Fungal Diversity

DOI 10.1007/s13225-016-0369-6

Families of Sordariomycetes
Sajeewa S. N. Maharachchikumbura 1,2,3 & Kevin D. Hyde 2 & E. B. Gareth Jones 4 &
E. H. C. McKenzie 5 & Jayarama D. Bhat 6,7 & Monika C. Dayarathne 2 & Shi-Ke Huang 2,8 &
Chada Norphanphoun 2 & Indunil C. Senanayake 2 & Rekhani H. Perera 1,2 & Qiu-Ju Shang 2 &
Yuanpin Xiao 2,8 & Melvina J. D’souza 2 & Sinang Hongsanan 2 & Ruvishika S. Jayawardena 2,9 &
Dinushani A. Daranagama 2,10 & Sirinapa Konta 2 & Ishani D. Goonasekara 2 & Wen-Ying Zhuang 10 &
Rajesh Jeewon 11 & Alan J. L. Phillips 12 & Mohamed A. Abdel-Wahab 4,13 & Abdullah M. Al-Sadi 3 &
Ali H. Bahkali 4 & Saranyaphat Boonmee 2 & Nattawut Boonyuen 14 & Ratchadawan Cheewangkoon 15 &
Asha J. Dissanayake 2,9 & Jichuan Kang 8 & Qi-Rui Li 8,16 & Jian Kui Liu 1 & Xing Zhong Liu 10 &
Zuo-Yi Liu 1 & J. Jennifer Luangsa-ard 17 & Ka-Lai Pang 18 & Rungtiwa Phookamsak 2 &
Itthayakorn Promputtha 19 & Satinee Suetrong 14 & Marc Stadler 20,21 &
Tingchi Wen 8 & Nalin N. Wijayawardene 2

Received: 13 February 2016 / Accepted: 14 April 2016

# School of Science 2016

Abstract Sordariomycetes is one of the largest classes of pathogens, as well as endophytes, saprobes, epiphytes, coproph-
Ascomycota that comprises a highly diverse range of fungi ilous and fungicolous, lichenized or lichenicolous taxa. They
characterized mainly by perithecial ascomata and inoperculate occur in terrestrial, freshwater and marine habitats worldwide.
unitunicate asci. The class includes many important plant This paper reviews the 107 families of the class

Electronic supplementary material The online version of this article

(doi:10.1007/s13225-016-0369-6) contains supplementary material,
which is available to authorized users.

* Zuo-Yi Liu 8
Engineering Research Center of Southwest Bio-Pharmaceutical
gzliuzuoyi@163.com Resources, Ministry of Education, Guizhou University,
Guiyang 550025, Guizhou Province, China
9
1 Institute of Plant and Environment Protection, Beijing Academy of
Guizhou Key Laboratory of Agricultural Biotechnology, Guizhou Agriculture and Forestry Sciences, No 9 of
Academy of Agricultural Sciences, Guiyang 550006, Guizhou, ShuGuangHuaYuanZhangLu, Haidian District Beijing 100097,
China China
2
Center of Excellence in Fungal Research, and School of Science, 10
State Key Laboratory of Mycology, Institute of Microbiology,
Mae Fah Luang University, Chiang Rai 57100, Thailand Chinese Academy of Sciences, Beijing 100101, China
3
Department of Crop Sciences, College of Agricultural and Marine 11
Department of Health Sciences, Faculty of Science, University of
Sciences, Sultan Qaboos University, P.O. Box 34, Al-Khod 123, Mauritius, Reduit, Mauritius
Oman
12
4 Faculty of Sciences, Biosystems and Integrative Sciences Institute
Department of Botany and Microbiology, College of Science, King (BioISI), University of Lisbon, Campo Grande,
Saud University, P.O. Box 2455, Riyadh 11451, Kingdom of Saudi 1749-016 Lisbon, Portugal
Arabia
13
5 Department of Botany, Faculty of Science, Sohag University,
Manaaki Whenua Landcare Research, Private Bag 92170, Sohag, Egypt
Auckland, New Zealand
14
6 Fungal Biodiversity Laboratory, BIOTEC, National Science and
No. 128/1-J, Azad Housing Society, Curca, P.O. Goa Velha, Technology Development Agency (NSTDA), 113 Thailand Science
403108 Goa, India Park, Thanon Phahonyothin, Tombon Khlong Nueng, Amphoe
7 Khlong Luang, Pathum Thani 12120, Thailand
Department of Botany, Goa University, Goa 403 206, India
 Fungal Diversity

Sordariomycetes and provides a modified backbone tree based 8. Batistiaceae Samuels & K.F. Rodrigues (Q Shang* &
on phylogenetic analysis of four combined loci, with a maxi- EBG Jones)
mum five representative taxa from each family, where available. 9. Beltraniaceae Nann. (SSN Maharachchikumbura*)
This paper brings together for the first time, since Barrs’ 1990 10. Bertiaceae Smyk (S-K Huang*)
Prodromus, descriptions, notes on the history, and plates or il- 11. Bionectriaceae Samuels & Rossman (RH Perera*)
lustrations of type or representative taxa of each family, a list of 12. Boliniaceae Rick (S-K Huang*)
accepted genera, including asexual genera and a key to these 13. Cainiaceae J.C. Krug (ID Goonasekara*)
taxa of Sordariomycetes. Delineation of taxa is supported where 14. Calosphaeriaceae Munk (S-K Huang* & EBG Jones)
possible by molecular data. The outline is based on literature to 15. Catabotrydaceae Petr. ex M.E. Barr (RH Perera*)
the end of 2015 and the Sordariomycetes now comprises six 16. Cephalothecaceae Höhn. (Q Shang*)
subclasses, 32 orders, 105 families and 1331 genera. The family 17. Ceratocystidaceae Locq. ex Réblová et al. (C
Obryzaceae and Pleurotremataceae are excluded from the class. Norphanphoun*)
18. Ceratostomataceae G. Winter (C Norphanphoun* &
EBG Jones)
Keywords Amplistromatales . Annulatascales . Boliniales .
19. Chadefaudiellaceae Faurel et al. (M Dayarathne*)
Calosphaeriales . Chaetosphaeriales . Coniochaetales .
20. Chaetomiaceae G. Winter (Q Shang*)
Conioscyphales . Cordanales . Coronophorales .
21. Chaetosphaerellaceae Huhndorf et al. (S-K Huang* &
Diaporthales . Falcocladiales . Glomerellales . Hypocreales .
EBG Jones)
Jobellisiales . Koralionastetales . Lulworthiales .
22. Chaetosphaeriaceae Réblová et al. (S-K Huang* &
Magnaporthales . Melanosporales . Meliolales .
EBG Jones)
Microascales . Ophiostomatales . Phylogeny .
23. Clavicipitaceae (Lindau) Earle ex Rogerson (Y Xiao*)
Phyllachorales . Pisorisporiales . Pleurotheciales .
24. Clypeosphaeriaceae G. Winter (S Konta*)
Sordariales . Savoryellales . Tirisporellales . Togniniales .
25. Coniocessiaceae Asgari & Zare (C Norphanphoun* &
Torpedosporales . Trichosphaeriales . Taxonomy .
EBG Jones)
Trichosphaeriales . Type species . Xylariales
26. Coniochaetaceae Malloch & Cain (Q Shang*)
Contents 27. Conioscyphaceae Réblová & Seifert#
1. Amphisphaeriaceae G. Winter (ID Goonasekara*) 28. Cordanaceae Nann. (ID Goonasekara*)
2. Amplistromataceae Huhndorf et al. (DA Daranagama*) 29. Cordycipitaceae Kreisel (Y Xiao*)
3. Annulatascaceae S.W. Wong et al. (MJ D’souza*) 30. Coronophoraceae Höhn. (S-K Huang*)
4. Apiosporaceae K.D. Hyde et al. (RH Perera* & EBG 31. Cryphonectriaceae Gryzenh. & M.J. Wingf. (IC
Jones) Senanayake*)
5. Armatellaceae Hosag. (S Hongsanan*) 32. Diaporthaceae Höhn. ex Wehm. (IC Senanayake*)
6. Australiascaceae Réblová & W. Gams (RS 33. Diatrypaceae Nitschke (Q Shang)
Jayawardena & Q Shang*) 34. Distoseptisporaceae K.D. Hyde & McKenzie#
7. Bartaliniaceae Wijayawardene et al. (ID Goonasekara*) 35. Etheirophoraceae Rungjindamai et al. (M Dayarathne
& EBG Jones*)
15
Department of Plant Pathology, Faculty of Agriculture, Chiang Mai 36. Falcocladiaceae Somrith. et al. (M Dayarathne*)
University, Chiang Mai 50200, Thailand 37. Flammocladiaceae Crous et al. (SSN
16
Department of Pharmacy, Guiyang Medical University, Maharachchikumbura*)
Guiyang, Guizhou 550025, China 38. Glomerellaceae Locq. ex Seifert & W. Gams (RS
17
Microbe Interaction Laboratory, BIOTEC, National Science and Jayawardena*)
Technology Development Agency (NSTDA), 113 Thailand Science 39. Gnomoniaceae G. Winter (IC Senanayake* & EBG
Park, Thanon Phahonyothin, Tombon Khlong Nueng, Amphoe
Khlong Luang, Pathum Thani 12120, Thailand
Jones)
18
4 0 . G o n d w a n a m y c e t a c ea e R é b l o v á e t a l . ( M
Institute of Marine Biology and Center of Excellence for the Oceans,
National Taiwan Ocean University, 2 Pei-Ning Road,
Dayarathne*)
Keelung 20224, Taiwan, Republic of China 41. Graphiaceae Z.W. de Beer (M Dayarathne*)
19
Department of Biology, Faculty of Science, Chiang Mai University,
42. Halosphaeriaceae E. Müll & Arx ex Kohlm (M
Chiang Mai 50200, Thailand Dayarathne* & EBG Jones*)
20
Department Microbial Drugs, Helmholtz-Zentrum für
43. Harknessiaceae Crous (IC Senanayake*)
Infektionsforschung GmbH, Inhoffenstrasse 7, 44. Helminthosphaeriaceae Samuels et al. (S-K Huang*)
38124 Braunschweig, Germany 45. Hispidicarpomycetaceae Nakagiri (M Dayarathne &
21
German Centre for Infection Research (DZIF), Partner Site EBG Jones*)
Hannover-Braunschweig, 38124 Braunschweig, Germany 46. Hypocreaceae De Not (RH Perera*)
Fungal Diversity

47. Hyponectriaceae Petr. (S Konta* & EBG Jones) 85. Reticulascaceae Réblová & W. Gams (RS
48. Iodosphaeriaceae O. Hilber (Q Li & KD Hyde*) Jayawardena & Q Shang*)
49. Jobellisiaceae Réblová (MJ D’souza*) 86. Robillardaceae Crous (SSN Maharachchikumbura*)
50. Juncigenaceae E.B.G. Jones et al. (M Dayarathne* & 87. Savoryellaceae Jaklitsch & Réblová (M Dayarathne*)
EBG Jones) 88. Schizoparmaceae Rossman (IC Senanayake*)
5 1 . K a t h i s t a c e a e M a l l o c h & M . B l a c k w. ( C 89. Scortechiniaceae Huhndorf et al. (S-K Huang*)
Norphanphoun) 90. Sordariaceae G. Winter (S-K Huang*)
52. Koralionastetaceae Kohlm. & Volkm.-Kohlm. (M 91. Spathulosporaceae Kohlm. (M Dayarathne*)
Dayarathne* & EBG Jones) 92. Sporocadaceae Corda (ID Goonasekara*)
53. Lasiosphaeriaceae Nannf. (S-K Huang*) 93. Sporidesmiaceae Fr. #
54. Lautosporaceae Kohlm. et al. (M Dayarathne*) 94. Stachybotryaceae L. Lombard & Crous (C
55. Lopadostomataceae Daranagama & K.D. Hyde (DA Norphanphoun*)
Daranagama* & KD Hyde) 95. Stilbosporaceae Link (IC Senanayake*)
56. Lulworthiaceae Kohlm. et al. (M Dayarathne* & EBG 96. Sydowiellaceae Lar.N. Vassiljeva (IC Senanayake*)
Jones) 97. Thyridiaceae O.E. Erikss & J.Z. Yue (RH Perera*)
57. Macrohilaceae Crous (IC Senanayake*) 98. Tilachlidiaceae L. Lombard & Crous (RH Perera)
58. Magnaporthaceae P.F. Cannon (C Norphanphoun* & 99. Tirisporellaceae Suetrong et al. (EBG Jones* & M
EBG Jones) Dayarathne)
59. Melanconidaceae G. Winter Chin (IC Senanayake* & 100. Togniniaceae Réblová et al. (IC Senanayake* & EBG
EBG Jones) Jones)
60. Meliolaceae G.W. Martin ex Hansf. (S Hongsanan*) 101. Torpedosporaceae E.B.G. Jones & K.L. Pang (M
61. Melogrammataceae G. Winter (Q Shang*) Dayarathne, KL Pang, EBG Jones*)
62. Microascaceae Luttr. ex Malloch (M Dayarathne*) 102. Trichosphaeriaceae G. Winter (C Norphanphoun*&
6 3 . M i c r o d o c h i a c e a e M . H e r n . - R e s t r. ( S S N EBG Jones)
Maharachchikumbura*) 103. Valsaceae Tul. & C. Tul. (IC Senanayake*)
64. Myelospermataceae K.D. Hyde & S.W. Wong (S. 104. Vialaeaceae P.F. Cannon (IC Senanayake*& EBG
Konta*) Jones)
65. Nectriaceae Tul. & C. Tul. (RH Perera*) 105. Xylariaceae Tul. & C. Tul. (DA Daranagama*)
66. Niessliaceae Kirschst. (S-K Huang* & EBG Jones) Excluded, doubtful, poorly known, or family previously
67. Nitschkiaceae Nannf. (S-K Huang* & EBG Jones) included in Sordariomycetes
68. Ophioceraceae Klaubauf et al. (C Norphanphoun*) 106. Obryzaceae Körb (RH Perera*)
69. Ophiocordycipitaceae G.H. Sung et al. (Y Xiao*) 107. Pleurotremataceae Walt. Watson (S-K Huang*)
70. Ophiostomataceae Nannf. (C Norphanphoun*) *These are the main contributing authors to the family
71. Papulosaceae Winka & O.E. Erikss. (MJ D’souza*) entry
#
72. Pestalotiopsidaceae Maharachch. & K.D. Hyde (SSN These were introduced in 2016 and thus no note are
Maharachchikumbura*) provided
73. Phaeochoraceae K.D. Hyde et al. (RH Perera*)
74. Phlogicylindriaceae Senan. & K.D. Hyde (ID
Goonasekara*) Introduction
75. Phyllachoraceae Theiss. & H. Syd. (M Dayarathne* &
EBG Jones) Sordariomycetes is the second largest class of Ascomycota
76. Pisorisporiaceae Réblová & J. Fourn. (MJ D’souza*) (Kirk et al. 2008; Hyde et al. 2013). Kirk et al. (2008) included
77. Plectosphaerellaceae W. Gams et al. (Q Shang* & 15 orders, 64 families, 1119 genera and 10,564 species in
EBG Jones) Sordariomycetes, while Lumbsch and Huhndorf (2010) in-
78. Pleurostomataceae Réblová et al. (S-K Huang*) cluded 18 orders, 63 families and 947 genera. The
79. Pleurotheciaceae Réblová & Seifert# Sordariomycetes have a cosmopolitan distribution and accom-
80. Pseudomassariaceae Senan. & K.D. Hyde (IC modates mostly terrestrial taxa, although several can be found
Senanayake*) in aquatic habitats (Hyde and Wong 2000; Samuels and
81. Pseudoplagiostomataceae Cheew. et al. (IC Blackwell 2001; Cai et al. 2006a; Jones et al. 2009a, b,
Senanayake*) 2015; Pratibha et al. 2014). Some are phytopathogens that
82. Pseudovalsaceae M.E. Barr (IC Senanayake*) cause leaf, stem and root diseases on a wide variety of hosts,
83. Pyriculariaceae Klaubauf et al. (C Norphanphoun*) while some cause diseases in arthropods and mammals (Sung
84. Requienellaceae Boise# et al. 2007; Prados-Rosale et al. 2012; Hyde et al. 2014).
 Fungal Diversity

Sordariomycetes are also commonly isolated as endophytes uncertain placement in Sordariomycetes was provided. In this
from various plants (Keim et al. 2014). Some taxa are paper, we deal with the family level classification of
fungicolous (PeiGui et al. 2000), while many persist as Sordariomycetes.
saprobes involved in decomposition and nutrient cycling
(Jaklitsch and Voglmayr 2012). Some species of
Sordariomycetes (i.e. Beauveria bassiana, Trichoderma Materials and methods
viride, T. harzianum) are economically important biocontrol
agents (Wraight et al. 1998; Kaewchai et al. 2009) and others Layout of the paper
produce a wide range of chemically diverse metabolites im-
portant in medicinal and other biotechnological industries Each family is treated with a family description, notes on its
(Semenova et al. 2012; Debbab et al. 2013; Xu et al. 2014). history and presence of any asexual morphs, and an account of
A large number of Sordariomycetes species are character- the genera, including phylogenetic data where applicable. The
ized by non-lichenized, perithecial ascomata and inoperculate type is listed along with a description of the type genus except
unitunicate asci (Zhang et al. 2006) or non-fissitunicate asci in cases where there is only a single genus in the family. Notes
(Kirk et al. 2008). Most members of the Xylariomycetidae on the type genus are provided along with full citations for the
and some of the Sordariomycetidae have dark perithecia, amy- type species. This is followed by accepted genera and their
loid asci, true paraphyses and periphysate ostioles, while most type species; important synonyms and occasional notes are
of the taxa of the Hypocreomycetidae have light coloured peri- given. Synonymies follow the basis of one name for a biolog-
thecia, non-amyloid apical rings in the asci when apical rings ical genus or species (see Hawksworth et al. 2011;
are present and the lack true paraphyses (Zhang et al. 2006). Hawksworth 2012) and follow published papers or Index
The class Sordariomycetes was introduced by Eriksson and Fungorum (2016). A key to the genera included in the family
Winka (1997) for taxa with perithecial ascomata, paraphysate is provided unless the number of genera is small. The type
hamathecium, periphysate ostioles and unitunicate or genus is illustrated with a representative plate of its key mor-
pseudoprotunicate asci. With the support of sequence data, phological features. For consistency, authorities and their ab-
Eriksson and Winka (1997) divided the class into three sub- breviations for all taxa and citations of place of publication
classes: Hypocreomycetidae, Sordariomycetidae and and abbreviations follow Index Fungorum (2016), even
Xylariomycetidae. The subclass Sordariomycetidae included though those are not always entirely consistent.
three orders (i.e. Diaporthales, Ophiostomatales and
Sordariales), while Xylariomycetidae was introduced to accom- Arrangement of sordariomycetes
modate a single order Xylariales (Eriksson and Winka 1997).
Eriksson and Winka (1997) separated Onygenales and The circumscription of the Sordariomycetes is based on the
Eurotiales from Sordariomycetes and placed them in the class previous arrangements given in Maharachchikumbura et al.
Eurotiomycetes based on morphology and phylogenetic (2015), which has been updated based on a consideration of
analyses. A comprehensive study of Sordariomycetes based recent publications and interpretation of genera from the liter-
on both morphological characters and SSU rDNA sequence ature (see Table 1). We consider this a working circumscrip-
data was carried out by Samuels and Blackwell (2001) and tion that will be further refined as more genera are studied both
Eriksson (2006). In the classification of Eriksson (2006), the at the morphological and molecular levels. Basionyms and
subclass Hypocreomycetidae comprised 4 orders (i.e. possible synonyms of genera and species are not listed as
Coronophorales, Halosphaeriales, Hypocreales and these can be found in Index Fungorum (2016).
Microascales). Coronophorales, Halosphaeriales and
Hypocreales are recognized as monophyletic and Examination of specimens
Microascales as paraphyletic. Samuels and Blackwell (2001)
excluded Erysiphales and Coryneliales sensu Barr (1990) from Specimens or slides were obtained from ANM, AR, B, BBH,
Sordariomycetes, while Eriksson (2006) placed Erysiphales in BCC, BISH, BP, BPI, BR, BRIP, CBS, DAOM, FH, G, GZU,
the class Leotiomycetes, a sister taxon of the Sordariomycetes. H, HAL, HHUF, HKU (M), IFRD, ILLS, IMI, K, L, LPS,
Eriksson (2006) also recognized Melanosporales as a distinct MFU, NY, P, PC, PREM, S, TRTC, UBC, UC, UCR, UPS,
order in the Hypocreomycetidae. In a recent revision URM, URM and W. Fruiting bodies were rehydrated in water
Maharachchikumbura et al. (2015) introduced three sub classes; and/or 5 % KOH prior to examination and sectioning. Hand
Diaporthomycetidae, Lulworthiomycetidae, and sections of the fruiting bodies were mounted in water for mi-
Meliolomycetidae based on morphology and sequence data. croscopic studies and photomicrography. The materials were
The recent Outlin e of the Sordariomycetes examined by a Nikon ECLIPSE 80i compound microscope
(Maharachchikumbura et al. 2015) included 28 orders, 90 and photographed by Canon 450D digital camera fitted to the
families and 1344 genera. In addition, a list of 829 genera with microscope. Measurements were made with the Tarosoft (R)
Fungal Diversity

Table 1 Sordariomycetes orders and included families Table 1 (continued)

Order Family Order Family

Amplistromatales Amplistromataceae Pyriculariaceae

Catabotrydaceae Melanosporales Ceratostomataceae
Annulatascales Annulatascaceae Meliolales Armatellaceae
Boliniales Boliniaceae Meliolaceae
Calosphaeriales Calosphaeriaceae Microascales Chadefaudiellaceae
Pleurostomataceae Ceratocystidaceae
Chaetosphaeriales Chaetosphaeriaceae Gondwanamycetaceae
Helminthosphaeriaceae Graphiaceae
Pleurotremataceae Halosphaeriaceae
Coniochaetales Coniochaetaceae Microascaceae
Conioscyphales Conioscyphaceae Ophiostomatales Kathistaceae
Cordanales Cordanaceae Ophiostomataceae
Coronophorales Bertiaceae Phyllachorales Phaeochoraceae
Chaetosphaerellaceae Phyllachoraceae
Coronophoraceae Pisorisporiales Pisorisporiaceae
Nitschkiaceae Pleurotheciales Pleurotheciaceae
Scortechiniaceae Savoryellales Savoryellaceae
Diaporthales Cryphonectriaceae Sordariales Chaetomiaceae
Diaporthaceae Lasiosphaeriaceae
Gnomoniaceae Sordariaceae
Harknessiaceae Spathulosporales Hispidicarpomycetaceae
Macrohilaceae Spathulosporaceae
Melanconidaceae Tirisporellales Tirisporellaceae
Pseudoplagiostomataceae Togniniales Togniniaceae
Pseudovalsaceae Torpedosporales Etheirophoraceae
Schizoparmaceae Juncigenaceae
Stilbosporaceae Torpedosporaceae
Sydowiellaceae Trichosphaeriales Trichosphaeriaceae
Valsaceae Xylariales Amphisphaeriaceae
Falcocladiales Falcocladiaceae Apiosporaceae
Glomerellales Australiascaceae Bartaliniaceae
Glomerellaceae Beltraniaceae
Reticulascaceae Cainiaceae
Plectosphaerellaceae Clypeosphaeriaceae
Hypocreales Bionectriaceae Coniocessiaceae
Clavicipitaceae Diatrypaceae
Cordycipitaceae Hyponectriaceae
Flammocladiaceae Iodosphaeriaceae
Hypocreaceae Lopadostomaceae
Nectriaceae Melogrammataceae
Niessliaceae Microdochiaceae
Ophiocordycipitaceae Myelospermataceae
Stachybotriaceae Pestalotiopsidaceae
Tilachlidiaceae Phlogicylindriaceae
Jobellisiales Jobellisiaceae Pseudomassariaceae
Koralionastetales Koralionastetaceae Requienellaceae
Lulworthiales Lulworthiaceae Robillardaceae
Magnaporthales Magnaporthaceae Sporocadaceae
Ophioceraceae Vialaeaceae
 Fungal Diversity

Table 1 (continued) among suboptimal trees from each run by comparing

Order Family likelihood scores under the GTR+GAMMA substitution
model. The resulting trees were printed with FigTree v.
Xylariaceae 1.4.0 (http://tree.bio.ed.ac.uksoftware/figtree/) and the
Diaporthomycetidae, families incertae sedis Distoseptisporaceae final layout was done with Adobe Illustrator CS v. 6.
Papulosaceae
Sporidesmiaceae Definitions of previously inconsistently defined terms used
Thyridiaceae in the study
Sordariomycetidae, families incertae sedis Batistiaceae
Sordariomycetes, families incertae sedis Cephalothecaceae We noticed that it was not always possible to be con-
Lautosporaceae sistent with terminology, especially because some terms
Excluded from Sordariomycetes Obryzaceae are not commonly accepted and some have been
Pleurotremataceae interpreted differently by different authors. Instead of
distinguishing between ascolocular pseudothecia and
ascohymenial perithecia, we used the neutral terms
perithecioid ascomata or else the term perithecia in a
Image Frame Work program (v. 0.9.0.7) and images used for broad sense, since by definition all ascomata in the
figures were processed with Adobe Photoshop CS6 software higher Ascomycota (Pezizomycotina) are homologous
(Adobe Systems, USA). Several type specimen were in poor and thus a differential terminology using the term
condition and very little information could be obtained. In pseudothecia for ascolocular perithecioid ascomata does
order to generate more information for this taxon, we relied not reflect phylogenetic relationships.
on the original publication. Hand drawings of these specimens
were made using drawing pens on parchment papers. Faces of
fungi and Index Fungorum numbers are as explained in Results and discussion
Jayasiri et al. (2015) and Index Fungorum (2016)
The combined LSU, SSU, TEF and RPB2 gene data set com-
prised 383 taxa, with Botryotinia fuckeliana, Dothidea
Phylogenetic analyses sambuci, Exophiala dermatitidis and Pyxidiophora
arvernensis as the outgroup taxa. The combined dataset com-
Sequences were obtained from GenBank mostly follow- prised 4367 characters including gaps. The best scoring
ing previous literature and are listed in supplementary RAxML trees are shown in Figs. 1 and 2. Bootstrap support
Table 1. This study used LSU, SSU, TEF and RPB2 values of ML (MLB) (equal to or above 50 %) are given at the
sequence data in the analyses. Multiple sequence align- nodes.
ments were generated with MAFFT v. 7 (http://mafft. In the phylogenetic trees (Figs. 1 and 2), the 380 strains of
cbrc.jp/alignment/server/) then manually corrected using Sordariomycetes included in the analysis cluster into six sub-
MEGA v. 6.06 (Kumar et al. 2012) to ensure alignment classes. Sordariomycetidae, Hypocreomycetidae and
and to minimize the number of uninformative gaps. The Xylariomycetidae as in the previous treatment of Lumbsch
datasets were produced to show families and order rela- and Huhndorf (2010), Meliolomycetidae as suggested by
tionships within the class Sordariomycetes. Kirk et al. (2001), as well as Diaporthomycetidae and
The combined alignments were split between the gen- Lulworthiomycetidae as suggested by
era to improve the robustness of the alignment across Maharachchikumbura et al. (2015). Figure 1 is a reduced
the four loci. Phylogenyies used Maximum Likelihood version of the tree in Fig. 2. Maharachchikumbura et al.
(ML) analyses. Ambiguously aligned regions were ex- (2015) placed the orders Coniochaetales and Cordanales
cluded from all analyses and gaps were treated as in the Diaporthomycetidae. However, given the inconsis-
“missing data” in the analysis. A Maximum Likelihood tency in the placement of the latter two orders, it would be
analysis was performed using RAxMLGUI v. 1.3 more appropriate to refer them to Sordariomycetes order
(Silvestro and Michalak 2011). The optimal ML tree incertae sedis The internal classification of Xylariomycetidae
search was conducted with 1000 separate runs, using is also somewhat problematic. The orders of the
the default algorithm of the program from a random Xylariomycetidae as circumscribed by Senanayake et al.
starting tree for each run. The final tree was selected (2015) were not supported by the phylogenetic analyses in
Fungal Diversity

