and based on the tribe Aulographées (Arnaud 1930) which is an invalid name (Index Fungorum 2013). Luttrell (1973) did not provide a Latin diagnosis and thus his family was also invalid, however this was later validated with a Latin diagnosis in Kirk et al. (2001). Subsequently, it was accommodated in Microthyriales based on the flattened thyriothecial ascomata (Kirk et al. 2008), but treated as a distinct family due to the opening by a longitudinal slit at maturity to expose the asci (George 1895; Luttrell 1973; Sierra 2006). Presently, the family comprises Aulographum and Polyclypeolina (Lumbsch and Huhndorf 2010). These genera occur on the surface of leaves (Fig. 16). In Polyclypeolina ascomata form irregularly, are clustered and fused, without a longitudinal slit, and asci and ascospores differ from Aulographum (Hansford 1945; Batista 1959, Fig. 3). Based on morphological characters this genus can be excluded and placed in Ascomycetes, genera incertae sedis. Molecular data is needed to confirm its correct placement. Wu et al. (2011a) added Lembosiella to Aulographaceae based on ascomatal structures resembling those of Aulographum. The ascomata are typically dimidiate, superficial thyriothecia and similar to Asterinaceae (Batista 1960; Ellis 1980). Ascomatal structures of Asterinaceae are generally but not always circular with outwardly radiating mycelium which dehisces by irregular fissures (Hansford 1946; Batista et al. 1958). In addition, they produce appressoria (Hofmann et al. 2010; Hosagoudar 2012), a character which may distinguish these families. Aulographum is similar to Aulographina and Lembosia in Asterinaceae and these genera may be synonyms. Aulographum may also be placed in Asterinaceae. For the proper placement of these taxa, molecular data is needed.

Aulographum hederae (Material examined: on leaves of Hedera sp., BPI1108478) a Thyriothecia. b-d Squash mount of thyriothecia. e-h Immature and mature asci. i-n Ascospores. Scale bars: a = 500 μm, d = 50 μm, e–n = 10 μm

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Type: Aulographum Lib., Pl. crypt. Arduenna, fasc. (Liège) 3: no. 272 (1834), MycoBank: MB 461 Fig. 16

Saprobic on dead leaves. Sexual state: Ascomata superficial, a thyriothecium, solitary, some fused, scattered, usually elongate, slightly convex or shield-like, mostly linear, some furcate or branched, or anastomosed and triangular in shape, dark brown to black, red brown at the margin, dehiscing by a longitudinal slit, mycelium appressed outwards from the base. Peridium composed of very tightly packed textura epidermoidea, somewhat brittle, easily broken. Hamathecium of asci, pseudoparaphyses lacking. Asci 8-spored, bitunicate, fissitunicate, broadly ellipsoid, subclavate-oblong, apex thickened, pedicel stumpy or lacking, ocular chamber not distinct. Ascospores 2–3-seriate, narrowly obovoid, clavate, 1-septate, not constricted at the septum, upper cell wider, hyaline, guttulate, smooth-walled. Asexual state: Unknown.

Notes: Aulographum was introduced by Libert (1834), and is typified by Aulographum hederae Lib.. The type specimen of Aulographum hederae was not available for study, therefore the illustration is derived from BPI 1108478 which appears to be an authentic specimen (Fig. 16). In the protologue by Libert (1834), thyriothecia are slightly linear, longitudinally splitting and produce oval-oblong, 1-septate, hyaline ascospores (Massee 1895). This genus comprises 75 epithets (Index Fungorum 2013). Several of the species however, were transferred to other families, such as Asterinaceae, Elsinoaceae, Leptopeltidaceae, or Lophiostomataceae. Aulographum hederae is characterized by longitudinally splitting ascomata and a peridial wall appressed by mycelium with bright-colours at the margin (Fig. 16a–d). These are different from Polyclypeolina, presently included in the same family (Hansford 1945; Batista 1959).

Type species: Aulographum hederae Lib., Pl. crypt. Arduenna, fasc. (Liège) 3: no. 272 (1834), MycoBank: MB 161393

=Aulographum vagum Desm. Annls Sci. Nat., Bot., sér. 2 19: 362 (1843)

Other genera included

Echidnodella Theiss. & Syd., Annls mycol. 15(6): 422 (1918) [1917]

Type species: Echidnodella linearis (Syd. & P. Syd.) Syd., Annls mycol. 15(6): 422 (1918) [1917]

Echidnodes Theiss. & Syd., Annls. Mycol. 15(6): 422 (1918) [1917]

Type species: Echidnodes liturae (Cooke) Theiss. & Syd. [as ‘litsurae’], Annls. Mycol. 15(6): 422 (1918) [1917]

Lembosina Theiss., Annls mycol. 11(5): 437 (1913)

Type species: Lembosina aulographoides (E. Bommer et al.) Theiss., Annls mycol. 11(5): 437 (1913)

Morenoina Theiss., Annls mycol. 11(5): 434 (1913)

Type species: Morenoina antarctica (Speg.) Theiss., Annls mycol. 11(5): 434 (1913).

Thyriopsis Theiss. & Syd.. Annls mycol. 13(3/4): 369 (1915).

Type species: Thyriopsis halepensis (Cooke) Theiss. & Syd., Annls mycol. 13(3/4): 369 (1915)

Key to genera of Aulographaceae

1. Ascospores hyaline or yellowish-brown at maturity .........2

1. Ascospores brown at maturity ...........................................4

2. Thyriothecia star-like, with thin upper layer of brick-like cells, opening with stellate fissures, on grasses ................................................................................Morenoina

2. Thyriothecia linear, or occasionally with three arms, cells of upper layer less distinct ..........................................................3

3. Ascospores conglobose, strongly constricted at the septum .................................................................................Thyriopsis

3. Ascospores ellipsoidal or subclavate, slighty constricted at the septum ...............................................................................4

4. Thyriothecia very long, always linear ..............Echidnodes

4. Thyriothecia relatively short, linear, or occasionally with three arms .........................................................Aulographum

5. Ascospores with protrusion at the base, light brown ................................................................................Lembosina

5. Ascospores conglobose, brown .....................Echidnodella

Bambusicolaceae D.Q. Dai & K.D. Hyde, fam. nov., MycoBank: MB 804293

Saprobic on decaying bamboo culms. Sexual state: Ascomata small, solitary, scattered, immersed, conical, ostiolate, coriaceous. Peridium thin, composed of brown and thick-walled cells of textura angularis, with the basal part composed of thinner, hyaline, smaller cells. Hamathecium of relatively thin, dense, anastomosing and branching interascal filaments. Asci 8-spored, bitunicate, fissitunicate, cylindrical, with a short furcate pedicel, apically rounded, with a shallow ocular chamber. Ascospores 2–3-seriate, slightly broad fusiform, 1-septate, narrowly rounded at both ends, hyaline, surrounded by a gelatinous sheath. Asexual state: Conidiomata small, pycnothyrial, acerous or subglobose, solitary, scattered, immersed to semi-immersed. Conidiophores indistinct. Conidiogenous cells holoblastic, annelidic, discrete, cylindrical, and smooth. Conidia pale brown to dark brown, 1–3-septate, cylindrical, straight or slightly curved, obtuse at the apex, with many guttules.

Typus: Bambusicola D.Q. Dai & K.D. Hyde

Notes: The family Bambusicolaceae is established to accommodate the genus Bambusicola which comprises three species. Morphologically, Bambusicolaceae species are similar to genera in Tetraplosphaeriaceae i.e. Tetraploasphaeria, Triplosphaeria and Polyplosphaeria (Tanaka et al. 2009), but Dai et al. (2012) phylogenetically showed Bambusicola species to belong in a different lineage. All the members in Tetraplosphaeriaceae have immersed to superficial, globose to subglobose ascomata, cylindrical to clavate asci and hyaline, fusiform ascospores surrounded by a mucilaginous sheath (Tanaka et al. 2009). Both Triplosphaeria and Bambusicola massarinia D.Q. Dai & K.D. Hyde have similar rim-like side walls (Tanaka et al. 2009; Dai et al. 2012). All genera in Tetraplosphaeriaceae have been linked to “Tetraploa”-like hyphomycetous states (Tanaka et al. 2009), however, coelomycetous states were observed in all species of Bambusicolaceae in culture or on host culms. Molecular data show this family to belong in Pleosporales, related to Massarinaceae and Montagnulaceae (Fig. 1)

Type: Bambusicola D. Q. Dai & K.D. Hyde, Cryptog. Mycol.33(3): 367 (2012), MycoBank: MB 801041 Fig. 17

Bambusicola massarinia (Material examined: THAILAND, Chiang Rai Province, Doi Fung, on decaying culm of bamboo, 4 May 2011, Dong-Qin Dai DDQ0003, MFLU12-0405, holotype). a Bamboo type specimen. b Habit of immersed ascomata on host. c, d Section through ostiole with periphyses. e Section of peridium. f Section of ascoma. g–i Hyaline and 1-septate ascospores. j Ascospore with gelatinous sheath. k Germinating ascospore. l Pseudoparaphyses above asci. m Asci with ascospores. n Fissitunicate ascus. o Apices of asci. p–r Conidiomata surrounded by mycelium. s, t, w Conidiophores and conidiogenous cells producing conidia., u, v Conidiogenous cells with 1–5 annellations. x–4 One septate conidia. Scale bars: a = 50 mm, b, p–r = 100 μm, c–f = 50 μm, g–k, s–4 = 5 μm, l–o = 10 μm

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Notes: Bambusicola includes four species (Dai et al. 2012), all from bamboo.

Type species: Bambusicola massarinia D.Q. Dai & K.D. Hyde, Cryptog. Mycol. 33(3): 370 (2012), MycoBank: MB 801045

Biatriosporaceae K.D. Hyde, fam. nov., MycoBank: MB 803357

Saprobic on decaying submerged intertidal mangrove wood. Sexual state: Ascomata relatively large, solitary to gregarious, pyreniform, immersed, lying horizontal to the host surface, ostiolate, periphysate, black, coriaceous, with ostiole recurving upwards to pierce the host surface. Peridium thin, composed of brown pseudoparenchymatous cells. Hamathecium of dense, thin (ca 1 μm diam) anastomosing, branching, trabeculate pseudoparaphyses embedded in a gelatinous matrix. Asci 8-spored, bitunicate, fissitunicate, cylindrical, with a relatively long pedicel, with a large, distinct ocular chamber. Ascospores overlapping uni-seriate, fusiform, dark brown, with hyaline swollen rounded ends, with 1–4 septa towards the ends, lacking septa in the centre part, smooth-walled, releasing mucilage from the ends. Asexual state: Unknown.

Typus: Biatriospora K.D. Hyde & Borse

Notes: This remarkable genus was introduced by Hyde and Borse (1986) for a mangrove inhabiting species and has since been collected many times and shown to have a wide ranging distribution and to occur on a wide range of hosts. The dark brown ascospores are unusual in that they are fusiform with hyaline, rounded swollen ends which release mucilage (Hyde and Jones 1989) and 1–4 septa are situated near the ends. In previous classifications this genus would have sat nicely in Melanommataceae, as it has trabeculate pseudoparaphyses. However, Liew et al. (2000) show this character to be unreliable in predicting family placement of genera in Dothideomycetes. In the phylogenetic tree (Figs. 1 and 2) this genus sits independently with Roussoella and Roussoellopsis strains with strong support (92 %). A new family, Biatriosporaceae in the Pleosporales is therefore introduced to accommodate this monotypic genus.

Type: Biatriospora K.D. Hyde & Borse, Mycotaxon 26: 263 (1986), MycoBank: MB 801041 Fig. 18

Biatriospora marina (Material examined: SEYCHELLES, on submerged wood of Rhizophora, 2 January 1985, K.D. Hyde, IMI 297768, holotype). a Ascomata semi-immersed on host. b Vertical section of ascoma. c Peridium. d Pseudoparaphyses stained with lactophenol cotton blue. e–f Bitunicate asci. h–i Ascospores. g Pedicle. Scale bars: b, c = 100 μm. e = 50 μm. d, f–i = 20 μm

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Type species: Biatriospora marina K.D. Hyde & Borse, Mycotaxon 26: 264 (1986), MycoBank: MB 801045

Notes: The genus is monotypic.

Botryosphaeriaceae Theiss. & Syd. [as ‘Botryosphaeriacae’], Annls mycol. 16(1/2): 16 (1918), MycoBank MB 80530

Biotrophic, necrotrophic and saprobic especially on wood in terrestrial habitats. Sexual state: Ascostromata uni- to multi-locular, with multi-layered walls, single or in clusters, with or without basal stroma, fully or partially erumpent at maturity. Hamathecium of hypha-like, branched or unbranched, septate pseudoparaphyses, which are constricted at the septum, frequently disappearing at maturity. Asci mostly 8-spored, bitunicate, fissitunicate, with thick endotunica, clavate to cylindro-clavate, short or long-pedicellate, apically rounded with an ocular chamber. Ascospores 2–3-seriate, hyaline to brown, smooth to verruculose, thin-walled, aseptate to septate, fusoid to ellipsoid or ovoid, with or without a mucoid sheath or rarely with appendages. Asexual states: with uni- to multi-locular pycnidial conidiomata, frequently embedded in stromatic tissue, with hyaline, phialidic conidiogenous cells. Conidia hyaline to pigmented, thin- to thick-walled, sometimes with mucoid appendages or sheaths, smooth to verruculose, sometimes striate, with or without germ slits.

Notes: The family Botryosphaeriaceae was introduced by Theissen and Sydow (1918) in Botryosphaeriales (Schoch et al. 2006). Hibbett et al. (2007) did not assign the Botryosphaeriales to any subclass, but Schoch et al. (2009a) accepted its position in the Pleosporomycetidae. Previously the family had been referred to various orders: Myrangiales (Theissen 1916), Dothideales (Miller 1928; Kirk et al. 2001), and Pseudosphaeriales/Pleosporales (Luttrell 1955). The family is well-defined morphologically and supported by molecular data, with a number of lineages strongly corresponding to asexual morphological genera, e.g. Diplodia, Fusicoccum, and Lasiodiplodia (Schoch et al. 2009a; Liu et al. 2012a). Species are widespread in the tropics and temperate zones, some are forest pathogens, and some may be host-specific, while generally they are saprobic or endophytic. They also cause important diseases in forestry and agriculture (e.g. on grape-vines, van Niekerk et al. 2004). Liu et al. (2012a) accepted 29 genera in the Botryosphaeriales and molecular data showed the clade for the family Botryosphaeriaceae comprising eight sub-clades corresponding to: A1 Diplodia, Neodeightonia and Lasiodiplodia; A2 Phaeobotryosphaeria, Phaeobotryon and Barriopsis; A3 Dothiorella, Spencermartinsia and Auerswaldia; A4 Botryobambusa; A5 Neofusicoccum; A6 Botryosphaeria including the type species of the genus, Neoscytalidium and Cophinforma (a new genus); A7 Pseudofusicocoum; and A8 Aplosporella (Liu et al. 2012a).

Type: Botryosphaeria Botryosphaeria Ces. & De Not., Comm. Soc. crittog. Ital.1(4): 211 (1863), MycoBank MB 635 Fig. 19

Botryosphaeria dothidea (PREM 57372, epitype) a Ascostromata on host substrate. b Section through ascostroma. c Peridium. d-e Asci. f-h Ascospores. Scale bars: b–c = 100 μm, d–e = 30 μm, f–h = 10 μm

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Hemibiotrophic or saprobic on leaves and wood in terrestrial habitats. Sexual state: Ascostromata erumpent through the bark, often forminga botryose aggregate, sometimes solitary, globose, brown to black, individual locules with a central ostiole, papillate or not. Peridium two-layered, outer layer composed of dark brown or brown, thick-walled cells of textura angularis, inner layer composed of hyaline, thin-walled cells of textura angularis, lining the locule. Hamathecium comprising numerous, 2–4 μm wide, septate, hypha-like pseudoparaphyses, rarely branched at the apex. Asci 8–spored, bitunicate, fissitunicate, clavate, short-pedicellate, apically rounded with a small ocular chamber. Ascospores bi-seriate, hyaline, aseptate, fusoid to ovoid, sometimes with tapered ends, giving a spindle-shaped appearance, smooth with granular contents, with or without a mucilaginous sheath. Asexual states: Conidiomata pycnidial in nature. Conidiogenous cells holoblastic, hyaline, subcylindrical, proliferating percurrently with 1–2 proliferations and periclinical thickening. Conidia hyaline, aseptate, narrowly fusiform, or irregularly fusiform, base subtruncate to bluntly rounded, rarely forming a septum before germination, smooth with granular contents (asexual morph description follows Slippers et al. 2004).

Notes: The genus Botryosphaeria was introduced in 1863 and emended by Saccardo (1877) and is based on the type species B. dothidea (Moug.: Fr.) Ces. & de Not. However, as the type material was immature, Slippers et al. (2004) epitypified the type species Botryosphaeria dothidea based on morphology and phylogenetic data. Species of Botryosphaeriaceae occur on woody branches, herbaceous leaves, stems, and twigs of a wide range of plants (monocotyledons, dicotyledons and gymnosperms) and have a cosmopolitan distribution (Crous et al. 2006). Species are frequently associated with dieback and cankers of woody tissues and other plant diseases.