Fig. 1 The best scoring RAxML Sordariomycetes tree (compressed bootstrap support values (MLB) are given at the nodes. The tree is
overview tree) from 383 taxa based on a combined dataset of LSU, rooted with Botryotinia fuckeliana, Dothidea sambuci, Exophiala
SSU, TEF and RPB2 genes with all lineages collapsed to family level dermatitidis and Pyxidiophora arvernensis
where possible. Orders are indicated in coloured blocks and RAxML

this study. Senanayake et al. (2015) re-validated the order the placement of the families within the subclass are not sta-
Amphisphaeriales based on both sequence data and morphol- ble. Therefore, we would like to keep the order Xylariales to
ogy. However, there is little support to validate this order and represent the taxa in the Xylariomycetidae.
 Fungal Diversity

Fig. 1 (continued)

Presently accepted orders of Sordariomycetes Annulatascales D’souza et al., in Maharachchikumbura

The Sordariomycetes comprise 32 orders. et al., Fungal Divers 72: 212 (2015)
A order within the class Sordariomycetes, subclass
Amplistromatales D’souza et al., in Maharachchikumbura Diaporthomycetidae. This order is characterized by its
et al., Fungal Divers 72: 212 (2015) typical freshwater habitat, growing on submerged
An order comprising families Amplistromataceae and woody substrates, with most genera distributed in the
Catabotrydaceae and was introduced by tropics. Significant characters of the taxa in
Maharachchikumbura et al. (2015). Two genera are included Annulatascales include cylindrical, thin-walled asci,
in Amplistromataceae namely Amplistroma Huhndorf et al. and with a massive, J-, refractive, apical ring, which assists
Wallrothiella Sacc. The genus Wallrothiella was redescribed by in active spore ejection (Tsui and Hyde 2003).
Réblová and Seifert (2004) along with neotypification of its Ascospores are usually equipped with appendages or
type species Wallrothiella congregata (Wallr.) Sacc. sheaths. These characters might be important in aquatic
Amplistroma has been segregated from Wallrothiella based on habitats where they aid in the attachment to substrates
the presence of stromatic ascomata. Catabotrydaceae is mono- (Shearer et al. 2007). The order presently comprises 20
typic with Catabotrys as its type genus. However, Catabotrys genera that belong to, or are referred to, the family
Theiss. & Syd. and Wallrothiella do not possess stromatic Annulatascaceae. To date, DNA sequences of many
ascomata, while Amplistroma is stromatic. LSU sequence data genera have not yet been analysed, and it is likely that
shows that Amplistromataceae has no relationship with the family as presently circumscribed is polyphyletic
Chaetosphaeriales and Magnaporthaceae (Huhndorf et al. (Campbell and Shearer 2004; Vijaykrishna et al. 2005;
2009). In phylogenetic analyses, Amplistromatales appear to Abdel-Wahab et al. 2011). Phylogenies herein reveals a
have a distant relationship with Meliolomycetidae and close relationship of the Annulatascaceae to the
Sordariomycetidae and thus are placed in Sordariomycetes or- Thyridiaceae and Ophiostomataceae with moderate sup-
der incertae sedis (Maharachchikumbura et al. 2015). port (Fig 2).
Fungal Diversity

Fig. 2 Maximum likelihood (ML) majority rule combined LSU, SSU, TEF above 50 %) are given at the nodes. The scale bar represents the expected
and RPB2 consensus tree for the analysed Sordariomycetes isolates. Families number of changes per site. The tree is rooted with Botryotinia fuckeliana,
are indicated in coloured blocks and RAxML bootstrap support values (MLB Dothidea sambuci, Exophiala dermatitidis and Pyxidiophora arvernensis

Boliniales P.F. Cannon, in Kirk et al., Ainsworth & Bisby’s An order of saprobic fungi within the subclass
Dictionary of the Fungi, Edn 9 (Wallingford): x (2001) Sordariomycetidae, which was introduced by Cannon (Kirk
 Fungal Diversity

Jobellisiales
73 Jobellisia guangdongensis GD14-4
96 100 Jobellisia luteola SMH2753 Jobellisiaceae
Jobellisia fraterna SMH2863
58
100 Jattaea prunicola STE-U 6201

Calosphaeriales
100 Jattaea mookgoponga STE-U 6184 Calosphaeriaceae
Calosphaeria pulchella JF 03200
84 Pleurostoma repens CBS H-7594
52 Pleurostoma repens CBS 294.39
Pleurostoma richardsiae CBS 270.33 Pleurostomataceae
100 Pleurostoma ootheca CMU 23858
Pleurostoma ochracea CBS 131321
50
89 Phaeoacremonium fraxinopennsylvanica M.R. 3064

Togniniales
Phaeoacremonium fraxinopennsylvanicum ATCC 26664
Phaeoacremonium africana STE U 6177 Togniniaceae
98 Phaeoacremonium griseo-olivacea STE U 5966
Phaeoacremonium novae-zealandiae WIN 113BI
Magnaporthe salvinii M21
74 Magnaporthe poae M47
89
Slopeiomyces cylindrosporus CBS 609.75 Magnaporthaceae
50 Bussabanomyces longisporus CBS 125232

Magnaporthales
Buergenerula spartinae ATCC 22848
94 Bambusicularia brunnea INA B 92 45
58
Proxipyricularia zingiberis HYZiM201-1-1-1
Pseudopyricularia kyllingae HYKB202 Pyriculariaceae
Pyricularia borealis CBS 461.65
98 62
Deightoniella cibiessia CPC18916
100 Ophioceras dolichostomum CBS 114926
95 Ophioceras dolichostomum HKUCC10113
Ophioceras aquaticus IFRDCC 3091 Ophioceraceae
100
79 Ophioceras commune M91

Amplistromatales
Ophioceras leptosporum CBS 894.70
100 Amplistroma erinaceum AH 43902
100 Amplistroma caroliniana BEO9923 Amplistromataceae
74
Amplistroma longicollis AH37870
Catabotrys deciduum SMH3436 Catabotrydaceae

Annulatascales
54 Pseudoproboscispora caudae A336 2D
Submersisphaeria aquatica A95-1B
Annulusmagnus triseptatus CBS 128831 Annulatascaceae
90
86 Ascitendus austriacus MR 2936
80
Annulatascus velatisporus HKUCC 3701
Thyridiaceae

Ophiostomatales
Thyridium vestitum AFTOL ID 172
76 92 Raffaelea canadensis C2233
Fragosphaeria purpurea CBS 133.34
50
Ophiostoma piliferum AFTOL ID 910 Ophiostomataceae
100
Ceratocystiopsis minuta CBS 116963
Trichosphaeriales

53 Ophiostoma stenoceras AFTOL-ID 1038

99 Cryptadelphia groenendalensis SMH3767
Trichosphaeriaceae
Cryptadelphia groenendalensis SH12
Papulosa amerospora AFTOL ID 748 Papulosaceae

Fig. 2 (continued)

et al. 2001). In our phylogenetic analysis (Figs 1 and 2), (Figs. 1 and 2). The taxa belonging to the order mainly
Boliniales is nested in between the Phyllachorales and comprise simple, dark perithecia, unitunicate asci, and
Sordariales, but appears to be more related to the hyaline to slightly pigmented, ellipsoid to allantoid as-
Phyllachorales. Boliniales comprises a single family, cospores (Réblová et al. 2004). The order forms a sis-
Boliniaceae, with nine genera. ter clade to the order Jobellisiales and Togniniales
(Figs. 1 and 2).
Calosphaeriales M.E. Barr, Mycologia 75(1): 11 (1983)
This order was introduced by Barr (1985). It presently Chaetosphaeriales Huhndorf et al., Mycologia 96(2):
comprises Calosphaeriaceae and Pleurostomataceae 378 (2004a)
Fungal Diversity

100 Ruzenia spermoides SMH4655

Chaetosphaeriales
97 Ruzenia spermoides SMH4606
74
3x Synaptospora plumbea SMH3962 Helminthosphaeriaceae
100
Helminthosphaeria hyphodermae SMH4192
2x
Echinosphaeria canescens SMH4791
73 69 87 Sporoschisma hemipsila SMH2125
Lecythothecium duriligni CBS 101317
71 Chaetosphaeria innumera SMH 2748 Chaetosphaeriaceae
100 Umbrinosphaeria caesariata CBS 102664
Exserticlava vasiformis TAMA 450
Chaetomidium galaicum CBS 113678
Chaetomium elatum IFO 6554
Corynascella inaequalis CBS 284.82 Chaetomiaceae
Achaetomium strumarium IMI 082624

Sordariales
Myceliophthora fergusii CBS 406.69
50 Cercophora mirabilis SMH4002
53 Bombardia bombarda AFTOL ID 967
Anopodium ampullaceum MJR 40 07 (UPS) Lasiosphaeriaceae
Lasiosphaeria ovina SMH4605
Jugulospora rotula ATCC 38359
78 Pseudoneurospora amorphoporcata CBS 626.80
100 Copromyces sp. CBS 386.78
100 Sordaria fimicola CBS 508.50 Sordariaceae
Neurospora crassa MUCL 19026
77
Gelasinospora tetrasperma CBS 178.33
Endoxyla operculata UAMH 11085

Boliniales
60 Camarops ustulinoides AFTOL ID 72
Camaropella pugillus SMH3846 Boliniaceae
59
Cornipulvina ellipsoides SMH1378
Apiorhynchostoma curreyi UAMH 11088

Phyllachorales
100 Coccodiella miconiae ppMP 1342
77 100 Coccodiella melastomatum CMU78543
95
Ascovaginospora stellipala P5-13A
Phyllachoraceae
Phyllachora graminis UME 31349

Cordanales
72 Cordana sp. FMR11828
Cordana inaequalis CBS 508.83
86 Cordanaceae
Cordana abramovii PE 0053-24a
Cordana pauciseptata CBS 121804

Coniochaetales
100
100 Coniochaeta ligniaria C8
89 Coniochaeta luteoviridis CBS 206.38
65 Coniochaeta ostrea AFTOL-ID 915
Coniochaetaceae
62 Barrina polyspora AWR9560A
94 Cephalotheca foveolata UAMH11631
89 Phialemonium atrogriseum CBS 604.67
96 Cephalothecaceae
Albertiniella polyporicola CBS 457.88
Cryptendoxyla hypophloia WM10 89
99 Irenopsis cornuta VIC 32058
Meliolales

Irenopsis vincensii VIC 31751

3x
100 Endomeliola dingleyae PDD 98304
Asteridiella obesa VIC 31239 Meliolaceae
Meliola centellae VIC 31244

Fig. 2 (continued)

This order was introduced in Sordariomycetidae Coniochaetales Huhndorf et al., Mycologia 96(2): 378
based on molecular analysis of LSU nrDNA sequence (2004a)
data by Huhndorf et al. (2004a). In our phylogenetic This order was introduced by Huhndorf et al. (2004a)
analysis (Fig. 1), Chaetosphaeriales is close to and incorporates a single family Coniochaetaceae with
Sordariales. The order comprises Chaetosphaeriaceae three genera. Coniochaetaceae differs from Sordariaceae
with 35 genera and Helminthosphaeriaceae with seven and related families in having ascospores with elongate
genera. germ slits (Malloch and Cain 1971). Cephalothecaceae
 Fungal Diversity

85 Bartalinia robillardoides CBS 122705

86 Zetiasplozna acaciae CPC 23421
100 Truncatella spartii MFLU 15-0721 Bartaliniaceae
Hyalotiella spartii MFLUCC 13-0397
70
Broomella vitalbae MFLUCC 15-0023
83 Robillarda sessilis CBS 114312
81
75 Robillarda roystoneae CBS 115445
64 Robillarda africana CBS 122.75 Robillardaceae
Robillarda terrae CBS 587.71
100
55 Robillarda sessilis CBS 101440
100
65 Seimatosporium cornii MFLUCC 14-0467
Sarcostroma restionis CBS 118154
99
66
Discostroma tosta HKUCC 1004 Sporocadaceae
Adisciso yakushimense KT1907
90
Discosia sp. 5 KT-2010

Xylariales
64 100 Pseudopestalotiopsis theae SAJ-0021
52 Pestalotiopsis sp. CGMCC 3.9103
70
Ciliochorella sp. MFLUCC12-0310 Pestalotiopsidaceae
76 Lepteutypa cupressi IMI 052255
92 Seiridium phylicae CPC 19962
100 Phlogicylindrium eucalyptorum CBS 111680
92 Phlogicylindrium eucalyptorum CBS 111689
83 Phlogicylindrium uniforme CBS 131312 Phlogicylindriaceae
100 Ciferriascosea rectamurum MFLUCC 15-0542
Ciferriascosea fluctamurum MFLUCC 15-0541
Beltrania pseudorhombica CPC 23656
Parapleurotheciopsis inaequiseptata MUCL 4108
94
Beltraniopsis neolitseae CPC 22168 Beltraniaceae
50
Beltraniella endiandrae CPC 22193
Subramaniomyces fusisaprophyticus CBS 418.95
100 Lepteutypa uniseptata HKUCC voucher
Lepteutypa uniseptata HKUCC 6349
100 71 Amphisphaeria sorbi MFLUCC 13-0721 Amphisphaeriaceae
Amphisphaeria umbrina AFTOL ID 1229
Amphisphaeria umbrina HKUCC 994
98 Arthrinium phaeospermum HKUCC 3395
79 Arthrinium montagnei AFTOL ID 951
99 Arthrinium hydei CBS 114990 Apiosporaceae
Arthrinium bambusae ICMP 6889
87 Arthrinium setosa ICMP 4207
Melogramma campylosporum MBU Melogrammataceae
100 Pseudomassaria chondrospora MFLUCC 15-0545
93 Pseudomassaria chondrospora PC1 Pseudomassariaceae
Pseudomassaria sepincoliformis PS
Hyponectria buxi UME 31430 Hyponectriaceae

Fig. 2 (continued)

and Cordanaceae are sister families with strong phyloge- this family inhabit freshwater and terrestrial habitats.
netic support. Maharachchikumbura et al. (2015) referred The mode of conidiogenesis is unique with multiple,
the Cordanales and Coni ochaetales to the conspicuous collarettes forming a multi-lamellar struc-
Diaporthomycetidae but recent publications reported that ture around the blastic conidiogenous locus of the in-
ordinal placement is still unresolved (Su et al. 2016; tercalary conidiogenous cells (Shearer and Motta
Yang et al. 2015). Therefore, in the present study, we treat 1973)
them under Sordariomycetes, orders incertae sedis.
Cordanales M. Hern.-Rest. & Crous, in Hernández-
Conioscyphales Réblová & Seifert, in Réblová et al., Restrepo et al., Phytotaxa 205(4): 233 (2015a)
Persoonia 37: 63 (2016) This order was introduced by Hernández-Restrepo
This order accommodates the family Conioscyphaceae et al. (2015a) to accommodate the family Cordanaceae with
with the monotypic genus Conioscypha. Members of a single genus Cordana Preuss. Morphologically it is easily
Fungal Diversity

89 Microdochium phragmitis CBS 423.78

86 Microdochium trichocladiopsis CBS 623.77
100 Idriella lunata CBS 204.56
Microdochiaceae
100 Selenodriella cubensis CBS 683.96
Selenodriella fertilis CBS 772.83
64 Coniocessia maxima CBS 593.74
100 Coniocessia nodulisporioides CBS 281.77 Coniocessiaceae
Coniocessia anandra Iran 1468C
100 Creosphaeria sassafras CM AT 018
100 Creosphaeria sassafras CBS 119001
95 Lopadostomataceae
Creosphaeria sassafras ANM 1978

Xylariales
Lopadostoma turgidum LT2
55 Diatrype whitmanensis ATCC MYA-4417
74 Eutypa lata CBS 208.87
94
Diatrype disciformis AFTOL ID 927 Diatrypaceae
71
Diatrype palmicola MFLUCC 11-0020
Monosporascus cannonballus FMR6682
74 Kretzschmaria deusta CBS 163.93
70 “Xylarioideae”
Xylaria hypoxylon CBS 122620
94 76 Podosordaria tulasnei CBS 128.80 Xylariaceae
99 Bisconiauxi anummularia MUCL 51395
Graphostroma platystoma AFTOL ID 1249 “Hypoxyloideae”
100 Vialaea mangifia MFLUCC 12-0808 Vialaeaceae
Vialaea minutella BRIP 56959
62 Amphibambusa bambusicola MFLUCC 11-0617
68 Arecophila bambusae HKUCC 4794
89 Seynesia erumpens SMH 1291
Cainiaceae
Cainia graminis CBS 136.62
Iodosphaeria tongrenensis FJS8 Iodosphaeriaceae

Melanosporales
100 Melanospora tiffanii ATCC 15515
100 Melanospora zamiae ATCC 96173
100 Melanospora zamiae ATCC 12340 Ceratostomataceae
5x
Melanospora singaporensis ATCC 38286
100 Vittatispora coorgii BICC 7817
Scortechiniellopsis leonensis GKM1269
91 Cryptosphaerella elliptica SMH4722
100 Biciliospora velutina GKM1268
2x
100 Neofracchiaea callista SMH2689 Scortechiniaceae
2x 58 Scortechinia acanthostroma SMH1143

Coronophorales
Coronophora gregaria ANM1555 Coronophoraceae
62 53 Bertia didyma SMH4719
Bertia ngongensis GKM1239
60 2x Bertiaceae
3x Bertia tropicalis SMH3513
100
Gaillardiella pezizoides GKM1245
Bertia moriformis SMH4320
100
4x
Acanthonitschkea argentinensis SMH1395
95 50 100 Acanthonitschkea tristis SMH4723
3x Nitschkia tetraspora SMH4787
63 100 Nitschkiaceae
Nitschkia tetraspora GKML148N
Fracchiaea broomeana SMH347
3x
100 Crassochaeta nigrita SMH1667
2x
3x Crassochaeta nigrita SMH2931
87 100
3x Chaetosphaerella phaeostroma SMH4257
Chaetosphaerellaceae
74
Falcocladiales

100 Chaetosphaerella fusca GKML124N

Spinulosphaeria nuda SMH1952
95 Falcocladium sphaeropedunculatum CBS 111292
95 Falcocladium thailandicum CBS 121717
2x
100 Falcocladium turbinatum BCC22055 Falcocladiaceae
Falcocladium multivesiculatum CBS 120386

Fig. 2 (continued)

distinguished from its sister order Coniochaetales of them constitute two distinct monophyletic groups
(Coniochaetaceae) by having a basal stroma, ascospores with- with reliable support.
out germ slits and polyblastic asexual morphs
(Hernández-Restrepo et al. 2015a). Our molecular data Coronophorales Nannf., Nova Acta R. Soc. Scient. upsal.,
also support the segregation of these two orders as both Ser. 4 8(no. 2): 54 (1932)
 Fungal Diversity

97 100 Juncigena adarca JK5548A

100 Juncigena adarca JK5235A

Torpedosporales
87
100 Moheitospora fruticosae EF14 Juncigenaceae
Marinokulati chaetosa BCRC FU30272
99 Marinokulati chaetosa BCRC FU30271
50
55 Etheirophora blepharospora JK5397A
76 Etheirophora blepharospora JK5289
100 98 Swampomyces armeniacus JK5325A Etheirophoraceae
Swampomyces triseptatus CY2802
Etheirophora unijubata JK5443B
100 Torpedospora radiata PP7763
100 Torpedospora radiata AFTOL ID 751
100 Torpedosporaceae
Glomerulispora mangrovis NBRC 105264
Torpedospora ambispinosa BCC16003
63 Hyperdermium pulvinatum P.C. 602
Cordyceps militaris OSC 93623
Ascopolyporus polychrous PC546 Cordycipitaceae
100
Cordyceps cardinalis OSC 93609
81 Torrubiella wallacei CBS 101237
52 100 Trichoderma viride GJS89 127
Trichoderma rufa DAOM JBT1003
59 Arachnocrea stipata TFC 97-43 Hypocreaceae
100
Escovopsis weberi ATCC 64542
Sphaerostilbella berkeleyanan CBS 102308
63 Hypocrella discoidea BCC 8237
80 Shimizuomyces paradoxus EFCC 6279
Balansia henningsiana AEG96-27 Clavicipitaceae

Hypocreales
Epichloe typhina ATCC 56429
50
Claviceps purpurea AEG 97 2
Flammocladiella aceris CPC 24422 Flammocladiaceae
95 Tolypocladium japonica OSC 110991
88 Tolypocladium capitata OSC 71233 Ophiocordycipitaceae
78 Ophiocordyceps sinensis YN09 64
Ophiocordyceps aurantiaca OSC 128578
Penicillifer diparietispora ATCC MYA 627
85 Gliocephalotrichum bulbilium ATCC 22228
86
Neonectria ramulariae CBS 151.29 Nectriaceae
97 Cosmospora coccinea AR2741
Nectria cinnabarina CBS 114055
52 Parasarcopodium ceratocaryi CBS 110664
Peethambara spirostriata CBS 110115
Myrothecium roridum ATCC 16297 Stachybotriaceae
85 Scopinella solani strain CBS 770.84
Stachybotrys chlorohalonata UAMH6417
Hydropisphaera peziza GJS92 101
81 Protocreopsis pertusa C.T.R. 72-184
74 Roumegueriella rufula GJS 91 64 Bionectriaceae
Clonostachys ochroleuca AFTOL ID 187
67
56 Stephanonectria keithii GJS92 133
100 Tilachlidium brachiatum CBS 505.67
Tilachlidiaceae
Tilachlidium brachiatum CBS 363.97
Niesslia exilis CBS 357.70
58
Niessliaceae
Niesslia exilis CBS 560.74

Fig. 2 (continued)

Taxa belonging to the order Coronophorales inhabit wood cases the ostiolar opening is either indistinct or lacking
and comprise Bertiaceae, Ceratostomataceae, (Mugambi and Huhndorf 2010).
Chaetosphaerellaceae, Coronophoraceae, Nitschkiaceae and
Scortechiniaceae. Taxa in the order are characterised by Diaporthales Nannf., Nova Acta R. Soc. Scient. upsal.,
mostly superficial ascomata, sometimes with an exten- Ser. 4 8(no. 2): 53 (1932)
sive hyphal subiculum or well-developed basal stroma The order Diaporthales comprises 12 families, which
that often becomes cupulate or collapsed, and in some are Cryphonectriaceae, Diaporthaceae, Gnomoniaceae,
Fungal Diversity

100 Aniptodera chesapeakensis ATCC 32818

94 Remispora maritima BBH28309
87
Nimbospora effusa JK 5104A
100 Halosphaeriaceae
Halosphaeria appendiculata CBS 197.60
Lignincola laevis JK50180
94 Microascus longirostris CBS 267.49

Microascales
100
100 Microascus trigonosporus AFTOL-ID 914

97
100 Doratomyces stemonitis AFTOL-ID 1380 Microascaceae
100 Pseudallescheria boydii CBS 108.54
Petriella setifera AFTOL ID 956
100 Graphium fimbriasporum CMW5605
57
Graphiaceae
Graphium penicillioides CBS 506.86
82 Gondwanamyces capensis CMW997
80
74 Gondwanamyces capensis AFTOL ID 1907
95 Gondwanamyces proteae (AF221011) Gondwanamycetaceae
100
Custingophora cecropiae CCF 3568
100 Custingophora olivacea CBS 335.68
63 Ambrosiella ferruginea CBS 460.82
100 Thielaviopsis thielavioides CBS 130.39 Ceratocystidaceae
Ceratocystis fimbriata C89
65 Colletotrichum fructicola LC0032
100 Colletotrichum asianum LC0037
Colletotrichum gloeosporioides LC0555 Glomerellaceae
53 61
Colletotrichum musae LC0962

Glomerellales
Colletotrichum brevisporum LC0600
54
68 Monilochaetes dimorphospora MUCL 40959
50 Monilochaetes queenslandica BRIP 24607
100
Monilochaetes infuscans CBS 869.96 Australiascaceae
100
Monilochaetes infuscans CBS 870.96
91
Monilochaetes guadalcanalensis CBS 346.76
87
99 Kylindria peruamazonensis CBS 838.91
Kylindria peruamazonensis CBS 421.95
100 99 Cylindrotrichum clavatus CBS 125239 Reticulascaceae
100 Cylindrotrichum clavatus CBS 125296
Sporoschismopsis sporoschismophora ATCC 42528
66 Plectosphaerella cucumerina DAOM 226828
Gibellulopsis nigrescens DAOM 226890
Verticillium dahliae ATCC 16535 Plectosphaerellaceae
100
Stachylidium bicolor DAOM 226658

Pleurotheciales
Acrostalagmus luteoalbus CBS 194.87
81 Helicoon farinosum DAOM 241947
81 Taeniolella rudis DAOM 229838
88 Pleurotheciella rivularia CBS 125238 Pleurotheciaceae
Carpoligna pleurothecii CBS 101580

Conioscyphales
100 Pleurothecium semifecundum CBS 131271
90 Conioscypha sp. FMR11245
100 Conioscypha lignicola CBS 335.93
100 Conioscypha japonica CBS 387.84 Conioscyphaceae
100 100 Conioscypha sp. ILL 41202
Conioscypha varius CBS 113653
Savoryellales

100 Canalisporium caribense SS038397

100 Canalisporium grenadoidia NB 2010a
98 Savoryella lignicola NF00204
100 Ascotaiwania lignicola NIL00005 Savoryellaceae
Ascotaiwania lignicola NIL00006

Fig. 2 (continued)

Harknessiaceae, Macrohilaceae, Melanconidaceae, perithecia with an elongate beak, often forming within
Pseudoplagiostomataceae, Pseudovalsaceae, stromatic tissues (Rossman et al. 2007). Asci generally
Schizoparmaceae, Stilbosporaceae, Sydowiellaceae and deliquesce at the base when mature and have a charac-
Valsaceae. Members of Diaporthales are pathogens, par- teristic refractive apical annulus.
asites, and endophytes of plants, human-animal patho-
gens, saprobes and soil inhabitants (Rossman et al. Falcocladiales R.H. Perera et al. in Maharachchikumbura
2007). The order Diaporthales is characterized by et al., Fungal Divers 72: 218 (2015)
 Fungal Diversity

Lulworthlales Pisorisporiales
100 Lindra thalassiae AFTOL ID 413
83 Cumulospora marina MF46

Koralionastetales
97
Lulworthia fucicola ATCC 64288s Lulworthiaceae
85 Hydea pygmea NBRC33069
100 Koralionastes ellipticus JK5769
Koralionastetaceae
Koralionastes ellipticus JK5771
100 Pisorisporium cymbiforme PRM 924378
100 Pisorisporium cymbiforme PRM 924377
100 Pisorisporium cymbiforme PRM 924379 Pisorisporiaceae
100 Ascocollumdensa aquatica HKUCC 3707
Achroceratosphaeria potamia JF 08139
92 Dothidea sambuci DAOM 231303
92 Botryotinia fuckeliana AFTOL ID 59
87 Exophiala dermatitidis AFTOL ID 668
Pyxidiophora arvernensis AFTOL-ID 2197

0.2

Fig. 2 (continued)

This monotypic order in the subclass Members of the Hypocreales are highly diverse
H y p o c re o m y c e t i d a e a c c o m m o d a t e t h e f a m i l y in the tropics, subtropics temperature regions
Falcocladiaceae. Members of this family are saprobes (Põldmaa 2011). Hypocreales is represented by
on leaf litter and leaves including Eucalyptus grandis Bionectriaceae, Clavicipitaceae, Cordycipitaceae,
and E. camaldulensis in tropical, terrestrial habitats Flammocladiaceae , Hypocreaceae, Nectriaceae,
(Crous et al. 1994; Jones et al. 2014). The order pres- Niessliaceae, Ophiocordycipitaceae, Stachybotriaceae
ently includes one hyphomycetous asexual genus and Tilachlidiaceae.
Falcocladium introduced by Crous et al. (1994). Jones
et al. (2014) introduced the monotypic family J o b e l l i s i a l e s D ’s o u z a & K . D . H y d e , i n
Falcocladiaceae based on SSU and LSU sequence data Maharachchikumbura et al., Fungal Divers 72: 219
to accommodate members of genus Falcocladium and (2015)
suggested further taxon sampling was needed to deter- Jobellisiales is a monotypic order in the class
mine its ordinal status. There is no recorded sexual Sordariomycetes, subclass Diaporthomycetidae. It includes a
morph for this order. Phylogenies recovered herein de- single family Jobellisiaceae characterized by saprobic
pict that the Falcocladiales is most closely related to lignicolous taxa found in terrestrial and freshwater hab-
the order Coronophorales (Figs. 1 and 2). itats. This order encompasses taxa that possess relatively
large, superficial ascomata, a three layered, thick perid-
Glomerellales Chadef. ex Réblová et al., Stud. Mycol. ium, cylindrical asci and one septate ascospores.
68(1): 170 (2011) Ascomata of Jobellisia luteola (Ellis & Everh.) M.E.
Chadefaud (1960) proposed the order “Glomérellales” Barr are peculiar in appearing orange to yellow brown,
but without a Latin diagnosis and the name was thus whereas those of J. barrii Huhndorf et al. and
invalid. The order Glomerellales was therefore validly J. viridifusca K.M. Tsui & K.D. Hyde have a bright
published by Réblová et al. (2011) in the class orange middle wall layer (Ranghoo et al. 2001, Liu
S o rd a r i o m y c e t e s a n d co m p r i s e d t h r e e f a m i l i e s et al. 2012). The family Jobellisiaceae was introduced
Australiascaceae, Reticulascaceae and Glomerellaceae. by Réblová (2008) as a monotypic family and currently
The introduction was based on analysis of ITS1, 5.8S includes seven species. Réblová (2008), using LSU
rDNA and ITS2 (ITS), nc28S (LSU) rDNA, and nc18S rDNA sequence analysis, showed that the closest rela-
(SSU) rDNA datasets, and a combined data set of LSU- tives to Jobellisiaceae are Calosphaeriales and
SSU-RPB2. Maharachchikumbura et al. (2015) included Togniniaceae. No asexual morph is reported for this
Plectosphaerellaceae in to this order based on a com- order.
bined data set of LSU-SSU-TEF-RPB2. Current analy-
ses clearly demonstrate that this order, represented by Koralionastetales Kohlm. et al., Mycol. Res. 113(3): 377
four families is monophyletic. (2009)
Based on the combined dataset of SSU and LSU rDNA
Hypocreales Lindau, in Engler & Prantl, Nat. sequences and morphological characters the genera
Pflanzenfam., Teil. I (Leipzig) 1(1): 343 (1897) Koralionastes and Pontogeneia were assigned to the new
Fungal Diversity

order Koralionastetales (C am pbell et al . 2009). member of Dothideomycetes (Lumbsch and Huhndorf

Koralionastetales shows phylogenetic affinities to 2007a).
Lulworthiales and Pisorisporiales.
Microascales Luttr. et al., Mycotaxon 12(1): 40 (1980)
Lulworthiales Kohlm. et al., Mycologia 92(3): 456 (2000) The order Microascales was introduced by Benny and
Kohlmeyer et al. (2000) assigned two genera Kimbrough (1980) to accommodate Chadefaudiellaceae,
Lulworthia and Lindra to this new order and the new Microascaceae and Pithoascaceae, and later expanded to in-
family (Lulworthiaceae). Abdel-Wahab et al. (2010) clude Ceratocystidaceae, Gondwanamycetaceae,
showed that a number of asexual morphs also belong Halosphaeriaceae and Graphiaceae (Réblová et al. 2011).
in this family, i.e. Halazoon, Hydea and Orbimyces.
Members of Lulworthiales lack an apical ring in the Ophiostomatales Benny & Kimbr., Mycotaxon 12(1): 48
asci and have ascospores with apical chambers from (1980)
which mucilage is released (Campbell et al. 2005). This order was introduced by Benny and Kimbrough
The Lulworthiaceae is a monophyletic lineage with high (1980) for the family Ophiostomataceae; while Kathistaceae
support and is a sister family to the Koralionastetaceae was added by Malloch and Blackwell (1990).
(Fig 2)
Phyllachorales M.E. Barr, Mycologia 75(1): 11 (1983)
Magnaporthales Thongk. et al., Fungal Diversity 34: The order Phyllachorales was introduced by Barr (1983)
166 (2009) comprising two families, viz. Phaeochoraceae and
A phylogenetic study of Pyricularia (Sacc.) Sacc. and Phyllachoraceae. This order is distinctive as the species are
related genera in Magnaporthales by Klaubauf et al. (2014) biotrophs on various hosts, where they form bright or black
resulted in the introduction of two new families, stroma (Pearce and Hyde 1994). Asci are unitunicate and as-
Ophioceraceae and Pyriculariaceae. Ophioceraceae com- cospores are hyaline or lightly pigmented (Barr 1983; Pearce
prises a single genus Ophioceras, containing species that and Hyde 1993a, b, 1994). There is a close phylogenetic as-
mostly occur on wood submerged in freshwater. sociation of the Phyllachorales to the Boliniales.
Magnaporthaceae was considered to include
Buergenerula, Bussabanomyces, Endopyricularia, Pisorisporiales Réblová & J. Fourn., Persoonia 34: 43
Gaeumannomyces, Harpophora, Magnaporthiopsis, (2015)
Nakataea, Omnidemptus, Pyriculariopsis and This monotypic order was introduced by Réblová et al.
Slopeiomyces with Nakataea as the correct name for the (2015) based on LSU, SSU and RPB2 sequence data. It
type of the family. The new genus Kohlmeyeriopsis was currently includes the family Pisorisporiaceae. In their
introduced based on molecular data and morphology. analyses, the Pisorisporiales nested in a weakly-
Pyriculariaceae was introduced and included supported clade sister to the orders Lulworthiales and
Deightoniella and Pyricularia, and the new genera Koralionastetales.
Bambusicularia, Barretomyces, Macgarvieomyces,
Neopyricularia, Proxipyricularia, Pseudopyricularia and Pleurotheciales Réblová & Seifert, in Réblová et al.,
Xenopyricularia (Klaubauf et al. 2014). Persoonia 37: 63 (2016)
Pleurotheciales accommodates the family
Melanosporales N. Zhang & M. Blackw., in Hibbett et al., Pleurotheciaceae, which comprises 11 genera. Members of
Mycol. Res. 111(5): 531 (2007) the order Pleurotheciales share dark, papillate, glabrous or
This order was suggested in Zhang et al. (2006) but not sparsely setose perithecia, upright or lying horizontally to
validly published; it was formally introduced by Hibbett et al. the host, asci with a distinct non-amyloid apical annulus, fili-
(2007) to accommodate Melanospora and Sphaerodes in form paraphyses that disintegrate partially at maturity and
Ceratostomataceae. fusiform to ellipsoidal, septate, hyaline ascospores (Réblová
et al. 2016).
Meliolales Gäum et al., Syst. Ascom. 5(1): 180 (1986)
This order was introduced by Hawksworth and Savoryellales Boonyuen et al., in Boonyuen et al.,
Eriksson (1986) and it accommodates a single family Mycologia 103(6): 1368 (2011)
Meliolaceae, which are the black mildews, and comprise The taxonomic placement of the genus Savoryella has been
eight genera. This group is unique in producing black widely debated and Jones et al. (2009a, b) referred it to the
web-like colonies on the host, comprising superficial Sordariales genera incertae sedis. Boonyuen et al. (2011), in a
brown to black mycelium with appressoria (Hosagoudar combined phylogenetic analysis of Savoryella species (LSU,
2004). Until recently the family was thought to be a SSU, 5.8S rRNA genes, RPB1, RPB2, TEF), showed that
 Fungal Diversity

they formed a monophyletic group in the Sordariomycetes, Torpedosporales E.B.G. Jones et al., in Jones et al., Fungal
but showed no affinities with other accepted orders. The order Diversity 73(1): 43 (2015)
Savoryellales was introduced to accommodate Savoryella Based on combined SSU and LSU rDNA sequence
species, along with the genera Ascotaiwania, Ascothailandia analysis Jones et al. (2015) showed that taxa in the families
(and its asexual morph Canalisporium), as they formed a dis- Etheirophoraceae, Juncigenaceae and Torpedosporaceae
tinct lineage in the Sordariomycetes (Boonyuen et al. 2011). form a highly supported clade in the Hypocreomycetidae
and a new order Torpedosporales was introduced. Similar
Sordariales Chadef. ex D. Hawksw. & O.E. Erikss., Syst. phylogenies are obtained in this study. Members of the order
Ascom. 5(1): 182 (1986) are saprobic on lignicolous substrates and leaves, in marine
This order was introduced by Hawksworth and Eriksson habitats. The Torpedosporales shows phylogenetic affinities
(1986) and comprises three families viz. Chaetomiaceae, to the orders Falcocladiales, Coronophorales and
Sordariaceae and Lasiosphaeriaceae sensu lato. Most species Melanosporales (Jones et al. 2015)
grow on either dung or decaying substrata, such as wood, or
are aquatic, growing on submerged wood (Zhang et al. 2006). Trichosphaeriales M.E. Barr, Mycologia 75(1): 11 (1983)
The Sordariales are characterized by membranous or coria- This order was introduced by Barr (1983) based on
ceous ascomata, and hyaline or brown ascospores often with Trichosphaeria pilosa (Pers.) Fuckel. The current classifica-
appendages or sheaths (Zhang et al. 2006). In most studies, a tion of Trichosphaeriales recognizes only one family, the ter-
close relationship between Sordariales and Chaetosphaeriales restrial Trichosphaeriaceae. Trichosphaeriaceae has affinities
has always been observed. with Annulatascaceae, Ophiostomataceae and
Papulosporaceae.
Spathulosporales Kohlm., Mycologia 65(3): 615 (1973)
The Spathulosporales is a marine order that includes the Xylariales Nannf., Nova Acta R. Soc. Scient. upsal., Ser. 4
family Hispidicarpomycetaceae and Spathulosporaceae. 8(no. 2): 66 (1932)
Some Spathulosporaceae species show affinities to the order The Xylariales is a large order of perithecial ascomy-
Lulworthiales (Inderbitzin et al. 2004), although the type spe- cetes with eight-spored unitunicate asci, with a J+, api-
cies (Spathulospora phycophila A.R. Caval. & T.W. Johnson) cal ring and ascospores with a prominent germ slit
has not been sequenced. Presently this order is classified under (Smith et al. 2003). Presently there are 22 families ac-
Sordariomycetes, order incertae sedis. cepted in Xylariales.

Tirisporellales Suetrong et al., in Jones et al., Fungal

Diversity: 73(1): 42 (2015) Taxonomy
Tirisporellales is a monotypic order in the class
Sordariomycetes, subclass Diaporthomycetidae and includes Outline of Sordariomycetes, 31 December 2015
a single family Tirisporellaceae. The family was introduced
●
by Suetrong et al. (2015) with two monotypic genera = sequence data in GenBank as at date on submission
Tirisporella and Thailandiomyces, based on SSU and LSU
rDNA sequences analysis and morphological observations. Class SORDARIOMYCETES sensu O.E. Erikss. & Winka
Tirisporella is a mangrove species found growing on the palm
Nypa fruticans while Thailandiomyces is a freshwater asco- Diaporthomycetidae Senan. et al.
mycete growing on the trunks of the palm Licuala
longicalycata. Annulatascales D’souza et al.
Annulatascaceae S.W. Wong et al.
Togniniales Senan. et al., in Maharachchikumbura et al., Annulatascus K.D. Hyde ●
Fungal Diversity: 10.1007/s13225-015-0331-z, [22] (2015) Annulusmagnus J. Campb. & Shearer ●
Togniniales is a monotypic order in the class Aqualignicola V.M. et al.
Sordariomycetes, subclass Diaporthomycetidae. Our analyses Aquaticola W.H. Ho et al. ●
also reveal a similar scenario and the order is basal to Ascitendus J. Campb. & Shearer●
the Diaporthales. The Togniniales includes a single fam- Ayria Fryar & K.D. Hyde
ily Togniniaceae containing plant pathogen and saprobic Cataractispora K.D. Hyde et al. ●
on dead wood. Togniniaceae comprising the sexual Chaetorostrum Zelski et al.
morph genera Conidiotheca and Togninia and the asex- Clohiesia K.D. Hyde ●
ual morph genus Phaeoacremonim (Réblová and Cyanoannulus Raja et al. ●
Mostert 2007). Dictyosporella Abdel-Aziz ●
Fungal Diversity

Diluviocola K.D. Hyde et al. Latruncellus M. Verm. et al. ●

Fusoidispora D. Vijaykrishna et al. ● Luteocirrhus C.F. Crane & T.I. Burgess ●
Longicollum Zelski et al. Mastigosporella Höhn. ●
Pseudoannulatascus Z.L. Luo et al. ● Microthia Gryzenh. & M.J. Wingf. ●
Pseudoproboscispora Punith. ● Prosopidicola Crous & C.L. Lennox ●
Rhamphoria Niessl. ● Rostraureum Gryzenh. & M.J. Wingf. ●
Submersisphaeria K.D. Hyde ● Ursicollum Gryzenh. & M.J. Wingf. ●
Torrentispora K.D. Hyde et al.
Vertexicola K.D. Hyde et al. ● Diaporthaceae Höhn. ex Wehm.
Allantoporthe Petr.
Calosphaeriales M.E. Barr Apioporthella Petr.
Calosphaeriaceae Munk ● Clypeoporthella Petr.
Calosphaeria Tul. & C Tul. ● Diaporthe Nitschke ●
Calosphaeriophora Réblová et al. ● = Phomopsis (Sacc) Bubák
Conidiotheca Réblová & L Mostert Diaporthella Petr. ●
Jattaea Berl ● Leucodiaporthe M.E. Barr & Lar.N. Vassiljeva
= Phragmocalosphaeria Petr. Mazzantia Mont. ●
= Wegelina Berl. = Mazzantiella Höhn.
Kacosphaeria Speg. Ophiodiaporthe Y.M. Ju et al. ●
Sulcatistroma A.W. Ramaley Pustulomyces D.Q. Dai et al. ●
Togniniella Réblová et al. ● Stenocarpella Syd. & P. Syd. ●
= Phaeocrella Réblová et al. ●
Tulipispora Révay & J. Gönczöl, in Révay et al. Gnomoniaceae G. Winter
Alnecium Voglmayr & Jaklitsch ●
Pleurostomataceae Réblová, L. Mostert, W. Gams & Ambarignomonia Sogonov ●
Crous Amphiporthe Petr. ●
Pleurostoma Tul. & C. Tul. ● Anisomyces Theiss. & Syd.
= Pleurostomophora Vijaykr.et al. Apiognomonia Höhn. ●
Apioplagiostoma M.E. Barr ●
Calosphaeriales genera, incertae sedis Asteroma DC. ●
Enchnoa Fr. ● Bagcheea E. Müll. & R. Menon
Clypeoporthe Höhn.
Diaporthales Nannf. Cryptosporella Sacc. ●
Cryphonectriaceae Gryzenh. & M.J. Wingf. Cylindrosporella Höhn.
Amphilogia Gryzenh. et al. ● Depazea Fr.
Aurantiosacculus Dyko & B. Sutton ● Diplacella Syd.
Aurapex Gryzenh. & M.J. Wingf. ● = Discosporium Höhn.
Aurifilum Begoude et al. ● = Discula Sacc.
Celoporthe Nakab. et al. ● Ditopella De Not. ●
Chromendothia Lar.N. Vassiljeva ● Ditopellopsis J. Reid & C. Booth ●
Chrysocrypta Crous & Summerell ● Gloeosporidina Petr.
Chrysofolia Crous & M.J. Wingf. ● Gnomonia Ces. & De Not.
Chrysoporthe Gryzenh. & M.J. Wingf. ● Gnomoniella Sacc. ●
= Chrysoporthella Gryzenh. & M.J. Wingf. ● Gnomoniopsis Berl. ●
Cryphonectria (Sacc.) Sacc. & D. Sacc. ● Mamiania Ces & De Not.
= Endothiella Sacc. Millerburtonia Cif .
Cryptometrion Gryzenh. & M.J. Wingf. ● Occultocarpon L.C. Mejía & Zhu L. Yang ●
Diversimorbus S.F. Chen & J. Roux ● Ophiognomonia (Sacc.) Sacc. ●
Endothia Fr. ● Phragmoporthe Petr. ●
Foliocryphia Cheewangkoon & Crous ● Phylloporthe Syd.
Holocryphia Gryzenh. & M.J. Wingf. ● Plagiostoma Fuckel ●
Immersiporthe S.F. Chen et al. ● = Cryptodiaporthe Petr.
Lasmenia Speg. ● = Diplodina Westend
 Fungal Diversity

Pleuroceras Riess. ● = Baeumleria Petr. & Syd.

= Linospora Fuckel Pilidiella Petr. & Syd. ●
Skottsbergiella Petr. = Schizoparme Shear
Sirococcus Preuss ●
Spataporthe Bronson et al. Stilbosporaceae Link
Uniseta Ciccar Crinitospora B. Sutton & Alcorn ●
Xenotypa Petr. Natarajania Pratibha & Bhat ●
Zythia Fr. ● Stilbospora Pers. ●
Stegonsporium Corda ●
Harknessiaceae Crous
Harknessia Cooke ● Sydowiellaceae Lar.N. Vassiljeva
Cainiella E. Müll. ●
Macrohilaceae Crous Calosporella J. Schröt ●
Macrohilum H.J. Swart ● Chapeckia M.E. Barr ●
Hapalocystis Auersw. ex Fuckel ●
Melanconidaceae G. Winter Lambro Racib.
Botanamphora Nograsek & Scheuer Rossmania Lar.N. Vassiljeva ●
Ceratoporthe Petr. Sillia P. Karst. ●
Cytomelanconis Naumov Stegophora Syd. & P. Syd.
Dicarpella Syd. & P. Syd. ● Sydowiella Petr. ●
Dictyoporthe Petr. Uleoporthe Petr.
Fremineavia Nieuwl. Winterella (Sacc.) Kuntze ●
Gibellia Sacc.
Hypophloeda K.D. Hyde & E.B.G. Jones Valsaceae Tul. & C. Tul.
Kensinjia J. Reid & C. Booth Amphicytostroma Petr.
Macrodiaporthe Petr. Chadefaudiomyces Kamat et al.
Massariovalsa Sacc. Cryptascoma Ananthap.
Mebarria J. Reid & C. Booth Cytospora Ehrenb. ●
Melanamphora Lafl. = Leucostoma (Nitschke) Höhn.
Melanconiella Sacc. ● = Valsa Fr.
Melanconiopsis Ellis & Everh. = ValsellaFuckel
Melanconis Tul. & C. Tul. ● = Valseutypella Höhn.
= Melanconium Link Ditopellina J. Reid & C. Booth
Phragmodiaporthe Wehm. Durispora K.D. Hyde
Plagiophiale Petr. Harpostroma Höhn.
Plagiostigme Syd. Hypospilina (Sacc) Traverso
Prostratus Sivan. et al. Kapooria J. Reid & C. Booth
Pseudovalsella Höhn. Leptosillia Höhn.
Wehmeyera J. Reid & C. Booth Maculatipalma J. Fröhlich & K.D. Hyde ●
Wuestneia Auersw. ex Fuckel ● Pachytrype Berl. ex M.E. Barr et al. ●
Wuestneiopsis J. Reid & Dowsett Paravalsa Ananthap.

Pseudoplagiostomataceae Cheew. et al. Diaporthales, genera incertae sedis

Pseudoplagiostoma Cheew. et al. ● ArgentinomycesN.I. Peña & Aramb.
Anisogramma Theiss. & Syd. ●
Pseudovalsaceae M.E. Barr Anisomycopsis I. Hino & Katum.
Apoharknessia Crous & S.J. Lee ● Apiosporopsis (Traverso) Mariani ●
Coryneum Nees = Sphaerognomonia Potebnia ex Höhn.
= Pseudovalsa Ces. & De Not. ● Apomelasmia Grove
Auratiopycnidiella Crous & Summerell ●
Schizoparmaceae Rossman Bagadiella Cheew. & Crous ●
Coniella Höhn. ● Caudospora Starbäck
Fungal Diversity

Chaetoconis Clem. ● Omnidemptus P.F. Cannon & Alcorn ●

Cryptoleptosphaeria Petr. Phomatospora Sacc. ●
Cryptonectriella (Höhn.) Weese Pseudophialophora J. Luo & N. Zhang ●
Cryptonectriopsis (Höhn.) Weese Pyriculariopsis M.B. Ellis ●
Diatrypoidiella Manohar et al. Slopeiomyces Klaubauf et al. ●
Disculoides Crous et al. ●
Dwiroopa Subram. & Muthumary Ophioceraceae Klaubauf et al.
Erythrogloeum Petr. ● Ophioceras Sacc. ●
Exormatostoma Gray
Greeneria Scribn. & Viala ● Pyriculariaceae Klaubauf et al.
Gyrostroma Naumov ● Bambusicularia Klaubauf et al. ●
Hercospora Fr. ● Barretomyces Klaubauf et al. ●
Hyalorostratum Raja & Shearer ● Deightoniella S. Hughes ●
Hypodermina Höhn. = Utrechtiana Crous & Quaedvl.
Keinstirschia J. Reid & C. Booth Macgarvieomyces Klaubauf et al. ●
Lollipopaia Inderbitzin ● Neocordana Hern.-Rest. & Crous ●
Mamianiella Höhn. ● Neopyricularia Klaubauf et al. ●
Pseudocryptosporella J. Reid & C. Booth Proxipyricularia Klaubauf et al. ●
Pseudothis Theiss. & Syd. Pseudopyricularia Klaubauf et al. ●
Rabenhorstia Fr. ● Pyricularia Sacc. ●
Savulescua Petr. Xenopyricularia Klaubauf et al. ●
Sphaerognomoniella Naumov & Kusnezowa
Stioclettia Dennis Magnaporthales, genera incertae sedis
Trematovalsa Jacobesco Pseudohalonectria Minoura & T. Muroi ●
Tubakia B. Sutton ●
Valsalnicola D.M. Walker & Rossman ● Ophiostomatales Benny & Kimbr.
Vismaya V.V. Sarma & K.D. Hyde Kathistaceae Malloch & M. Blackw. ●
Kathistes Malloch & M. Blackw. ●
Jobellisiales D’souza & K.D. Hyde Mattirolella S. Colla
Jobellisiaceae Réblová Termitariopsis M. Blackw. et al.
Jobellisia M.E. Barr ●
Ophiostomataceae Nannf.
Magnaporthales Thongk. et al. Ceratocystiopsis H.P. Upadhyay & W.B. Kendr. ●
Magnaporthaceae P.F. Cannon Fragosphaeria Shear ●
Budhanggurabania P. Wong et al. ● Hyalobelemnospora Matsush.
Buergenerula Syd. ● Hyalorhinocladiella H.P. Upadhyay & W.B. Kendr. ●
Bussabanomyces Klaubauf et al. ● Klasterskya Petr.
Ceratosphaerella Huhndorf et al. ● Leptographium Lagerb. & Melin ●
Ceratosphaeria Niessl. ● Ophiostoma Syd. & P. Syd. ●
Clasterosphaeria Sivan. Pesotum JL Crane & Schokn. ●
Clasterosporium Schwein Phialographium H.P. Upadhyay & W.B. Kendr. ●
Clavatisporella K.D. Hyde Raffaelea Arx & Hennebert ●
Gaeumannomyces Arx & D.L. Olivier ● Spumatoria Massee & E.S. Salmon
Harpophora W. Gams ● Subbaromyces Hesselt. ●
Herbampulla Scheuer & Nograsek
Kohlmeyeriopsis Klaubauf et al. ● Ophiostiomatales, genera incertae sedis
Magnaporthe R.A. Krause & R.K. Webster ● Lanspora K.D. Hyde & E.B.G. Jones ●
Magnaporthiopsis J. Luo & N. Zhang ●
Muraeriata Huhndorf et al. ● Tirisporellales Jones et al.
Mycoleptodiscus Ostaz. ● Tirisporellaceae Suetrong et al. ●
Nakataea Hara ● Bacusphaeria Norlailatul et al. ●
Neogaeumannomyces D.Q. Dai & K.D. Hyde ● Tirisporella E.B.G. Jones et al. ●
 Fungal Diversity

Thailandiomyces Pinruan et al. ● Conioscyphales Réblová & Seifert

Conioscyphaceae Réblová & Seifert
Togniniales Senan. et al. Conioscypha Höhn. ●
Togniniaceae Réblová et al. ● = Conioscyphascus Réblová & Seifert
Conidiotheca Réblová & L. Mostert Coronophorales Nannf.
Phaeoacremonium W. Gams et al. ● Bertiaceae Smyk
= TogniniaBerl. ● Bertia De Not. ●
Gaillardiella Pat. ●
Trichosphaeriales M.E. Barr
Trichosphaeriaceae G. Winter Chaetosphaerellaceae Huhndorf et al.
Acanthosphaeria Kirschst. Chaetosphaerella E. Müll. & C. Booth ●
Brachysporium Sacc. Crassochaeta Réblová ●
Collematospora Jeng & Cain Oedemium Link
Coniobrevicolla Réblová = Veramycina Subram.
Cresporhaphis M.B. Aguirre Spinulosphaeria Sivan. ●
Cryptadelphia Réblová & Seifert ●
Eriosphaeria Sacc. Coronophoraceae Höhn.
Fluviostroma Samuels & E. Müll. Coronophora Fuckel ●
Kananascus Nag Raj ●
Khuskia H.J. Huds. ● Nitschkiaceae (Fitzp) Nannf.
Koorchaloma Subram. ● Acanthonitschkea Speg. ●
Neorehmia Höhn. Biciliosporina Subram. & Sekar
Oplothecium Syd. Botryola Bat. & J.L. Bezerra
Rizalia Syd. & P. Syd. Fracchiaea Sacc. ●
Schweinitziella Speg. Groenhiella Jørg. Koch et al.
Setocampanula Sivan. & W.H. Hsieh Janannfeldtia Subram. & Sekar
Trichosphaeria Fuckel ● Lasiosphaeriopsis D. Hawksw. & Sivan.
Unisetosphaeria Pinnoi et al. Loranitschkia Lar.N. Vasiljeva
Neochaetosphaerella Lar.N. Vassiljeva et al.
Diaportheomycetidae, families incertae sedis Neotrotteria Sacc. ●
Distoseptisporaceae K.D. Hyde & McKenzie Nitschkia G.H. Otth ex P. Karst.
Distoseptispora K.D. Hyde et al.● Rhagadostoma Körb.
Rhagadostomella Etayo
Papulosaceae Winka & O.E. Erikss. Tortulomyces Lar.N. Vassiljeva et al.
Brunneosporella V.M. Ranghoo & K.D. Hyde
Fluminicola S.W. Wong et al. Scortechiniaceae Huhndorf et al.
Papulosa Kohlm & Volkm-Kohlm ● Biciliospora Petr. ●
Coronophorella Höhn. ●
Sporidesmiaceae Fr. Cryptosphaerella Sacc. ●
Sporidesmium Link ● Euacanthe Theiss. ●
Neofracchiaea Teng ●
Thyridiaceae O.E. Erikss & J.Z. Yue Scortechinia Sacc. ●
Mattirolia Berl. & Bres. Scortechiniella Arx & E. Müll. ●
= Balzania Speg. Scortechiniellopsis Sivan. ●
= Thyronectroidea Seaver Tympanopsis Starbäck ●
Pleurocytospora Petr.
Thyridium Nitschke ● Coronophorales, genera incertae sedis
Pseudocatenomycopsis Crous & L.A. Shuttlew. ●
Diaportheomycetidae, genera incertae sedis
Platytrachelon Réblová ● Falcocladiales R.H. Perera et al.
Falcocladiaceae Somrithipol et al.
Subclass Hypocreomycetidae O.E. Erikss. & Winka Falcocladium S.F. Silveira et al. ●
Fungal Diversity

Glomerellales Chadef. ex Réblová et al. Ijuhya Starbäck ●

Australiascaceae Réblová & W. Gams Kallichroma Kohlm. & Volkm.-Kohlm. ●
Monilochaetes Halst. ex Harter ● Lasionectria (Sacc) Cooke ●
= Australiasca Sivan. & Alcorn Mycocitrus Möller
Hyalocylindrophora J.L. Crane & Dumont Nectriella Nitschke ex Fuckel
Nectriopsis Maire ●
Glomerellaceae Locq. ex Seifert & W. Gams Ochronectria Rossman & Samuels ●
Colletotrichum Corda ● Ovicuculospora Etayo ●
= Glomerella Spauld. & Schrenk Paranectria Sacc.
Peristomialis (W. Phillips) Boud. ●
Plectosphaerellaceae W. Gams et al. Pronectria Clem.
Acrostalagmus Corda ● Protocreopsis Yoshim Doi ●
Chordomyces Bilanenko, M.L et al. ● Rhopalocladium Schroers et al.
Gibellulopsis Bat. & H. Maia ● Roumegueriella Speg. ●
Lectera P.F. Cannon ● Selinia P. Karst. ●
Musicillium Zare & W Gams ● Spicellum Nicot & Roquebert ●
Plectosphaerella Kleb. ● Stephanonectria Schroers & Samuels ●
= Plectosporium M.E. Palm et al. Stilbocrea Pat. ●
= Spermosporina U. Braun Stromatocrea W.B. Cooke
Sodiomyces A.A. Grum-Grzhim. et al. ● Stromatonectria Jaklitsch & H. Voglmayr ●
Stachylidium Link ● Trichonectria Kirschst. ●
Verticillium Nees ● Vesicladiella Crous & M.J. Wingf. ●
Verrucostoma Hirooka et al. ●
Reticulascaceae Réblová & W. Gams Virgatospora Finley ●
Cylindrotrichum Bonord. ●
= Reticulascus Réblová & W. Gams Clavicipitaceae (Lindau) Earle ex Rogerson
Kylindria DiCosmo et al. ● Aciculosporium I. Miyake ●
Sporoschismopsis Hol-Jech. & Hennebert ● = Albomyces I. Miyake
= Porosphaerellopsis Samuels & E. Müll. = Mitosporium Clem. & Shear
Amphichorda Fr.
Glomerellales, genera, incertae sedis Atkinsonella Diehl. ●
Ascocodinaea Samuels et al. ● Balansia Speg. ●
= Dothichloë G.F. Atk. ●
Hypocreales Lindau Cavimalum Yoshim. Doi et al.
Bionectriaceae Samuels & Rossman Chamaeleomyces Sigler ●
= Spicariaceae Nann. Claviceps Tul. ●
Acremonium Link ● Collarina Giraldo et al. ●
Anthonectria Döbbeler Conoideocrella D. Johnson et al. ●
Aphanotria Döbbeler Corallocytostroma Y.N. Yu & Z.Y. Zhang ●
Battarrina (Sacc.) Clem. & Shear Diploöspora Grove
Bryocentria Döbbeler ● Dussiella Pat. ●
Clonostachys Corda ● Ephelis Fr. ●
= Bionectria Speg. ● Epichloë (Fr.) Tul. & C. Tul. ●
Clibanites (P. Karst.) P. Karst. Epicrea Petr.
Didymostilbe Henn ● Helminthascus Tranzschel
Dimerosporiella Speg. Heteroepichloë E. Tanaka et al. ●
Gliomastix Guég. ● Hypocrella Sacc. ●
Globonectria Etayo = Aschersonia Mont. ●
Gracilistilbella Seifert Konradia Racib.
Halonectria E.B.G. Jones Loculistroma F. Patt & Charles
Heleococcum P.M. Jørg. ● Metacordyceps G.H. Sung et al. ●
Hydropisphaera Dumort ● Metarhiziopsis D.W. Li et al. ●
 Fungal Diversity

Metarhizium Sorokīn ● = Trichodermataceae Fr.