Botryosphaeria is characterized by ascostromata, bitunicate asci with a thick endotunica, stalked or sessile, clavate with a well-developed apical chamber, with hyaline pseudoparaphyses, constricted at the septa, and aseptate, hyaline ascospores, fusoid to ellipsoid or ovoid, generally lacking mucoid sheaths or appendages. Botryosphaeria asexual morphs have been assigned to some 18 coelomycete genera based on molecular data (Crous et al. 2006; Liu et al. 2012a; Wijayawardene et al. 2012).

Type species: Botryosphaeria dothidea (Moug: Fr.) Ces. & de Not., Comment Soc. crittog. Ital. 1: 212 (1863)

≡ Sphaeria dothidea Moug., in Fries, Syst. mycol. (Lundae) 2(2): 423 (1823)

Other genera included

Aplosporella Speg., Anal. Soc. cient. argent. 10(5–6): 157 (1880)

Type species: Aplosporella chlorostroma Speg., Anal. Soc. cient. argent. 10(5–6): 158 [no. 117, reprint page 35] (1880)

Auerswaldia Sacc., Syll. Fung. (Abellini) 2: 626 (1883)

Type species: Auerswaldia examinans (Mont. & Berk.) Sacc., Syll. Fung. 2: 266 (1883)

Auerswaldiella Theiss. & Syd., Ann. mycol. 12(3): 278 (1914)

Type species: Auerswaldiella puccinioides (Speg.) Theiss. & Syd., Ann. mycol. 12(3): 278 (1914)

Barriopsis A.J.L. Phillips et al., Persoonia 21: 39 (2008)

Type species: Barriopsis fusca (N.E. Stevens) A.J.L. Phillips et al., Persoonia 21: 39 (2008)

Botryobambusa R. Phookamsak et al., Fungal Divers. 57: 166 (2012)

Type species: Botryobambusa fusicoccum R. Phookamsak et al., Fungal Divers. 57: 166 (2012)

Cophinforma Doilom, J.K. Liu & K.D. Hyde, in Liu et al. Fungal Divers. 57: 174 (2012)

Type species: Cophinforma eucalyptus Doilom, J.K. Liu & K.D. Hyde, in Liu et al. Fungal Divers. 57: 174 (2012)

Dichomera Cooke, Nuovo G. bot. ital. 10: 24 (1878)

Type species: Dichomera saubinetii (Mont.) Cooke, Nuovo G. bot. ital. 10: 24 (1878)

Diplodia Fr., in Montagne, Annls Sci. Nat., Bot., sér. 2 1: 302 (1834)

Type species: Diplodia mutila (Fr.) Mon., Annls Sci. Nat., Bot., sér. 2 1: 302 (1834)

Dothiorella Sacc., Michelia 2 (no. 6): 5 (1880)

Type species: Dothiorella pyrenophora Berk. ex Sacc., Michelia 2 (no. 6): 5 (1880)

Endomelanconiopsis E.I. Rojas & Samuels, in Rojas et al., Mycologia 100 (5): 770 (2008)

Type species: Endomelanconiopsis endophytica E.I. Rojas & Samuels, in Rojas et al., Mycologia 100 (5): 770 (2008)

Lasiodiplodia Ellis & Everh., Bot. Gaz. 21: 92 (1896)

Type species: Lasiodiplodia theobromae (Pat.) Griffon & Maubl., Bull. Soc. mycol. Fr. 25: 57(1909)

Macrophomina Petr., Annls. mycol. 21(3/4): 314 (1923)

Type species: Macrophomina phaselina (Tassi) Gorf., Annali Sper. agr.n.s. 1: 457 (1947)

Microdiplodia Allesch., Rabenh. Krypt.-Fl., Edn 2 (Leipzig) 1(7): 78 (1901) [1903]

Type species: Microdiplodia conigena Allesch., Rabenh. Krypt.-Fl., Edn 2 (Leipzig) 1(7): 79 (1901) [1903]

Macrovalsaria Petr., Sydowia 15: 298 (1962) [1961]

Type species: Macrovalsaria leonensis (Deighton) Petr., Sydowia 15(1–6): 298 (1962) [1961]

Neodeightonia C. Booth, in Punithalingam, Mycol. Pap. 119: 17 (1970) [1969], MycoBank MB 3450 Figs. 20 and 21

Neodeightonia subglobosa (Material examined: MFLU 11–0199); THAILAND, Lampang Province., Jae Hom District, Mae Yuag Forestry Plantation, on dead culms of Bambusa sp., 19 August 2010, R. Phookamsak, RP0079, MFLU 11–0199)). a Ascomata on host tissue. b Section through uni-loculate ascoma. c Section through peridium. d Pseudoparaphyses. e, f Asci g Asci with pseudoparaphyses. h-k Ascospores with a prominent sheath. Scale bars: a = 500 μm, b = 200 μm, c, d, g = 50 μm, e, f = 20 μm, h–k = 10 μm

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Neodeightonia palmicola - asexual state (Material examined: THAILAND, Chiang Rai Province., Muang District, Khun Korn Waterfall, on dead leaves of Arenga westerhoutii, 18 Dec 2009, J.K. Liu, JKA0022, MFLU 10–0407, holotype); Chiang Rai Prov., Muang District, Khun Korn Waterfall, on living leaves of Caryota urens., 22 Jul 2009, R. Phookamsak, RP0004 (MFLU 100409 a, b Conidiomata on pine needles. c Section of conidioma. d, e Conidiogenous cells. f, g Young conidia. h, i Mature conidia with septa. Scale bars: a, b = 500 μm, c = 100 μm, d, e = 30 μm, g–j = 10 μm

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Saprobic on dead wood and leaves of monocotyledons. Sexual state: Ascostromata brown to dark brown, uni-loculate, immersed to erumpent, globose to subglobose. Ostiole circular, central. Peridium of dark, brown walled cells of textura angularis. Hamathecium of hypha-like, branched or unbranched, septate pseudoparaphyses constricted at the septa. Asci 8-spored, bitunicate, fissitunicate, clavate to cylindrical-clavate, apically rounded with an ocular chamber. Ascospores uni-seriate or irregularly bi-seriate, hyaline, aseptate, ellipsoidal-fusiform, or fusiform, surrounded or not surrounded by a complex sheath. Asexual state: Pycnidia uni-loculate or multi-locular, semi-immersed, solitary, globose, covered by mycelium, wall composed of dark brown thick-walled textura angularis, becoming thin-walled and hyaline towards the inner region. Paraphyses hyaline, cylindrical. Conidiogenous cells holoblastic, hyaline, aseptate, cylindrical to subcylindrical. Conidia initially hyaline, aseptate, ellipsoid to obovoid, thick-walled with granular content, rounded at apex, occasionally truncate at base. Aged conidia becoming cinnamon to sepia, and 1-septate, brown to dark brown.

Notes: Neodeightonia was introduced by Booth (in Punithalingam 1969). However, von Arx and Müller (1975) transferred the type of the genus, N. subglobosa C. Booth,, to Botryosphaeria, reducing Neodeightonia to synonymy. Phillips et al. (2008) reinstated this genus which is distinguishable from Botryosphaeria morphologically (based on the dark, 1-septate conidiospores) and phylogenetically (Phillips et al. 2008; Abdollahzadeh et al. 2009) and described a new species N. phoenicum A.J.L. Phillips & Crous. Liu et al. (2010) added the fourth species; N. palmicola based on morphological studies of the sexual and asexual morphs and phylogenetic data, which is illustrated here (Fig. 3).

Type species: Neodeightonia subglobosa C. Booth, in Punithalingam, Mycol. Pap. 119: 19 (1970) [1969]

Neofusicoccum Crous et al., Stud. Mycol. 55: 247 (2006)

Type species: Neofusicoccum parvum (Pennycook & Samuels) Crous et al., Stud. Mycol. 55: 248 (2006)

Neoscytalidium Crous & Slippers, Stud. Mycol. 55: 244 (2006)

Type species: Neoscytalidium dimidiatum Crous & Slippers, Stud. Mycol. 55: 244 (2006)

Phaeobotryon Theiss. & Syd., Ann. Mycol. 13 (5/6): 664 (1915)

Type species: Phaeobotryon cercidis (Cooke) Theiss. & Syd., Annls. mycol. 13: 664. 1915

Phaeobotryosphaeria Speg., Anal. Mus. nac. B. Aires, Ser. 3 17(10): 120 (1908)

Type species: Phaeobotryosphaeria yerbae Speg., Anal. Mus. nac. B. Aires, Ser. 317(10): 120(1908)

Phyllachorella Syd., Annls. mycol. 12: 489 (1914)

Type species: Phyllachorella micheliae Syd., Annls. mycol. 12(5): 489 (1914)

Pseudofusicoccum Mohali et al., Stud. Mycol. 55: 249 (2006)

Type species: Pseudofusicoccum stromaticum (Mohali et al.) Mohali et al., Stud. Mycol.55: 249 (2006)

Pyrenostigma. Syd., Annls. mycol. 24: 370 (1926)

Type species: Pyrenostigma siparunae Syd., Annls. mycol.24: 370 (1926)

Sivanesania W.H. Hsieh & Chi Y. Chen, Mycol. Res. 98 (1): 44 (1994)

Type species: Sivanesania rubi W.H. Hsieh & Chi Y. Chen, Mycol. Res. 100(9): 1106 (1996)

Spencermartinsia A.J.L. Phillips et al., Persoonia 21: 51 (2008)

Type species: Spencermartinsia viticola (A.J.L. Phillips & J. Luque) A.J.L. Phillips et al., Persoonia 21: 51 (2008)

Tiarosporella Höhn., in Weese, Mitt. bot. Inst. tech. Hochsch. Wien 1 (3): 82 (1924)

Type species: Tiarosporella paludosa (Sacc. & Fiori ex P. Syd.) Höhn., in Weese, Mitt. bot. Inst. tech. Hochsch. Wien 1 (3): 82 (1924)

Thyrostroma Höhn., Sber. Akad. Wiss. Wien, Math.-naturw.Kl., Abt. 1 120: 472 [94 repr.] (1911)

Type species: Thyrostroma compactum (Sacc.) Höhn., Sber. Akad. Wiss. Wien, Math.-naturw. Kl., Abt. 1 120: 472 [94 repr.] (1911)

Vestergrenia Rehm, Hedwigia 40: 100 (1901)

Type species: Vestergrenia nervisequia Rehm, Hedwigia 40: 101 (1901)

Key to sexual genera of Botryosphaeriaceae

1. Ascostromata unilocular .....................................................2

1. Ascostromata multi-locular ................................................4

2. Ascostromata superficial ....................................................3

2. Ascostromata immersed to erumpent ...........Cophinforma

3. Asci cylindrical to clavate with a small ocular chamber ..........................................................................Phyllachorella

3. Asci clavate to ovoid with a long pedicel, apically rounded with an ocular chamber .......................................Vestergrenia

4. Ascostromata superficial, immersed with a foot-like hypostroma ............................................................................5

4. Ascostromata immersed, covered by cuticle or epidermis ................................................................................................7

5. Asci cylindro-clavate, with a short pedicel, apically rounded with a small ocular chamber .............................................6

5. Asci broadly clavate with short narrow furcate pedicel and large ocular chamber .........................................Pyrenostigme

6. Ascostromata superficial on upper surface of the leaf, four to numerous locules ............................................Auerswaldia

6. Ascostromata superficial on lower surface of the leaf, subglobose to globose .....................................Auerswaldiella

7. Ascostromata globose to ovoid .........................................8

7. Ascostromata flask shaped, ascospores with hyaline apiculus .................................................Phaeobotryosphaeria

8. Ascospores septate .............................................................9

8. Ascospores aseptate .........................................................11

9. Ascospores 1-septate .......................................................10

9. Ascospores 2-septate ....................................Phaeobotryon

10. Ascospores uni-seriate, elliptical fusoid, slightly constricted at septum, skull cap-like germ apparatus at the lower end .........................................................Macrovalsaria

10. Ascospores uniseptate with an apiculus at each end, hyaline to brown ...............................................Spencermartinsia

11. Ascospores brown when mature ....................................12

11. Ascospores hyaline when mature ...................................13

12. Ascostromata immersed, discoid to pulvinate or hemisphaerical ........................................................Barriopsis

12. Ascostromata immersed to erumpent, subglobose to ovoid, ascospores filiform, with simple appendage................. ..............................................................................Sivanesania

13. Ascostromata erumpent, subglobose................................... ........................................................................Botryobambusa

13. Ascostromata immersed, globose ..................................14

14. Ascospores spindle shape .........................Botryosphaeria

14. Ascospores ellipsoidal to fusiform .................................15

15. Ascospores with prominent sheath ............Neodeightonia

15. Ascospores without mucilaginous sheath....................... .........................................................................Neofusicoccum

Key to asexual genera of Botryosphaeriaceae

1. Conidiomata superficial .....................................................2

1. Conidiomata immersed ......................................................5

2. Conidiophores reduced to conidiogenous cells, holoblastic, cylindrical ..............................................................................3

2. Conidia occurring in arthric chains in aerial mycelium .........................................................................Neoscytalidium

3. Paraphyses hyaline, thin walled, usually aseptate .................................................................................Barriopsis

3. Paraphyses absent ..............................................................4

4. Conidia hyaline, fusiform to ellipsoidal.......................... ........................................................................Botryobambusa

4. Conidia brown, broadly ellipsoidal to subcylindrical ................................................................Endomelanconiopsis

5. Conidiomata uni-locular .....................................................8

5. Conidiomata multi-locular .................................................6

6. Paraphyses septate, Conidiogenous cells holoblastic, smooth walled .....................................................Aplosporella

6. Paraphyses aseptate ............................................................7

7. Conidia aseptate............................................Neodeightonia

7. Conidia septate ..............................................Phaeobotryon

8. Conidia septate .................................................................11

8. Conidia aseptate ...............................................................13

11. Conidia brown when attached to conidiophores ...........12

11. Conidia hyaline when attached to conidiophores, becoming brown when mature .............................................Diplodia

12. Mycelium immersed or superficial, branched, septate, melanised ..............................................................Dothiorella

12. Conidiogenous cells lining inner surface of conidiomata .....................................................................Spencermartinsia

13. Matured conidia hyaline ...........................Botryosphaeria

13. Matured conidia brown ..................................................14

14. Conidia with undulating striations on the surface .........15

14. Conidia without undulating striations on the surface ..............................................................................................16

15. Conidia with granular content .....................Lasiodiplodia

15. Conidia without granular content .................Auerswaldia

16. Conidia enclosed in a mucous sheath.............................. ........................................................................Macrophomina

16. Conidia not enclosed in a mucous sheath .....................17

17. Conidia apex round, base flat ..................Neofusicoccum

17. Conidia clavate, straight, thick-walled .......................................................................................Phaeobotryosphaeria

Brefeldiellaceae E. Müll. & Arx, in Müller & von Arx, Beitr. Kryptfl. Schweiz 11(no. 2): 148 (1962), MycoBank: MB 80534

Foliar epiphytes or parasites on leaves of various hosts worldwide. Sexual state: Thallus relatively large, circular or irregular, or spreading as root-like structures across the leaf surface, brown to black, extremely thin, covering thyriothecia, and covering host tissue with a skin-like layer; cells of thallus consisting of a single layer of somewhat radially arranged cylindrical to cuboid cells, each cell distinct. Thyriothecia circular in outline, flattened, thin-walled, developing under thallus, with numerous asci. Ostioles central. Hamathecium comprising asci, pseudoparaphyses not observed. Asci 8-spored, bitunicate, fissitunicate, saccate, short-pedicellate or sessile, ocular chamber not obvious. Ascospores 2–3-seriate, oblong-ellipsoidal or clavate, hyaline, 1-septate, trans-septate or muriform, smooth-walled. Asexual state: see in notes.

Notes: According to Cooke and Hawksworth (1970) the name Brefeldiellaceae is illegitimate. Eriksson (1981) however, mentioned that Müller and von Arx (1962) referred to a legitimate description by Theissen (of subfam. Brefeldiineae) and suggested the name is therefore legitimate. Eriksson (1981) also mentioned the status of the family is uncertain. In this study we accept Brefeldiellaceae and add several genera. The characters of Brefeldiellaceae are rather confusing and fresh material and molecular data is needed to establish the taxonomic placement of this family and the various genera that we accommodate in this family. The spreading thallus which covers ascomata with a parallel arrangement of cells is typical of the arrangement of cells in the thyriothecia found in Asterinaceae and Microthyriaceae. The asci are however, different to those in Asterinaceae, saccate to cylindro-clavate in Brefeldiellaceae and globose in Asterinaceae. The Brefeldiellaceae may therefore be a sister family to Microthyriaceae. In this monograph we include Acrogenotheca, Saccardinula, Trichopeltina and Trichopeltum in the family mainly because of thallus characteristics and saccate to cylindro-clavate asci. Acrogenotheca (type A. pulcherrima Bat. & Cif.) was described from undetermined leaves collected in Cuba. Unfortunately more than one taxon is represented on the type material. Saccardinula (type S. guaranitica Speg.) is a less convincing member of the family as it has globose asci and muriform spores, while the thallus is less developed. Trichopeltella (type T. montana (Racib.) Höhn.) was described on leaves from Java, Indonesia and although we were only able to find asexual structures, the thallus is typical of Brefeldiella. The conidia are produced on small conidiogenous cells that line the conidiomata wall and conidia are cylindrical, hyaline and unicellular. Trichopeltella may be a synonym of Brefeldiella. Trichopeltum (type T. hawaiiense Bat. & C.A.A. Costa) was described on leaves of Smilax from Hawaii. This species has a spreading root-like thallus which superficially look similar to those in Teratosphaeriaceae but are made up of cells typical of Brefeldiellaceae.