Metapochonia Kepler et al. ● Aphysiostroma Barrasa et al. ●
Moelleriella Bres. ● Arachnocrea Z. Moravec. ●
Mycomalus Möller Dialhypocrea Speg.
Myriogenospora G.F. Atk. ● Escovopsis J.J. Muchovej & Della Lucia ●
Neobarya Lowen ● Escovopsioides H.C. Evans & J.O. Augustin ●
Neoclaviceps J. White et al. ● Hypocreopsis P. Karst. ●
Neocordyceps Kobayasi Hypomyces (Fr.) Tul. ●
Neotyphodium Glenn et al. ● = Cladobotryum Nees
Nigrocornus Ryley & Langdon ● Lichenobarya Etayo et al. ●
Nomuraea Maubl. ● Mycogone Link ●
Orbiocrella D. Johnson et al. ● Payoshaeria W.F. Leong ●
Parepichloë F.J. White Jr. & Reddy ● Protocrea Petch ●
Periglandula U. Steineret al. ● Pseudohypocrea Yoshim. Doi
Pochonia Bat. & O.M. Fonseca ● Rogersonia Samuels & Lodge
Pseudogibellula Samson & H.C. Evans ● Sepedonium Link ●
Pseudomeria G.L. Barron Sibirina G.R.W. Arnold
Regiocrella Chaverri & K.T. Hodge ● Sphaerostilbella (Henn.) Sacc. & D. Sacc ●
Romanoa Thirum. = Gliocladium Corda
Rotiferophthora G.L. Barron ● Sporophagomyces K. Põldmaa & Samuels ●
Samuelsia Chaverri & K.T. Hodge ● Stephanoma Wallr ●
Shimizuomyces Kobayasi ● Trichoderma Pers. ●
Sphacelia Lév. ● = Hypocrea Fr.
Sphaerocordyceps Kobayasi = Sarawakus Lloyd
Stereocrea Syd. & P. Syd.
Tyrannicordyceps Kepler & Spatafora ● Nectriaceae Tul. & C. Tul.
Ustilaginoidea Bref. ● = Tuberculariaceae Fr.
= Villosiclava E. Tanaka & C. Tanaka Albonectria Rossman & Samuels ●
Allantonectria Earle ●
Cordycipitaceae Kreisel ex G.M. Sung et al. Allonectella Petr.
Akanthomyces Lebert ● Antipodium Piroz.
Ascopolyporus Möller ● Aquanectria L. Lombard & Crous ●
Beejasamuha Subram. & Chandrash. Atractium Link ●
Beauveria Vuill. ● Baipadisphaeria Pinruan ●
Coremiopsis Sizova & Suprun Bisifusarium L. Lombard et al. ●
Engyodontium de Hoog ● Calonectria De Not. ●
Cordyceps (Fr.) Link ● = Cylindrocladium Morgan
Gibellula Cavara ● Calostilbe Sacc. & Syd.
Granulomanus de Hoog & Samson = Calostilbella Höhn. ●
Hyperdermium J. White et al. ● Campylocarpon Halleen et al. ●
Isaria Pers. ● Chaetonectrioides Matsush. ●
Lecanicillium W. Gams & Zare ● Chaetopsina Rambelli ●
Microhilum H.Y. Yip & A.C. Rath ● = Chaetopsinectria J Luo & WY Zhuang
Pseudogibellula Samson & H.C. Evans ● Coccinonectria Lombard & Crous ●
Rotiferophthora G.L. Barron ● Corallomycetella Henn. ●
Simplicillium W. Gams & Zare ● Corallonectria C. Herrera & P. Chaverri ●
Syspastospora P.F. Cannon & D. Hawksw. ● Cosmospora Rabenh. ●
Torrubiella Boud. ● = Dialonectria (Sacc.) Cooke
Curvicladiella Decock & Crous
Flammocladiaceae Crous et al. Curvicladium Decock & Crous ●
Flammocladiella Crous et al. ● Cyanochyta Höhn.
Cyanonectria Samuels & Chaverri ●
Hypocreaceae De Not. Cyanophomella Höhn.
Fungal Diversity

Cylindrocladiella Boesew. ● = Chaetodochium Höhn.

= Nectricladiella Crous & C.L. Schoch = Volutellonectria J. Luo & W.Y. Zhuang
Cylinrocarpostylus Xenoacremonium Lombard & Crous ●
Cylindrodendrum Bonord. ● Xenocalonectria Crous & C.L. Schoch ●
Dacryoma Samuels = Xenocylindrocladium Decock et al. ●
Dactylonectria L. Lombard & Crous ● Xenogliocladiopsis Crous & W.B. Kendr. ●
Dematiocladium Allegr. et al. ● Xenonectriella Weese
Fusarium Link ●
= Gibberella Sacc. Niessliaceae Kirschst.
Fusicolla Bonord ● Atronectria Etayo
Geejayessia Schroers et al. ● Circinoniesslia Samuels & M.E. Barr
Gliocephalotrichum J.J. Ellis & Hesselt. ● Eucasphaeria Crous ●
= Leuconectria Rossman et al. Hyaloseta A.W. Ramaley ●
Gliocladiopsis S.B. Saksena ● Malmeomyces Starb.
= Glionectria Crous & C.L. Schoch Melanopsamma Niessl ●
Ilyonectria P. Chaverri & C. Salgado ● Melchioria Penz. & Sacc.
Macroconia (Wollenw.) Gräfenhan et al. ● Miyakeomyces Hara
Mariannaea G. Arnaud ex Samson ● Myrmaeciella Lindau ●
Microcera Desm. ● Niesslia Auersw. ●
= Pseudomicrocera Petch Paraniesslia K.M. Tsui et al.
Nalanthamala Subram. ● Pseudonectriella Petr.
= Rubrinectria Rossman & Samuels Pseudorhynchia Höhn.
Nectria (Fr.) Fr. ● Rosasphaeria Jaklitsch Voglmayr ●
= Tubercularia Tode Taiwanascus Sivan. & H.S. Chang
Nectricladiella Crous & C.L. Schoch ● Trichosphaerella E. Bommer et al. ●
Neonectria Wollenw. ● Valetoniella Höhn.
= Cylindrocarpon Wollenw.
= Heliscus Sacc Ophiocordycipitaceae G.H. Sung et al.
Neocosmospora E.F. Sm. ● Drechmeria W. Gams & H.B. Jansson ●
= Haematonectria Samuels & Nirenberg = Haptocillium W. Gams & Zare
Ophionectria Sacc. ● Harposporium Lohde ●
Paracremonium L. Lombard & Crous ● = Atricordyceps Samuels
Payosphaeria W.F. Leong ● = Podocrella Seaver
Penicillifer Emden ● = Polyrhina Sorokin
= Viridispora Samuels & Rossman Ophiocordyceps Petch ●
Persiciospora P.F. Cannon & D. Hawksw. ● Polycephalomyces Kobayasi ●
Pleogibberella Sacc. = Blistum B. Sutton
Pleonectria Sacc. ● Purpureocillium Luangsa-ard et al. ●
= Zythiostroma Höhn ex Falck Tolypocladium W. Gams ●
Pleurocolla Petr. ● = Chaunopycnis W. Gams
Pseudocosmospora C. Herrera & P. Chaverri ● = Elaphocordyceps G.H. Sung & Spatafora
Pseudonectria Seaver ●
Rectifusarium Lombard et al. ● Stachybotriaceae L. Lombard & Crous
Rugonectria P. Chaverri & Samuels ● Albosynnema E.F. Morris ●
Sarcopodium Ehrenb. ● Myrothecium Tode ●
= Actinostilbe Petch Parasarcopodium Melnik et al. ●
= Lanatonectria Samuels & Rossman Peethambara Subram. & Bhat ●
Stachybotryna Tubaki & T. Yokoy Sarcopodium Ehrenb. ●
Stalagmites Theiss. & Syd. Scopinella Lév. ●
Stylonectria Höhn. ● Stachybotrys Corda ●
Thelonectria P. Chaverri & C.G. Salgado ● = Memnoniella Höhn.
Thyronectria Sacc. ● = Ornatispora K.D. Hyde et al.
Volutella Fr. ● = Melanopsamma Niessl.
 Fungal Diversity

= Valsonectria Speg. Gonatobotrys Corda

Melanospora Corda ●
Tilachlidiaceae Lombard & Crous = Proteophiala Cif.
Septofusidium W. Gams ● Pteridiosperma J.C. Krug & Jeng
Tilachlidium Preuss ● Pustulipora P.F. Cannon
Rhytidospora Jeng & Cain
Hypocreales, genera incertae sedis Setiferotheca Matsush.
Acremoniopsis Giraldo et al. ● Vittatispora P. Chaudhary et al. ●
Alfaria Crous et al. ●
Berkelella (Sacc.) Sacc. Melanosporales genera, incertae sedis
Bulbithecium Udagawa & T Muroi ● Sphaerodes Clem. ●
Emericellopsis J.F.H. Beyma ● Papulaspora Preuss ●
Fecundostilbum T.P. Devi & Chowdhry
Geosmithia J. Pitt ● Microascales Luttr. ex Benny & Kimbr.
Gynonectria Döbbeler = Halosphaeriales Kohlm.
Hapsidospora Malloch & Cain ● Chadefaudiellaceae Faurel & Schotter ex Benny & Kimbr.
Haptospora G.L. Barron Chadefaudiella Faurel & Schotter
Harzia Costantin ● Faurelina Locq-Lin. ●
Illosporiopsis D. Hawksw.
Illosporium Mart. ● Ceratocystidaceae Locq. ex Réblova et al.
Leucosphaerina Arx ● Ambrosiella Brader ex Arx & Hennebert ●
Metadothella Henn. Ceratocystis Ellis & Halst. ●
Microcyclephaeria Bat. ● Chalaropsis Peyronel ●
Munkia Speg. Davidsoniella Z.W. de Beer et al. ●
Mycoarachis Malloch & Cain ● Endoconidiophora Münch ●
Neomunkia Petr ● Huntiella Z.W. de Beer et al. ●
Nigrosabulum Malloch & Cain ● Thielaviopsis Went. ●
Peloronectria Möller Gondwanamycetaceae Réblová et al.
Perennicordyceps Matočec & I. Kušan ● Custingophora Stolk et al. ●
Pseudoacremonium Crous ● Gondwanamyces Marais & M.J. Wingf. ●
Pseudoidriella Crous & R.G. Shivas ●
Pseudomeliola Speg. Graphiaceae De Beer
Pseudomicrodochium B. Sutton Graphium Corda ●
Rodentomyces Doveri et al. ●
Roselliniella Vain ● Halosphaeriaceae E. Müll & Arx ex Kohlm.
Sarocladium W. Gams & D. Hawksw. ● Alisea J. Dupont & E.B.G. Jones ●
Sedecimiella K.L. Pang et al. ● Aniptodera Shearer & M. Miller ●
Septomyrothecium Matsush. Anisostagma K.R.L. Petersen & Jørg. Koch
= Sporothrix Hektoen & C.F. Perkins Antennospora Meyers ●
Stanjemonium W. Gams et al. ● Appendichordella R.G. Johnson et al.
Stilbella Lindau ● Arenariomyces Höhnk ●
Ticonectria Döbbeler Ascosacculus J. Campbell et al. ●
Tilakidium Vaidya et al. Bathyascus Kohlm.
Trichothecium Link ● Carbosphaerella I. Schmidt ●
Valetoniellopsis Samuels & M.E. Barr ● Ceriosporopsis Linder ●
= Bovicornua Jørg Koch & E.B.G. Jones
Melanosporales N. Zhang & M. Blackw. Chadefaudia Feldm.-Maz.
Ceratostomataceae G. Winter Corallicola Volkm.-Kohlm. & Kohlm.
Acrospeira Berk & Broome Corollospora Werderm ●
Arxiomyces P.F. Cannon & D. Hawksw. = Halosigmoidea Nakagiri et al.
Erythrocarpon Zukal = Peritrichospora Linder
Fungal Diversity

Cucullosporella K.D. Hyde & E.B.G. Jones ● Toriella Sakay. et al. ●

Ebullia K.L. Pang ● Trailia G.K. Sutherl.
Fluviatispora K.D. Hyde ● Trichomaris Hibbits et al.
Gesasha Abdel-Wahab & Nagah. ● Tubakiella Sakay. et al. ●
Haiyanga K.L. Pang & E.B.G. Jones ● Tunicatispora K.D. Hyde
Haligena Kohlm. ● = Buxetroldia K.R.L. Petersen & Jørg Koch
Halosarpheia Kohlm. & E. Kohlm. ●
Halosphaeria Linder ● Microascaceae Luttr. ex Malloch
Halosphaeriopsis T.W. Johnson ● Anekabeeja Udaiyan & Hosag.
= Culcitalna Meyers & R.T. Moore Brachyconidiellopsis Decock et al. ●
Havispora K.L. Pang & Vrijmoed ● Canariomyces Arx ●
Iwilsoniella EBG Jones Cephalotrichum Link
Kitesporella Jheng & K.L. Pang ● Doratomyces Corda ●
Kochiella Sakay. et al. ● Echinobotryum Corda
Lautisporiopsis E.B.G. Jones et al. ● Enterocarpus Locq.-Lin. ●
Lignincola Höhnk ● Kernia Nieuwl. ●
Limacospora Jørg. Koch & E.B.G. Jones Knoxdaviesia M.J. Wingf. et al. ●
Luttrellia Shearer Lophotrichus R.K. Benj. ●
Magnisphaera J. Campb. et al. ● Microascus Zukal ●
= Matsusphaeria K.L. Pang & E.B.G. Jones Parascedosporium Gilgado et al. ●
Marinospora A.R. Caval. ● Petriella Curzi ●
= Ceriosporella (Kohlm.) A.R. Caval. Pseudallescheria Negroni & I. Fisch. ●
Moana Kohlm. & Volkm.-Kohlm. = Petriellopsis Gilgado et al.
Morakotiella Sakay. ● Pseudoscopulariopsis M. Sandoval-Denis et al. ●
Nais Kohlm. ● Scedosporium Sacc. ex Castell. & Chalm. ●
Natantispora J. Campb. et al. ● Scopulariopsis Bainier ●
Nautosphaeria E.B.G. Jones ● Tinhaudeus K.L. Pang et al. ●
Neptunella K.L. Pang & E.B.G. Jones ● Wardomyces F.T. Brooks & Hansf. ●
Nereiospora EBG Jones et al. ● Wardomycopsis Udagawa & Furuya ●
= Peritrichospora Linder
Nimbospora Jørg. Koch ● Microascales, genera incertae sedis
Nohea Kohlm. & Volkm.-Kohlm. ● Bisporostilbella Brandsb. & E.F. Morris
Oceanitis Kohlm. ● Cephalotrichiella Crous ●
= Ascosalsum J. Campb. et al. Cornuvesica C.D. Viljoen et al. ●
= Falcatispora K.L. Pang & E.B.G. Jones Gabarnaudia Samson & W. Gams●
Ocostaspora E.B.G. Jones et al. ● Sphaeronaemella P. Karst. ●
Okeanomyces K.L. Pang & E.B.G. Jones ● Sporendocladia G. Arnaud ex Nag Raj & W.B. Kendr. ●
Ondiniella E.B.G. Jones et al. ● Trichurus Clem. ●
Ophiodeira Kohlm. & Volkm.-Kohlm. ● Viennotidia Negru & Verona ex Rogerson
Phaeonectriella R.A. Eaton & E.B.G. Jones ● Vermiculariopsiella Bender●
Praelongicaulis Jones et al. ●
Panorbis J. Campb. et al. ● Pleurotheciales Réblová & Seifert
Pileomyces K.L. Pang & Jheng ● Pleurotheciaceae Réblová & Seifert
Pseudolignincola Chatmala & E.B.G. Jones Adelosphaeria Réblová ●
Remispora Linder ● Brachysporiella Bat. ●
Saagaromyces K.L. Pang & E.B.G. Jones ● Helicoön Morgan ●
= Littispora J. Campb. et al. Melanotrigonum Réblová ●
Sablicola E B.G. Jones et al. ● Phaeoisaria Höhn. ●
Thalassogena Kohlm. & Volkm.-Kohlm. ● Phragmocephala E.W. Mason & S. Hughes ●
Thalespora Chatmala & E.B.G. Jones ● Plagiascoma Réblová & J. Fourn. ●
Tirispora E.B.G. Jones & Vrijmoed ● Pleurotheciella Réblová ●
 Fungal Diversity

Pleurothecium Höhn. ● Lulworthia G.K. Sutherl ●

Sterigmatobotrys Oudem. ● Matsusporium E.B.G. Jones & K.L. Pang ●
Taeniolella S. Hughes ● Moleospora Abdel-Wahab et al. ●
Moromyces Abdel-Wahab et al. ●
Savoryellales Boonyuen et al. Orbimyces Linder ●
Savoryellaceae Jaklitsch & Réblová Rostrupiella Jørg Koch et al. ●
Ascotaiwania Sivan. & H.S. Chang● Spathulospora A.R. Caval. & T.W. Johnson ●
Canalisporium Nawawi & Kuthub. ●
= Ascothailandia Sri-indr. et al. Pisorisporiales Réblová & J. Fourn.
Helicoon Morgan● Pisorisporiaceae Réblová & J. Fourn.
Monotosporella S. Hughes● Achroceratosphaeria Réblová et al. ●
Savoryella E.B.G. Jones & R.A. Eaton● Pisorisporium Réblová & J. Fourn.

Subclass Meliolomycetidae P.M. Kirk & K.D. Hyde

Savoryellales, genera incertae sedis
Carpoligna F.A. Fernández & Huhndorf●
Conioscypha Höhn. ● Meliolales Gäum. ex D. Hawksw. & O.E. Erikss.#
Armatellaceae Hosag.
Flammispora Pinruan et al. ●
Armatella Theiss. & Syd.
Hypocreomycetidae, families incertae sedis
Meliolaceae G.W. Martin ex Hansf.
Etheirophoraceae Rungjindamai et al.
Amazonia Theiss.
Etheirophora Kohlm. & Volkm.-Kohlm. ●
Appendiculella Höhn. ●
Swampomyces Kohlm. & Volkm. ●
Asteridiella McAlpine ●
Cryptomeliola S. Hughes & Piroz. ●
Juncigenaceae E.B.G. Jones et al. Endomeliola S. Hughes & Piroz. ●
Juncigena Kohlm et al. ●
Irenopsis F. Stevens ●
Fulvocentrum E.B.G. Jones & Abdel-Wahab ●
Meliola Fr. ●
Marinokulati E.B.G. Jones & K.L. Pang ●
Moheitospora Abdel-Wahab et al. ●
Subclass Sordariomycetidae O.E. Erikss & Winka

Torpedosporaceae E.B.G. Jones & K.L. Pang Boliniales P.F. Cannon

Glomerulispora Abdel-Wahab & Nagah. ● Boliniaceae Rick
Torpedospora Meyers ● Apiocamarops Samuels & J.D. Rogers
Hypocreomycetidae, genera incertae sedis Apiorhynchostoma Petr. ●
Myrmecridium Arzanlou et al. ● Bolinia (Nitschke) Sacc.
Camaropella Lar.N. Vassiljeva ●
Subclass Lulworthiomycetidae Dayarathne et al. Camarops P. Karst. ●
Cornipulvina Huhndorf et al. ●
Koralionastetales Kohlm. et al. Endoxyla Fuckel ●
Koralionastetaceae Kohlm. & Volkm.-Kohlm. Mollicamarops Lar.N. Vassiljeva
Koralionastes Kohlm. & Volkm.-Kohlm. ● Neohypodiscus J.D. Rogers et al. ●
Pontogeneia Kohlm. ● Pseudovalsaria Spooner ●

Lulworthiales Kohlm. et al. Chaetosphaeriales Huhndorf et al.

Lulworthiaceae Kohlm. et al. Chaetosphaeriaceae Réblová et al.
Cumulospora I. Schmidt ● Ascochalara Réblová
Halazoon Abdel-Aziz et al. ● Brunneodinemasporium Crous & R.F. Castañeda ●
Haloguignardia A. Cribb & J. Cribb ● Catenularia Grove
Hydea K.L. Pang & E.B.G Jones ● Chaetosphaeria Tul. & C. Tul. ●
Kohlmeyeriella E.B.G. Jones et al. ● Chloridium Link ●
Lindra I. Wilson ● Codinaea Maire ●
Lulwoana Kohlm. et al. ● Codinaeopsis Morgan-Jones ●
Lulwoidea Kohlm. et al. ● Craspedodidymum Hol-Jech.
Fungal Diversity

Cryptophiale Piroz. Serenomyces Petr. ●

Dendrophoma Sacc. ●
Dinemasporium Lév. ● Phyllachoraceae Theiss. & H. Syd.
Dictyochaeta Speg. ● Acerviclypeatus Hanlin
Dictyochaetopsis Aramb. & Cabello Apiosphaeria Höhn.
Exserticlava S. Hughes ● Ascovaginospora Fallah et al. ●
Gonytrichum Nees & T. Nees Brobdingnagia K.D. Hyde & P.F. Cannon
Hemicorynespora M.B. Ellis Camarotella Theiss. & Syd.
Infundibulomyces Plaingam et al. ● Coccodiella Hara ●
Kionochaeta P.M. Kirk & B. Sutton ● Cyclodomus Höhn.
Lecythothecium Réblová & Winka ● Deshpandiella Kamat & Ullasa
Melanopsammella Höhn. ● Diachora Müll. Arg.
Menispora Pers. ● = Diachorella Höhn.
Miyoshiella Kawam. Diatractium Syd. & P. Syd. ●
Neopseudolachnella A. Hashim. & Kaz. Tanaka ● Erikssonia Penz. & Sacc.
Phaeostalagmus W. Gams Fremitomyces P.F. Cannon & H.C. Evans
Phialogeniculata Matsush. Geminispora Pat.
Pseudobotrytis Krzemien. & Badura ● Gibellina Pass. Ex Roum.
Pseudodinemasporium A. Hashim. & Kaz. Tanaka ● Imazekia Tak. Kobay. & Y. Kawabe
Pseudolachnea Ranoj. ● Isothea Fr.
Pyrigemmula D. Magyar & R. Shoemaker ● Lichenochora Hafellner
Rattania Prabhug. & Bhat ● Lindauella Rehm
Sporoschisma Berk. & Broome ● Linochora Höhn.
= Melanochaeta E. Müll. et al. Lohwagia Petr.
Striatosphaeria Samuels & E. Müll. ● Maculatifrondes K.D. Hyde
Tainosphaeria F.A. Fernández & Huhndorf ● Malthomyces K.D. Hyde & P.F. Cannon
Thozetella Kuntze ● Muelleromyces Kamat & Anahosur
Umbrinosphaeria Réblová ● Mycohypallage B. Sutton
Zanclospora S. Hughes & W.B. Kendr. Neoflageoletia J. Reid & C. Booth
Zignoëlla Sacc. ● Ophiodothis Sacc.
Ophiodothella (Henn.). Höhn. ●
Helminthosphaeriaceae Samuels et al. Orphnodactylis Malloch & Mallik
Echinosphaeria A.N. Mill. & Huhndorf ● Oswaldina Rangel
Endophragmiella B. Sutton Oxodeora K.D. Hyde & P.F. Cannon
Helminthosphaeria Fuckel ● Parberya C.A. Pearce & K.D. Hyde
Hilberina Huhndorf & A.N. Mill. ● Petrakiella Syd.
Ruzenia O. Hilber ● Phaeochorella Theiss. & Syd.
Synaptospora Cain ● Phycomelaina Kohlm.
Tengiomyces Réblová Phyllachora Nitschke ex Fuckel ●
Phylleutypa Petr.
Chaetosphaeriales genera incertae sedis Phyllocrea Höhn.
Caudatispora J. Fröhl. & K.D. Hyde ● Polystigma DC. ●
Erythromada Huhndorf et al. ● = Polystigmina Sacc.
Lasiosphaeriella Sivan. ● Pseudothiella Petr.
Leptosporella Penz. & Sacc. ● Pseudothiopsella Petr.
Nawawia Marvanová ● Pterosporidium W.H. Ho & K.D. Hyde
Rimaconus Huhndorf et al. ● Rehmiodothis Theiss. & Syd.
Retroa P.F. Cannon
Phyllachorales M.E. Barr Rhodosticta Woron.
Phaeochoraceae K.D. Hyde et al. Rikatlia P.F. Cannon
Cocoicola K.D. Hyde Schizochora Syd. & P. Syd.
Phaeochora Höhn. Sphaerodothella C.A. Pearce & K.D. Hyde
Phaeochoropsis K.D Hyde & P.F. Cannon Sphaerodothis (Sacc. & P. Syd.) Shear ●
 Fungal Diversity

Stigmatula (Sacc.) Syd. & P. Syd. Cercophora Fuckel ●

Stigmochora Theiss. &Syd. Cladorrhinum Sacc. & Marchal ●
Stromaster Höhn. Diffractella Guarro et al.
Telimena Racib. Emblemospora Jeng & J.C. Krug
Telimenella Petr. Eosphaeria Höhn.
Telimenochora Sivan. Fimetariella N. Lundq. ●
Trabutia Sacc. & Roum. Immersiella A.N. Mill. & Huhndorf ●
Tribulatia J.E. Taylor et al. Jugulospora N. Lundq. ●
Uropolystigma Maubl. Lacunospora Cailleux
Vitreostroma P.F. Cannon Lasiosphaeria Ces. & De Not. ●
Zimmermanniella Henn. Mammaria Ces. ex Rabenh. ●
Melanocarpus Arx ●
Phyllachorales genera incertae sedis Periamphispora J.C. Krug
Marinosphaera K.D. Hyde ● Podospora Ces. ●
Mangrovispora K.D. Hyde & Nakagiri ● Pseudocercophora Subram. & Sekar
Phycomelaina Kohlm. ● Rinaldiella Deanna A. Sutton et al. ●
Schizothecium Corda ●
Sordariales Chad. ex D. Hawksw. & O.E. Erikss. Strattonia Cif. ●
Chaetomiaceae G. Winter Thaxteria Sacc.
Achaetomium J.N. Rai et al. ● Triangularia Boedijn ●
Bommerella Marchal Tripterosporella Subram. & Lodha
Boothiella Lodhi & Mirza Zygopleurage Boedijn ●
Botryotrichum Sacc. & Marchal ● Zygospermella Cain ●
Chaetomidium (Zopf) Sacc. ●
Corynascella Arx & Hodges ● Sordariaceae G. Winter
Chaetomiopsis Mustafa & Abdul-Wahid Copromyces N. Lundq. ●
Chaetomium Kunze ● Effetia Bartoli et al.
Crassicarpon Y. Marín et al. ● Gelasinospora Dowding ●
Emilmuelleria Arx ● Guilliermondia Boud.
Farrowia D. Hawksw. ● Neurospora Shear & B.O. Dodge ●
Guanomyces M.C. Gonzáles et al. ● = Chrysonilia Arx
Humicola Traaen ● Pseudoneurospora Dania García et al. ●
Madurella Brumpt ● Sordaria Ces & De Not. ●
Myceliophthora Costantin ● Stellatospora T. Ito & A. Nakagiri
= Corynascus Arx
Staphylotrichum J.A. Mey. & Nicot ● Sordariales, genera incertae sedis
Subramaniula Arx Abyssomyces Kohlm
Taifanglania Z.Q. Liang et al. ● Acanthotheciella Höhn.
Thielavia Zopf ● Ascolacicola Ranghoo & K.D. Hyde ●
Asterosporium Kunze ●
Lasiosphaeriaceae Nannf. Bombardiella Höhn.
Angulimaya Subram & Lodha Cancellidium Tubaki ●
Anopodium Lundq. ● Coronatomyces Dania García et al. ●
Apiosordaria Arx & W. Gams ● Corylomyces Stchigel et al. ●
Apodospora Cain & J.H. Mirza ● Cuspidatispora A. Mill. ●
Apodus Malloch & Cain ● Globosphaeria D. Hawksw.
Arniella Jeng & J.C. Krug Isia D. Hawksw & Manohar
Arnium Nitschke ex G. Winter ● Lasiosphaeris Clem. ●
Bellojisia Réblová ● = Lasiadelphia Réblová & W Gams
Biconiosporella Schaumann Lockerbia K.D. Hyde
Bombardia (Fr.) P. Karst. ● Madurella Brumpt ●
Bombardioidea C. Moreau ex N. Lundqv. ● Nitschkiopsis Nannf. & R. Sant.
Camptosphaeria Fuckel Onygenopsis Henn.
Fungal Diversity

Phaeosporis Clem. Dictyoarthrinium S. Hughes ●

Ramophialophora M. Calduch et al. ● Endocalyx Berk. & Broome ●
Reconditella Matzer & Hafellner Scyphospora L.A. Kantsch. ●
Rhexodenticula W.A. Baker & Morgan-Jones ● Spegazzinia Sacc. ●
Rhexosporium Udagawa & Furuya
Roselliniomyces Matzer & Hafellner Bartaliniaceae Wijayawardene et al.
Roselliniopsis Matzer & Hafellner Bartalinia Tassi ●
Sporidesmiopsis Subram. & Bhat ● Broomella Sacc ●
Utriascus Réblova Dyrithiopsis L. Cai et al. ●
Ypsilonia Lév. Hyalotiella Papendorf ●
Truncatella Steyaert ●
Sordariomycetidae, families incertae sedis Zetiasplozna Nag Raj ●

Batistiaceae Samuels & K.F. Rodrigues$ Beltraniaceae Nann.