The arrangement of Brefeldiellaceae as treated here is certainly polyphyletic. Reynolds and Gilbert (2005) also treated this family with similar wide ranging ascal and thallus characters and the genera certainly need recollecting and sequencing to establish a better phylogeny.

Type: Brefeldiella Speg., Boln Acad. nac. Cienc. Córdoba 11(4): 558 (1889), MycoBank: MB 648 Figs. 22, 23 and 24

Brefeldiella brasiliensis (Material examined: BRAZIL, Apiahy, on leaves of Bambusaceae , J. Puiggari, no 2356, 1888, holotype). a, b Appearance of thallus on substrate c Squash of thallus. d Arrangment of cells of thallus. Scale bars: c–d = 100  μm

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Brefeldiella brasiliensis ascospores, asci and structure of thallus (redrawn from Spegazzini’s drawing on the herbarium packet ( http://www.cybertruffle.org.uk/spegazzini/eng/001400a_.htm )

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Trichopeltina labecula (Material examined: FRENCH GUYANA, Gabaret River, on coriaceous leaf, September 1835, Coll. no. 664, PC0084645, syntype). a Herbarium specimens. b Thallus on host substrate. c Thyriothecia within thin superficial thallus in 70 % lactic acid. d Cells of thallus and thyriothecium ostiole. e Cells of thallus. f Hamathecium in Melzer’s reagent. g Ascus when immature in Melzer’s reagent. h Ascus in Melzer’s reagent. i Ascus in lactophenol cotton blue reagent. j Ascospore in 70 % lactic acid. k Ascospore in Melzer’s reagent. l Ascospore in Lactophenol cotton blue reagent. Scale bars: c = 100 μm. e–i = 20 μm. j–l = 10 μm

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Foliar epiphytes or parasites on leaves of various hosts. Sexual state: Thallus relatively large, circular or irregular, or spreading as root-like structures across leaf surface, brown to black, extremely thin, covering some ascomata, and covering host tissue with a skin-like layer; cells of thallus consisting of a single layer of somewhat radially arranged cylindrical to cuboid cells, each cell distinct. Thyriothecia circular in outline, flattened, thin-walled, developing under thallus, with numerous asci. Ostioles central. Hamathecium comprising of asci, pseudoparaphyses not observed. Asci 8-spored, bitunicate, fissitunicate, cylindric-clavate, short-pedicellate, apically rounded with small ocular chamber. Ascospores 2–3-seriate, hyaline, oblong-ellipsoidal, 1-septate, upper cell wider, shorter, and more rounded than thinner and longer lower cell, thin-walled, smooth-walled. Asexual state: Unknown.

Notes: The genus comprises four species (Index Fungorum 2013). Eriksson (1981) examined the type collection of B. brasiliensis Speg. (LPS) which contained numerous ascomata, but which are immature. This genus may have been described on more than one occasion and needs recollecting.

Type species: Brefeldiella brasiliensis Speg., Boln Acad. nac. Cienc. Córdoba 11(4): 558 [no. 344] (1889), MycoBank: MB 230084

Other genera included

Acrogenotheca Cif. & Bat., in Batista & Ciferri, Saccardoa 2: 51 (1963)

Type species: Acrogenotheca pulcherrima Bat. & Cif., Saccardoa 2: 52 (1963)

Pycnoderma Syd. & P. Syd., Annls. mycol 12(6): 563 (1914)

Type species: Pycnoderma bambusinum Syd. & P. Syd. Annls. mycol 12(6): 563 (1914)

Saccardinula Speg., Anal. Soc. cient. argent. 19(6): 257 (1885)

Type species: Saccardinula guaranitica Speg., Anal. Soc. cient. argent. 19(6): 258 (1885)

Trichopeltella Höhn., Sber. Akad. Wiss. Wien, Math.-naturw. Kl., Abt. 1 119: 458 [66 repr.] (1910)

Type species: Trichopeltella montana (Racib.) Höhn., Sber. Akad. Wiss. Wien, Math.-naturw. Kl., Abt. 1 119: 458 [66 repr.] (1910)

Trichopeltina Theiss., Beih. bot. Zbl., Abt. 2 32: 3 (1914)

Type species: Trichopeltina labecula (Mont.) Theiss., Zentbl. Bakt. ParasitKde, Abt. II 39: 630 (1914)

Trichopeltum Bat. et al., in Batista et al., Publções Inst. Micol. Recife 90: 20 (1957)

Type species: Trichopeltum hawaiiense Bat. & C.A.A. Costa [as ‘hawaiiensis’], in Batista et al., Publções Inst. Micol. Recife 90: 21 (1957)

Key to genera of Brefeldiellaceae

1. Thallus appearing as root-like structures growing on the host surface ........................................................Trichopeltum

1. Thallus spreading as a regular outgrowths, often rounded or sheet-like on host surface .......................................................2

2. Ascospores muriform .........................................................3

2. Ascospores with transverse septa only ..............................4

3. Ascospores with slightly constricted at septate, pale brown, gelatinous sheath .................................................Pycnoderma

3. Ascospores with hardly constricted at septate, hyaline .............................................................................Saccardinula

4. Ascospores 2-celled ............................................................5

4. Ascospores 3-celled ....................................Acrogenotheca

5. Ascospores pale brown ..................................Trichopeltina

5. Ascospores hyaline ...........................................Brefeldiella

Capnodiaceae Höhn. ex Theiss., Verh. zool.-bot. Ges. Wien 66: 363 (1916), MycoBank: MB 82008

Saprobic on honey dew excretions from insects, usually forming blackened sooty-like regions on green healthy leaves, stems, bark and even rocks and often in association with numerous other species and fungicolous taxa. Thallus comprising mycelium with black sooty growth on the host surface, producing superficial colonies with septate, cylindrical, brown hyphae. Sexual state: Ascomata arise from the mycelial mass, subglobose to globose, coriaceous, with or without setae, dark brown, with a central ostiole. Peridium brown, relatively thin, comprising cells of textura angularis. Hamathecium lacking pseudoparaphyses. Asci 8-spored, bitunicate, saccate, with a short pedicel, lacking an ocular chamber. Ascospores overlapping bi-seriate, multi-septate or muriform, hyaline to brown. Asexual states: pycnidial, mostly elongated, with short or long narrow necks, and with or without a swelling where the conidia are produced. Ostiole prominent at the apex of the pycnidial neck. Conidia occur on typical capnodiaceous hyphae within the pycnidia, small, one-celled, ellipsoidal, hyaline.

Notes: The family Capnodiaceae was introduced by von Höhnel (1909a) and validated by Theissen (1916) according to von Arx and Müller (1975) and Hughes (1976), and is circumscribed as dothideomycetous sooty moulds with mostly pycnidial asexual states (Reynolds 1982). Kirk et al. (2008) stated the Capnodiaceae include 26 genera and 117 species, while Lumbsch and Huhndorf (2010) mentioned 13 genera. Chomnunti et al. (2011) revisited the Capnodiaceae and removed many genera to other families based on morphological characters of type specimens, thus only five genera are now accepted in Capnodiaceae.

The taxa of this family can be recognized by black mycelial growth spreading on the host surface which produces superficial colonies with septate, cylindrical, dark-brown hyphae. Ascomata arise from the mycelial mass and are subglobose to globose (Sivanesan 1984; Barr 1987a). Their asexual states form elongated pycnidia that develop from a superficial mycelium on living plant surfaces. Pycnidia have short or long narrow necks with a conspicuous oval swelling which produce minute, unicellular and hyaline conidia from near the base, middle or apex (Chomnunti et al. 2011).

In the past some members of the family had been known only from asexual states, and these pycnidia had been assigned to one or other sexual state genera and families depending on whether the researchers gave more emphasis to Saccardoan generic concepts such as spore morphology and colour, or colony hyphal development, hyphal morphology and ascomatal and pycnidial formation (as with Hughes 1976). The use of molecular techniques has now enabled us to confirm their relationships in the Capnodiaceae. Crous et al. (2009b) used molecular methods to evaluate the monophyly of the order Capnodiales, yet they did not discuss the family in detail, although concluding that it probably contained diverse lineages. Chomnunti et al. (2011) used a RAxML maximum likelihood tree based on combined LSU and SSU genes to show that Capnodium, Leptoxyphium, Phragmocapnias and Scorias are resolved genera in Capnodiaceae.

Type: Capnodium Mont. Annls Sci. Nat., Bot., sér. 311: 233 (1849), MycoBank: MB 809 Figs. 25 and 26

Capnodium tiliae (Material examined: GERMANY, Biebrich, on branch of Tilia parviflora, Herbier Fuckel no. 899 1894, G, Herbier Barbey Boissier, exsiccata of Fumago tiliae). a Label data and specimen. b Vertical section through ascoma. c Dark brown mycelium. d Peridium. e, f. Peridium with cells of textura angularis. g–i Cylindrical to cylindric-clavate asci. j. Ascospores 3−4-septate. Scale bars: f = 200 μm, b–d = 100 μm, e, g–i = 20 μm, j = 10 μm

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Capnodium coartatum (Material examined: THAILAND, Chiang Rai, on living leaf of Bischofia javanica, 27 April 2011, Putarak Chomnunti 001, MFLU11−1155, holotype) a–c Appearance of sooty moulds on host. d–g Elongate pycnidia. e Globose, septate mycelium. h Conical pycnidium. i Ostiole surroundedby hyaline hyphae. j Conidia. Scale bars: d, e, g = 50 μm, f, h = 20 μm, i, j = 10 μm

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Saprobic on sugary exudates from insects growing on the surface of leaves, fruits, stems and other non plant objects. Thallus a loose or dense network of pale brown, superficial hyphae or a thick pseudoparenchymatous stroma, with sexual and asexual states often growing together. Sexual state: Ascomata superficial on mycelium of thallus, brown to dark brown or black, globose to ellipsoidal, short-stalked or sessile, ostiolate at maturity, scattered or in groups, without setae. Peridium comprising dark brown to pale brown, thick-walled cells forming a textura angularis. Hamathecium lacking pseudoparaphyses. Asci 8-spored, bitunicate, clavate, ovoid or saccate, apedicellate, lacking an ocular chamber. Ascospores overlapping bi-seriate, brown, oblong or ovoid and some reniform, transversely septate with one or more vertical septa or without vertical septa. Asexual state: “Polychaeton” spp. Pycnidia elongate, often with long stalks, dark brown. Ostiole at apex of pycnidia with hyphae continued upwards to the tapered neck, terminating in an ostiole which is surrounded by dull hyphal round ends. Conidia hyaline, 1-celled.

Notes: Chomnunti et al. (2011) revisited the sooty mould genera in Capnodiaceae based on examination of types and a RAxML maximum likelihood tree based on combined LSU and SSU genes, using 15 specimens of sooty moulds from Thailand; Phragmocapnias and Scorias were also epitypified (Chomnunti et al. 2011). In addition, it was suggested that Capnodium should be used rather than Polychaeton, because it is better known, has widespread use and is the sexual state and type of the family and order. Hawksworth (2012) gave Capnodium as the example for widely used names of fungi that could be proposed for use, against the older lesser used asexual name (Polychaeton). It is expected that Capnodium will be defined for future use in the lists of accepted names to be approved by the General Committee under Art. 14.13. The sexual state of Capnodium is illustrated here based on Capnodium tiliae Berk. & Desm., and the asexual state is illustrated based on a recent collection of Capnodium coartatum Chomnunti & K.D. Hyde,. Acanthorus Bat. & Cavalc. was reported in Abolisiaceae which is pycnidial (Batista and Cavalcante 1967) and an invalid family. Kirk et al. (2008) mention that Acanthorus is the asexual state of Capnodiaceae, but this has not been followed by other researchers.

Type species: Capnodium citri Berk. & Desm., in Berkeley, Journal of the Royal Horticultural Society 4: 11 (1849), MycoBank: MB 164508

Other genera included

Leptoxyphium Speg., Physis, B. Aires 4: 294 (1918)

Type species: Leptoxyphium graminum (Pat) Speg., Physis. B. Aires 4: 294 (1918)

Phragmocapnias Theiss. & Syd., Annls mycol. 15(6): 480 (1918) [1917]

Type species: Phragmocapnias betle (Syd., P. Syd & E.J. Butler) Theiss. & Syd., Annls mycol. 15(6): 480 (1918) [1917]

Scorias Fr. Syst. mycol. 3(2): 269, 290 (1832)

Type species: Scorias spongiosa (Schwein.) Fr., Syst. mycol. 3(2): 291 (1832)

Scoriadopsis J.M. Mend., in Stevens, Annls mycol. 28 (5/6): 365 (1930)

Type species: Scoriadopsis miconiae J.M. Mend., in Stevens, Annls mycol. 28 (5/6): 365 (1930)

Key to genera of Capnodiaceae

1. Pycnidia with a black stalk, and an upper brown swollen region producing conidia; ascospores hyaline ................................................................Phragmocapnias

1. Pycnidia without a black stalk or an upper brown swollen region producing conidia; ascospores brown or hyaline ................................................................................................2

2. Pycnidia lacking a basal bulbous part; ascospores brown ...............................................................................Capnodium

2. Pycnidia with basal bulbous part; ascospores other than above ......................................................................................3

3. Ascomycetous state unknown, pycnidia with narrow bulbous base, apex cup-like which produces conidia ..........................................................................Leptoxyphium

3. Pycnidia with swollen bulbous base; ascomata present, gregarious on blacked mycelial mat ......................................4

4. Ascospores mostly 3-septate ...................................Scorias

4. Ascospores mostly 1-septate ...........................Scoriadopsis

Cladosporiaceae Nann., Repert. mic. uomo: 404 (1934), MycoBank: MB 80600

Synonym:

Davidiellaceae C.L. Schoch, Spatafora, Crous & Shoemaker, in Schoch, Shoemaker, Seifert, Hambleton, Spatafora & Crous, Mycologia 98(6): 1048 (2007) [2006], MycoBank: MB 504453

Saprobic or phytopathogenic, also endophytic and involved in diseases of humans and animals. Sexual state: Ascomata pseudothecial, gregarious or scattered, inconspicuous, black to red-brown, globose to subglobose, uni-loculate, inconspicuous and immersed in the substratum to superficial, in plant pathogenic species beneath stromata, situated on a reduced stroma, with 1(−3) short, periphysate ostiolar necks; periphysoids frequently growing down into cavity. Ostiole necks, with numerous periphysoids. Peridium consisting of 3–6 layers of dark brown cells, usually textura angularis. Hamathecium of hyaline, septate, subcylindrical pseudoparaphyses, present in mature ascomata. Asci 8-spored, bitunicate, fissituicate, sessile to short-stalked, obovoid to broadly ellipsoid or subcylindrical, straight to slightly curved. Ascospores bi- to multi-seriate, or overlapping, hyaline, obovoid to ellipsoid-fusiform, with irregular luminar inclusions, mostly thick-walled, straight to slightly curved; with age frequently becoming brown and verruculose in asci, 1-septate, not or somewhat constricted at the septum, at times covered by a mucilaginous sheath. Asexual state: hyphomycetous. Conidiophores macronematous, mononematous, simple or branched, brown. Conidiogenous cells integrated, terminal and intercalary, sympodial or synchronous, holoblastic, mostly polyblastic, conidiogenous loci conspicuous, mostly protuberant, darkened-refractive, coronate, i.e. composed of a convex central dome (which is the slightly bulging half of the original septum, delimiting the conidium and the conidiogenous cell, after being cleft) and a raised periclinal rim. Conidia solitary or mostly in branched or unbranched acropetal chains, amero-, didymo- to phragmosporous, subhyaline to brown, smooth to verrucose or echinulate, sometimes forming ramoconidia, dry, conidium secession schizolytic (Schoch et al. 2006; Seifert et al. 2011; Bensch et al. 2012).