Acrostroma Seifert Beltrania Penz. ●
Batistia Cif. ● Beltraniella Subram. ●
= Acrostroma Seifert Beltraniomyces Manohar.
Beltraniopsis Bat. & J.L. Bezerra ●
Sordariomycetidae, genera incertae sedis Parapleurotheciopsis P.M. Kirk ●
Arecacicola Joanne E. Taylor et al. Porobeltraniella Gusmão
Barbatosphaeria Réblová Pseudobeltrania Henn. ●
Bullimyces A. Ferrer et al. ● Subramaniomyces Varghese & V.G. Rao ●
Ceratolenta Réblová ●
Ceratostomella Sacc. ● Cainiaceae J.C. Krug
Chaetosphaerides Matsush. Amphibambusa D.Q. Dai & K.D. Hyde ●
Conlarium F. Liu & L. Cai ● Arecophila K.D. Hyde ●
Hanliniomyces Raja & Shearer Atrotorquata Kohlm. & Volkm.-Kohlm. ●
Hydromelitis A. Ferrer et al. ● Cainia Arx & E. Müll. ●
Lentomitella Höhn. ● Seynesia Sacc. ●
Mirannulata Huhndorf et al. ●
Menisporopascus Matsush. Clypeosphaeriaceae G. Winter
Merugia Rogerson & Samuels Apioclypea K.D. Hyde ●
Mycomedusiospora G.C. Carroll & Munk Brunneiapiospora K.D. Hyde et al. ●
Nigromammilla K.D. Hyde & J. Fröhl. Clypeosphaeria Fuckel ●
Phaeotrichosphaeria Sivan. Crassoascus Checa et al. ●
Phragmodiscus Hansf. Palmomyces K.D. Hyde et al. ●
Plagiosphaera Petr.
Rhodoveronaea Arzanlou et al. ● Coniocessiaceae Asgari & Zare
Riomyces A. Ferrer et al. ● Coniocessia Dania García et al. ●
Spadicoides S. Hughes ●
Xylomelasma Réblová ● Diatrypaceae Nitschke
Woswasia Jaklitsch et al. Anthostoma Nitschke ●
Cryptosphaeria Ces & De Not. ●
Subclass Xylariomycetidae O.E. Erikss & Winka Cryptovalsa Ces & De Not. ex Fuckel ●
Xylariales Nannf. Diatrype Fr. ●
Diatrypella (Ces & De Not.) De Not. ●
Amphisphaeriaceae G. Winter Diatrypasimilis J.J. Zhou & Kohlm. ●
Amphisphaeria Ces & De Not. ● Echinomyces Rappaz
Lepteutypa Petr. ● Eutypa Tul. & C. Tul. ●
Eutypella (Nitschke) Sacc. ●
Apiosporaceae K.D. Hyde et al. Leptoperidia Rappaz
Appendicospora K.D. Hyde ● Monosporascus Pollack & Uecker
Arthrinium Kunze ● Pedumispora K.D. Hyde & E.B.G. Jones ●
 Fungal Diversity

Peroneutypa Berl. ● Leiosphaerella Hohn. ●

Quaternaria Tul. & C. Tul. Pseudomassaria Jacz. ●
Requienellaceae Boise
Hyponectriaceae Petr. Acrocordiella O.E. Erikss. ●
Apiothyrium Petr. Requienella Fabre ●
Arecomyces K.D. Hyde Robillardaceae Crous
Arwidssonia B. Erikss. Robillarda Sacc. ●
Cesatiella Sacc. Sporocadaceae Corda
Chamaeascus L. Holm et al. = Discosiaceae Maharachch. & K.D. Hyde
Charonectria Sacc. Adisciso Kaz. Tanaka ●
Discosphaerina Höhn. ● Discosia Lib. ●
Exarmidium P. Karst. Discostroma Clem. ●
Frondicola K.D. Hyde Sarcostroma Cooke ●
Hyponectria Sacc. ● Seimatosporium Corda ●
Micronectria Speg. Strickeria Körb ●
Papilionovela Aptroot
Pellucida Dulym. et al. ● Vialaeaceae P.F. Cannon
Physalospora Niessl ● Vialaea Sacc. ●
Phragmitensis M.K.M. Wong et al.
Rhachidicola K.D. Hyde & J. Fröhl. Xylariaceae Tul. & C. Tul.
Xenothecium Höhn. “Hypoxyloideae”
Annulohypoxylon Y.M. Ju et al.●
Iodosphaeriaceae O. Hilber Anthocanalis Daranagama et al. ●
Iodosphaeria Samuels ● Biscogniauxia Kuntze ●
Calceomyces Udagawa & S Ueda ●
Lopadostomataceae Daranagama & K.D. Hyde Camillea Fr. ●
Creosphaeria Theiss. ● Daldinia Ces. & De Not. ●
Lopadostoma (Nitschke) Traverso ● Durotheca Læssøe et al. ●
Entonaema Möller ●
Melogrammataceae G. Winter Graphostroma Piroz. ●
Melogramma Fr. ● Hypoxylon Bull. ●
Induratia Samuels et al. ●
Microdochiaceae Hern.-Restr., Crous & J.Z. Groenew. Nodulisporium Preuss ●
Idriella P.E. Nelson & S. Wilh. ● Obolarina Pouzar ●
Microdochium Syd. ● Phylacia Lév. ●
Selenodriella R.F. Castañeda & W.B. Kendr. ● Pyrenomyxa Morgan ●
Rhopalostroma D. Hawksw. ●
Rostrohypoxylon J. Fourn. & M. Stadler ●
Myelospermataceae K.D. Hyde & S.W. Wong Ruwenzoria J. Fourn. et al. ●
Myelosperma Syd. & P. Syd. Thamnomyces Ehrenb. ●
Theissenia Maubl. ●
Pestalotiopsidaceae Maharachch. & K.D. Hyde Thuemenella Penz. & Sacc. ●
Ciliochorella Syd. ● Vivantia J.D. Rogers et al.
Monochaetia (Sacc.) Allesch. ●
Neopestalotiopsis Maharachch. et al. ● “Xylarioideae”
Pestalotiopsis Steyaert ● Amphirosellinia Y.M. Ju et al.●
Pseudopestalotiopsis Maharachch. et al. ● Arthroxylaria Seifert & W Gams ●
Seiridium Nees ● Ascotricha Berk. ●
Phlogicylindriaceae Senan. & K.D. Hyde Astrocystis Berk. & Broome ●
CiferriascoseaSenanayake et al. ● Coniolariella Dania García et al. ●
Phlogicylindrium Crous et al. ● Collodiscula I. Hino & Katum. ●
Pseudomassariaceae Senan. & K.D. Hyde Emarcea Duong et al. ●
Fungal Diversity

Entoleuca Syd. ● Striatodecospora D.Q. Zhou et al.

Euepixylon Füisting ● Stromatoneurospora S.C. Jong & E.E. Davis
Halorosellinia Whalley et al. ● Virgaria Nees ●
Helicogermslita Lodha & D. Hawksw. Wawelia Namysl.
Hypocopra (Fr) J. Kickx f ● Whalleya J.D. Rogers et al. ●
Hypocreodendron Henn. Xylocrea Möller
Kretzschmaria Fr. ●
Leprieuria Laessøe et al. Xylariales, genera incertae sedis
Nemania Gray ● Adomia S. Schatz
Ophiorosellinia J.D. Rogers et al. Ascotrichella Valldos. & Guarro
Podosordaria Ellis. & Holw. ● Castanediella Hern.-Restr. & Crous ●
Poroleprieuria M.C. González et al. ● Diamantinia A.N. Mill. et al. ●
Poronia Willd. ● Fassia Dennis
Rosellinia De Not. ● Idriellopsis Hern.-Restr. & Crous ●
Stilbohypoxylon Henn. ● Lanceispora Nakagiri et al. ●
Xylaria Hill ex Schrank ● Lasiobertia Sivan. ●
Xylotumulus J.D. Rogers et al. Leiosphaerella Höhn. ●
Linocarpon Syd. & P. Syd. ●
Other Xylariaceae genera with conidial states not belong to Muscodor Worapong et al. ●
nodulisporium–like or geniculosporium–like or with Neoidriella Hern.-Restr. & Crous ●
unkown conidial states Neolinocarpon K.D. Hyde ●
Anthostomella Sacc. ● Oxydothis Penz. & Sacc. ●
Appendixia B.S. Lu & K.D. Hyde Palmicola K.D. Hyde
Areolospora S.C. Jong & E.E. Davis Paraidriella Hern.-Restr. & Crous ●
Barrmaelia Rappaz ● Pidoplitchkoviella Kiril. ●
Brunneiperidium Daranagama et al. ● Plectosphaera Theiss. ●
Cannonia J.E. Taylor & K.D. Hyde Polyancora Voglmayr & Yule ●
Chaenocarpus Rebent. Pulmosphaeria Joanne E. Taylor et al.
Chlorostroma A.N. Mill. et al. Sporidesmina Subram. & Bhat
Cyanopulvis J. Fröhl. & K.D. Hyde ● Tristratiperidium Daranagama et al. ●
Engleromyces Henn. Subramaniomyces Varghese & V.G. Rao ●
Fasciatispora K.D. Hyde ● Yuea O.E. Erikss.
Gigantospora B.S. Lu & K.D. Hyde
Guestia G.J.D. Sm. & K.D. Hyde Sordariomycetes, orders incertae sedis
Jumillera J.D. Rogers et al. ●
Kretzschmariella Viégas Amplistromatales D’souza et al.
Leptomassaria Petr. Amplistromataceae Huhndorf et al.
Libertella Desm. ● Acidothrix Hujslová & M. Kolařík ●
Lunatiannulus Daranagama et al. ● Amplistroma Huhndorf et al. ●
Myconeesia Kirschst Wallrothiella Sacc. ●
Nipicola K.D. Hyde = Pseudogliomastix W. Gams
Occultitheca J.D. Rogers & Y.M. Ju = Zignoina Cooke
Pandanicola K.D. Hyde
Paramphisphaeria F.A. Fernández et al. Catabotrydaceae Petr. ex M.E. Barr
Paucithecium Lloyd Catabotrys Theiss. & Syd. ●
Pyriformiascoma Daranagama et al. ●
Sabalicola K.D. Hyde Coniochaetales Huhndorf et al.
Sarcoxylon Cooke● Coniochaetaceae Malloch & Cain
Seynesia Sacc. Barrina A.W. Ramaley ●
Spirodecospora B.S. Lu et al. Coniochaeta (Sacc.) Cooke ●
Squamotubera Henn.
Steganopycnis Syd. & P. Syd. Cordanales M. Hern.-Rest. & Crous
 Fungal Diversity

Cordanaceae Nann. Caleutypa Petr.

Cordana Preuss ● Calosphaeriopsis Petr.
= Porosphaerella E. Müll. & Samuels Caproniella Berl.
Chaetoamphisphaeria Hara
Spathulosporales Kohlm. Ciliofusospora Bat. & J.L. Bezerra
Hispidicarpomycetaceae Nakagiri Calcarisporium Preuss
Hispidicarpomyces Nakagiri Clypeoceriospora Sousa da Câmara
Clypeosphaerulina Sousa da Câmara
Spathulosporaceae Kohlm. Crinigera Schmidt
Retrostium Nakagiri & Tad Ito Cryptoascus Petri
Spathulospora A.R. Caval. & T.W. Johnson ● (only in part) Cryptomycella Höhn.
Cryptomycina Höhn.
Sordariomycetes, families incertae sedis Cucurbitopsis Bat. & Cif.
Cephalothecaceae Höhn. Curvatispora V.V. Sarma & K.D. Hyde
Albertiniella Kirschst. ● Dasysphaeria Speg.
Cephalotheca Fuckel ● Delpinoëlla Sacc.
Cryptendoxyla Malloch & Cain ● Diacrochordon Petr.
Phialemonium W. Gams & McGinnis ● Digicatenosporium S.M. Leão et al.
Dryosphaera Jørg. Koch & E.B.G. Jones
Lautosporaceae Kohlm. et al. Duradens Samuels & Rogerson●
Lautospora K. D. Hyde & E.B.G. Jones Endoxylina Romell
Esfandiariomyces Ershad
Sordariomycetes, genera incertae sedis Frondisphaera K.D. Hyde
Acerbiella Sacc. Glabrotheca Chardόn
Acrospermoides Miller & G.E. Thomps. Hapsidascus Kohlm. & Volkm.-Kohlm.
Ameromassaria Hara Heliastrum Petr.
Amphisphaerellula Gucevič Hyaloderma Speg.
Amphisphaerina Höhn. Hydronectria Kirschst.
Amphorulopsis Petr. Hypotrachynicola Etayo
Amylis Speg. Immersisphaeria Jaklitsch.
Anaexserticlava T.S. Santa Izabel et al. Iraniella Petr.
Anthostomaria (Sacc.) Theiss. & Syd. Imicles Shoemaker & Hambl.
Anthostomellina L.A. Kantsch. Konenia Hara
Apharia Bonord. Kravtzevia Schwartzman
Apodothina Petr. Kurssanovia Kravtzev
Apogaeumannomyces Matsush. Lecythiomyces Doweld
Aquadulciospora Fallah & Shearer = Lecythium Zukal
Aquasphaeria K.D. Hyde Leptosacca Syd.
Aropsiclus Kohlm. & Volkm.-Kohlm. Leptosphaerella Speg.
Ascorhiza Lecht.-Trnka Leptosporina Chardón
Ascoyunnania L. Cai & K.D. Hyde Liberomyces Pažoutová et al. ●
Assoa Urries Linocarpon Syd. & P. Syd. ●
Atrogeniculata J.S. Monteiro et al. Lyonella Syd.
Aulospora Speg. Mangrovispora K.D. Hyde & Nagakiri
Azbukinia Lar.N. Vassiljeva Marisolaris Jørg. Koch & E.B.G. Jones
Bactrodesmiastrum Hol.-Jech. ● Melanographium Sacc.
Bactrosphaeria Penz. & Sacc. Melomastia Nitschke ex Sacc.
Biflua J.Koch & EBG Jones Microcyclephaeria Bat.
Biporispora J.D. Rogers et al. Monotosporella S. Hughes ●
Bombardiastrum Pat. Mycothermus D.O. Natvig et al.
Brenesiella Syd. Naumovela Kravtzev
Byrsomyces Cavalc. Natantiella R blová ●
Byssotheciella Petr. Neocryptospora Petr.
Fungal Diversity

Neoeriomycopsis Crous & M.J. Wingf. ● Tunstallia Agnihothr.

Neolamya Theiss. & Syd. Vleugelia J. Reid & C. Booth
Neolinocarpon K.D. Hyde ● Zalerion R.T. Moore & Meyers ●
Neophysalospora Crous & M.J. Wingf. ●
Neothyridaria Petr. Descriptions and notes on families
Nigrospora Zimm. ● Amphisphaeriaceae G. Winter [as ‘Amphisphaerieae’],
Ophiomassaria Jacz. Rabenh. Krypt.-Fl., Edn 2 (Leipzig) 1.2: 259 (1885)
Orcadia GK Sutherl Facesoffungi number: FoF 00673
Paoayensis Cabanela et al. Saprobic, hemibiotrophic or necrotrophic on leaves, twigs
Paramicrodochium Hern.-Restr. & Crous ● and branches of deciduous shrubs and trees, conifers or
Pareutypella Y.M. Ju & J.D. Rogers monocotyledons, appearing as slightly raised, black dots
Phialemoniopsis Perdomo et al. ● on host surface, often surrounded by a darkened area.
Phomatosporella Tak. Kobay & K. Sasaki Sexual morph: Pseudostromata when present made up
Phyllocelis Syd. of host cells and brown to black fungal hyphae, solitary,
Pleocryptospora J. Reid & C. Booth uni- to bi-loculate, hemisphaerical, initially appearing as
Pleosphaeria Speg. raised, pale brown areas, with small, black dots at the
Porodiscus Lloyd center, becoming dark at maturity, glabrous. Ascomata
Protocucurbitaria Naumov perithecial, scattered to clustered, immersed in host cortex,
Pulvinaria Bon. becoming raised, subglobose to lenticular, glabrous, dark
Pumilus Viala & Marsais brown to black, vegetative hyphae surrounding the locules,
Rehmiomycella E. Müll. ostioles individual, central. Papilla erumpent through host
Rhamphosphaeria Kirschst. surface, internally covered by hyaline, filamentous
Rhizophila K.D. Hyde & E.B.G. Jones periphyses. Peridium of unequal thickness, thinner at the
Rhopographella (Henn.) Sacc. & Trotter base, thicker towards the mid-upper section of the
Rhynchosphaeria (Sacc.) Berl. ascomata, comprising several layers of dark brown pseu-
Rivulicola K.D. Hyde doparenchymatous cells, outer layer of thick-walled, brown
Romellina Petr. cells of textura prismatica, inner layer of flattened, hyaline
Saccardoëlla Speg. cells of textura prismatica. Hamathecium comprising nu-
Sarcopyrenia Nyl. merous, septate, filamentous paraphyses. Asci 8-spored,
Sartorya Vuill. unitunicate, cylindrical, short pedicellate, apex rounded,
Scharifia Petr. with J+ or J- apical ring. Ascospores overlapping
Scoliocarpon Nyl. uniseriate, light to dark brown, ellipsoidal to fusiform, 1-
Scotiosphaeria Sivan. septate. Asexual morph: Coelomycetous. Conidiomata
Selenosporella G. Arnaud ex MacGarvie solitary or aggregated, globose, dark brown. Peridium
Servazziella J. Reid & C. Booth comprising thick-walled, septate, brown mycelium.
Sporoctomorpha J.V. Almeida & Sousa da Câmara Conidiophores dichotomously branched, septate, thick-
Stanjehughesia Subram. ● walled, smooth, hyaline. Conidiogenous cells phialidic,
Stearophora L. Mangin & Viala elongated, wide at the base and narrow at the tip, thin-
Stegophorella Petr. walled, hyaline. Conidia hyaline, 1-celled, smooth-walled,
Stellosetifera Matsush. elongate to fusiform, narrow at both ends.
Stereosphaeria Kirschst. ● Type: Amphisphaeria Ces. & De Not.
Stomatogenella Petr. Notes: The family Amphisphaeriaceae was introduced by
Stromatographium Höhn. Winter (1885) as ‘Amphisphaerieae’ and later established as
Sungaiicola Fryar & K.D. Hyde Amphisphaeriaceae to accommodate the type genus
Synsphaeria Bon. Amphisphaeria (typified by A. umbrina. Ces. & De Not) and
Tamsiniella S.W. Wong et al. similar genera, viz. Ohleria, Trematosphaeria, Caryospora,
Tectonidula Réblová Winteria and Strickeria (Winter 1887) The family is charac-
Teracosphaeria R blová & Seifert ● terized by partially immersed or erumpent, perithecial
Thelidiella Fink. ascomata, unitunicate, cylindrical asci, with J+ or J- apical
Thyridella (Sacc.) Sacc. rings and pale to dark brown, 1-septate ascospores (Kang
Thyrotheca Kirschst. et al. 1998; Senanayake et al. 2015). The species of the family
Trichospermella Speg. were subsequently included in various other families
Trichosphaeropsis Bat. & Nasc. (Clements and Shear 1931; von Arx and Müller 1954;
 Fungal Diversity

Müller and von Arx 1973), leading to a doubt about its exact However, Senanayake et al. (2015) illustrated the sexual
taxonomic position (Munk 1957; Kang et al. 1999a). and asexual morphs of Amphisphaeria sorbi, the former
Hence, Amphisphaeriaceae was reintroduced by Müller being introduced in Liu et al. (2015).
and von Arx (1962) based on the presence of a small Amphisphaeria species are thought to be widely dis-
iodine positive ring or disc in the apex of the asci and tributed among both tropical and temperate regions hav-
in having ascomata immersed under a clypeus (Kang et al. ing mostly been recorded in Europe, and others across
1999a; Maharachchikumbura et al. 2015). Samuels et al. the Americas, South East Asia and South India (Liu
(1987) suggested that the genera linked to Pestalotia-like et al. 2015; Farr and Rossman 2016). However, since
asexual morphs, Amphisphaeria, Broomella, Discostroma, the genus is quite confused, its distribution needs to be
Lepteutypa and Pestalosphaeria should be kept as confirmed by species identifications supported by mo-
Amphisphaeriaceae (sensu stricto). Hawksworth et al. lecular data.
(1995) listed Amphisphaeriaceae as a heterogeneous sexu-
al morph family with 36 genera. Of these, Kang et al. Amplistromataceae Huhndorf et al., Mycologia 101(6):
(1999a) accepted only Amphisphaeria, Broomella, 905 (2009).
Discostroma, Ellurema, Griphosphaerioma, Neobroomella, Facesoffungi number: FoF 00618
Blogiascospora, Lepteutypa, Paracainiel la and Saprobic on bark or wood. Sexual morph: Stromata su-
Pestalosphaeria based on a study of molecular and mor- perficial, turbinate, obovoid to irregularly pulvinate, texture
phological data. As a result of various studies, previously soft or firm; or stromata absent and perithecia separate to
included genera in Amphisphaeriaceae have been moved clustered, with hyphal subiculum or absent. Ascomata globose
to other taxonomic groups such as Cainiaceae or subglobose, polystichous or monostichous, with long
Clypeosphaeriaceae, Phyllachoraceae, Pleurotremataceae necks. Hamathecium comprising hyaline, abundant, filamen-
and Xylariaceae (Krug 1977; Kang et al. 1998, 1999b). tous paraphyses. Asci 8–spored, numerous, unitunicate, cylin-
Senanayake et al. (2015) accepted only the type genus drical to clavate, apical ring not bluing in Melzer’s reagent.
Amphisphaeria excluding all the other genera based on Ascospores uniseriate, hyaline, globose. Asexual morph:
their morphology and phylogeny. Some were transferred Undetermined.
to Bartaliniaceae, Discosiaceae, Iodosphaeriaceae, Type: Amplistroma Huhndorf et al.
Pestalotiopsidaceae and Phlogicylindriaceae, while most Notes: This family was introduced by Huhndorf et al.
genera were listed as Amphisphaeriales genera incertae (2009) to accommodate Amplistroma and Wallrothiella.
sedis. Amphisphaeriaceae is currently classified in the or- Two genera were recognized in the family based on
der Xylariales and is known to share a close relationship molecular and morphological data (Huhndorf et al.
with Cainiaceae, Clypeosphaeriaceae, and 2009). Wallrothiella congregata (Wallr.) Sacc., and spe-
Hyponectriaceae (Jeewon 2001), as well as with cies of Amplistroma share cylindro-clavate, stipitate asci,
Apiosporaceae (Crous and Groenewald 2013; Senanayake minute, globose ascospores and wide paraphyses that
et al. 2015). As the family is presently monotypic, a fam- are long-tapering above the asci. Amplistroma and
ily description only is provided. Wallrothiella differ mainly in the size amount and tex-
Amphisphaeria Ces. & De Not., Comm. Soc. crittog. Ital. ture of stromatal structures (Huhndorf et al. 2009).
1(4): 223 (1863) Phylogenetic analyses of 28S rDNA group these taxa
Facesoffungi number: FoF 02099; Figs 3, 4 in a well-supported clade distinct from known orders
Type species: Amphisphaeria umbrina (Fr.) De Not., Sfer. within Sordariomycetidae. Amplistromataceae resolves
Ital.: 69 (1863) as a monophyletic clade with strong bootstrap and
Notes: Amphisphaeria was introduced by Cesati and Bayesian support with LSU data, but its sister relation-
De Notaris (1863), with no generic type designated ship with Chaetosphaeriales is not supported. The fam-
(Wang et al. 2004). Later Petrak (1923) proposed ily was placed within Sordariomycetidae incertae sedis
A. umbrina (previously introduced as Sphaeria umbrina) (Huhndorf et al. 2009). Maharachchikumbura et al.
as the lectotype of this genus. The characterization of (2015) showed that Amplistroma clustered with
Amphisphaeria and subsequent epitypification of the ge- Wallrothiella as a sister group to Catabotrys deciduum
neric type A. umbrina, and its current phylogenetic sta- in Catabotrydaceae, thus they introduced a new order
tus has been reviewed in Senanayake et al. (2015). Amplistromatales to accommodate these families.
Most of the confusion regarding the genus arises from Hujslová et al. (2014) included Acidothrix, an acidophil-
the lack of sexual-asexual morph connections (Kang ic, asexual, soil fungus into this family. The morpholo-
et al. 1998). The asexual morph Bleptosporium has been gy of Acidothrix acidophila Hujslová & M. Kolařík and
linked to Amphisphaeria (Nag Raj 1977; Hyde et al. the acrodontium-like asexual morphs occurring in
2011), but has not been proven based on sequence data. Amplistroma species have similar characters. However
Fungal Diversity