Notes: Schoch et al. (2006) introduced the new family Davidiellaceae in Capnodiales to accommodate Davidiella with its Cladosporium asexual morphs (type species Davidiella tassiana (De Not.) Crous & U. Braun, asexual state Cladosporium herbarum (Pers.: Fr.) Link,). Previously, Ellis (1971, 1976) and Sivanesan (1984) had reported Cladosporium as the asexual states of Mycosphaerella and placed the genus in Mycosphaerellaceae. Braun et al. (2003) clearly showed that Mycosphaerella sensu stricto and Cladosporium herbarum, the type of Cladosporium, belong to different phylogenetic lineages. At the same time they showed several “Mycosphaerella”-like sexual states grouped with C. herbarum that possessed ascomata which were very similar to Mycosphaerella sect. Tassiana. Hence they introduced the new genus Davidiella for “Mycosphaerella”-like sexual morphs connected to Cladosporium asexual morphs. This was confirmed by Schoch et al. (2006) who introduced the family Davidiellaceae. Previous concepts of Cladosporium had been very broad, covering a wide range of dematiaceous hyphomycetes with conidia formed in acropetal chains (Dugan et al. 2004). Braun et al. (2003) and Crous et al. (2007b) treated Cladosporium sensu stricto along with Davidiella, and discussed the circumscription of this genus and its delimitation from morphologically similar, often confused genera. Crous et al. (2007b) provided a key to cladosporioid genera, which is in slightly modified form also included in Seifert et al. (2011). Numerous “Cladosporium”-like species, previously referred to as Cladosporium sensu lato but not congeneric, belong elsewhere and have been reallocated to different genera. For example, C. malorum Rühle (Pleosporales) was first moved to Alternaria (Braun et al. 2003) and later to Chalastospora (Crous et al. 2009b). Other cladosporoid fungi with unthickened, non-coronate conidiogenous loci, belonging to the Venturiaceae, represent Fusicladium asexual states of Venturia (Schubert et al. 2003; Crous et al. 2007b). Species of the genera Graphiopsis, Rachicladosporium, Toxicocladosporium and Verrucocladosporium are morphologically cladosporioid, except for lacking coronate conidiogenous loci, but phylogenetically distinct. They cluster in adjacent position but do not belong to the Cladosporiaceae clade sensu stricto (Crous et al. 2007b: 36, Fig. 1). Based on the new ICN, Hawksworth (2012) proposed to use Cladosporium as holomorph genus as it is the oldest and most widely used name. Bensch et al. (2012), who published a comprehensive monograph of Cladosporium, also used the latter name in this sense and reintroduced the old family name Cladosporiaceae instead of Davidiellaceae. We also suggest using Cladosporiaceae hereafter, as it is older than Davidiellaceae and reflects the current use of Cladosporium as holomorph name. This family is homotypic, i.e. it comprises only the genus Cladosporium, including holomorphs with asexual hyphomycetous and their sexual “Davidiella” states and “Davidiella” teleo-holomorphs, without any asexual states and asexual states which may be ana-holomorphs.

Type: Cladosporium Link, Mag. Gesell. naturf. Freunde, Berlin7: 37. 1816 [1815], MycoBank: MB 7681

Figure 27

Cladosporium herbarum (CPC 11600), asexual and sexual morphs (from the host and CPC 12181). a, b Macronematous conidiophores. c Micronematous conidiophores. d Microcyclic conidiogenesis. e Conidial chain. f Ascomata on the leaf. g Ascomata formed in culture on nettle stems. h–i Asci on the host. j–k Ascospores in culture. l Asci in culture. Scale bars: a, e, h, j–l = 10 μm f–g, l = 200 μm

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Acrosporella Riedl & Ershad, Sydowia 29(1–6): 166 (1977) [1976]Davidiella Crous & U. Braun, in Braun et al., Mycol. Progr. 2(1): 8 (2003)Notes: The genus Cladosporium was introduced by Link (1816) [1815] and it is considered as one of the largest and most heterogeneous genera of hyphomycetes (Dugan et al. 2004), currently comprising 773 epithets (Index Fungorum 2013). Bensch et al. (2012) however, included 139 names of ‘‘Heterosporium’’ (reduced ‘‘Heterosporium’’ under Cladosporium) in their monograph of Cladosporium sensu lato. Hence the total number of epithets in Cladosporium sensu lato is 993 (854 in Cladosporium and 139 in ‘‘Heterosporium’’) (Bensch et al. 2012). Cladosporium sensu stricto (asexual state of ‘Davidiella’) is characterised by having coronate conidiogenous loci and conidial hila, i.e., with a convex central dome surrounded by a raised periclinal rim (Bensch et al. 2012).

Type species: Cladosporium herbarum (Pers.) Link, in Willdenow, Willd., Sp. pl., Edn 4 6(1): 556 (1816), MycoBank: MB 231458

Other genera included

Acroconidiella J.C. Lindq. & Alippi, Darwiniana 13(2–4): 612 (1964)

Type species: Acroconidiella tropaeoli (T.E.T. Bond) J.C. Lindq. & Alippi, Darwiniana 13(2–4): 613 (1964)

Graphiopsis Trail, Scott., N.S. 4 (‘10’): 75 (1889)

Type species: Phaeoisaria clematidis (Fuckel) S. Hughes, Can. J. Bot.36: 794 (1958)

Hoornsmania Crous, in Crous et al., Fungal Planet 11: [2] (2007)

Type species: Hoornsmania pyrina Crous, Fungal Planet 11–21: 11: [2] (2007)

Rachicladosporium Crous, U. Braun & C.F. Hill, in Crous et al., Stud. Mycol. 58: 38 (2007)

Type species: Rachicladosporium luculiae Crous, U. Braun & C.F. Hill, in Crous, Braun, Schubert & Groenewald, Stud. Mycol. 58: 39 (2007)

Toxicocladosporium Crous & U. Braun, in Crous, Braun, Schubert & Groenewald, Stud. Mycol. 58: 39 (2007)

Type species: Toxicocladosporium irritans Crous & U. Braun, in Crous, Braun, Schubert & Groenewald, Stud. Mycol.58: 39 (2007)

Verrucocladosporium K. Schub., Aptroot & Crous, in Crous, Braun, Schubert & Groenewald, Stud. Mycol. 58: 41 (2007)

Type species: Verrucocladosporium dirinae K. Schub., Aptroot & Crous, in Crous, Braun, Schubert & Groenewald, Stud. Mycol. 58: 41 (2007)

Key to the asexual genera of Cladosporiaceae

1. Setae unbranched, brown .............................Acroconidiella

1. Setae absent ........................................................................2

2. Conidiophores compact, dense ........................Graphiopsis

2. Conidiophores solitary ........................................................3

3. Conidiogenous cell cylindrical .....................Cladosporium

3. Conidiogenous cell elongate-ellipsoid to fusoid............. ...........................................................................Hoornsmania

Coccoideaceae Henn. ex Sacc. & D. Sacc., Syll. fung. (Abellini) 17: 860 (1905), MycoBank: MB 80616

Parasitic on living leaves. Sexual state: Ascostromata large, up to 2 mm diam, single to solitary, or scattered, mostly superficial, slightly immersed in host tissue, circular to subcircular, discoid, cushion-like, semicircular in section, edge entire, slightly convex at the top, raised from the base, black, thickened at the base, soft, with numerous locules, in a layer in the upper part the ascostromata. Cells of ascostromata comprising heavily, pigmented cells of textura angularis. Locules completely immersed in ascostromata, globose, subglobose or obpyriform, with apical ostiole. Hamathecium of filiform, anastomosing, branched, septate, hyaline, relatively wide pseudoparaphyses. Asci 8-spored, bitunicate, fissitunicate, cylindrical to subclavate, pedicellate, apically rounded, with a distinct, wide, ocular chamber. Ascospores 2-seriate, apiosporous, ellipsoidal-obovoid, slightly oval, subglobose, yellowish to light brown, tapering towards the narrow, lower end, 1-septate at the lower end, not constricted at the septum, darkened at the septa, smooth-walled. Asexual state: Unknown.

Notes: The family Coccoideaceae was invalidly introduced by Hennings (1904a) and was subsequently validly published in Saccardo and Saccardo (1905). These are mainly parasites on the lower surface of living leaves and have superficial, circular or discoid and darkly pigmented, ascostromata (Saccardo and Saccardo 1905). The family Coccoideaceae now includes three genera and is placed under Dothideales in Kirk et al. (2008). Lumbsch and Huhndorf (2010) however, list two genera, i.e. Coccoidea and Coccoidella placed in Dothideomycetes incertae sedis. There is no molecular data to confirm these placements. The family is based on the type species C. quercicola Henn. & Shirai from Japan, which we have not been able to locate. von Höhnel (1909b) added a second genus Coccoidella, having circular and flattened ascostromata and bitunicate, oblong asci (Berkeley 1876). We have examined this which mainly differs in having 2-celled ascospores. Both genera are placed in the same family based on only morphological classification and the genera need to be re-studied.

Type: Coccoidea Henn., Bot. Jb. 28: 275 (1900), MycoBank: MB 1144 Fig. 28

Coccoidea quercicola (Material examined: INDIA, Sukliapokhari, Alt. 7,000 ft., on living leaves of Quercus lamellosa Sm., 11 May 1967, M.K. Maity PCC 1252, BPI 643971). a, b Ascostromata. c Section of ascostroma (Note ostioles - arrowed). d Pseudoparaphyses. e–g Immature and mature asci, with a wide ocular chamber. Note: Fissitunicate ascus when releasing ascospores. h–j Ascospores. Scale bars: a = 5 mm, b = 1 mm, c = 200 μm, d, f–g = 50 μm, h–j = 10 μm

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Possible synonyms:

Apiodothina Petr. & Cif., Annls mycol. 30(3/4): 153 (1932)

Coccodiscus Henn., Hedwigia 43: 144 (1904)

Parasitic on living leaves of Quercus lamellosa. Sexual state: Ascostromata large, up to 2 mm diam, single to solitary, or scattered, mostly superficial, slightly immersed in host tissue, circular to subcircular, discoid, cushion-like, semicircular in section, edge entire, slightly convex at the top, raised from the base, black, thickened at the base, soft, with numerous locules, in a layer in the upper part the ascostromata. Cells of ascostromata comprising heavily pigmented cells of textura angularis. Locules of ascomata completely immersed in ascostromata, globose to subglobose or obpyriform, with upper ostiole. Hamathecium of filiform, anastomosing, branched, septate, hyaline, relatively wide pseudoparaphyses. Asci 8-spored, bitunicate, fissitunicate, cylindrical-clavate, with short to long pedicels, apically rounded, with a wide ocular chamber. Ascospores 1–2-seriate, ellipsoidal-obovoid, slightly oval, subglobose, tapering towards narrow, nipple-like base, hyaline and 1-celled when immature, at maturity, 1-septate near the lower end, not constricted and dark brown at the septum, yellowish to light brown, smooth-walled. Asexual state: Unknown.

Notes: Coccoidea was introduced by Hennings (1900), for species with superficial, circular-shaped and black ascostromata. The locules are reported to be in a layer at the top of the ascostromata. Besides the type species it contains three other epithets (Müller and Sanwal 1954; Sawada 1959; Müller and von Arx 1962; Kobayashi et al. 1992; Kishi 1998), one being the type of Coccoidella. Both genera have been placed in Venturiaceae, Pseudosphaeriales (Müller and von Arx 1962). The specimen illustrated here is from India (Kar and Maity 1971) and is different to the Japanese species in having many locules scattered throughout the larger ascostromata. The genus and family need to be restudied.

Type species: Coccoidea quercicola Henn. & Shirai, in Hennings, Bot. Jb. 28: 275 (1900) MycoBank: MB 167300

Other genera included

Coccoidella Höhn., Sber. Akad. Wiss. Wien, Math.-naturw. Kl., Abt. 1 118: 847 [35 repr.] (1909)

Type species: Coccoidella scutula (Berk. & M.A. Curtis) Höhn., Sber. Akad. Wiss. Wien, Math.-naturw. Kl., Abt. 1 118: 847 [35 repr.] (1909)

Key to genera of Coccoideaceae

1. Asci slightly oval, septate at the lower end, yellowish-brown to light brown ..............................................Coccoidea

1. Asci ellipsoidal-oblong to clavate, septate at the centre, hyaline-pale brown ...............................................Coccoidella

Coniothyriaceae W.B. Cooke, Revta Biol., Lisb. 12: 289 (1983), MycoBank: MB: 80635

Sexual state: Unknown. Pathogens of leaves or bark. Asexual state: Conidiomata pycnidial, separate, globose, dark or pale brown, immersed, uni-locular, thin-walled; wall of brown, thick-walled cells of textura angularis or globulosa. Ostiole circular, central, sometimes papillate. Conidiophores absent. Conidiogenous cells holoblastic, annellidic, indeterminate, discrete, doliiform to cylindrical, hyaline or pale brown, smooth, with 1- close or widely separated annellations, formed from the inner cells of the pycnidial wall. Conidia hyaline to brown, thick-walled, 0–1-euseptate, verruculose, apex obtuse, base truncate (sometimes with a marginal frill), cylindrical, sphaerical, elliptical or broadly clavate.

Notes: Cooke (1983) introduced this family to accommodate Coniothyrium spp. However, Kirk et al. (2008) synonymized Coniothyriaceae with Leptosphaeriaceae. This was followed by several researchers (de Gruyter et al. 2009; Schoch et al. 2009a, b; Aveskamp et al. 2010). However, de Gruyter et al. (2012) showed that C. palmarum Corda has distinct phylogenetic relationships with Leptosphaeriaceae. Therefore they reinstated the family Coniothyriaceae in Pleosporales. In their study they transferred some Phoma spp. to Coniothyrium as they group in clade Coniothyriaceae. Although C. palmarum is placed in Coniothyriaceae, C. minitans W.A. Campb. and C. sporulosum (W. Gams & Domsch) Aa grouped in Montagnulaceae hence they were moved to a new genus, Paraconiothyrium (Verkley et al. 2004). Coniothyrium zuluense M.J. Wingf. was shown to be accommodated in Mycosphaerellaceae (Cortinas et al. 2004), but Cortinas et al. (2006) and Crous et al. (2009c) transferred C. zuluense to Colletogloeopsis and showed it best accommodated in Teratosphaeriaceae. However, C. concentricum (Desm.) Sacc. belongs in Phaeosphaeriaceae.

Type: Coniothyrium Corda, Icon. fung. (Prague) 4: 38 (1840), MycoBank: MB 7765

Figure 29

Coniothyrium palmarum (Material examined: CBS 400–71). a, b Conidiomata on Chamaerops humilis (Palmae). c, d Developing conidia attach to conidiogenous cells. e Immature conidia with 0–1 septa. f Mature conidia with one septum. Scale bars. c–f = 10 μm

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Type species: Coniothyrium palmarum Corda, Icon. fung. (Prague) 4: 38 (1840), MycoBank: MB 170942

Cookellaceae Höhn. ex Saccardo & Trotter, in Saccardo, Syll. Fung. (Abellini) 22: 585 (1913), MycoBank: MB 80637

Possible synonym:

Uleomycetaceae Moreau [as ‘Uléomycétacées’], Les Champignons 2: 1384 (1954)

Parasitic on leaves in terrestrial habitats, some possibly fungicolous. Sexual state: Ascostromata superficial, subglobose or irregular, pulvinate to effuse, cushion-shaped, surface rough, flat or applanate at the base, scattered, pigmented, containing locules with individual asci, non-papillate, opening by breaks in the upper surface; soft-textured, cells of ascostromata comprising textura angularis to subglobulosa, with a basal hypostroma, developing in the host tissue, appearing as stromal blocks and swollen cells, of compressed packages of textura angularis to prismatica. Locules scattered throughout ascostromata, small, globose-subglobose, containing single asci, wall not obvious. Hamathecium lacking pseudoparaphyses. Asci 8-spored, bitunicate, fissitunicate, globose to subglobose or saccate, apedicellate, lacking an ocular chamber. Ascospores scattered in 3–5-seriate overlapping rows, muriform, ellipsoid to oblong, hyaline when immature, becoming brown at maturity, straight to slightly curved, ends rounded, with 3-transverse septa and 1-longitudinal septum in each cell, constricted at the septa, smooth-walled. Asexual state: (see notes under genus).

Notes: The family Cookellaceae was introduced by von Höhnel (1909b) as Cookellaceen, and formally published by Saccardo and Trotter (1913). The family is characterized by superficial pulvinate to effuse, cushion-shaped, pigmented ascostromata, without pseudoparaphyses, forming hypostroma in the host tissues. Asci are arranged in individual locules, irregular and mostly globose to subglobose. Ascospores are hyaline to brown, septate and muriform, and forcible discharge of ascospores occurs at maturity (von Höhnel 1909b; von Arx 1963; von Arx and Müller 1975; Barr 1979a, 1987a). von Höhnel (1909a) initially placed Cookella in Myriangiaceae, based on the superficial, disk-like ascostromata with flattened bases, and asci produced in locules in a hyphal tissue. von Höhnel (1909b, 1911) later considered that ascomata with a flattened base, and bright thin-walled peridium, were characteristics that differed from the Myriangiaceae and therefore introduced the family Cookellaceae. von Arx (1963) placed Cookellaceae in Myriangiales based on the superficial, stromatic ascomata and parasitic life form on leaves or other fungi. Later, the family was excluded from Myriangiales and placed in Dothideomycetes of uncertain order (Eriksson 2005). The Cookellaceae presently comprise three genera, Cookella, Pycnoderma and Uleomyces (Lumbsch and Huhndorf 2010; Kirk et al. 2008). Uleomyces is a very poorly known genus that may be similar to Cookella and has pulvinate ascostromata, globose asci and hyaline to brown ascospores (Inácio and Cannon 2008) and also occurs on oak leaves. Uleomyces is said to be fungicolous (Hennings 1895). Pycnoderma is more like a member of Brefeldiellaceae. No molecular data is available for any of these taxa and their placement in the Myriangiales cannot be confirmed.