Fig. 3 Amphisphaeria sorbi (Material examined: ITALY, Province of pseudostroma with ostioles c Biloculate pseudostromata d Section of
Trento [TN], Dimaro, Folgarida, on branch of Sorbus aucuparia ascoma e Peridium f Paraphyses g–i Immature asci j–l Mature asci m–o
(Rosaceae), 2 August 2013, Erio Camporesi IT 1400 (MFLU 14-0797, Ascospores p Sheath surrounding ascospore. Scale bars: a–c = not to
holotype) a Appearance of pseudostroma on host b Close up of scale, d, e = 10 μm, f = 20 μm, g–p = 50 μm

in the phylogenetic analysis Acidothrix acidophila Facesoffungi number: FoF 00618; Fig. 5
placed outside both Amplistroma and Wallrothiella, Saprobic on bark or wood. Sexual morph: Stromata su-
therefore Hujslová et al. (2014) introduced this as a perficial, turbinate, obovoid to irregularly pulvinate, texture
new genus. soft or firm. Ascomata globose or subglobose, arranged in
Amplistroma Huhndorf et al., Mycologia 101(6): 907 multiple layers, with long necks. Peridium outwardly com-
(2009) prising thin-walled, light brown cells of textura intricata and
 Fungal Diversity

Fig. 4 Asexual morph of Amphisphaeria sorbi (introduced in Senanayake et al. 2015) a Conidiomata on MEA b–d Conidia attached to conidiophores
e-g Conidia. Scale bars: a = 1000 μm, b–g = 20 μm

inwardly comprising thin-walled, hyaline cells of textura necks, that are either erumpent from the stromatal surface or
intricata. Hamathecium comprising hyaline, abundant, fila- form bumps or protuberances. Amplistroma was placed in
mentous paraphyses. Asci 8-spored, unitunicate, cylindrical Amplistromataceae by Huhndorf et al. (2009) based on mo-
to clavate, pedicellate, ascus apical ring minute to inconspic- lecular and morphological data.
uous, not staining in Melzer’s reagent. Ascospores uniseriate,
hyaline, globose, thick-walled. Asexual morph:
Hyphomycetous, acrodontium-like in culture and occurring
on young stromata and near base of older stromata. Other genera included
Conidiophores arising vertically, sometimes horizontally, hy-
aline to light brown, not strongly differentiated, with whorls of Acidothrix Hujslová & M. Kolařík, in Hujslova et al., Mycol.
secondary and tertiary branching, arising vertically along the Progr. 13(3): 824 (2014)
length. Conidiogenous cells arising in verticils or whorls on Type species: Acidothrix acidophila Hujslová & M.
the branches, lageniform to ampulliform, apex forming elon- Kolařík in Hujslova et al., Mycol. Progr. 13(3): 824 (2014)
gate rachis, proliferating sympodially, flexuous, with denticu- Wallrothiella Sacc., Syll. fung. (Abellini) 1: 455 (1882).
late, blunt, conidiogenous pegs. Conidia light brown, pyri- Type species: Wallrothiella congregata (Wallr.) Sacc., Syll.
form to obovoid, with basal frill (Huhndorf et al. 2009). fung. (Abellini) 1: 455 (1882).
Type species: Amplistroma carolinianum Huhndorf et al.,
Mycologia 101(6): 907 (2009).
Notes: Amplistroma was described as a new genus for Fig. 5 Amplistroma carolinianum (Material examined: USA, North„
A. carolinianum Huhndorf et al. as the type along with Carolina, Macon County, Ellicott Rock Trail, off Bull Pen Road, on
bark, 14 October 1990, Y. Doi, A.Y. Rossman, G.J. Samuels (BPI
A. diminutisporum Huhndorf et al., A. guianense Huhndorf
878925, holotype) a Herbarium package b Stromata c Black ostiolar
et al., A. hallingii Huhndorf et al., A. ravum Huhndorf et al., necks in stromata surface d Cross section through stromata e
A. tartareum Huhndorf & Samuels and A. xylarioides (Pat.) Arrangement of perithecia into layers in stromatal tissue f Embedded
Huhndorf & Samuels (Huhndorf et al. 2009). Amplistroma perithecia in stromatal tissue g Ascoma h Peridium i, j Mature asci k
Ascospores l Apex of the asci lacking apical ring m, n Dried culture.1,
erinaceum Checa et al. was recently added (Checa et al.
o Conidia p, q Dried culture 2 s Conidia t Culture details. Scale bars:
2014). Species of Amplistroma are distinguished by large stro- b = 2000 μm, c, d = 1000 μm e, f = 300 μm, g = 100 μm, h–j = 20 μm, k,
mata of textura intricata with polystichous ascomata and long o, s = 5 μm l = 10 μm
Fungal Diversity

Key to genera of Amplistromataceae 1. Reported with sexual morph ……………………….. 2

1. Ascomata arranged in multiple layers with long necks,
1. Reported with only asexual morph ………….. Acidothrix that are either erumpent from the stromatal surface or form
 Fungal Diversity

protuberances, forming large stromata of textura intricata supported by molecular analyses using LSU rDNA datasets
…………..…………..…………..………….. Amplistroma (Campbell and Shearer 2004; Huhndorf et al. 2004a). The
2. Ascomata with long-necks and develop individually or family Annulatascaceae is polyphyletic (Raja et al. 2003;
are gregarious on the substrate, but do not form large stromata Campbell and Shearer 2004; Huhndorf et al. 2004a;
………….…….…….…….…….……...…….. Wallrothiella Vijaykrishna et al. 2005). Clohiesia was placed in
Annulatascaceae by Tsui et al. (1998) based on morphology,
Annulatascaceae S.W. Wong et al., Syst. Ascom. 16(1–2): but was shown to have phylogenetic affinity with
18 (1998) Lasiosphaeriaceae and Sordariaceae (Raja et al. 2003;
Facesoffungi number: FoF 01212 Duong et al. 2004). The genus Chaetorostrum produces a
Saprobic on submerged wood, sometimes on bamboo and taeniolella-like asexual morph in culture and is the first record
on other substrates in terrestrial habitats. Sexual morph: of an asexual morph for Annulatascaceae (Zelski et al. 2011).
Ascomata perithecial, solitary or gregarious, superficial, The family currently comprises 18 genera and members of this
erumpent, semi-immersed or immersed, black, pale brown to family are reported from temperate and tropical regions and in
brown, or reddish brown, unilocular, rarely clypeate, globose both terrestrial as well as freshwater habitats. All genera need
to subglobose, ellipsoidal. Necks black or hyaline, glabrous, further study at the molecular level to clarify phylogenetic
rarely with setae, hyphae or hairs. Peridium coriaceous or relatedness and possibly determine their evolution.
membranous composed of textura angularis, textura Annulatascus K.D. Hyde, Aust. Syst. Bot. 5(1): 118
intricata, textura epidermoidea, textura prismatica or pseudo- (1992)
parenchymatous cells or compressed cells. Hamathecium with Facesoffungi number: FoF 01213; Figs 6, 7
numerous, septate, branched or unbranched, tapering paraph- Saprobic on submerged wood. Sexual morph: Ascomata
yses. Asci 8-spored, unitunicate, cylindrical, pedicellate, usu- clustered or separate, brown to black, superficial, semi-
ally with a massive J-, refractive, discoid, or wedge-shaped, immersed or immersed, solitary or gregarious, globose or
apical ring. Ascospores usually uniseriate, sometimes overlap- subglobose, immersed or semi-immersed, carbonaceous or
ping, hyaline to brown, unicellular or septate, septa mostly coriaceous, ostiolate, periphysate. Peridium comprising sev-
transverse, sometimes with longitudinal and transverse septa. eral layers of cells of textura angularis, or pseudoparenchy-
Appendages, mucilaginous sheath and germ pores may be matous, or melanized angular cells. Hamathecium comprising
present. Asexual morph: Hyphomycetous. taeniolella-like paraphyses which are hyaline, septate and taper distally. Asci
for Chaetorostrum where Conidiophores are micronematous, 8-spored, unitunicate, cylindrical, pedicellate, with a massive
mononematous. Conidia monoblastic, elongate cylindrical, bipartite apical ring. Ascospores unicellular to multi-septate,
trans-septate, euseptate, brown, paler near apex. Conidial uniseriate to overlapping biseriate, ellipsoidal or fusiform or
secession schizolytic lunate or citriform, hyaline to pale brown, verrucose or
Type: Annulatascus K.D. Hyde smooth, guttulate or non-guttulate, with or without sheath
Notes: The family Annulatascaceae was introduced by and appendages. Asexual morph: Undetermined.
Wong et al. (1998) to accommodate saprobic, lignicolous, Type species: Annulatascus velatispora K.D. Hyde, Aust.
freshwater genera of ascomycetes viz. Annulatascus, Syst. Bot. 5(1): 118 (1992).
Ascotaiwania, Clohiesia, Frondicola, Proboscispora and Notes: This genus is cosmopolitan, and has been described
Submersisphaeria. Wong et al. (1998) considered that on submerged substrata, such as submerged decaying wood,
Annulatascus could be best placed in Lasiosphaeriaceae due stems of Phragmites australis, and dead petioles of Licuala
to the presence of tapering paraphyses and within the order ramsayi and bamboo. Currently Annulatascus comprises 16
Sordariales. The family is characterized by immersed or su- species. Annulatascus is known to be polyphyletic according
perficial, coriaceous, usually dark-walled ascomata, long, cy- to LSU rDNA sequence data (Raja et al. 2003; Campbell and
lindrical asci with a relatively massive, J-, refractive, apical Shearer 2004; Abdel-Wahab et al. 2011). Tsui and Hyde
ring. Using LSU rDNA molecular data, Ranghoo et al. (1999) (2003) provided a key to species and a synoptic table of 12
showed that Annulatascaceae is a separate family, which has Annulatascus species based on morphological characteristics.
relationships with Sordariales. Ho and Hyde (2000) reviewed Boonyuen et al. (2012) provided comparative table of mor-
the ultrastructural studies on Annulatascaceae and concluded phological features for 16 Annulatascus species.
that more taxa need to be sequenced and analyzed to
strengthen data for this family, and that the ultrastructural
data supports descriptions of new genera. Based on LSU
rDNA sequence data, Réblová and Winka (2001) showed that Other genera included
that Annulatascaceae had affinities outside Sordariales. Kirk
et al. (2001, 2008) placed Annulatascaceae in Annulusmagnus J. Campb. & Shearer, Mycologia 96(4): 826
Sordariomycetidae families incertae sedis, which was further (2004)
Fungal Diversity

Fig. 6 Annulatascus velatispora (Material examined: AUSTRALIA. substrate d, e Vertical section f Section showing peridium and neck
North Queensland: Milaa Milaa Falls, on submerged wood in a river, region g Squash mount of asci h, i asci j Apical ring k Apical ring in
July 1990, K.D. Hyde, (BRIP 17373, holotype) a Herbarium label and Melzer’s reagent l Ascospores. Scale bars: d-e = 100 μm, f-i = 50 μm,
specimen of Annulatascus velatispora b Host material c Ascomata on l = 10 μm, j–k = 5 μm
 Fungal Diversity

Cataractispora K.D. Hyde et al., Mycol. Res. 103(8): 1019

(1999)
Type species: Cataractispora aquatica K.D. Hyde et al.,
Mycol. Res. 103 (8): 1020 (1999)
Chaetorostrum Zelski et al., Mycosphere 2(5): 594 (2011)
Type species: Chaetorostrum quincemilensis Zelski et al.,
Mycosphere 2 (5): 595 (2011)
Clohiesia K.D. Hyde, Nova Hedwigia 61(1–2): 125 (1995)
Type species: Clohiesia corticola K.D. Hyde, Nova
Hedwigia 61 (1–2): 126 (1995)
Cyanoannulus Raja et al., Mycotaxon 88: 8 (2003)
Type species: Cyanoannulus petersenii Raja et al.,
Mycotaxon 88: 11 (2003)
Dictyosporella Abdel-Aziz, in Ariyawansa et al., Fungal
Diversity: 10.1007/s13225-015-0346-5, [117] (2015)
Type species: Dictyosporella aquatica Abdel-Aziz, in
Ariyawansa et al., Fungal Diversity: 10.1007/s13225-015-
0346-5, [119] (2015)
Diluviocola K.D. Hyde et al., Fungal Diversity Res. Ser. 1:
141 (1998)
Type species: Diluviicola capensis K.D. Hyde et al.,
Fungal Diversity 1: 134 (1998)
Fusoidispora Vijaykr. et al., Sydowia 57(2): 272 (2005)
Type species: Fusoidispora aquatica Vijaykr. et al.,
Sydowia 57 (2): 272 (2005)
Longicollum Zelsk et al., in Zelski et al., Mycosphere 2(5):
540 (2011)
Type species: Longicollum biappendiculatum Zelski et al.,
Mycosphere 2 (5): 540 (2011)
Pseudoannulatascus Z.L. Luo et al., in Z.L. Luo et al.
Fig. 7 Taeniolella-like asexual morph of Chaetorostrum quincemilensis.
(Redrawn from Zelski et al. 2011) a, b Conidia. Scale bars: a, b = 20 μm Phytotaxa 239(2): 179 (2015)
Type species: Pseudoannulatascus biatriisporus (K.D.
Hyde) Z.L. Luo et al., in Z.L. Luo et al. Phytotaxa 239(2):
179 (2015)
Type species: Annulusmagnus triseptatus (S.W. Wong Pseudoproboscispora Punith., Kew Bull. 54(1): 234
et al.) J. Campb. & Shearer, Mycologia 96 (4): 826 (2004) (1999)
Aqualignicola Ranghoo et al., Mycol. Res. 105(5): 628 Type species: Pseudoproboscispora aquatica (S.W. Wong
(2001) & K.D. Hyde) Punith., Kew Bulletin 54 (1): 234 (1999)
Type species: Aqualignicola hyalina Ranghoo et al., Rhamphoria Niessl, Verh. nat. Ver. Brünn 14: 204, 206
Mycol. Res. 105 (5): 628 (2001) (1876)
Aquaticola W.H. Ho, et al., Fungal Diversity Res. Ser. 3: Type species: Rhamphoria delicatula Niessl, Verh. nat.
88 (1999) Ver. Brünn 14: 206 (1876)
Type species: Aquaticola hyalomura W.H. Ho et al., Submersisphaeria K.D. Hyde, Nova Hedwigia 62(1–2):
Fungal Diversity 3: 88 (1999) 172 (1996)
Ascitendus J. Campb. & Shearer, Mycologia 96(4): 829 Type species: Submersisphaeria aquatica K.D. Hyde,
(2004) Nova Hedwigia 62: 172 (1996)
Type species: Ascitendus austriacus (Réblová et al.) J. Torrentispora K.D. Hyde et al., Mycol. Res. 104(11): 1399
Campb. & Shearer, Mycologia 96 (4): 829 (2004) (2000)
Ayria Fryar & K.D. Hyde, Cryptog. Mycol. 25(3): 248 Type species: Torrentispora fibrosa K.D. Hyde et al.,
(2004) Mycol. Res. 104 (11): 1399 (2000)
Type species: Ayria appendiculata Fryar & K.D. Hyde, Vertexicola K.D. Hyde et al., Mycologia 92(5): 1019
Cryptog. Mycol. 25: 248 (2004) (2000)
Fungal Diversity

Type species: Vertexicola caudatus K.D. Hyde et al., 17. Ascospores with polar conical caps attached at
Mycologia 92: 1020 (2000) each end ………………...............……Diluviocola
17. Ascospores without polar conical caps
……………….…............................... Annulatascus
Key to the genera of Annulatascaceae
Apiosporaceae K.D. Hyde et al., in Hyde et al., Sydowia
1. Ascospores muriform ………..…………….. Rhamphoria 50(1) 23 (1998)
1. Ascospores lacking longitudinal septa ……………… 2 Facesoffungi number: FoF 01364
2. Ascomata formed under a clypeus ………… Clohiesia Pathogenic or saprobic on leaves, stems and roots of
2. Ascomata without clypeus…………….…………….. 3 Arecaceae (palms), Cyperaceae (sedges) and Poaceae
3. Necks possessing setae or hyphae or hairs ..…………. 4 (grasses and bamboo), or endophytic on plant tissues, lichens,
3. Necks glabrous …………………………….…….…. 6 and marine algae, occasionally infecting humans, or isolat-
4. Necks hyaline, with stiff hairs ………… Chaetorostrum ed from soil. Sexual morph: Pseudostromata immersed or
4. Necks black, setae present on necks ...……………… 5 semi-immersed in host epidermis, becoming erumpent,
5. Peridium membranous, ascospores 1-celled, lacking through a longitudinal split in the host. Ascomata perithe-
s h e a t h s o r s h e a t h o n on e e n d o f a s co s po r e cial, globose, densely arranged, immersed in
...……………………………………… Aqualignicola Pseudostromata, with papillate ostioles. Peridium com-
5. Peridium coriaceous, ascospores 3-septate, sheath pres- posed of small, brown cells of textura angularis Asci 8-
ent ……..……..… Annulusmagnus spored, unitunicate, clavate to broadly cylindrical, short
6. Ascospores thick-walled, distoseptate ……..... Vertexicola pedicellate. Paraphyses hyphae-like, broad, septate,
6. Ascospores not distoseptate ………………...………. 7 branched. Ascospores overlapping 1–3-seriate to irregularly
7. Ascospores brown except in Annulatascus arranged, 1-septate, apiosporous, ellipsoidal, in equilateral,
aquatorba…….…….…….…….…….....….………. 8 straight or curved, hyaline, smooth-walled, with or without
7. Ascospores hyaline …………......................………… 9 gelatinous sheath. Asexual morph: Coelomycetous or hy-
8. Ascospores with brown central cells, end cells hyaline, phomycetous. Conidiomata sporodochial, solitary to gre-
with longitudinal striations ………….....…. Ascitendus garious, immersed, erumpent from host tissue when ma-
8. Ascospores with polar pad-like appendages ture, irregular, black, carbonaceous, coriaceous.
…………..……………………... Submersisphaeria Conidiomata basal stroma composed of several layers of
9. Ascomata pale reddish brown with long necks dark brown to hyaline thick-walled cells at the side, thin upper
……………………………………..... Cyanoannulus wall cells of textura angularis. Setae lacking, or present occa-
9. Ascomata not as above …………………..…………. 10 sionally, if present intermingled among conidiophores.
10. Ascomata membranous …………………............... 11 Conidiophores arising from hyphae or aggregated in a brown
10. Ascomata coriaceous or carbonaceous stroma, forming black sporodochia, forming conidia terminally
……………………………….............………… 12 and laterally, brown to dark brown. Setae present or absent,
11. Necks long, ascospores uniseriate with bipolar append- erect, sparsely septate, pale brown to brown, smooth-walled.
ages ……..….…………………………. Longicollum Conidiogenous cells discrete, basauxic, doliiform to
11. Necks short, ascospores overlapping uniseriate or ampulliform or cylindrical, brown, smooth to finely verruculose.
biseriate, lacking appendages ….. Ayria Conidia unicellular, brown to dark brown, smooth-walled or
12. Ascospores without appendages ………....………. 13 with minute wall ornamentations, with a truncate basal scar,
12. Ascospores with or without appendages ….…......…. 14 guttulate to granular, frequently with an equatorial germ slit.
13. Ascospores aseptate or septate, guttulate Type: Arthrinium Kunze
…………………………………..…....... Aquaticola Notes: This family was introduced by Hyde et al. (1998)
13. Ascospores aseptate, eguttulate with Apiospora montagnei Sacc. as the type species. Earlier
………………………...........…..…. Torrentispora the genus Apiospora had been assigned to Lasiosphaeriaceae
14. Ascospores ellipsoid .....……… Pseudoproboscispora by Barr (1990). Hyde et al. (1998) considered that the sexual
14. Ascospores fusoid ……...............………………… 15 and asexual morphs (Arthrinium, Cordelia and Pteroconium)
15. Ascomata ampulliform ……..………… Fusoidispora of Apiospora differed from those in Lasiosphaeriaceae and
15. Ascomata not as above …...........................………. 16 proposed the family Apiosporaceae to accommodate
1 6. A scospo re app endag es unfu rlin g in water Apiospora. Hyde et al. (1998) did not suggest any order place-
………………………….…..........… Cataractispora ment for Apiosporaceae. Smith et al. (2003) analyzed LSU
16. Ascospore appendages not as above and SSU sequence data and showed that members of
……………………………….........…………… 17 Apiosporaceae cluster within the order Xylariales, but
 Fungal Diversity

relationships to other families of Xylariales was uncertain. Conidiogenous cells basauxic, cylindrical, smooth-walled
Lumbsch and Huhndorf (2010) listed Apiosporaceae in or verrucose. Conidia subglobose to globose, dark brown,
Sordariomycetes family incertae sedis. Crous and smooth-walled or with minute wall ornamentations, with a
Groenewald (2013) used phylogenetic analysis to confirm truncate basal scar.
Apiosporaceae as a family within Xylariales, and sister to Type species: Arthrinium caricicola Kunze & J.C.
Amphisphaeriaceae. Hyde et al. (1998) considered Schmidt, Mykologische Hefte (Leipzig) 1: 9 (1817)
Dictyoarthrinium, Endocalyx, Scyphospora and Spegazzinia Notes: Apiospora was identified as a genus in the family
as other possible genera of this family, but molecular data is Amphisphaeriaceae based on morphology by Müller and von
lacking to confirm this (Crous and Groenewald 2013). Arx (1962). Later it was transferred to Hyponectriaceae by
Arthrinium was confirmed as the asexual morph of Barr (1976a). Barr (1990) moved Apiospora to
Apiospora by molecular data (Crous and Groenewald 2013; Lasiosphaeriaceae and this was followed by Barr and
Senanayake et al. 2015) and Apiospora, Cordella and Cannon (1994) and Hawksworth et al. (1995). Hyde et al.
Pteroconium were reduced to synonymy with Arthrinium (1998) introduced a new family Apiosporaceae to accommo-
(Crous and Groenewald 2013). Maharachchikumbura et al. date the genus Apiospora, with A. montagnei as the type
(2015) listed six genera under Apiosporaceae. species. Phylogenetic analyses by many authors accepted
Arthrinium Kunze, in Kunze & Schmidt, Mykologische Apiospora as a distinct group in the order Xylariales
Hefte (Leipzig) 1: 9 (1817) (Huhndorf et al. 2004a; Zhang et al. 2006; Jaklitsch and
Facesoffungi number: FoF 02100; Figs 8, 9 Voglmayr 2012; Dai et al. 2014a). Index Fungorum (2016)
Saprobic or pathogenic on monocotyledons, especially lists 55 epithets under genus Apiospora. Crous and
grasses, or endophytic on plant tissues, lichens, and ma- Groenewald (2013) synonymized the sexual name
rine algae, occasionally infecting humans, or isolated from Apiospora under the asexual name Arthrinium, since
soil. Sexual morph: Pseudostromata visible as raised, lin- Arthrinium is the older and more commonly encountered,
ear, blackened areas on the host surface, with neck and and more frequently used name in the literature. The link
upper surface visible through slits in the host tissue, most- between Arthrinium and Apiospora has also been con-
ly gregarious, fusiform, ellipsoid to irregular, black, cells firmed by molecular data by Crous and Groenewald
between ascomata and darkened layer above usually thick, (2013), and this was followed by Maharachchikumbura
composed of brown cells of textura angularis. Ascomata et al. (2015) and Senanayake et al. (2015). Crous and
solitary or usually gregarious in linear groups, immersed Groenewald (2013) synonymised Pteroconium and
in pseudostromata, subglobose to globose, membranous, Cordella under Arthrinium.
papillate, ostiolate. Ostiole periphysate. Peridium com-
posed of several layers, of small, brown or reddish brown
to hyaline cells of textura angularis. Hamathecium com- Other genera included
posed of dense, hypha-like, long, broad, septate, branched,
paraphyses. Asci 8-spored, unitunicate, broad cylindrical to Appendicospora K.D. Hyde, Sydowia 47(1): 31 (1995)
clavate or subglobose, short pedicellate, without an apical Type species: Appendicospora coryphae (Rehm) K.D.
ring. Ascospores overlapping 1–3-seriate to irregularly ar- Hyde, Sydowia 47(1): 32 (1995)
ranged, hyaline, apiosporous, with a large, straight or Dictyoarthrinium S. Hughes, Mycol. Pap. 48: 29 (1952)
curved, upper cell and smaller lower cell, usually Type species: Dictyoarthrinium quadratum S. Hughes,
surrounded by a gelatinous sheath. Asexual morph: Mycol. Pap. 48: 30 (1952)
Coelomycetous or hyphomycetous. Conidiomata Endocalyx Berk. & Broome, J. Linn. Soc., Bot. 15(1): 84
sporodochial, solitary to gregarious, immersed, erumpent (1876) [1877]
from host tissue when mature, irregular, black, carbona- Type species: Endocalyx thwaitesii Berk. & Broome, J.
ceous, coriaceous. Conidiomata basal stroma composed Linn. Soc., Bot. 15(1): 84 (1876) [1877]
of several layers of dark brown to hyaline cells of textura Scyphospora L.A. Kantsch., Bolêz. Rast. 17: 87 (1928)
angularis, with thick side wall, thin at upper and lower Type species: Scyphospora phyllostachydis Kantsch. [as
walls. Setae lacking, or occasionally present, intermingled ‘phyllostachidis’], Bolêz. Rast. 17: 88 (1928)
among conidiophores. Conidiophore verrucose, flexuous. Spegazzinia Sacc., Spegazzinia: [1] (1879)
Conidiophore mother cells ampulliform, with a verrucose Type species: Spegazzinia ornata Sacc., Michelia 2(no. 6):
wall, producing a single hyphoid, cylindrical, 1–2-septate. 172 (1880)
Fungal Diversity

Fig. 8 Arthrinium yunnanum (Material examined: CHINA, Yunnan, surface c-e Vertical section through pseudostroma f Paraphyses g, h Asci i
Kunming, Kunming Institute of Botany, Chinese Academy of Science, Ascospores j Ascospore with sheath. Scale bars: c = 200 μm, d,
on dead culm of Phyllostachys nigra (Poaceae), 7 July 2014, Dong-Qin e = 100 μm, f–i = 5 μm
Dai DDQ00279 (MFLU 15-0382, holotype) a, b Pseudostromata on host
 Fungal Diversity

Fig. 9 Arthrinium yunnanum (asexual morph, holotype) a Conidiomata on PDA b Vertical section through conidioma c Conidiomatal wall d Conidia
e Conidium f, g Conidia with basauxic conidiogenous cells. Scale bars: b = 200 μm, c, d = 50 μm, e = 20 μm, f, g = 50 μm

Key to the genera with sexual morph 5. Conidiomata cup-shaped to funnel-shaped, surrounded
by setae, conidia dark brown, smooth-walled or with
1. Ascomata stromatic, asci clavate to broadly cylindrical, minute wall ornamentations …………........ Endocalyx
ascospores ellipsoidal without appendages 5. Conidiomata not cupulate or funnel-shaped, setae
…………………………………….................. Apiospora absent or occasionally present, conidia hyaline be-
1. Ascomata astromatic, asci clavate, ascospores clavate coming dark brown at maturity, smooth, verrucose,
with appendages at one end ................ Appendicospora ornamented ………………………….... Arthrinium

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

Facesoffungi number: FoF 00723
Key to the genera with asexual morph Epiphytes on the surface of leaves. Superficial hyphae
dense, branched, septate, brown to reddish, with hyphopodia,
1. Coelomycetous or hyphomycetous, conidiomata acervular, hyphal setae lacking. Hyphopodia single stellate to sublobate
pycnidial, sporodochial or synnematous, conidia 1-celled on stalk cells, alternate on hyphae, 2-celled, brown to reddish.
…………………………………………………………. 2 Sexual morph: Ascomata superficial on surface of hosts,
1. Hyphomycetous, conidiomata synnematous or scattered, flattened when immature, globose to subglobose at
sporodochial, conidia 4 or 16-celled, muriform or often maturity, developing on hyphae, surface verrucose, covered
septate …………………………………………….... 3 with tuberculate projections, ascomatal setae and appendages
3. Conidia 4-celled, smooth to spinose …….. Spegazzinia lacking. Peridium comprising two strata, outer stratum amor-
3. Conidia 4 or16-celled, conidiomata verrucose or tuber- phous and black, inner stratum, comprising reddish to brown,
culate ……………………………… Dictyoarthrinium scleroparenchymatous cells of textura angularis to globulosa.
4. Conidiomata acervular, conidia, smooth-walled Hamathecium with paraphyses, evanescent. Asci 4–8-spored,
………………………………………..... Scyphospora unitunicate, ovoid to clavate. Ascospores 2–3-seriate, hyaline
4. Conidiomata, smooth, verrucose, ornamented or with to light brown, ellipsoidal to oblong, 1-septate. Asexual
minute wall ornamentations ……………………........ 5 morph: Undetermined.
Fungal Diversity