Type: Cookella Sacc., Michelia 1(no. 4): 407 (1878), MycoBank: MB 1225 Fig. 30

Cookella microscopica (a–i redrawn from PAD: no. 2509; b–d redrawn from Arnaud 1925; e redrawn from von Arx 1963). a Reproduction of notes on herbarium envelope of Cookella microscopica. b, c Ascostromata on substrate. d, e Section through ascostroma to show distribution of locules. f, h Immature and mature asci. i Immature and mature ascospores

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Possible synonyms:

Ascomycetella Peck, Bull. Torrey bot. Club 8: 50 (1881)

Phaneroascus Baudyš, Glasnik zemalyste Muzy Basni Hercegou. 30: 320 (1919)

Parasitic on leaves in terrestrial habitats. Sexual state: Ascostromata superficial, subglobose, cushion-shaped, surface rough, flat or applanate at the base, scattered, pigmented, containing locules with individual asci, non-papillate, opening by breaks in the upper surface; cells of ascostromata comprising textura angularis to subglobulosa, with a basal hypostroma, developing in the host tissue, appearing as stromal blocks with swollen cells, compressed packages of textura angularis to prismatica. Locules scattered throughout ascostromata, small, globose to subglobose, peridial wall composed of textura angularis to subglobulosa. Hamathecium lacking pseudoparaphyses. Asci 8-spored, bitunicate, fissitunicate, globose to subglobose or saccate, apedicellate, lacking an ocular chamber. Ascospores scattered in 3–5-seriate overlapping rows, muriform, ellipsoid to oblong, hyaline when immature, becoming brown at maturity, straight to slightly curved, ends rounded, with 3-transverse septa and 1-longitudinal septum in each cell, constricted at the septa, smooth-walled. Asexual state: see notes.

Type species: Cookella microscopica Sacc., Michelia 1(no. 4): 407 (1878), MycoBank: MB 218808

Notes: The type specimen obtained from PAD on oak leaves lacking ascostromata. Therefore, we provide an account of Cookella based on the protologue and drawings in Arnaud (1925) and von Arx (1963) and from the drawing with the PAD type material. Cookella was established by Saccardo (1878) based on a description of Cookella microscopica. Five species of Cookella are listed in Index Fungorum (2013) and are typically parasites on leaves (von Höhnel 1909b; Barr 1987a). Saccardo (1878) found a coelomycetous taxon with muriform conidia, named Stigmella on leaves around ascostromata of C. microscopica, and considered that this might be the asexual state. The fruiting bodies of the two taxa are however dissimilar (ascostromata versus immersed individual conidiomata). The relationship of these two morphs, which are based on being found on the same material, must be treated with caution.

Other genera included

Pycnoderma Syd. & P. Syd., Annls mycol. 12(6): 563 (1914)

Type species: Pycnoderma bambusinum Syd. & P. Syd. Annls mycol. 12(6): 563 (1914)

Uleomyces Henn., Hedwigia 34: 107 (1895)

Type species: Uleomyces parasiticus Henn., Hedwigia 34: 107 (1895)

Key to genera of Cookellaceae

1. Ascostromata superficial, cushion or disk-shaped .............2

1. Thyriothecia superficial, disk-shaped, .............Pycnoderma

2. Ascospores brown .................................................Cookella

2. Ascospores hyaline ............................................Uleomyces

Corynesporascaceae Sivan., Mycol. Res. 100(7): 786 (1996), MycoBank: MB 81981

Pathogenic and saprobic on leaves. Sexual state: Ascomata cleistothecial with conidiophores arising form cleistothecial walls, sphaerical, solitary to aggregated, superficial on mycelia, or immersed when grown in agar. Peridium thin, composed of pale brown, compressed cells, forming a texura globosa or angularis, covered by mycelia and conidiophores. Hamathecium comprising paraphysoids, with branched, septate, apically free, cylindrical hyphae, which break up when asci mature. Asci 8-spored, thick-walled, bitunicate, obovoid, deliquescent, short-stalked, fasciculate and arising to form a hymenium among deliquescent paraphysoids. Ascospores 2–3-seriate, oblong, pale to dark brown, 1-euseptate at the centre and indistinctly 1-distoseptate in the upper and lower cell, constricted at the median septum, smooth, with a somewhat slightly longer basal cell, which possesses an indistinct, hyaline area, at or near its centre, with three lenticular to globose, granular lumena, the central lumen occupying both the central cells. Asexual state: Conidiophores formed on the superficial, floccose mycelium and cleistothecial wall, macronematous, mononematous, single or caespitose, erect, simple, straight or flexuous, subhyaline to pale brown, none to few septate, smooth, cylindrical. Conidiogenous cells monotretric, integrated, terminal, percurrently proliferating, subhyaline to pale brown, smooth, cylindrical, becoming spathulate towards the apex. Conidia solitary, obclavate, wide at the truncate base, tapering towards the apex, subhyaline to pale brown, acrogenous, unbranched, smooth-walled, straight to curved, multi to distoseptate, germinating from the apical end of the terminal cell by means of 1–4 germ tubes (Sivanesan 1996).

Notes: The family Corynesporascaceae was introduced by Sivanesan (1996) with Corynesporasca caryotae Sivan. as the type species, isolated from a decaying leaf of Caryota urens collected in Sri Lanka. Sivanesan (1996) linked the sexual (Corynesporasca caryotae) and asexual (Corynespora) state in culture. The Corynesporascaceae differ from Testudinaceae by its cephalothecioid peridium and hamathecium with cellular pseudoparaphyses (Hawksworth and Booth 1974; Hawksworth 1979). Didymosphaeria resembles Corynesporasca in having brown, 1-septate ascospores, but are rather small, smooth to echinulate and uni-seriately arranged inside the persistent, usually cylindrical asci, which are provided with a narrow, ocular chamber. The ascomata are immersed and clypeate unlike those in Corynesporasca. Corynesporasca has asci with a wide ocular chamber and lacks a refractive ring. Immature ascospores are 1-septate almost in the middle and appear to be faintly distoseptate in each cell. The ascospore cells have lenticular to globose lumina. The ascospores lack a mucilaginous sheath. While Massariaceae have asci with a wide ocular chamber but often surrounded by refractive, non-amyloid ring; the ascospores are usually distoseptate in some state of their development, often with lenticular to globose lumina and surrounded by a mucilaginous sheath.

Pyrenulaceae and Requienellaceae are similar with Corynesporasca in having distoseptate and euseptate ascospores with lumina, but are distinguishable from Corynesporasca by their lichenized habit and peridium comprised of textura intricata in Pyrenulaceae, or cellular pseudoparaphyses in Requienellaceae (Aptroot 1991a, b). The Trypetheliaceae differ from Corynesporasca in having lichenized ascomata, which may be aggregated in a pseudostroma, a peridium of textura intricata and a pycnidial asexual state (Aptroot 1991a, b). Tubeufiaceae differ from Corynesporasca in having a hamathecium with cellular pseudoparaphyses and submedianly 1-septate, hyaline to yellowish brown, sheathed ascospores (Rodrigues and Samuels 1994).

Type: Corynesporasca Sivan. Mycol. Res. 100(7): 786 (1996), MycoBank: MB 27579

Figure 31

Corynesporasca caryotae (Material examined: SRI LANKA, Kandy, isolate from living leaf of Caryota urens, 18 March 1994, J.M. Waller 4058d, IMI362840a, holotype). a Dried culture of herbarium specimen. b-d Vertical section through cleistothecium. e Surface view of cleistothecium wall. f Wall with conidiophores. g, h Peridium. i, j Asci. k-m Ascospores. Scale bars: b, c = 100 μm, d, e, h = 50 μm, f, g, j = 20 μm, i, k, l, m = 10 μm

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Notes: Corynespora includes 89 species and the genus has a widespread distribution (Kirk et al. 2008), or 173 species epithets (Index Fungorum 2013) and are saprobes, pathogens, and endophytes on woody and herbaceous plants, other fungi, nematodes, and human skin (Dixon et al. 2009). The type of Corynespora is Corynespora mazei Güssow (current name C. cassiicola (Berk. & M.A. Curtis) C.T. Wei). Corynespora cassiicola is an important fungus causing target-spot on a wide host range in tropical and subtropical countries, especially Hevea brasiliensis (rubber) in Sri Lanka and other countries (de Liyanage et al. 1986). In Schoch et al. (2009b, Fig. 2a) C. cassiicola and C. smithii (Berk. & Broome) M.B. Ellis cluster in Clade K in Pleosporales which is basal to Morosphaeriaceae, while C. olivacea (Wallr.) M.B. Ellis clustered with Massarinaceae. Corynespora is obviously polyphyletic and requires detailed molecular analysis. We cannot therefore synonymize Corynesporasca under Corynespora (the older name) at this time, as the types of the two genera may be unrelated. Gemination of ascospores occurs with the germ tube arising from the end of the basal cell. A single germ tube may emerge from the base, but single or multiple germ tubes may be produced from any surface area of the upper cell (Sivanesan 1996).

Type species: Corynesporasca caryotae Sivan., Mycol. Res. 100(7): 786 (1996), MycoBank: MB 415571

Cucurbitariaceae G. Winter [as ‘Cucurbitarieae’], Rabenh. Krypt.-Fl., Edn 2 (Leipzig) 1.2: 308 (1885), MycoBank: MB 80667

Necrotrophic or saprobic on woody plants. Sexual state: Ascomata immersed, semi-immersed, becoming erumpent, to nearly superficial, scattered, or clustered on basal hypostroma, base not easy to remove from the substrate, usually containing host particles, globose to subglobose, black, ostiolate. Ostiole black, papillate, ostiolar canal filled with hyaline cells. Peridium composed of several cell layers of textura angularis, light brown to reddish-brown, smooth to rough, or hairy. Hamathecium of dense, filiform, hyaline, filamentous, septate pseudoparaphyses. Asci 8-spored, bitunicate, fissitunicate, cylindrical to clavate, with furcate pedicel and minute ocular chamber. Ascospores uni-seriate, or partially overlapping, ellipsoidal, golden brown to dark brown, multi-septate, muriform, constricted at the septum. Asexual state: “Coniothyrium”-like, “Phaeoseptoria”-like, Pyrenochaeta and Pyrenochaetopsis coelomycetous states are included in Cucurbitariaceae.

Notes: The family Cucurbitariaceae was introduced by Winter (1885) with Cucurbitaria berberidis (Pers.) Gray, as the type species,based on aggregated ostiolate, ascomata on a basal stromatic structure, fissitunicate and cylindrical asci and pigmented, phragmosporous or muriform ascospores. Barr (1987a) included Cucurbitariaceae in the order Pleosporales, with Cucurbitaria, Cucurbidothis, Otthia, Rhytidiella and Syncarpella. Schoch et al. (2009b), based on molecular data, have shown that Otthia groups in the Botryosphaeriaceae, although this may have not been a good representative strain. Little molecular data isavailable for taxa assigned to the family (Schoch et al. 2006; Kruys and Wedin 2009). In the dataset by Zhang et al. (2012a), based on sequences of four genes, the clade including the type species is poorly supported and labeled Cucurbitariaceae/Didymosphaeriaceae? Thus indicating that further taxon sampling is required to resolve its position in the Pleosporales. Based on the multi-gene analysis Pyrenochaeta, and Pyrenochaetopsis were shown to be accommodated in Cucurbitariaceae (de Gruyter et al. 2010, 2012). Wijayawardene et al. (2012) listed “Camarosporium”-like, “Coniothyrium”-like and “Phaeoseptoria”-like asexual states in Cucurbitariaceae. The type species of Cucurbitaria (C. berberidis) is linked to Pyrenochaeta berberidis (Sacc.) Brunaud (Sivanesan 1984; Farr et al. 1989; de Gruyter et al. 2010) hence we accept that Cucurbitaria has “Pyrenochaeta”-like asexual states. However, Doilom et al. (2013) show that Pyrenochaeta nobilis De Not. which is the type of Pyrenochaeta, groups away from Cucurbitaria berberidis, thus we do not synnonimize Pyrenochaeta under Cucurbitaria (Doilom et al. 2013) and that this is a well-supported family.

Type: Cucurbitaria Gray, Nat. Arr. Brit. Pl. (London) 1: 508, 519 (1821), MycoBank: MB 1348 Fig. 32

Cucurbitaria elongata (Fr.) Grev. (Material examined: ITALY, Forlì-Cesena, Corniolo, on dead wood of Cytisus scoparius, 06 December 2011, E Campestri, MFLU13-0085). a Ascomata on the host stems, partly erumpent. b Close up of the ascomata. c Section of an ascoma. d Close up of the peridium. e Hamathecium comprising asci and filamentous pseudoparaphyses. f Hyaline, filamentous pseudoparaphyses. g 8-spored asci with short pedicels. h Ascus tip showing apical apparatus. i–l Golden brown to dark brown, ellipsoidal to broadly, muriform ascospores. Scale bars: c = 200 μm, d = 20 μm, e = 10 μm, f–h = 30 μm, i–l = 10 μm

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Saprobic on woody branches. Sexual state: Ascomata black, semi-immersed, scattered or clustered on a basal hypostroma beneath the host periderm or on decorticated wood, fully or partly erumpent, globose, smooth to rough or hairy, with a papillate ostiole. Hypostroma cells brown, angular to globose, thickened at the base. Peridium comprising 5–6 cell layers of light brown to reddish-brown cells of textura angularis. Hamathecium of 2–3 μm wide, numerous, filiform, hyaline pseudoparaphyses. Asci 8-spored, bitunicate, thick-walled, cylindrical, short-pedicellate, clusters arising from the base. Ascospores uni-seriate, initially hyaline, becoming golden brown to dark brown, ellipsoidal to broadly, muriform, constricted at the septum, with obtuse or acute ends. Asexual state: “Pyrenochaeta”-like, Pycnidia superficial on wood in bark fissures, scattered or aggregated, (sub)globose or obpyriform, unilocular, dark brown to black, thick-walled, with setae, pseudoparenchymatous walled. Setae dark brown, thick-walled, sometimes light-brown and thin-walled, ends narrowly rounded and broadly at the base, septate, numerous, crowded. Conidiogenous cells enteroblastic, phialidic, hyaline, smooth. Conidia agglutinated in masses, oblong to ellipsoid, hyaline, aseptate, guttulate, smooth-walled (Sutton 1980).

Notes: Cucurbitaria is one of the oldest pyrenomycete genera separated from Sphaeria Haller in the sense of Tode (1790) and Persoon (1801), while Fries (1823) considered it under Pyrenomycetes along with the genus Sphaeria. Cucurbitaria was described by Gray (1821) and C. berberidis is considered the type of the genus. The original generic diagnosis of Cucurbitaria was “Thallus spreading irregular, thecae in tufts, placed on the thallus.” but the scope of the genus was unlimited. Therefore, Greville (1824, 1825a, b), Tulasne and Tulasne (1863) Saccardo (1883), Winter (1887), Ellis and Everhart (1892) and Berlese (1900), have described the genus in more detail but still the delimitation of the genus remains inconclusive. Winter (1887) placed Cucurbitaria in the family Cucurbitariaceae with other taxa having aggregated fruiting bodies. The subsequent monographic study by Welch (1926) had a narrow concept for the genus, and accepted only C. berberidis, C. caraganae P. Karst., C. laburni (Pers.) De Not., C. elongata (Fr.) Grev. and C. arizonica Ellis & Everh. Other species were regarded as doubtful species, or excluded species.

Type species: Cucurbitaria berberidis (Pers.) Gray, Nat. Arr. Brit. Pl. (London) 1: 508, 519 (1821), MycoBank MB 239072

≡ Sphaeria berberidis Pers., Neues Mag. Bot. 1: 83 (1794)

Other genera included

Curreya Sacc., Syll. fung. (Abellini) 2: 651 (1883)

Type species: Curreya conorum (Fuckel) Sacc., Syll. fung. (Abellini) 2: 651 (1883)

Pyrenochaeta De Not., Mem. R. Accad. Sci. Torino, Ser. 2 10: 348 (1849)

Type species: Pyrenochaeta nobilis De Not., Mem. R. Accad. Sci. Torino, Ser. 2 10: 347 (1849)

Pyrenochaetopsis Gruyter et al., Mycologia 102(5): 1076 (2010)

Type species: Pyrenochaetopsis leptospora (Sacc. & Briard) Gruyter et al., Mycologia 102(5): 1076 (2010)

Rhytidiella Zalasky, Can. J. Bot. 46(11): 1383 (1968)

Type species: Rhytidiella moriformis Zalasky, Can. J. Bot. 46(11): 1383 (1968)

Syncarpella Theiss. & Syd., Annls mycol. 13(5/6): 631 (1915)

Type species: Syncarpella tumefaciens (Ellis & Harkn.) Theiss. & Syd., Annls mycol. 13(5/6): 633 (1915)

Key to sexual genera of Cucurbitariaceae

1. Ascospores muriform .........................................................2

1. Ascospores filiform ............................................................3

2. Asci 183 × 17 μm, cylindrical .........................Cucurbitaria

2. Asci 77 × 28 μm, cylindrical-clavate to broadlyclavate .....................................................................................Curreya

3. Ascospores with 3 transverse septa, slightly constrict at the septa ......................................................................Syncarpella

3. Ascosporeswith more than 7 transverse septa, sigmoid to curved .....................................................................Rhytidiella

Key to asexual genera of Cucurbitariaceae

1. Conidiomata with setae (in most cases), conidia wider than 2 μm ..................................................................Pyrenochaeta

1. Conidiomata lack of setae, conidia narrower than 2 μm .....................................................................Pyrenochaetopsis

Dacampiaceae Körb. [as ‘Dacampieae’], Syst. Lich. Germ.: 322 (1855), MycoBank: MB 80680

Possible synonym:

Pyrenidiaceae Zahlbr., in Engler, Syllabus, Edn 2 (Berlin): 46 (1898)

Lichenicolous, lichenized or saprobic. Sexual state: Ascomata perithecioid, mostly uni-locular, with a central ostiole, some genera forming a clypeus around the ostiole, lacking setae, black, immersed to superficial, mostly subglobose to obpyriform, rarely elongate or irregularly-shaped, solitary and scattered over the substratum, or grouped. Peridium composed of several layers of angular pseudoparenchymatous, radially compressed, thick-walled cells of textura angularis, reddish to dark brown, or pale brown to colorless, surrounded by a dark brown involucrellum. Hamathecium of numerous, usually branched and anastomosing, trabeculate or cellular pseudoparaphyses, I-, more rarely K/I + bluish. Asci up to 8-spored, bitunicate, fissitunicate, elongate-clavate to short-cylindrical, short-pedicellate, with a thick-walled apex and a small, sometimes indistinct ocular chamber, I-, more rarely K/I + bluish. Ascospores 1–2-seriate, 1- or more septate to muriform, in most species brown to dark brown, more rarely hyaline to pale brown, ellipsoid to fusiform, often constricted at the septa, smooth to verrucose, perispore present in some species. Asexual state: pycnidial states reported in a few genera, e.g. in Eopyrenula (colourless, simple microconidia and brown, 1- or more septate macroconidia), Munkovalsaria (“Lecythophora”-like asexual state), Polycoccum (“Cyclothyrium”-like asexual state of P. peltigerae, “Phoma”-like asexual state of P. rubellianae).