Type: Armatella Theiss. & Syd. ascomata with raised conoid cells, and 3–4-septate, hyaline
Notes: The family Armatellaceae was introduced by to dark brown ascospores (Hongsanan et al. 2015).
Hosagoudar (2003), with the generic type Armatella, which
is a monotypic genus (Hosagoudar 2003; Hosagoudar et al. Armatella Theiss. & Syd., Annls mycol. 13(3/4): 235
2012; Hongsanan et al. 2015). Hongsanan et al. (2015) accept- (1915)
ed Armatellaceae as a separate family from Meliolaceae, Facesoffungi number: FoF 00723; Fig. 10
which is followed in this study because it has superficial hy- Epiphytes on the surface of leaves. Superficial hyphae
phae without phialides, ascomata covered with tuberculate dense, branched, septate, brown to reddish, with hyphopodia,
projections, and comprising scleroparenchymatous cells, hyphal setae lacking. Hyphopodia single stellate to sublobate
aseptate to 1-septate, hyaline to light brown ascospores, while on stalk cells, alternate on hyphae, 2-celled, brown to reddish.
Meliolaceae has superficial hyphae mostly with phialides, Sexual morph: Ascomata superficial on surface of hosts,

Fig. 10 Armatella litseae (Material examined: JAPAN, Province Awa, of young ascomata when viewed in squash mounts g, h Hyphopodia on
Tokushima, Kigasumi, on leaves of Litsea glauca (Lauraceae), 25 hyphae i Immature ascus in Melzer’s reagent j, k Mature ascospores in
December 1897, S. Kusano, (SF70331, holotype). a Herbarium packet Melzer’s reagent. Scale bars: e = 100 μm, f = 20 μm, i–k = 10 μm, g,
b-d Colony on leaves e Vertical section through ascoma f Upper cell walls h = 10 μm
 Fungal Diversity

scattered, flattened when immature, globose to subglobose at with the use of molecular data. Rong and Gams (2000)
maturity, developing on hyphae, surface verrucose, covered used the branching of the conidiophores and the shapes
with tuberculate projections, ascomatal setae and appendages and dimensions of conidia to distinguish Monilochaetes
lacking. Peridium comprising two strata, outer stratum amor- from the similar genera Dischloridium and Exochalara.
phous and black, inner stratum, comprising reddish to brown, ITS and LSU phylogenies confirmed that Dischloridium
scleroparenchymatous cells of textura angularis to globulosa. and Monilochaetes are congeneric, hence Dischloridium
Hamathecium with paraphyses, evanescent. Asci 4–8-spored, became a generic synonym of Monilochaetes. The con-
unitunicate, ovoid to clavate. Ascospores 2–3-seriate, hyaline nection between the sexual (Australiasca) and the asex-
to light brown, ellipsoidal to oblong, aseptate when immature ual (Monilochaetes) morph has been experimentally
and 1-septate at maturity, constricted at the septum, ends established (Sivanesan and Alcorn 2002; Réblová et al.
rounded, smooth-walled. Asexual morph: Undetermined. 2011).
Type species: Armatella litseae (Henn.) Theiss. & Syd.,
Annls mycol. 13(3/4): 235 Monilochaetes Halst. ex Harter, J. Agric. Res., Washington
Notes: No Armatellaceae species have been obtained in 5: 791 (1916)
culture and no sequence data is available in GenBank Facesoffungi number: FoF 01093; Fig. 11
(Hongsanan et al. 2015). Index Fungorum (2016) lists 19 ep- Type species: Monilochaetes infuscans Harter, J. Agric.
ithets in Armatella. However most of these species need to be Res., Washington 5: 791 (1916)
re-evaluated. Notes: The asexual genus Monilochaetes comprises seven
Australiascaceae Réblová & W. Gams, in Réblová et al., species (Index Fungorum 2016). Dischloridium and
Stud. Mycol. 68(1): 171 (2011) Monilochaetes are similar in morphology and molecular phy-
Facesoffungi number: FoF 01110 logenies confirm that these two genera are congeneric.
Pathogenic and saprobic on terrestrial plant leaves, Monilochaetes infuscans causes scurf disease or soil stain of
branches, spathes and stipes. Sexual morph: Stromata absent. sweet potato in many countries (Rong and Gams 2000).
Ascomata gregarious to solitary, brown to black, conical to Monilochaetes basicurvata (Matsush.) Réblová & Seifert,
obpyriform, glabrous or with setae, ostiole periphysate. M. dimorphospora Réblová & W. Gams, M. laeénsis
Setae scant, acute, thick-walled, septate, dark brown. (Matsush.) Réblová, W. Gams & Seifert and M. regenerans
Peridium 18–22 μm wide, becoming thick towards the base, (Bhat & W.B. Kendr.) Réblová & Seifert are reported as
2-layered, fragile, of textura epidermoidea to prismatica in saprobes while M. camelliae (Alcorn & Sivan.) Réblová, W.
surface view. Hamathecium comprising septate, persistent, Gams & Seifert has been recorded as a pathogen (Rong and
branching paraphyses. Asci 8-spored, unitunicate, cylindrical Gams 2000; Réblová et al. 2011). Rong and Gams (2000)
to clavate, short-pedicellate, apex truncate, with a distinct, proposed the combination of M. guadalcanalensis, which
shallow, J- apical ring. Ascospores overlapping biseriate, hy- was also accepted in Réblová et al. (2011). A recent study
aline, oblong to ellipsoidal, apiculate at both ends, septate, by Martínez-Rivera et al. (2014) also has accepted this con-
smooth-walled. Asexual morph: Hyphomycetous, cept. However, in Index Fungorum (http://www.
Conidiophores macronematous, pale to dark brown or black, indexfungorum.org/Names/Names.asp2016), M.
unbranched, septate. Conidiogenous cells monophialidic, guadalcanalensis has been synonymised under Exochalara
ampulliform to cylindrical, subhyaline, with a minute, flared guadalcanalensis (Matsush.) W. Gams & Hol.-Jech. Further
collarette. Conidia ellipsoid to cylindrical–ellipsoidal, clarification of this species taxonomic placement is required.
smooth-walled, hyaline to greenish, septate, aggregated in The sex ual-ase xual c onne ctions of Australiasca
slime or in straight to curled pseudo-chains. queenslandica with Monilochaetes camelliae and
Type: Monilochaetes Halst. ex Harter Australiasca laeénsis with Monilochaetes laeénsis have been
Notes: This family was established by Réblová et al. experimentally established (Sivanesan and Alcorn 2002;
(2011) to accommodate the holomorphic genus Réblová et al. 2011). Australiasca includes only two species,
Australiasca and asexual morph Monilochaetes namely A. laeénsis Réblová & W. Gams and A. queenslandica
(Maharachchikumbura et al. 2015). The sexual morph Sivan & Alcorn. Although these species are morphologically
of this family mimics Chaetosphaeria which have al- similar, molecular analyses of ITS and LSU sequence data
most indistinguishable ascomata. The asexual morph is confirmed that they are distinct. The released ascospores in
also similar to the asexual morphs of Chaetosphaeria Australiasca were often observed to be 1–3-septate (Réblová
(Réblová et al. 2011). Monilochaetes was established et al. 2011). The ascospores of Australiasca queenslandica
for a single dematiaceous hyphomycete species become muriform at maturity, but this character has not been
M. infuscans Harter (Halsted 1890). Rong and Gams observed in Australiasca laeénsis (Réblová et al. 2011). When
(2000) added M. guadalcanalensis to this genus. This it comes to the use of one name one fungus concept, we
addition was later confirmed by Réblová et al. (2011) recommend the use of Monilochaetes as this generic name is
Fungal Diversity

Fig. 11 Australiasca queenslandica (Material examined: g Setae h Conidiophores i Conidiogenous cells j Chains of conidia k
AUSTRALIA, Queensland, Malanda, on branch of Camellia sinensis Conidia (conidiogenous cells and conidia chains of Monilochaetes
(L.) Kuntze (Theaceae), 26 March 1988, J. L. Alcorn, BRIP 25190a, infuscans redrawn from Rong and Gams 2000). Scale bars: b-
holotype). a Label and herbarium material b Ascoma on the host c c = 100 μm, e–f, g = 50 μm, g, i–k = 10 μm
Cells of peridium d Immature asci e Asci with ascospores f Ascospores

older (introduced in 1916) than the generic name Australiasca Bartaliniaceae Wijayawardene et al., in Senanayake et al.,
(introduced in 2002). Also, Monilochaetes (7) has higher Fungal Divers. 73: 13 (2015)
number of species than Australiasca (2). Furthermore, Facesoffungi number: FoF 00667
Monilochaetes is a well-established name in plant pathologi- Saprobic, endophytic or pathogenic on leaves, twigs and
cal community. branches of a variety of hosts. Sexual morph: Pseudostromata
 Fungal Diversity

immersed, solitary, scattered. Ascomata black, immersed to (2015) to accommodate the genera Bartalinia, Broomella,
semi-immersed, solitary or gregarious, globose to subglobose. Dyrithiopsis, Hyalotiella, Truncatella and Zetiasplonza based
Ostiole central, circular. Papilla periphysate. Peridium composed on both morphological characters and phylogenetic analysis.
of an outer layer, of brown to dark brown cells of textura It is phylogenetically closely related to the families
angularis or textura prismatica, with thin inner layer of hyaline Sporocadaceae, Pestalotiopsidaceae and Robillardaceae.
cells of textura angularis. Hamathecium comprising numerous,
hyaline, septate, filamentous paraphyses. Asci 8-spored, Bartalinia Tassi, Bulletin Labor. Orto Bot. de R. Univ.
unitunicate, cylindrical, short pedicellate, apex rounded, with Siena 3: 4 (1900)
J+ or J-, subapical ring. Ascospores overlapping uniseriate, pale Facesoffungi number: FoF 00659; Fig. 12
brown to brown, ellipsoid to fusiform, 3–4-septate. Asexual Endophytic or saprobic on plants. Sexual morph:
morph: Coelomycetous. Conidiomata acervular to pycnidial to Undetermined. Asexual morph: Conidiomata pycnidial
irregular, solitary or gregarious, superficial to sub-immersed, uni- or variable, solitary to gregarious, subepidermal,
locular, globose to subglobose, dark brown to black. Ostiole erumpent at maturity, globose, unilocular, brown to
apapillate. Conidiomata wall comprising a thick outer layer of black. Conidiomata wall two layered, outer layer com-
dark brown cells of textura angularis, and thin inner wall of prising dark brown to black cells of textura angularis,
hyaline to sub-hyaline cells of textura angularis. inner wall comprising thin, hyaline cells of textura
Conidiophores present or reduced to conidiogenous cells, when angularis. Conidiophores reduced to conidiogenous
present cylindrical, hyaline, sparsely septate, smooth-walled. cells. Conidiogenous cells ampulliform, holoblastic, dis-
Conidiogenous cells holoblastic, ampulliform, integrated or dis- crete, determinate, hyaline, forming from the inner layer
crete, determinate, hyaline, smooth-walled. Conidia fusiform, of the pycnidium wall. Conidia hyaline to pale brown,
straight to slightly curved, subhyaline to brown, bearing an apical subcylindrical, trans-septate, with appendages, basal cell
appendage or both apical and basal appendages. with truncate base, obconic, hyaline, with single un-
Type: Bartalinia Tassi branched appendage; 3 median cells subcylindrical, hy-
Notes: The family Bartaliniaceae (order aline to pale brown, apical cell conical, almost hyaline,
Amphisphaeriales) was introduced by Senanayake et al. with three branched appendage.

Fig. 12 Bartalinia robillardoides (Material examined: THAILAND, 1, MFLU 13-0084) a Conidiomata on host b, c Section of conidiomata d
Chiang Rai, Mae Fah Luang University ground, on leaves of Conidia arising from conidiogenous cells e-h Conidia. Scale bars: b–
Eucalyptus sp., 30 June 2012, Nalin N. Wijayawardene, NNW 120630- c = 150 μm, d = 20 μm, e–h = 25 μm
Fungal Diversity

Type species: Bartalinia robillardoides Tassi, Bulletin globulosa to textura angularis, inwardly merging with rela-
Labor. Orto Bot. de R. Univ. Siena 3: 4 (1900) tively thin-walled, colourless cells. Conidiophores reduced to
Notes: Bartalinia was introduced by Tassi (1900) with conidiogenous cells. Conidiogenous cells integrated, cylindri-
Bartalinia robillardoides as the type species. Morgan-Jones cal, phialidic, percurrently proliferating 1–2-times, hyaline,
et al. (1972) and Sutton (1980) accepted nine species in the smooth. Conidia pale brown or brown, fusiform to aciculate,
genus, while emphasizing the need for a taxonomic revision of with acute ends, straight or slightly curved, 3-septate, con-
the genus. von Arx (1981) listed Bartalinia under stricted at septa, verruculose, thick walled, bearing up to 5
Seimatosporium. Nag Raj (1993) rejected this classification stat- appendages at the apex and a single appendage at the base.
ing the differences in conidial appendages and retained the type Type species: Broomella vitalbae (Berk. & Broome) Sacc.,
B. robillardoides and five other species, B. bischofiae Nag Raj, Syll. fung. (Abellini) 2: 558 (1883)
B. lateripes (Ellis & Everh.) Nag Raj, B. laurina (Mont. ex Notes: The genus Broomella, introduced by Saccardo (1883)
Desm.) Nag Raj, B. pistacina (J.L. Maas) Nag Raj and and is characterized by having unitunicate, cylindrical-elongate
B. tamarindi Nag Raj, as belonging to the genus Bartalinia. asci, with a J-, discoid apical ring and 3-septate, ellipsoid-
New Bartalinia species have been added as a result of various fusiform ascospores, with brown median cells and lighter termi-
studies (Anderson and Bianchinotti 1996; Xi et al. 2000; Chi nal cells, bearing a single, centric appendage at each end
et al. 2002; Andrianova and Minter 2007; Marincowitz et al. (Shoemaker and Müller 1963; Li et al. 2015a). Broomella has
2010; Senanayake et al. 2015) and currently 19 epithets are listed been linked to a pestalotiod-like asexual morph (Shoemaker et al.
in Index Fungorum (2016). Bartalinia species are characterized 1989; Yuan and Zhao 1992; Kang et al. 1999b). Currently, 20
by having three to four septate conidia. A key to ten Bartalinia epithets are listed under Broomella in Index Fungorum (2016).
species, including the six listed by Nag Raj (1993), has been However, they lack sequence data. Broomella species and their
provided by Andrianova and Minter (2007). In a study of truncatella-like asexual morphs differ in many ways from the
appendaged coelomycetes, Crous et al. (2014a) designated an type B. vitalbae (Berk. & Broome) Sacc. and its asexual morph
epitype for B. robillardoides, collected on Leptoglossus and therefore may not be congeneric. In a study based on both
occidentalis. Bartalinia species have been recorded on a wide LSU sequence data and morphological data by Li et al. (2015a),
range of hosts - especially flowering shrubs and trees, and caus- Broomella appeared to be a distinct, natural group in
ing leaf spots on economically important plants across Australia, Amphisphaeriaceae. Based on the phylogenetic analysis by
China, Europe, India, South America and South East Asia Senanayake et al. (2015) Broomella was placed in
(Wong et al. 2003; Farr and Rossman 2016). Bartaliniaceae, Amphisphaeriales along with Bartalinia,
Dyrithiopsis, Hyalotiella, Truncatella and Zetiasplozna. The
asexual morph was described in Senanayake et al. (2015).
Other genera included Dyrithiopsis L. Cai et al., in Jeewon et al., Mycologia
95(5): 912 (2003)
Broomella Sacc., Syll. fung. (Abellini) 2: 557 (1883) Type species: Dyrithiopsis lakefuxianensis L. Cai et al., in
Facesoffungi number: FoF 00626; Fig. 13 Jeewon et al., Mycologia 95(5): 913 (2003)
Saprobic on various dicots, monocots such as bamboo and Hyalotiella Papendorf, Trans. Br. mycol. Soc. 50(1): 69
pine. Sexual morph: Ascomata solitary to gregarious, (1967)
uniloculate, glabrous, globose to subglobose, papillate. Ostiole Type species: Hyalotiella transvalensis Papendorf, Trans.
centrally located, comprising of longitudinal cells, internally lined Br. mycol. Soc. 50(1): 69 (1967)
with hyaline periphyses. Peridium composed of light brown, Truncatella Steyaert, Bull. Jard. bot. État Brux. 19: 293
thick-walled cells of textura prismatica in the upper part, and (1949)
thin-walled, hyaline to pale brown cells in other parts. Type species: Truncatella truncata (Lév.) Steyaert, Bull.
Hamathecium comprising of cylindrical, septate, paraphyses. Jard. bot. État Brux. 19: 295 (1949)
Asci 8-spored, unitunicate, cylindrical to cylindrical-clavate, short Zetiasplozna Nag Raj, Coelomycetous Anamorphs with
pedicel, apically rounded, with a J- apical ring. Ascospores Appendage-bearing Conidia (Ontario): 996 (1993)
biseriate or overlapping bi- or tri-seriate, fusiform, median cells Type species: Zetiasplozna caffra Matsush. [as ‘caffera’],
pale yellowish brown to greyish brown, end cells grey or hyaline, Matsush. Mycol. Mem. 9: 29 (1996)
glabrous, straight or inequilaterally curved, 3-septate, constricted
at the mid septa, thick-walled, each end cell bearing a simple,
centric, tubular appendage. Asexual morph: Coelomycetous.
Conidiomata stromatic, pycnidioid, scattered to gregarious, Key to sexual genera of Bartaliniaceae
immersed to semi-immersed, oval or elongated, black, uniloc-
ular, papillate, glabrous. Pycnidial wall outer layers comprised 1. Asci with J-, apical ring, having 3-septate ascospores
of thick-walled, pale brown to brown cells of textura ………………………………. Broomella
 Fungal Diversity

Fig. 13 Broomella vitalbae (Material examined: ITALY. Province of up of ascomata d Section of ascoma e Ostiole f Peridium g Paraphyses h-
Forlì-Cesena [FC], Modigliana, Montebello, on dead stem of Clematis k Asci with ascospores l-n Ascospores. Scale bars: a = 500 μm, b–
vitalba (Ranunculaceae), 23 February 2013, Erio Camporesi, IT-1079, c = 200 μm, d–f = 50 μm, g–n = 20 μm
MFLU 15-0065, epitype) a Appearance of ascomata on host b, c Close

1. Asci with J+, subapical ring, having muriform asco- 2. Spores smooth-walled, median cells hyaline or almost
s po r e s ( 3- t r a n s v e r s e, 1– 2 l o ng i t u d i n al s ep - hyaline to pale brown.................................................... 3
ta)……………………………………… Dyrithiopsis 2. Spores verruculose, median cells
brown………………………………….. Truncatella
3. Conidia with single basal append-
Key to asexual genera of Bartaliniaceae age……………………………….......….. Bartalinia
3. Conidia lacking a basal append-
1. Spores bearing single apical appendage, branched age……………………………………... Hyalotiella
……………………………………………………….... 2
1. Spores bearing several apical appendages, un- Batistiaceae Samuels & K.F. Rodrigues, Mycologia 81(1):
branched..……………………………… Zetiasplozna 54 (1989)
Fungal Diversity

Facesoffungi number: FoF 01321 spaced annellations, comprising thick, brown cells of textura
S a p rob i c o n w o o d. Se x u a l m o r ph : A s c o m at a prismatica, cap globose, black, carbonaceous, surface formed
cleistothecial, stipitate-capitate, solitary to gregarious, super- of thick polyhedral plates, plates in greatest dimension, de-
ficial, black, globose, long stipitate, carbonaceous, without an hiscing along the lines separating the plates. Opening widely
ostiole. Peridium cephalothecoid. Asci 8-spored, unitunicate, as petals of a flower to expose the brown, powdery mass of
globose, lacking a visible discharge ring and deliquescent. ascospores, without an ostiole. Peridium cephalothecoid,
Ascospores irregularly arranged, brown, ellipsoidal to oblong, composed of several layers of brown to dark brown, thick cells
1-celled, without germ slits or germ pores, smooth-walled. of textura angularis. Asci 8-spored, unitunicate, subglobose to
Asexual morph: Hyphomycetous. Synnemata determinate, globose, sessile, lacking a visible discharge apparatus and del-
dark brown to black. Hyphae of stipe parallel, dark brown, iquescent. Ascospores irregularly arranged, translucent brown,
septa simple. Conidiophores forming a compact, grey pali- ellipsoidal to oblong, 1-celled, without germ slits or germ
sade. Capitulum terminal, globose, in vertical section com- pores, smooth-walled. Asexual morph: Hyphomycetous.
posed of a darkly pigmented dome of angular cells giving rise Synnemata, determinate, caespitose or solitary, cylindrical-
to loose textura intricata of hyaline, more or less swollen capitate, subulate-capitate, sinuous or straight, of medium stat-
hyphae, a compact subhymenium of similar hyphae and then ure, slender or very slender, unbranched, dark brown to black,
a conidiogenous hymenium. Conidiogenous cells phialidic, slightly lighter immediately beneath the capitulum, smooth or
single or in terminated pairs, light brown, cylindrical, with appearing inconspicuously annellated when dry, annellations
an abruptly terminated apex and cylindrical collarette conspicuous when rehydrated. Hyphae of stipe parallel, dark
smooth-walled. Conidia in dry chains without connectives, brown, septa simple. Conidiophores forming a compact, grey
pale olivaceous, light brown to grey in mass, ellipsoidal, glo- palisade. Capitulum terminal, globose, in vertical section
bose, subglobose or oblong, 1-celled, with thin, smooth-walls. composed of a darkly pigmented dome of angular cells giving
(descriptions based on Seifert 1987; Samuels and Rodrigues rise to loose textura intricata of hyaline, more or less swollen
1989). hyphae, a compact subhymenium of similar hyphae and then a
Notes: Batistiaceae was introduced by Samuels and conidiogenous hymenium. Conidiogenous cells phialidic, sin-
Rodrigues (1989) as a monotypic family classified in gle or in terminated pairs, cylindrical, light brown, with an
Sordariales, and is typified by Batistia. Kirk et al. abruptly terminated apex and cylindrical, smooth-walled
(2001) removed the family from the order Sordariales to collarette. Conidia in dry chains without connectives, pale
Sordariomycetidae incertae sedis (Huhndorf et al. 2004a; olivaceous, light brown to grey in mass, ellipsoidal, globose,
Lumbsch and Huhndorf 2007b). Based on analysis of subglobose or oblong, 1-celled, with thin, smooth wall (Seifert
LSU sequence data, Huhndorf et al. (2004a) showed the 1987; Samuels and Rodrigues 1989).
genus to belong outside the perithecial ascomycetes and Type species: Batistia annulipes (Mont.) Cif., Atti Ist bot
placed it in the Pezizomycotina incertae sedis. Lumbsch Univ Lab crittog Pavia, Ser. 5 15: 166 (1958)
and Huhndorf (2010) and Wijayawardene et al. (2012) Notes: Batistia is typified by Batistia annulipes, and was
followed this classification. However, sequence data pub- originally described as a species of Thamnomyces
lished by Huhndorf has been shown to be contaminated (Xylariaceae) by Montagne (1834) (Benny and Kimbrough
(Maharachchikumbura et al. 2015). Batistiaceae was clas- 1980; Samuels and Rodrigues 1989). Samuels and
sified in Sordariomycetidae family incertae sedis and there Rodrigues (1989) provided a detailed description and illustra-
is presently no reliable sequence data available for this tion for Batistia and its asexual morph and linked to
family in GenBank (Kirk et al. 2008; B. annulipes with the type species of Acrostroma
Maharachchikumbura et al. 2015; Index Fungorum 2016). (A. annellosynnema) by culture methods. Kirk et al. (2008),
This family is characterized by stipitate ascomata that Seifert et al. (2011) and Wijayawardene et al. (2012) accepted
form on wood and with a relatively thick peridium this placement. We re-examined the type material of both
(Benny and Kimbrough 1980). Batistia and its asexual Acrostroma and Batistia, and found the morphology matched
morph, Acrostroma are presently placed in the family that reported by Samuels and Rodrigues (1989). Thus, in this
(Wijayawardene et al. 2012; Maharachchikumbura et al. paper, we synonymize the genus Acrostroma under Batistia.
2015). The genus Acrostroma presently has three records in Index
Fungorum (2016).
Batistia Cif., Atti Ist. bot. Univ. Lab. crittog. Pavia, Ser. 5
15: 166 (1958) Beltraniaceae Nann., Repert. mic. uomo: 498. 1934.
Facesoffungi number: FoF 01322, Figs 14, 15 emend.
S a p rob i c o n w o o d. Se x u a l m o r ph : A s c o m at a Facesoffungi number: FoF 01899
cleistothecial, solitary to gregarious, superficial, stipitate-cap- Saprobic on plant tissues. Sexual morph: Ascomata pale
itate, stipitate sinuous or straight, unbranched, with evenly yellow, solitary to aggregated on OA and PDA, globose to
 Fungal Diversity

Fig. 14 Batistia annulipes (Material examined: BRAZIL, Rio de longitudinal section g Stipitate tissues from a longitudinal section h
Janeiro, Corcovado, on bark, Montagne, 1831–1833, MNHN-PC- Peridium i-l Asci m-n Ascospores. Scale bars: c = 2000 μm,
PC0167686, MC10053b, holotype) a Herbarium material b Host c d = 200 μm, e = 50 μm, f = 100 μm, g = 30 μm, h = 10 μm, i–l = 3 μm,
Ascomata on the host d Ascoma e Peridium with asci f Ascoma in m–n = 2 μm
Fungal Diversity

Fig. 15 Batistia annulipes (Material examined: VENEZUELA, Dto. d Synnemata from longitudinal section e Stem tissues from a longitudinal
Fed. Ca. 7 km. S. of Chichiriviche, on road between El Portachuelo and section f Detail of capitulum tissues from longitudinal section g
Chichiriviche, on unidentified wood, K.P. Dumont, R.F. Cain, G.J. Conidiogenous cells h–i Conidia. Scale bars: c = 500 μm, d = 50 μm,
Samuels and B. Manara, 30 July 1972, NY 00936852 (Dumont-VE e = 30 μm, f = 20 μm, g = 10 μm, h–j = 3 μm
6722), holotype) a Herbarium material b Host c Synnemata on the host
 Fungal Diversity

somewhat papillate, with central ostiole; wall of 3–4 layers of

subhyaline textura angularis to intricata. Pseudoparaphyses
hyaline, septate, cellular, anastomosing, distributed
among asci. Asci 8-spored, sessile, unitunicate, hyaline,
subcylindrical. Ascospores tri- to multi-seriate, obovoid,
hyaline, granular, smooth, aseptate with non-persistent
mucoid sheath. Asexual morph: Hyphomycetous.
Mycelium immersed to superficial, composed of
subhyaline to brown, thin-walled hyphae. Stromata usu-
ally present, parenchymatous to pseudoparenchymatous,
hyaline to brown, often confined to epidermal cells.
Setae present or absent, straight, thick-walled, dark
brown, smooth or verrucose, with radially lobed basal
cell, tapering to acute apex. Conidiophores simple,
erect, septate, pale brown, arising from the base of setae
or separate. Conidiogenous cells pale brown, integrated,
denticulate (based on Crous et al. 2015c).
Type: Beltrania Penz.
Notes: Beltraniaceae was introduced by Nannizzi in 1934,
but, the family was not commonly used. Crous et al. (2015c)
validated the family with the support of sequence data and
also included Beltraniella, Beltraniomyces, Beltraniopsis,
Parapleurotheciopsis, Porobeltraniella, Pseudobeltrania and Fig. 16 Beltrania rhombica (redrawn from Seifert et al. 2011)
Subramaniomyces. In earlier studies, based on sequence data,
Beltraniella, Pseudomassaria carolinensis and allied hypho-
mycetes formed an independent clade in the Xylariales line- Other genera included
age, closely related to Amphisphaeriaceae (Shirouzu et al.
2010) Species in Beltraniaceae are generally Beltraniella Subram., Proc. Indian Acad. Sci., Sect. B 36: 227
hyphomycetous and commonly isolated as saprobes. Crous (1952)
et al. (2015c) introduced Pseudobeltrania ocoteae, which Type species: Beltraniella odinae Subram., Proc. Indian
was the first recorded sexual morph for the family. Acad. Sci., Pl. Sci. 36: 224 (1952)
Beltraniomyces Manohar., D.K. Agarwal & Rao, Indian
Beltrania Penz., Nuovo G. bot. ital. 14: 72 (1882) Phytopath. 56(4): 418 (2003)
Facesoffungi number: FoF 02101; Fig. 16. Type species: Beltraniomyces lignicola Manohar. et al.,
Saprobic on leaf litter. Sexual morph: Undetermined. Indian Phytopath. 56(4): 420 (2003)
Asexual morph: Setae erect, numerous, dark to medium Beltraniopsis Bat. & J.L. Bezerra, Publicações Inst. Micol.
brown, thick-walled, septate, straight or flexuous, tapering Recife 296: 4 (1960)
to an acute apex, septate, basal cell lobed. Conidiophores Type species: Beltraniopsis esenbeckiae Bat. & J.L.
erect or mononematous, unbranched, medium brown, Bezerra, Publicações Inst. Micol. Recife 296: 7 (1960)
smooth, septate several times. Conidiogenous cells termi- Parapleurotheciopsis P.M. Kirk, Trans. Br. mycol. Soc.
nal, discrete, pale to dark brown, polyblastic, globe to 78(1): 63 (1982)
subglobose, usually aseptate. Conidia solitary, biconic, Type species: Parapleurotheciopsis inaequiseptata
colourless to dark brown, aseptate, with or without apical (Matsush.) P.M. Kirk, Trans. Br. mycol. Soc. 78(1): 65 (1982)
appendages. Porobeltraniella Gusmão, Mycologia 96(1): 151 (2004)
Notes: This genus is typified by Beltrania rhombica Penz. Type species: Porobeltraniella porosa (Piroz. & S.D. Patil)
which was isolated from Citrus limonum in Sicily. Generally Gusmão, Mycologia 96(1): 151 (2004)
in Beltrania and related genera, conidiogenous cells are inte- Pseudobeltrania Henn., Hedwigia 41: 310 (1902)
grated, polyblastic and often associated with almost colourless Type species: Pseudobeltrania cedrelae Henn., Hedwigia
to pale brown separating cells that usually become detached 41: 310 (1902)
with the conidia (Bhat and Kendrick 1993). Presently there are Subramaniomyces Varghese & V.G. Rao, Kavaka 7: 83
18 epithets for Beltrania (Index Fungorum 2016). (1980) [1979]
Type species: Beltrania rhombica Penz., Michelia 2(no. 8): Type species: Subramaniomyces indicus Varghese & V.G.
474 (1882) Rao, Kavaka 7: 83 (1980) [1979]
Fungal Diversity