Notes: The family Dacampiaceae, as currently understood, is certainly heterogeneous, and the inclusion of several genera is tentative, as the lack of molecular data does not support or reject their inclusion. The core of the family, as suggested by Crivelli (1983) and Hawksworth and Diederich (1988) comprises the type, Dacampia, and several other lichenicolous genera, such as Clypeococcum, Polycoccum, Pyrenidium and Weddellomyces. Most species included in these genera share blackish, perithecioid ascomata, bitunicate, fissitunicate asci, similar pseudoparaphyses and medium to dark brown, thick-walled ascospores. However, there have been few ontogenetic studies concerning the nature of the hamathecium, and such studies would probably have an influence on the circumscription of the family. Very few asexual morphs have been reported within these genera, and almost no species have been isolated in pure culture. No molecular data is available yet for the genus Dacampia, and very few sequences have been published from other genera currently referred to the Dacampiaceae. Munkovalsaria appendiculata Aptroot was included in a phylogenetic analysis by Wang et al. (2007) and the species referred to the Dacampiaceae, but as no Dacampia species has been sequenced, interpretations of the phylogenetic position of the family are speculative.

Most species of the Dacampiaceae are lichenicolous, and almost all seem to be host-specific. Pyrenidium actinellum Nyl. was often considered as an exception, being found on unrelated hosts, but in reality represents an assemblage of many distinct, mostly undescribed species. The type of Dacampia is remarkable in being lichenized, and a further genus often included in the Dacampiaceae, Eopyrenula, is a facultative lichen. Several poorly known genera, some with aberrant characters, such as hyaline ascospores and unbranched hamathecial filaments, mainly growing on plants, have provisionally been included in the Dacampiaceae and are listed and keyed out below. The family includes ten genera and 110 species. Aaosphaeria is a monotypic, non-lichenicolous genus with a Microsphaeropsis anamorph that “could be close to Polycoccum in the Dacampiaceae” (Aptroot 1995a). Clypeococcum includes nine lichenicolous species. Species of Eopyrenula are bark-inhabiting, facultatively lichenized with a Trentepohlia photobiont; six species were keyed out by Aptroot (2012) who treated Eopyrenula as part of Pyrenulaceae (Eurotiomycetes); further studies are needed to clarify the position of this genus. Leptocucurthis is a monotypic, non-lichenicolous genus included in Dacampiaceae by Aptroot and van Iperen (1998), but differing by unbranched pseudoparaphyses. Munkovalsaria is a non-lichenicolous genus with an anamorph resembling Lecythophora “best accommodated in the Dacampiaceae” (Aptroot 1995a), with two species. Polycoccum includes 56 lichenicolous species. Pseudonitschkia is a poorly known, monotypic lichenicolous genus “sharing certain features with the Dacampiaceae” (Coppins and Kondratyuk 1995). Pyrenidium includes nine species, while Weddellomyces includes 12 lichenicolous species, of which eight are keyed out by Calatayud and Navarro-Rosinés (1998).

The conidial taxa of Dacampiaceae need to be resolved. Halici and Hawksworth (2008) speculated whether Dacampia muralicola Halici & D. Hawksw. has asexual states in Phoma as D. muralicola is associated with Phoma spp. However this link has not been proven by any physical connection (Halici and Hawksworth 2008). Most of these associated lichenicolous “Phoma”-like spp. (Phoma sensu strict restricted to Didymellaceae) must be grouped in Phaeosphaeriaceae (Lawrey et al. 2012); however, it is not yet possible to reject with certainty any link between Dacampia and associated Phoma spp. Wijayawardene et al. (2012) listed Cyclothyrium as an asexual state of Polycoccum, but Sutton (1980) stated that Cyclothyrium juglandis (Schumach.) B. Sutton, the type of the genus Cyclothyrium, is the asexual state of Thyridaria rubronotata (Berk. & Broome) Sacc. This link was proven by Verkley et al. (2004) using molecular phylogeny. However Thyridaria was not placed in Dacampiaceae (Schoch et al. 2009b; Zhang et al. 2012a); hence the link between Cyclothyrium sensu strictoand Polycoccum is questionable.

Type: Dacampia A. Massal., Sulla Lec. hook. Schaer.: 7 (1853), MycoBank: MB 1401

Figures 33 and 34

Dacampia hookeri (Material examined: UK, Scotland, Mid Perthshire, on dead mosses on the micaceous soil of Ben Lawers, on thallus of lichen, W.J. Hooker, K(M) 140027, isotype?). a Herbarium label b Specimen of Dacampia hookeri. c Ascomata on white thallus. d Section through ascoma. e Section through peridium. f Asci with pseudoparaphyses. g Young ascus with pale brown, immature ascospores. h-j Mature asci. k-o Ascospores stained by Lactoglycerol. Scale bars: c = 1 mm, d = 100 μm, f = 50 μm, e, g, h, i, j = 20 μm, k–o = 10 μm

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Lichenicolous members of Dacampiaceae. a Dacampia engeliana (Material examined: GERMANY, Bayern, Bad Tölz-Wolfratshausen, 1978, Feuerer), black perithecia immersed in a thallus of Solorina saccata. b Clypeococcum cladonema (Material examined: BELGIUM, Houffalize, 2010, Diederich 17134), black clypei with immersed perithecia on a thallus of Xanthoparmelia pulla. c Ascospore of same specimen. d Polycoccum trypethelioides (Material examined: CANADA, British Columbia, Kispiox, 1995, Goward 95–282), black perithecia inducing galls on Stereocaulon tomentosum. e Weddellomyces epicallopisma (Material examined: FRANCE, Vaucluse, Gordes, 1995, Diederich 12327), black perithecia immersed in a partly decolorized thallus of Caloplaca aurantia. f Pyrenidium baeomycearium (Material examined: Santesson, Fungi Lich. Exs. 331, sub P. actinellum), black perithecia inducing galls on Baeomyces rufus. g Pyrenidium sp. (Material examined: LUXEMBOURG, Baschleiden, on Verrucaria, 2007, Kuborn), 3-septate ascospores with an apically paler wall. Scale bars: a–b, d–f = 200 μm, c, g = 10 μm

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Possible synonyms:

Decampia Mudd, (1861)

Xenosphaeria Trevis., Conspect. Verruc.: 18 (1860)

Lichenicolous or with a lichenized thallus. Sexual state: Ascomata perithecioid, with a central ostiole, lacking setae, black, immersed to semi-immersed, subglobose to obpyriform, solitary, sometimes clustered, scattered over the host thallus or apothecia. Peridium composed of several layers of angular pseudoparenchymatous, radially compressed, thick-walled cells of textura angularis, reddish to dark brown. Hamathecium of numerous, branched and anastomosing, cellular pseudoparaphyses, I-. Asci 8-spored, bitunicate, elongate-clavate to short-cylindrical, short-pedicellate, with a thick-walled apex and a small, sometimes indistinct, ocular chamber. Ascospores 1–2-seriate, muriform (but transverse septa may be missing when immature), initially pale yellowish to pale brown, becoming brown to dark brown, ellipsoid to fusiform, often constricted at the septa, smooth, without visible perispore. Asexual state: Unknown.

Notes: Unlike all other known species of the genus, which are lichenicolous and non-lichenized, the type Dacampia hookeri (Borrer) A. Massal., shows an unusual and for a long time misunderstood biology. Ascomata of D. hookeri, develop over a white, terricolous, lichenized thallus, often associated when young with Solorina thalli. Henssen (1995) demonstrated that this white thallus belongs to D. hookeri, which is therefore the only known lichenized member of the genus. Other species are parasitic and form necrotic patches on the host thallus or tend to be commensalistic. A total of 13 species are currently known and a key to seven of them is given by Halici and Hawksworth (2008).

Type species: Dacampia hookeri (Borrer) A. Massal., Sulla Lec. hook. Schaer.: 7 (1853)

MycoBank: MB 119080

≡ Verrucaria hookeri Borrer, in Smith & Sowerby, Engl. Bot., Suppl. 1: pl. 2622, Fig. 1 (1831)

Other genera included

Aaosphaeria Aptroot, Nova Hedwigia 60(3–4): 329 (1995)

Type species: Aaosphaeria arxii (Aa) Aptroot, Nova Hedwigia 60(3–4): 329 (1995)

Clypeococcum D. Hawksw., J. Linn Soc., Bot. 75: 196 (1977)

Type species: Clypeococcum cladonema (Wedd.) D. Hawksw., J. Linn. Soc., Bot. 75(2): 197 (1977)

Eopyrenula R.C. Harris, Michigan Bot. 12(1): 19 (1973)

Type species: Eopyrenula leucoplaca (Wallr.) R.C. Harris, Michigan Bot. 12(1): 19 (1973)

Leptocucurthis Aptroot, in Aptroot & Iperen, Nova Hedwigia 67: 485 (1998)

Type species: Leptocucurthis quadrata Aptroot, in Aptroot & Iperen, Nova Hedwigia 67: 485 (1998)

Munkovalsaria Aptroot, Nova Hedwigia 60(3–4): 346 (1995)

Type species: Munkovalsaria donacina (Niessl) Aptroot, Nova Hedwigia 60(3–4): 346 (1995)

Polycoccum Saut. ex Körb., Parerga lichenol. (Breslau): 470 (1865)

Type species: Polycoccum sauteri Körb., Parerga lichenol. (Breslau): 470 (1865)

Pseudonitschkia Coppins & S. Y. Kondr., Arch. Protistenk. 52(2): 232 (1995)

Type species: Pseudonitschkia parmotrematis Coppins & S.Y. Kondr., Edinb. J. Bot. 52: 232 (1995)

Pyrenidium Nyl., Flora, Jena 48: 210 (1865)

Type species: Pyrenidium actinellum Nyl., Flora, Jena 48: 210 (1865)

Weddellomyces D. Hawksw., Notes R. bot. Gdn Edinb. 43(3): 511 (1986)

Type species: Weddellomyces epicallopisma (Wedd.) D. Hawksw. [as ‘epicallopismum’], Notes R. bot. Gdn Edinb. 43(3): 512 (1986)

Key to genera of Dacampiaceae

1. Pseudoparaphyses unbranched, ostiole slit-like to stellate, with 3–6 radiate splits, ascospores hyaline, elongate fusiform to subcylindrical, 1-septate ..............................Leptocucurthis

1. Pseudoparaphyses branched, ostiole punctiform ...............2

2. Ascomatal surface deeply cracked and rugulose with 10–17 μm warts, ascospores hyaline, becoming brownish when over mature, fusiform, 1(−3)-septate, 40–49 × 6.5–7.5 μm .......................................................................Pseudonitschkia

2. Ascomatal surface smooth, not distinctly cracked or rugulose, ascospores pale to dark brown, length/breadth ratio usually smaller ..............................................................................3

3. Ascospores 1-septate ..........................................................4

3. Ascospores 3–7-septate to muriform .................................7

4. Ascomata discrete, not united in a clypeus ........................5

4. Ascomata united in groups by a common black clypeus .................................................................................................6

5. Ascomata on lichens ........................................Polycoccum

5. Ascomata on plants .........................................Aaosphaeria

6. Ascomata on lichens ....................................Clypeococcum

6. Ascomata on stems and bark of various plants .........................................................................Munkovalsaria

7. Peridium in upper part made of cephalothecioid plates, breaking up irregularly, ascospores 3-septate to muriform .........................................................................Weddellomyces

7. Peridium not cephalothecioid ............................................8

8. Ascomata non-lichenicolous, usually with an immersed, inconspicuous or whitish corticolous thallus that may be lichenized with a Trentepohlia photobiont, often with a dark brown involucrellum, frequently accompanied by a micro- or macroconidial pycnidial asexual morph, ascospores 3–7-septate, apices often paler ..........................................Eopyrenula

8. Ascomata lichenicolous, more rarely lichenized with a thick, white thallus with a Coccomyxia photobiont and external cephalodia with Nostoc ....................................................9

9. Ascospores 3-septate, wall at both apices paler and thinner, ascomata lichenicolous .........................................Pyrenidium

9. Ascospores muriform, wall uniformly thickened and pigmented, ascomata lichenicolous or lichenized with a thick, white thallus with a Coccomyxia photobiont and external cephalodia with Nostoc ...........................................Dacampia

Delitschiaceae M.E. Barr, Mycotaxon. 76: 109 (2000), MycoBank: MB 82092

Terrestrial, hyper-saprobic on herbivore dung, or rarely on aged wood or plants. Sexual state: Ascomata uni-locular pseudothecia, solitary or scattered, sometimes gregarious, immersed to erumpent, brown or black, globose to subglobose, or pyriform, membranous to coriaceous, with a papilla or a well differentiated ostiole, outer wall smooth or hairy, periphysate. Peridium pseudoparenchymatous exostratum, consists with textura angularis cells. Hamathecium of dense, long, branching, anastomosing, cellular pseudoparaphyses. Asci 8-spored, sometimes 4 or poly-spored, bitunicate, fissitunicate, cylindrical to cylindric-clavate, pedicellate, with a non-amyloid, refractive apical apparatus. Ascospores uni- or bi-seriate, rarely tri-seriate, ovoid or ellipsoid, pale to dark brown, unicellular or bi- to multi-cellular, often constricted at the septum, resulting in fragmenting cells, wall smooth, with a full length germ slit in each cell, surrounded by a gel coat. Asexual state: not reported.

Notes: The family Delitschiaceae (Pleosporales) was introduced to accommodate three genera of the family Sporormiaceae, that were clearly different from that family in having a periphysate ostiolum and asci with a well-developed and refractive apical chamber (Barr 2000a). Delitschia, the type genus of this family, was introduced by Auerswald (1866) and assigned to Sphaeriaceae. It was considered to be closely related to Sordariaceae and Amphisphaeriaceae. Winter (1887) assigned Delitschia under Sordariaceae and this placement is followed in several subsequent studies (Griffiths 1901; Kirschstein 1911). Cain (1934) suggested that Delitschia might belong in Pleosporaceae and this proposal was supported by Moreau (1953) and Dennis (1968). In his key to coprophilous Sphaeriales, Cain (1934) separated the fissitunicated bitunicate fungi in to three genera, Sporormia, Delitschia, and Pleophragmia on the basis of ascospore septation. Munk (1957) established Sporormiaceae and Delitschia was assigned therein together with other coprophilous genera, i.e. Sporormia (inclusive of Sporormiella), Perisporium (= Preussia), and Trichodelitschia. Barr (1987b) included the Sporormiaceae in Phaeotrichaceae, but later (Barr 1990b) regarded them as independent families, misinterpreting Phaeotrichaceae as having unitunicate asci and accommodating them in Sordariales. Finally (Barr 2000a) subdivided coprophilous bitunicate fungi in three families, assigning those with ascospores with germ pores to Phaeotrichaceae, those with germ slits or no germinal mechanism to Delitschiaceae or Sporormiaceae, the latter aperiphysate and with a narrow ascal endotunica lacking a refractive apical apparatus. Phylogenetic molecular studies (Kruys et al. 2006; Schoch et al. 2009b) showed Phaeotrichaceae, Sporormiaceae and Delitschiaceae to form separate monophyletic groups.

Type: Delitschia Auersw., Hedwigia 5: 49 (1866), MycoBank: MB 1443 Fig. 35

Delitschia didyma (Material examined: ITALY, F. Doveri, MFLU 12–2217). a, b Ascomata on natural substrate. c Cross section of ascoma. d Ostiole. e Peridium f Pseudoparaphyses. g, h Asci in water. i Apical chamber. j, k Ascospores in water. Scale bars: a = 1 mm, b = 500 μm, c, d = 100 μm, e–h = 50 μm, i–k = 20 μm

Full size image

Possible synonyms:

Delitschiella Sacc. & D. Sacc., in Saccardo & Saccardo, Syll. fung. (Abellini) 17: 688 (1905)

Pachyspora Kirschst., Verh. bot. Ver. Prov. Brandenb. 48: 48 (1907) [1906]

Habitat terrestrial. Saprobic on old herbivore dung. Sexual state: Ascomata solitary or scattered, immersed to semi-immersed, subglobose to subpyriform, brown or black, membranous, relatively large, often covered with dense, hyphoid hairs, neck blackish, central, usually coriaceous, with a wide opening and a smooth or tuberculate to hairy surface. Peridium thick at the apex, thinner at the base, well differentiated with an outer layer of thick-walled, dark pigmented cells, and an inner layer of larger, thin-walled, lightly pigmented cells. Hamathecium of dense, very long, anastomosing and branching, filiform pseudoparaphyses. Asci 8-spored, cylindrical to cylindro-clavate, short, narrow pedicellate, which are rounded at the base, apex with a wide ocular chamber. Ascospores obliquely uni-seriate and partially overlapping, ellipsoid, reddish brown, 1-septate, slightly constricted at the septum, smooth-walled, cells often easily separable from each other, each cell with a full length germ slit. Asexual state: Unknown.

Notes: Luck-Allen and Cain (1975) reviewed the genus Delitschia, which they definitively regarded as usually fimicolous, with bitunicate asci, and pigmented, 1-septate ascospores, with an elongated germ slit in each cell and surrounded by a gelatinous sheath. They described several new taxa and accepted 46 species. Since then about 15 new Delitschia species have been described, some as wood-inhabiting (Eaton and Jones 1971; Romero and Samuels 1991; Hyde and Steinke 1996). Kirk et al. (2008) recognized 51 species, whereas 81 taxa are mentioned in Index Fungorum (2013).

Type species: Delitschia didyma Auersw., Hedwigia 5: 49 (1866), MycoBank: MB 177056

Other genera included

Ohleriella Earle, Bulletin of the New York Botanical Garden 2: 349 (1902)

Type species: Ohleriella neomexicana Earle, Bulletin of the New York Botanical Garden 2: 349 (1902)

Semidelitschia Cain & Luck-Allen, Mycologia 61: 581 (1969)

Type species: Semidelitschia agasmatica Cain & Luck-Allen, Mycologia 61: 581 (1969)

Key to genera in Delitschiaceae

1. Ascospores 1-celled, fimicolous ..................Semidelitschia

1. Ascospores septate, also on other substrata .......................2

2. Ascospores 2-celled, fimicolous, rarely lignicolous ..................................................................................Delitschia

2. Ascospores more than 2-celled, on woody material ..................................................................................Ohleriella

Diademaceae Shoemaker & C.E. Babc., Can. J. Bot. 70(8): 1618 (1992), MycoBank: MB 81955

Parasitic or saprobic in stems and leaves. Sexual state: Ascomata subepidermal or subcuticular and later become superficial, dark brown to black, globose, opening via a flat, circular lid. Peridium thin, consisting of small, pigmented, thick-walled cells of textura angularis. Hamathecium of dense, cellular pseudoparaphyses. Asci 8-spored, bitunicate, fissitunicate, clavate or ellipsoidal, with short, broad, furcate pedicel, apically rounded with an ocular chamber. Ascospores bi-seriate, partially overlapping, fusiform, brown, with 3 or more transverse septa, with or without longitudinal septa, applanate or rarely terete, with a thick sheath most of the time. Asexual state: Unknown.

Notes: Shoemaker and Babcock (1992) introduced Diademaceae which they considered to be a unique family based on the ascomata opening by a flat, circular lid and comprising Clathrospora, Comoclathris, Diadema, Diademosa and Macrospora Shoemaker and Babcock (1992). Macrospora was excluded by Lumbsch and Huhndorf (2010) and included in the family Pleosporaceae, as it was considered to be a synonym for Pyrenophora. Other than the main feature of ascoma opening via a flat, circular lid, as an adaptation to the alpine habitat (Shoemaker and Babcock 1989), ascomata are superficial, globose to subglobose, brown to dark brown, with 8-spored, clavate or ellipsoidal, bitunicate asci. Ascospores are fusiform, brown, with 3 or more transverse septa with or without longitudinal septa, applanate or rarely circular in section, but narrowing to one end (Diademosa) with a thick sheath most of the time (Shoemaker and Babcock 1992; Zhang et al. 2011b). Platyspora has been added to this family by various authors (Zhang et al. 2011b), or was considered a synonym of Graphyllium (Zhang et al. 2011b) or as a synonym of Comoclathris (Index Fungorum 2013). Lumbsch and Huhndorf (2010) included Graphyllium in the family Diademaceae, but Zhang et al. (2011b) referred it the Hysteriaceae due to its hysterothecium-like ascomata, forming a longitudinal opening. The final placement of Graphyllium will rely on molecular data (Zhang et al. 2011b). No asexual state has been linked to most of the members of this family, but an “Alternaria”-like asexual morph has been linked to Clathrospora and Comoclathris (Zhang et al. 2011a, b). No new genera have been included in the family since Diademaceae was introduced by Shoemaker and Babcock (1992). Molecular data is only available in GenBank for putative strains of Clathrospora elynae Rabenh. (CBS 196.54) and C. diplospora (Ellis & Everh.) Wehm. (IMI 68086), which clustered within the family Pleosporaceae.

Species of the order Pleosporales with applanate ascospores can be separated into three families (Diademaceae, Hysteriaceae, Pleosporaceae) and differ in the way the ascomata open. Ascomata opening by a flat, circular lid are characteristic of the family Diademaceae, while species of Hysteriaceae open via a long, narrow slit and species of Pleosporaceae open by a central pore (Shoemaker and Babcock 1992). The placement of Diademaceae is not supported by molecular data and studies are required to resolve its phylogenetic relationship in the Pleosporales.

Type: Diadema Shoemaker & C.E. Babc., Can. J. Bot. 67(5): 1349 (1989), MycoBank:

MB25293 Fig. 36

Diadema tetramerum (Material examined: USA, California, Mt. Shasta, ridge south of Horse Camp, elevation 8,250 ft, on culms of Trisetum spicatum (L.) Richter, 2 July 1947 W.B. Cooke 20223, DAOM, holotype). a Ascomata on substrate opening via a flat circular lid. b Vertical section of ascoma. c Close up of the peridium. d Hyaline, cellular, pseudoparaphyses. e–i Asci with short orbicular pedicel. j–l Reddish-brown ascospores with very broad sheath. Scale bars: b = 100 μm, c = 10 μm, d–g = 60 μm, h–j = 30 μm

Full size image

Saprobic on culms of grasses (Poaceae). Sexual state: Ascomata scattered, immersed, intra-epidermal, globose to subglobose, black to brown, smooth-walled and opening via a flat circular lid. Peridium 1-layered, composed of small, pigmented, thick walled, compressed cells, base consisting of small, pigmented, thick-walled cells of textura angularis. Hamathecium of dense, numerous, septate, hyaline, cellular pseudoparaphyses. Asci 8-spored, numerous, bitunicate, fissitunicate, broadly-clavate, with a short, orbicular pedicel, rounded at apex, without an ocular chamber. Ascospores obliquely bi-seriate, broadly fusiform, 3-trans-septate, without longitudinal septa, brown to reddish-brown with guttules, smooth-walled, finely punctate, with wide mucilaginous sheath. Asexual state: Unknown.

Notes: Diadema is characterized by having large ascospores, without longitudinal septa and ascomata with a circular, lid-like opening (Shoemaker and Babcock 1989). Eight species of Diadema are listed in Index Fungorum (2013). Six species were included when the genus was introduced and another two species (Diadema ahmadii, Kaz. Tanaka & S.H. Iqbal, Diadema sieversiae (Peck) Huhndorf) were later added (Shoemaker and Babcock 1989; Zhang et al. 2011b.

Type species: Diadema tetramerum Shoemaker & C.E. Babc. [as ‘tetramera’], Can. J. Bot. 67(5): 1354 (1989), MycoBank: MB 136222

Other genera included

Comoclathris Clem., Gen. fung. (Minneapolis), 37: 173 (1909)

Type species: Comoclathris lanata Clem. [as ‘Comochlatris’], Gen. fung. (Minneapolis): 1–227 (1909)

Diademosa Shoemaker & C.E. Babc., Can. J. Bot. 70(8): 1641 (1992)

Type species: Diademosa californiana (M.E. Barr) Shoemaker & C.E. Babc. [as ‘californianum’], Can. J. Bot. 70(8): 1641 (1992)

Key to genera of Diademaceae

1. Ascospores with transverse septa, without longitudinal septa ..........................................................................Diadema

1. Ascospores with transverse septa, with longitudinal septa ................................................................................2

2. Ascospores cylindrical .......................................Diademosa

2. Ascospores flattened ......................................Comoclathris

Didymellaceae Gruyter, Aveskamp & Verkley, Mycol. Res. 113(4): 516 (2009), MycoBank: MB 508292

Parasites or saprobes on wood and dead herbaceous stems or leaves. Sexual state: Ascomata pseudothecia, separate or gregarious, or scattered, immersed, erumpent, or rarely superficial, brown to black, ostiole central. Peridium relatively thin, composed of 2–3 cell layers of textura angularis. Hamathecium lacking pseudoparaphyses. Asci 8-spored, bitunicate, fissitunicate, cylindric-clavate to clavate, short-pedicellate or apedicellate, with an ocular chamber. Ascospores overlapping uni-seriate or bi-seriate, upper cell wider than lower cell, hyaline 1-septate, constricted at the septum, smooth-walled, asymmetrical. Asexual states: coelomycetous. Conidiomata pycnidial, immersed, or semi-immersed, sometimes becoming erumpent, uni-locular, pale to medium brown, globose, thin or thick-walled, peridium with cells of textura angularis. Ostiole central, circular, papillate or not. Conidiophores mostly absent, only present in Phoma cava Schulzer and P. tracheiphila Petri) L.A. Kantsch. & Gikaschvili and then either filiform, septate, and branched, or short, irregularly branched, and ramified respectively. Conidiogenous cells enteroblastic, phialidic, doliiform to lageniform, ampulliform or cylindrical, hyaline, thin walled, smooth-walled. Conidia hyaline or pale brown, septate or aseptate, thin-walled, guttulate, ellipsoid, cylindrical, fusiform, pyriform or globose.

Notes: The family Didymellaceae was introduced by de Gruyter et al. (2009) in order to accommodate the type species Didymella exigua, together with some Phoma or Phoma-like genera which formed a robust familial clade in the phylogenetic tree. Zhang et al. (2009a) included the family in the order Pleosporales within the suborder Pleosporineae. De Gruyter et al. (2009) included the sexual morphs of Didymella and Leptosphaerulina in Didymellaceae. Wijayawardene et al. (2012) and Zhang et al. (2012a) listed the asexual states of Didymellaceae as Ampelomyces, Ascochyta, Boeremia, Chaetasbolisia, Dactuliochaeta, Epicoccum, Microsphaeropsis, Peyronellaea, Phoma, Piggotia and Pithoascus. Among them Ascochyta and Phoma are well known pathogens that are significantly important in quarantine (Kaiser et al. 2008; Aveskamp et al. 2010; de Gruyter et al. 2012, in press). Much research has been carried out since Saccardoan times to distinguish Ascochyta and Phoma according to their morphological characters, such as their habitat or substrate and morphological characters, such as presence or absence of septa (Van der Aa et al. 1990). However molecular evidence shows that there are significant differences between Ascochyta and Phoma (de Gruyter et al. 2009). Previously Ascochyta was considered to be the asexual state of Mycosphaerella (Stone 1912; van Warmelo 1966). Peever et al. (2007) and Chilvers et al. (2009) showed that Didymella pinodes (Berk. & A. Bloxam) Petr. is the sexual state of Ascochyta pinodes L.K. Jones, and D. pisi Chilvers et al. (2009) is the sexual state of A. pisi Lib. The latter species of Ascochyta is the type species; hence Ascochyta is confirmed as Didymellaceae. Recently, Su et al. (2012) showed A. anemones Lib. to have a Didymella sexual state and named as Didymella anemones. Since, Ascochyta pisi is the type species of Ascochyta it is confirmed as Didymellaceae. However, some Phoma species are also linked with Didymella species (Aveskamp et al. 2008; Woudenberg et al. 2013). Therefore the Ascochyta-Didymella-Phoma complex needs to be resolved.

Boerema (1997) and Boerema et al. (2004) proposed nine sections for Phoma based on morphological and physiological characters; i.e., Phoma, Heterospora, Paraphoma, Peyronellaea, Phyllostictoides, Sclerophomella, Plenodomus, Macrospora and Pilosa. de Gruyter et al. (2009) discussed the confusion regarding this classification system, and de Gruyter et al. (2009, 2010) proposed that Phoma must be restricted only to Didymellaceae. Therefore they proposed new names for all other species which group outside Didymellaceae (i.e. in Cucurbitariaceae, Coniothyriaceae, Leptosphaeriaceae and Montagnulaceae) and accepted the entire Phoma spp. group inside Didymellaceae along with P. herbarum, the type species. Ampelomyces quisqualis the type species of Ampelomyces was shown to belong in Phaeosphaeriaceae (de Gruyter et al. 2009; Wijayawardene et al. 2013) and at the same time A. quercinus grouped in Didymellaceae (de Gruyter et al. 2009). This contradiction was discussed by Aveskamp et al. (2010) and they moved all Ampelomyces spp. grouped in Didymellaceae to Phoma. Herein we do not accept Ampelomyces as Didymellaceae. Aveskamp et al. (2010) showed that Phoma exigua is a separate clade, for which they established the new genus Boeremia. Chaetasbolisia, Epicoccum, Microsphaeropsis and Peyronellaea (Phoma glomerata) was shown to belong in Didymellaceae (de Gruyter et al. 2009; Aveskamp et al. 2010). Epicoccum was confirmed to be a dimorphic genus which includes species with either hyphomycetous or “Phoma”-like synanamorphs (Aveskamp et al. 2010; Seifert et al. 2011). Peyronellaea is one of the sections of Phoma, so it is better to exclude it from the list of asexual genera of Didymellaceae. However, the taxonomic placements of Dactuliochaeta, Piggotia and Pithoascus need to be confirmed by a molecular study. Currently six genera are accepted in the family (Zhang et al. 2012a) and here we accept 13 genera. Even though this family is recognized as a separate family in Pleosporales, further studies on morphology (sexual morphs as well as their asexual morphs combined with evolutionary linkages) and molecular analysis are needed to confirm relationships.

Type: Didymella Sacc., Michelia 2(no. 6): 57 (1880), MycoBank MB1548 Fig. 37

Didymella exigua (Material examined: FRANCE, Memise sur Thollon, on Rumex arifolius (Polygonaceae), isolated by E. Müller, 2 Nov 1953, deposited by E. Müller, May 1955, CBS H-20123, culture ex-neotype CBS 183.55. Notes: the CBS Culture is sterile, but the original specimen is in Zurich) a Ascomata on host. b Surface view of ascoma. c-g Asci with ascospores. h Hyaline, ascospores. Scale bars: a–h = 10 μm

Full size image

Possible synonyms:

Arcangelia Sacc., Bull. Soc. mycol. Fr. 5: 115 (1890)

Haplotheciella Höhn., Ber. dt. bot. Ges. 36: 314 (1918)

Mycosphaerellopsis Höhn., Annls mycol. 16(1/2): 157 (1918)

Pathogens of many important crops, cortex of stems or in bracts of dead inflorescences, some putatively named species are marine fungi. Sexual state: Ascomata pseudothecia, separate or gregarious, or scattered, immersed, erumpent, or rarely superficial, brown to black, ostiole central. Peridium comprising two layers, out layer composed of 4–5 layers of blackened, thick-walled cells of textura angularis, inner layer hyaline, thin-walled. Hamathecium lacking pseudoparaphyses. Asci 8-spored, bitunicate, fissitunicate, cylindric-clavate to clavate, short-pedicellate or apedicellate, apically rounded, with an ocular chamber. Ascospores overlapping uni-seriate or bi-seriate, hyaline, ovoid to ellipsoidal, asymmetrical, upper cell wider than lower cell, 1-septate, constricted at the septum, smooth-walled. Asexual states: coelomycetous. Conidiomata pycnidial, amphigenous, separate, globose, brown, immersed, uni-locular, of thin-walled cells of textura angularis. Ostiole central, circular, slightly papillate. Conidiophores absent. Conidiogenous cells enteroblastic, phialidic, determinate, discrete, doliiform to lageniform, hyaline, smooth-walled, formed from the inner cells of the pycnidial wall. Conidia hyaline, medianly 1-septate, continuous or constricted, thin-walled, smooth-walled, ± guttulate, cylindrical to irregular (Sutton 1980).

Notes: Didymella exigua (Niessl) Sacc. is the type of the genus Didymella (Corlett 1981). Didymella contains some very important and serious plant pathogenic species, as well as species that are endophytic and saprobic on a wide range of crops (Aveskamp et al. 2010). Sivanesan (1984) mentioned that Didymella and Mycosphaerella were originally described in the Mycosphaerellaceae. Later, Didymella was placed in the Pleosporales (Pleosporaceae), Phaeosphaeriaceae (Barr 1979a; Silva-Hanlin and Hanlin 1999), Venturiaceae (Reddy et al. 1998), or considered as a genus incertae sedis (Lumbsch and Huhndorf 2010). More recently the Didymellaceae was introduced to accommodate this genus (de Gruyter et al. 2009). The herbarium material of the holotype of D. exigua was not present in M or BRNU, where the main original collections of Niessl von Mayendorf are preserved, and appears to have been lost. A neotype was designated for Didymella exigua de Gruyter et al. 2009 and is a dried culture CBS H-20123 isolated from this taxon on Rumex arifolius (Polygonaceae) France, with a living ex-neotype culture (CBS 183.55).

Type species: Didymella exigua (Niessl) Sacc., Syll. fung. (Abellini) 1: 553 (1882)

MycoBank: MB 218525

≡ Didymosphaeria exigua Niessl, Öst. bot. Z. 25(4): 165 (1875)

Other genera included

Ascochyta Lib., Pl. crypt. Arduenna, fasc. (Liège) 1(Praef.): 8 (1830)

Type species: Ascochyta pisi Lib., Pl. crypt. Arduenna, fasc. (Liège) 1: no. 59 (1830)

Boeremia Aveskamp et al., in Aveskamp et al., Stud. Mycol. 65: 36 (2010)

Type species: Boeremia exigua (Desm.) Aveskamp et al., in Aveskamp et al., Stud. Mycol. 65: 37 (2010)

Chaetasbolisia Speg., Physis, B. Aires 4: 293 (1918)

Type species: Chaetasbolisia erysiphoides Griffon & Maubl., Physis, Rev. Soc. Arg. Cienc. Nat. 4: 293 (1918)

Dactuliochaeta G.L. Hartm. & J.B. Sinclair, Mycologia 80(5): 697 (1988)

Type species: Dactuliochaeta glycines (R.B. Stewart) G.L. Hartm. & J.B. Sinclair, Mycologia 80(5): 699 (1988)

Epicoccum Link, Mag. Gesell. naturf. Freunde, Berlin 7: 32 (1816) [1815]

Type species: Epicoccum nigrum Link, Mag. Gesell. naturf. Freunde, Berlin 7: 32 (1816) [1815]

Leptosphaerulina McAlpine, Fungus Diseases of stone-fruit trees in Australia: 103 (1902).

Type species: Leptosphaerulina australis McAlpine, Fungus Diseases of stone-fruit trees in Australia: 103 (1902)

Macroventuria Aa, Persoonia 6(3): 359 (1971)

Type species: Macroventuria anomochaeta Aa, Persoonia 6(3): 362 (1971)

Microsphaeropsis Höhn., Hedwigia 59: 267 (1917)

Type species: Microsphaeropsis olivacea (Bonord.) Höhn. [as ‘olivaceus’], Hedwigia 59: 267 (1917)

Monascostroma Höhn., Annls mycol. 16(1/2): 160 (1918)

Type species: Monascostroma innumerosum (Desm.) Höhn. [as ‘innumerosa’], Annls mycol. 16(1/2): 160 (1918)

Phoma Sacc., Michelia 2(no. 6): 4 (1880)

Type species: Phoma herbarum Westend., Bull. Acad. R. Sci. Belg., Cl. Sci., sér. 2 19(3): 118 (1852)

Piggotia Berk. & Broome, Ann. Mag. nat. Hist., Ser. 2 7: 95 (1851)

Type species: Piggotia astroidea Berk. & Broome, Ann. Mag. nat. Hist., Ser. 2 7: no. 503 (1851)

Pithomyces Berk. & Broome, J. Linn. Soc., Bot. 14(no. 74): 100 (1873) [1875]

Type species: Pithomyces flavus Berk. & Broome, J. Linn. Soc., Bot. 14(no. 74): 100 (1873) [1875]

Key to sexual genera of Didymellaceae

1. Ascospores muriform ..............................Leptosphaerulina

1. Ascospores 2-celled, rarely 3- to 4-celled .........................2

2. Ascospores constricted at the central septum ....................3

2. Ascospores septate near the lower end ..............Platychora

3. Ascospores narrowly inequilateral ............Monascostroma

3. Ascospores ovoid to ellipsoidal ..........................................4

4. Ascomata with hairs or setae ......................Macroventuria

4. Ascomata without hairs or setae .........................Didymella

Key to asexual genera of Didymellaceae

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

1. Hyphomycetous asexual states ...........................................8

2. Pycnidia having dark brown setae, verrucose, conidia ellipsoid .........................................................................................3

2. Pycnidia lacking setae ........................................................4

3. Conidiophores hyaline to sub hyaline, ampulliform to lageniform .......................................................Dactuliochaeta

3. Conidiophores hyaline, doliiform to ampulliform ..........................................................................Chaetasbolisia

4. Conidiophores present ...........................................Piggotia

4. Conidiophores absent .........................................................5

5. Conidia brown, globose, pyriform, cylindrical or ellipsoid .....................................................................Microsphaeropsis

5. Conidia hyaline ..................................................................6

6. Conidiogenous cells doliiform to lageniform, hyaline, conidia hyaline, cylindrical to irregular ......................Ascochyta

6. Conidiogenous cells ampulliform to doliiform ..................7

7. Conidia ellipsoid, cylindrical fusiform, pyriform or globose, often guttulate ...............................................................Phoma

7. Conidia irregular shape, large conidia (15 × 5 μm),1(−2)-septate ......................................................................Boeremia

8. Dark sporodochia, with branched (sub) hyaline conidiophores ....................................................................Epicoccum

8. Conidiophores brown, denticles .......................Pithomyces

Didymosphaeriaceae Munk, Dansk bot. Ark. 15(no. 2): 128 (1953), MycoBank: MB 80702

Terrestrial, saprobic or parasitic on woody branches and herbaceous stems and leaves, also parasitic on other fungi. Sexual state: Ascomata solitary, scattered, or in small groups, immersed to erumpent or superficial, globose to ovoid, ostiolate. Ostiole papillate, ostiolar canal filled with hyaline cells (periphyses). Peridium thin, composed of brown, thick-walled pseudoparenchymatous cells. Hamathecium of narrow, dense, trabeculate pseudoparaphyses, anastomosing above the asci, in a gelatinous matrix. Asci 2–4-spored or 8-spored, bitunicate, fissitunicate, cylindrical, pedicellate, with a small ocular chamber. Ascospores uni-seriate or partially overlapping, ellipsoid or oblong, brown, 1-septate. Asexual states: “Fusicladium”-like and “Phoma”-like

Notes: Based on the distoseptate ascospores and trabeculate pseudoparaphyses, mainly anastomosing above the asci, Munk (1953) introduced a new family Didymosphaeriaceae, typified by the genus Didymosphaeria, in the order Pleosporales. Barr (1990c) included this family in the order Melanommatales based on its trabeculate pseudoparaphyses. Familial placement of the Didymosphaeriaceae is still uncertain because several authors refer the family to different higher taxa; i.e., von Arx and Müller (1975) treated Didymosphaeriaceae as a synonym of the Pleosporaceae, Lumbsch and Huhndorf (2007) assigned it to the Montagnulaceae, while Zhang et al. (2012a) tentatively included Didymosphaeriaceae as a separate family in the Pleosporales under the suborder Pleosporineae. In the same study it was shown that Didymosphaeria futilis (Berk. & Broome) Rehm is closely related to the Cucurbitariaceae (Zhang et al. 2012a, b). Barr (1990c) included Neotestudina under this family by considering the morphology of ascospores. Kohlmeyer and Volkmann-Kohlmeyer (1990) assigned Coronopapilla under Didymosphaeriaceae, while Lojkania enalia (Kohlm.) M.E. Barr (= Didymosphaeria enalia Kohlm.) was also included in the family, due to the same character. Zhang et al. (2012a) accepted Appendispora, Didymosphaeria and Phaeodothis in the family. Correct placement of this family depends on further molecular and morphological data based with correctly identified and verified strains.

Sivanesan (1984) stated that Didymosphaeria has Ascochyta and Periconia asexual states, while Keissleriella has Dendrophoma asexual states. However, Schoch et al. (2009b) and Zhang et al. (2012a) showed that Keissleriella belongs to Lentitheciaceae. Crous et al. (2013) showed that Dendrophoma cytisporoides Sacc., the type of Dendrophoma groups with Chaetosphaeria hebetiseta Réblová & W. Gams and C. callimorpha (Mont.) Sacc., hence we exclude Dendrophoma from Didymosphaeriaceae. Ellis (1971) also reported that Didymosphaeria has asexual states in Periconia, but according to Seifert et al. (2011), this record was a misidentification of Lophiostoma as Didymosphaeria. Placing Ascochyta in Didymosphaeriaceae also met with some contradictions, as Ascochyta pisi Lib., the type was shown to belong in Didymellaceae (de Gruyter et al. 2009). Wijayawardene et al. (2012) stated that Roussoëlla has a Cytoplea asexual state and this was phylogenetically linked by Kang et al. (1998) and in culture (Hyde et al. 1996). However the phylogenetic placement of Roussoëlla is also uncertain (Zhang et al. 2012a) as in their study it grouped in Arthopyreniaceae while in this study is groups near Bioatriosporaceae and is accommodated in a new family Roussoellaceae (Liu et al. 2013).

Type: Didymosphaeria Fuckel, Jb. nassau. Ver. Naturk. 23–24: 140 (1870) [1869–70], MycoBank: MB 1562 Fig. 38

Didymosphaeria futilis (Material examined: ITALY, Forlì-Cesena, Ravaldino, on dead cane of Rubus ulmifolius, 14 January 2012, E. Camporesi, MFLU12-2215). a Immersed ascomata on the host surface. b Close up of the ascomata. c Section of an ascoma. d Peridium. e Trabeculate, anastomosing and branching pseudoparaphyses. f Eight spored asci with pedicels which are rounded at their bases. g Brown, 1-septate ascospores with spinulose ornamentation. Scale bars: c = 200 μm, d = 20 μm, e = 10 μm, f–h = 30 μm, i–l = 10 μm

Full size image

Terrestrial, saprobic or parasitic on woody branches and herbaceous stems and leaves, also parasitic on other fungi. Sexual state: Ascomata solitary, scattered, or in small groups, immersed to erumpent, globose to ovoid, ostiolate. Ostiole papillate with a pore-like opening, ostiolar canal filled with hyaline cells (periphyses). Peridium 1-layered, thin, composed of brown pseudoparenchymatous cells of textura angularis. Hamathecium of dense, trabeculate, pseudoparaphyses, anastomosing mostly above the asci. Asci 2–4-spored, or 8-spored, bitunicate, fissitunicate, cylindrical, with a furcated pedicel, apically rounded with an indistinct ocular chamber. Ascospores uni-seriate, ellipsoid, brown, 1-septate, wall smooth or roughed. Asexual state: see notes below.

Notes: Didymosphaeria was introduced by Fuckel (1870) for three species with two-celled ascospores. This genus now comprises species having a wide distribution and a broad host range. Saccardo (1882) restricted species only with brown spores to the genus (Aptroot 1995a). More than 100 species have also been excluded from this genus (Barr 1989a, b, 1990b, 1992a, b, 1993a, b; Hawksworth 1985a, b; Hawksworth and Boise 1985; Hawksworth and Diederich 1988; Scheinpflug 1958), while Aptroot (1995b) included over 400 epithets of Didymosphaeria in his monograph of the genus after examining over 3,000 species. Hawksworth and David (1989) proposed to conserve the genus with a lectotype specimen, in Fungi Rhenani 1770 wchich we follow here, while Didymosphaeria epidermidis (Fr.) Fuckel was chosen as the lectotype species by Aptroot (1995b). In his monograph Aptroot (1995b) listed seven species which were closely related with the type of Didymosphaeria without considering differences of host or country of origin. Many taxa were excluded from the Didymosphaeriaceae: Aaosphaeria, Amphisphaeria, Astrosphaeriella, Dothidotthia, Flagellosphaeria, Kirschsteiniothelia, Megalotremis, Montagnula, Munkovalsaria, Mycomicrothelia, Parapyrenis or Phaeodothis. Aptroot (1995a) concluded that Didymosphaeria belonged in the family Didymosphaeriaceae (Pleosporales) based on a peridium consisting of flattened or irregular cells, a hamathecium consisting of narrow, trabeculate pseudoparaphyses, richly anastomosing above the asci; and brown, thinly distoseptate ascospores. Currently there are around 500 species epithets listed in Index Fungorum (2013), while GenBank has 21 hits for the genus including putative strains of Didymosphaeria futilis (HKUCC 5834) and Verruculina enalia (Kohlm.) Kohlm. & Volkm.-Kohlm (AFTOL-ID 1601) (a genus now referred to the Testudinaceae, Schoch et al. 2006). Didymosphaeria has been accepted as a well-established genus represented by D. futilis. However few molecular investigations of Didymosphaeria have been undertaken when compared to the morphological studies.

Type species: Didymosphaeria futilis (Berk. & Broome) Rehm, Hedwigia 18: 167 (1879), MycoBank: MB 223613

≡ Sphaeria futilis Berk. & Broome, Ann. Mag. nat. Hist., Ser. 2 9: 326 (1852)

Other genera included

Appendispora K.D. Hyde, Anal. Soc. cient. argent. 46(1): 29 (1994)

Type: Appendispora frondicola K.D. Hyde, Sydowia 46(1): 30 (1994)

Phaeodothis Syd. & P. Syd., Annls mycol. 2(2): 166 (1904)

Type: Phaeodothis tricuspidis Syd. & P. Syd., Annls mycol. 2(2): 166 (1904)

Key to genera of Didymosphaeriaceae

1. Hamathecium consisting of narrow, trabeculate pseudoparaphyses ..................................................................2

1. Hamathecium consisting of thin filaments, often sparse pseudoparaphyses ...............................................Phaeodothis

2. Asci with faint ring and short pedicels .........Appendispora

2. Cylindrical asci with a furcate pedicel ......Didymosphaeria

Dissoconiaceae Crous & de Hoog, in Crous et al., Stud. Mycol. 64: 36 (2009), MycoBank: MB 514699

Mycoparasitic on powdery mildew, pathogens on Eucalyptus species. Sexual state: Ascomata pseudothecial, immersed, globose, uni-locular, papillate, ostiolate, canal periphysate. Peridium consisting of 3–4 layers of brown textura angularis; inner layer of fattened, hyaline cells. Pseudoparaphyses absent. Asci 8-spored, bitunicate. Ascospores fasciculate, ellipsoid-fusoid, 1-septate, hyaline, with or without mucoid sheath. Asexual state: Mycelium internal and external, consisting of branched, septate, smooth-walled, hyaline to pale brown hyphae. Conidiophores separate, arising from hyphae, subcylindrical, subulate or lageniform to cylindrical, tapering to a bluntly rounded or truncate apex, straight to once geniculate, smooth-walled, medium brown to brown, multi-septate; loci terminal and lateral, visible as slightly thickened, darkened scars on a rachis. Conidiogenous cells terminal or lateral, integrated, tapering towards an acutely rounded apex, pale to brown, subcylindrical, proliferating sympodially, forming a rachis with slightly thickened and darkened, circular. Conidia solitary, pale olivaceous-brown, smooth-walled, ellipsoid to obclavate or globose, 0–1-septate; hila somewhat darkened. Secondary conidia present or absent; developing adjacent to primary conidia, pale olivaceous to subhyaline, aseptate, pyriform; conidium discharge active or passive. Dissoconium has secondary conidia but Ramichloridium does not produce secondary conidia (Crous et al. 2009b; Li et al. 2012).

Notes: The asexual typified family Dissoconiaceae (Capnodiales), was introduced by Crous et al. (2009b). The type is Dissoconium aciculare de Hoog et al. and species of Dissoconium have “Mycosphaerella”-like asexual states (Crous et al. 2004). Crous et al. (2004) showed that Dissoconium aciculare clusters with Mycosphaerella communis Crous & Mansilla, indicating Mycosphaerella as the sexual state. However, Crous et al. (2009b) and Li et al. (2012) clearly showed that Dissoconium aciculare, D. australiensis Crous & Summerell, D. commune Crous & Mansilla, Ramichloridium apiculatum (J.H. Mill. et al.) de Hoog, Uwebraunia commune (J.H. Mill. et al.) de Hoog and U. dekkeri (de Hoog & Hijwegen) Crous group together, but separately as the sister clade of Mycosphaerellaceae. The characteristic feature of the genus is conidia which are formed in pairs that are forcefully discharged, which is quite unique in the Capnodiales (de Hoog et al. 1983). Li et al. (2012) accepted Dissoconium, Pseudoveronaea, Ramichloridium and Uwebraunia in Dissoconiaceae. There is evidence that some species such as U. dekkeri (= D. dekkeri de Hoog & Hijwegen) could be plant pathogenic (Jackson et al. 2004), while some species occur as commensalists, associated with pathogenic species of Capnodiales. The type species of Ramichloridium, R. apiculatum (J.H. Mill. et al.) de Hoog, is placed in Dissoconiaceae, while R. musae Stahel (EU041857), R. australiense Arzanlou & Crous (EU041852), R. strelitziae Arzanlou et al. (EU041860) and R. cerophilum (Tubaki) de Hoog (GU214485) group in Mycosphaerellaceae.