Key to genera of Beltraniaceae order Coronophorales based on multi-gene analysis and trans-
ferred the genus Gaillardiella (typified by G. pezizoides Pat.)
1. Conidiophores lacking radial lobes at base to this family. Gaillardiella has superficial, brown ascomata
……………………………………….... Beltraniomyces with roughened papulose peridium (Miller and Huhndorf
1. Conidiophores with radially lobed basal cells…….......2 2009). Maharachchikumbura et al. (2015), used a multigene
2. Conidia lacking transverse, pale band ..........................3 sequence dataset to confirm the placement of Bertia and
2. Conidia with transverse, pale band................................4 Gaillardiella in Bertiaceae in a highly supported clade in the
3. Conidiogenous cells ellipsoidal, not integrated in conid- Coronophorales.
iophores……………………………... Porobeltraniella
3. Conidiogenous cells cylindrical, Bertia De Not., G. bot. ital. 1(1): 334 (1844)
integrated.......................................Subramaniomyces Facesoffungi number: FoF 01112; Fig. 17
4. Setae absent ......................................... Pseudobeltrania Saprobic on wood in freshwater and terrestrial habitats.
4. Setae present ………………………………………...5 Sexual morph: Ascomata perithecial, solitary or gregarious,
5. Conidiophores septate, or arriving from, basal cells of superficial or erumpent, globose to subglobose, dark brown to
setae……..…..…..…..…..…..…..…..…..…..…...….. 6 black, carbonaceous to membranaceous, tuberculate or
5. Conidiophores arising from all cells of setae………....7 smooth, collabent or collapsing laterally or not collapsing,
6. Conidia biconic…………………………..… Beltrania with a short papilla. Peridium thick (100–250 μm), composed
6. Conidia fusoid…..................…… Parapleurotheciopsis of three layers, outer layer comprising dark tissues, thin, car-
7. Conidia biconic, proximal end rounded, distal free end bonaceous; middle layer comprising dark brown to brown
rostrate…….….….….….…..….……….. Beltraniopsis cells of textura angularis, thick, membranaceous, munk pores
7. Conidia lageniform, proximal end rostrate, distal free present and the inner layer comprising hyaline cells of textura
end truncate................................................. Beltraniella prismatica, thin, membranaceous. Hamathecium composed of
few, inflated, branched and hyaline paraphyses. Asci 8-spored,
Bertiaceae Smyk, Ukr. bot. Zh. 38(6): 47 (1981) unitunicate, clavate, with long tapering pedicel, apical ring
Facesoffungi number: FoF 01111 indistinct or absent. Ascospores irregularly arranged, hyaline
Saprobic on wood in freshwater and terrestrial habitats. to brown, cylindrical or fusiform, aseptate to multi-septate,
Sexual morph: Ascomata perithecial, dark brown to black, slightly curved, smooth-walled, mostly with guttules.
solitary or in small to large groups, superficial to erumpent, Asexual morph: Undetermined.
cupuliform, globose to subglobose, carbonaceous to Type species: Bertia moriformis (Tode) De Not., G. bot.
membranaceous, turbinate or tuberculate or smooth, collabent ital. 1(1): 335 (1844)
or collapsing laterally or not collapsing, ostiolate. Peridium Notes: The genus Bertia was introduced by De Notaris
thick, munk pores present or absent, outer layer (1844) and 47 epithets are listed in Index Fungorum (2016).
(pseudoparenchymatic cells) hard, composed of dark tissues, The type species, B. moriformis, is characterized by tubercu-
basal part mixed with host cells; inner layer composed of late ascomata, clavate asci and hyaline ascospores. The
brown to dark brown to hyaline cells of textura angularis. ascomata are mostly tuberculate, when ascomata are smooth
Hamathecium composed of few, inflated, branched and hya- the ascospores are consistently brown (Mugambi and
line paraphyses. Asci 8-spored, unitunicate, clavate, long ped- Huhndorf 2010).
icellate, apical ring indistinct or absent. Ascospores 2–3-seri-
ate or irregularly arranged, hyaline to brown, cylindrical to
fusiform, aseptate to multi-septate, smooth-walled, mostly
with guttules. Asexual morph: Undetermined. Other genus included
Type: Bertia De Not., G. bot. ital. 1(1): 334 (1844)
Notes: The family Bertiaceae was introduced by Smyk Gaillardiella Pat., in Patouillard & Lagerheim, Bull. Soc.
(1981) based on superficial, black, turbinate ascomata and mycol. Fr. 11(4): 226 (1895)
clavate asci with hyaline, fusiform ascospores, irregularly ar- Type species: Gaillardiella pezizoides Pat., in Patouillard &
ranged in the upper region of ascus, and typified by Bertia (De Lagerheim, Bull. Soc. mycol. Fr. 11(4): 226 (1895)
Notaris 1844). Nannfeldt (1975) concluded that Nitschkiaceae
was composed of three separate groups, but that the differ-
ences between them were not distinct enough for separate
families. However, he expected that Bertia should have an Key to genera of Bertiaceae
distinct position in the family. Smyk (1981) agreed with this
perspective and introduced the family Bertiaceae based on 1. Ascomata tuberculate, ascospores hyaline (if the ascomata
morphology. Mugambi and Huhndorf (2010) reevaluated the smooth, ascospores brown) ……………..................… Bertia
 Fungal Diversity

Fig. 17 Bertia moriformis (Material examined: ITALY, Riva Valdobbia, arrowed h-j Asci k-n Ascospores. Notes: j, m, n stained in Melzer’s
on dead wood of Rhododendron sp., 30 April 1859, Carestia, S F298) a reagent, f, i, k, l in KOH. Scale bars: c = 1 mm, d = 200 μm,
Label and herbarium material b Herbarium material c Black ascomata on e = 100 μm, f = 50 μm, h–k = 20 μm, g. l–n = 10 μm
host surface d Ascoma in vertical section e-f Peridium g Munk pores
Fungal Diversity

1. Ascomata tuberculate, ascospores brown = Bionectria Speg., Boln Acad. nac. Cienc. Córdoba 23(3–
......…………….……………..........…… Gaillardiella 4): 563 (1919) [1918]
Facesoffungi number: FoF 02102; Figs 18, 19
Bionectriaceae Samuels & Rossman, Stud. Mycol. 42: 15 Herbicolous, corticolous or fungicolous. Sexual morph:
(1999) Ascomata forming on a well-developed, erumpent stroma, rare-
Facesoffungi number: FoF 01367 ly superficial on the substratum, densely crowded in large
Herbicolous, corticolous, lichenicolous or fungicolous, numbers, sometimes solitary, perithecial, globose, in orange
found in terrestrial, freshwater and marine habitats. Sexual hues (yellowish orange, pale orange, light orange, brownish
morph: Ascomata embedded in a weakly or well-developed orange), without changing colour in KOH or lactic acid,
erumpent stroma, or superficial on the substratum, solitary or slightly papillate, without a neck, smooth to rough or warted.
densely aggregated, crowded, Ascomata perithecial, rarely Peridium composed of two or three strata, outer strata com-
cleistothecial, if perithecial, globose, subglobose to pyriform, posed of angular to subglobose cells, merging into the stro-
if cleistothecial, globose, white, yellow, orange, reddish brown, ma and into the cells of warts (when present), central strata
greenish or purple to violet, without changing colour in KOH or consisting of intertwined hyphae forming a textura intricata
lactic acid, smooth to rough, or warted, ostiolate or non- in surface view, inner strata composed of layers of lobed
ostiolate. Peridium composed of 1–3 layers. Periphyses present cells, with conspicuous pseudopores in the cell wall.
or absent. Asci (2–)8-spored, uniseriate, biseriate, multi-seriate Periphyses persisting or not. Asci 8-spored, unitunicate, cla-
or irregular, unitunicate, clavate, saccate, cylindrical, sessile or vate, apex rounded, flat, or with prominent edges, with or
short pedicellate, rarely evanescent, apex simple or with an without a refractive ring. Ascospores 0–1-septate, ellipsoid,
inconspicuous or distinct, J-, apical ring. Ascospores unicellular not disarticulating into part-spores, hyaline, smooth-walled,
to multi-septate, sometimes muriform, globose, fusiform, ellip- spinulose to tuberculate or striate. Asexual morph:
soid or broadly ellipsoid, hyaline, smooth-walled, spinulose to Hyphomycetous. Conidiophores dimorphic or monomorphic,
tuberculate or striate. Asexual morph: Hyphomycetous, mononematous or forming sporodochia, hyaline, smooth-
acremonium- or gliocladium-like. Conidiophores dimorphic walled, generally differentiated into a stipe and a branched
or monomorphic, commonly sporodochial or synnematous, part (penicillus). Stipes arise erect from a submerged
[?] hyaline, subhyaline to brown or blackish brown, smooth- supporting hypha, a solitary aerial hypha or aerial hyphal
walled to finely echinulate. Conidiogenous cells phialidic. fascicles, ropes or strands, consisting of one to several cells
Phialides mostly cylindrical to flask-shaped. Conidia unicellu- and is always delimited by a septum near the base.
lar to multi-septate, ellipsoidal, fusiform to subfusiform, some- Penicillus consists of single whorl of phialides
times with papillate or truncate ends, hyaline to greenish hya- (monoverticillate), several whorls of phialides arising from
line or olivaceous grey, smooth or striate walled. intercalary cells of the main axis (2– or more-level verticil-
Type: Bionectria Speg. (= Clonostachys Corda) late, or consists of successive branches that form whorls of
Notes: The family Bionectriaceae was introduced by additional supporting cells or whorls of phialides
Rossman et al. (1999) to accommodate 26 genera including five (biverticillate, terverticillate). Conidiogenous cells phialidic.
cleistothecial genera and with Bionectria Speg. as the type genus. Phialides cylindrical to flask-shaped, with or without trun-
Although it included five cleistothecial genera, four of them were cate apex in secondary conidiophores. Conidial masses hya-
accepted in the family according to analysis of molecular data line, salmon, yellowish, green or hardly pigmented, arrange
(Rossman et al. 2001). Rossman et al. (2013) suggested synon- in imbricate chains. Conidia aseptate or 1-septate, ellipsoidal
ymizing Spicariaceae, which has rarely been cited, under to subfusiform, slightly curved, frequently having a some-
Bionectriaceae. This family is characterized by uniloculate peri- what flattened side, and a laterally displaced hilum, rarely
thecial or rarely cleistothecial ascomata, that are white, yellow, straight with an almost median hilum or entirely symmetrical
orange to tan or brown, not changing colour in KOH or lactic without a visible hilum, hyaline or greenish hyaline, smooth
acid, and are generally superficial, lacking a stroma, or immersed or very rarely ornamented.
in the substratum (Rossman et al. 1999). Many of the genera Type species: Clonostachys araucaria Corda, in Lombard,
were based on species initially described in Nectria (Rossman van der Merwe, Groenewald & Crous 2015
et al. 1999). Phylogenetic analysis by Rossman et al. (2001) Notes: Spegazzini (1919) proposed the genus Bionectria
including those genera and related asexual morph taxa has con- (based on characters of the asexual morph) for species of
firmed that Bionectriaceae is monophyletic within Hypocreales. Nectria that occur on living plant material. He originally in-
Currently 39 genera are listed under this family cluded one species based on a single specimen, which was
(Maharachchikumbura et al. 2015). characterized by flesh-coloured to orange perithecia, crowded
on a well-developed stroma, and fusiform, apically rounded
Clonostachys Corda, Pracht-Fl. Eur. Schimmelbild.: 31 asci, with 1-septate ascospores. Subsequent authors consid-
(1839) ered Bionectria a synonym of Nectria (Müller and von Arx
 Fungal Diversity

Fig. 18 Clonostachys compactiuscula (sexual morph) (Material 00966769, holotype). a Herbarium material b, c Ascomata on host
examined: USA, North Carolina, Jackson Co., Nantahala Natl. Forest, substrate d, e Transverse section through ascomata f Peridium g, h Asci
Bull Pen Rd. to Chattooga River, Ellicott Rock Trail from Fowler i Ascus in Melzer’s reagent j-l Ascospores m Ascospore in Melzer’s
Creek, on bark of recently dead Fagus sp., 28 Sept 1989, G.J. Samuels, reagent. Scale bars: b–d = 500 μm, e = 50 μm, f = 100 μm, g–
C.T. Rogerson, R.C. Harris, W.R. Buck, Det. G.J. Samuels, NY h = 20 μm, i = 20 μm, j–m = 10 μm

1962; Samuels 1988). Bionectria forms a monophyletic clade given priority because it is the older asexual genus and the
based on analyses of DNA sequence data (Schroers 2001; name Clonostachys rosea (the generic type) is commonly
Maharachchikumbura et al. 2015). used in biocontrol studies (Rossman et al. 2013;
The asexual morphs of species in Bionectria are character- Maharachchikumbura et al. 2015).
ized by penicillate, frequently sporodochial, and, in many
cases, dimorphic conidiophores (Schroers 2001). Schroers
(2001) reported a possible link between Clonostachys and Other genera included
Bionectria, and Rossman et al. (2013) have synonymized
Bionectria under Clonostachys. This link was followed in Acremonium Link, Mag. Gesell. naturf. Freunde, Berlin 3(1–
Maharachchikumbura et al. (2015). Clonostachys has been 2): 15 (1809)
Fungal Diversity

Fig. 19 Clonostachys compactiuscula (asexual morph) (Material Conidiophores with multiple conidial columns e Conidiophores in
examined: Dry culture on CMA, NY 00966769) a Dry culture b Congo red f Conidial column stained with Congo red g Conidia Scale
Sporulation on CMA c Conidiophores stained in Congo red d bars: b = 500 μm, c = 100 μm, d = 50 μm, e = 20 μm, f = 50, g = 20 μm

Type species: Acremonium alternatum Link, Mag. Gesell. Type species: Battarrina inclusa (Berk. & Broome) Clem.
naturf. Freunde, Berlin 3(1–2): 15 (1809) & Shear, Gen. fung., Edn 2 (Minneapolis): 279 (1931)
Anthonectria Döbbeler, Mycologia 102(2): 405 (2010) Bryocentria Döbbeler, Mycol. Progr. 3(3): 247 (2004)
Type species: Anthonectria mammispora Döbbeler, Type species: Bryocentria brongniartii (P. Crouan & H.
Mycologia 102(2): 405 (2010) Crouan) Döbbeler, Mycol. Progr. 3(3): 248 (2004)
Aphanotria Döbbeler, Mycol. Res. 111(12): 1408 (2007) Clibanites (P. Karst.) P. Karst., Bidr. Känn. Finl. Nat. Folk 19:
Type species: Aphanotria paradoxa Döbbeler, Mycol. Res. 14, 167 (1871)
111(12): 1408 (2007) Type species: Clibanites paradoxa (P. Karst.) P. Karst.,
Battarrina (Sacc.) Clem. & Shear, Gen. fung., Edn 2 Bidr. Känn. Finl. Nat. Folk 19: 14 (1871)
(Minneapolis): 279 (1931) Didymostilbe Henn., Hedwigia 41: 148 (1902)
 Fungal Diversity

Type species: Didymostilbe coffeae Henn., Hedwigia 41: Type species: Ovicuculospora parmeliae (Berk. & M.A.
148 (1902) Curtis) Etayo, Bull. Soc. linn. Provence 61: 83–128 (2010)
Dimerosporiella Speg., Revta Mus. La Plata 15: 10 (1908) Paranectria Sacc., Michelia 1(no. 3): 317 (1878)
Type species: Dimerosporiella paulistana Speg., Revta Type species: Paranectria affinis (Grev.) Sacc., Michelia
Mus. La Plata 15(2): 11 (1908) 1(no. 3): 317 (1878)
Gliomastix Guég., Bull. Soc. mycol. Fr. 21: 240 (1905) Peristomialis (W. Phillips) Boud., Hist. Class. Discom. Eur.
Type species: Gliomastix chartarum (Cooke) Guég., Bull. (Paris): 116 (1907)
Soc. mycol. Fr. 21: 240 (1905) Type species: Peristomialis berkeleyi Boud., Hist. Class.
Globonectria Etayo, Biblthca Lichenol. 84: 47 (2002) Discom. Eur. (Paris): 116 (1907)
Type species: Globonectria cochensis Etayo, Biblthca Pronectria Clem., in Clements & Shear, Gen. fung., Edn 2
Lichenol. 84: 47 (2002) (Minneapolis): 78, 282 (1931)
Gracilistilbella Seifert, Stud. Mycol. 45: 18 (2000) Type species: Pronectria lichenicola (Cooke) Clem., in
Type species: Gracilistilbella clavulata (Mont.) Seifert, in Clements & Shear, Gen. fung., Edn 2 (Minneapolis): 282 (1931)
Seifert & Samuels, Stud. Mycol. 45: 18 (2000) Protocreopsis Yoshim. Doi, Kew Bull. 31(3): 551 (1977)
Halonectria E.B.G. Jones, Trans. Br. Mycol. Soc. 48(2): 287 Type species: Protocreopsis zingibericola Yoshim. Doi,
(1965) Kew Bull. 31(3): 552 (1977)
Type species: Halonectria milfordensis E.B.G. Jones, Rhopalocladium Schroers et al., in Schroers et al., Mycologia
Trans. Br. Mycol. Soc. 48(2): 287 (1965) 91(2): 375 (1999)
Heleococcum C.A. Jørg., Bot. Tidsskr. 37: 417 (1922) Type species: Rhopalocladium myxophilum Schroers
Type species: Heleococcum aurantiacum C.A. Jørg., Bot. et al., in Schroers et al., Mycologia 91(2): 375 (1999)
Tidsskr. 37: 417 (1922) Roumegueriella Speg., in Roumeguère, Revue mycol.,
Hydropisphaera Dumort., Comment. bot. (Tournay): 89 Toulouse 2(no. 1): 18 (1880)
(1822) Type species: Roumegueriella muricospora Speg., in
Type species: Hydropisphaera peziza (Tode) Dumort., Roumeguère, Revue mycol., Toulouse 2(no. 1): 18 (1880)
Comment. bot. (Tournay): 90 (1822) Selinia P. Karst., Meddn Soc. Fauna Flora fenn. 1: 57 (1876)
Ijuhya Starbäck, Bih. K. svenska VetenskAkad. Handl., Afd. Type species: Selinia pulchra (G. Winter) Sacc., in
3 25(no. 1): 30 (1899) Saccardo, Syll. fung. (Abellini) 2: 457 (1883)
Type species: Ijuhya vitrea Starbäck, Bih. K. svenska Spicellum Nicot & Roquebert, Revue Mycol., Paris 39(4):
Vetensk Akad. Handl., Afd. 3 25(no. 1): 30 (1899) 272 (1976) [1975]
Kallichroma Kohlm. & Volkm.-Kohlm., Mycol. Res. 97(6): Type species: Spicellum roseum Nicot & Roquebert,
759 (1993) Revue Mycol., Paris 39(4): 272 (1976) [1975]
Type species: Kallichroma tethys (Kohlm. & E. Kohlm.) Stephanonectria Schroers & Samuels, in Schroers et al.,
Kohlm. & Volkm.-Kohlm., Mycol. Res. 97(6): 759 (1993) Sydowia 51(1): 116 (1999)
Lasionectria (Sacc.) Cooke, Grevillea 12(no. 64): 111 (1884) Type species: Stephanonectria keithii (Berk. & Broome)
Type species: Lasionectria mantuana (Sacc.) Cooke, Schroers & Samuels, Sydowia 51(1): 116 (1999)
Grevillea 12(no. 64): 112 (1884) Stilbocrea Pat., Bull. Soc. mycol. Fr. 16: 188, 186 (1900)
Mycocitrus Möller, Bot. Mitt. Trop. 9: 297 (1901) Type species: Stilbocrea dussii Pat., Bull. Soc. Mycol. Fr.
Type species: Mycocitrus aurantium Möller, Bot. Mitt. 16: 186 (1900)
Trop. 9: 297 (1901) Stromatocrea W.B. Cooke, Mycologia 44(2): 248 (1952)
Nectriella Nitschke ex Fuckel, Jb. nassau. Ver. Naturk. 23–24: Type species: Stromatocrea cerebriformis W.B. Cooke [as
175 (1870) [1869–70] ‘cerebriforme’], Mycologia 44(2): 248 (1952)
Type species: Nectriella fuckelii Nitschke ex Fuckel, Jb. Stromatonectria Jaklitsch & Voglmayr, Mycologia 103(2):
nassau. Ver. Naturk. 23–24: 176 (1870) [1869–70] 435 (2011)
Nectriopsis Maire, Annls mycol. 9(4): 323 (1911) Type species: Stromatonectria caraganae (Höhn.)
Type species: Nectriopsis violacea (J.C. Schmidt ex Fr.) Jaklitsch & Voglmayr, Mycologia 103(2): 435 (2011)
Maire, Annls mycol. 9(4): 323 (1911) Trichonectria Kirschst., Verh. bot. Ver. Prov. Brandenb. 48:
Ochronectria Rossman & Samuels, in Rossman et al., Stud. 60 (1907) [1906]
Mycol. 42: 53 (1999) Type species: Trichonectria aculeata Kirschst., Verh. bot.
Type species: Ochronectria calami (Henn. & E. Nyman) Ver. Prov. Brandenb. 48: 60 (1907) [1906]
Rossman & Samuels, in Rossman et al., Stud. Mycol. 42: 53 Vesicladiella Crous & M.J. Wingf., in Crous et al., Mycotaxon
(1999) 50: 454 (1994)
Ovicuculospora Etayo, Bull. Soc. linn. Provence 61: 83–128 Type species: Vesicladiella capitata (B. Sutton) Crous &
(2010) M.J. Wingf. [as ‘capitatum’], Mycotaxon 50: 456 (1994)
Fungal Diversity

Verrucostoma Hirooka et al., in Hirooka et al., Mycologia 10. Ascospores subsphaerical, with granulose
102(2): 422 (2010) ornamentation . . . . . . . . . . . . . . . . . . .Globonectria
Type species: Verrucostoma freycinetiae Hirooka et al., in 10. Ascospores fusiform, ovoid or ellipsoid, verruculose or
Hirooka et al., Mycologia 102(2): 422 (2010) smooth walled . . . . . . . . . . . . . . . . . . . . . Pronectria
Virgatospora Finley, Mycologia 59(3): 538 (1967) 11. Ascomata immersed in herbaceous tissue, bark or
Type species: Virgatospora echinofibrosa Finley, wood, rarely fungicolous . . . . . . . . . . . . . . Nectriella
Mycologia 59(3): 538 (1967) 11. Ascomata immersed in thalli of liverworts or mosses,
not fungicolous . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
12. Ascomata immersed in leaves of Polytrichadelphus sp.,
globose with an apical papilla, or pyriform, pale to
Key to the sexual/ and asexual morph genera golden-yellow or pale orange, ascospores composed
of Bionectriaceae of two parts, fertile and sterile . . . . . . . Anthonectria
12. Ascomata immersed in perianth of Drepanolejeunea,
1. Ascomata cleistothecial, globose, asci clavate ………… 2 colourless, ascospores fusiform, colourless, with five
1. Ascomata perithecial, subglobose to pyriform, asci elon- transverse septa . . . . . . . . . . . . . . . . . . . . Aphanotria
gate, very rarely clavate ..…….…….…….……….… 4 13. Ascospores with long attenuated ends, on lichens or
2. Ascospores ellipsoid, 1-septate, smooth to ornamented algae . . . . . . . . . . . . . . . . . . . . . . . . . . . .Paranectria
with wings …………………......……… Heleococcum 13. Ascospores with rounded ends, on lichens, algae, bryo-
2. Ascospores globose, aseptate, ornamented with sharp, phytes, fungi or other substrata. . . . . . . . . . . . . . . . 14
pointed spines ………………….………...……….… 3 14. Ascomata with erect solitary hairs, ascospores 1–multi-
3. Ascomata on dung and well-rotten debris, ascospores septate . . . . . . . . . . . . . . . . . . . . . . . . . Trichonectria
with sparse echinulations, asexual morph Gliocladium- 14. Ascomata without hairs, or if present, hairs fasciculate
like …...…...…...…...…………….......Roumegueriella or flexuous, sometimes with few short setae or tooth
3. Ascomata on fruiting bodies of Tuber, ascospores dense- like structures surrounding ostiolar region, ascospores
ly echinulate, a sexual morph U ndetermined generally 1-septate . . . . . . . . . . . . . . . . . . . . . . . . . 15
.….….….….….….….….….….….….….… Battarrina 15. Ascomata superficial on a thin stroma . . . . . . . . . . 16
4. Ascomata immersed in the substratum or in a stroma that 15. Ascomata superficial on the substratum or immersed in
may itself be immersed in the substratum.….….….… 5 a stroma . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
4. Ascomata superficial or immersed in a hyphal subiculum 17. Ostiolum surrounded by a crown-like structure made
or thin stromata 13 up of angular to oblong, tooth-like or sometimes
5. Ascomata immersed in a stromata that may itself be hypha-like cells, asexual morph Myrothecium-
immersed in the substratum, on dung, corticolous, like . . . . . . . . . . . . . . . . . . . . . . . . . Stephanonectria
herbicolous, not lichenicolous or fungicolous, asco- 17. Ostiolum not surrounded by a crown-like structure,
spores 0–1-septate ………………………………….. 6 asexual morph Acremonium-like . . . . . Verrucostoma
5. Ascomata immersed in substratum, non stromatic, 18. Ascomata immersed in a stroma or superficial with
corticolous, herbicolous, lichenicolous, fungicolous or white to tan or green hyphae covering the ascomatal
bryophilous, not on dung, ascospores non to multi- wall . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
septate or muriform ………………………………… 8 18. Ascomata superficial, seated directly on the substratum,
6. Ascospores aseptate, on dung ………………… Selinia and without white to tan hyphae covering the ascomatal
6. Ascospores 1-septate, corticolous or herbicolous …….7 wall . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22
7. Stromata yellow orange to red or purple, asci fusoid or 19. Ascomata immersed, loosely united in a thin, pseudo-
clavate ……………………………… Stromatonectria parenchymatous stroma, ascospores narrowly cylindri-
7. Stromata buff to rufous, asci cylindrical . . . Mycocitrus cal, smooth walled . . . . . . . . . . . . . . . . . . . Clibanites
8. On algae or wood in marine habitats, ascomata with a 19. Ascomata immersed in an effused hyphal stroma or
long neck. . . . . . . . . . . . . . . . . . . . . . . . . Halonectria superficial with a covering of white to tan or green
8. Terrestrial or in freshwater habitats, ascomata without a hyphae, ascospores ellipsoid to fusiform, occasionally
long neck . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 obovoid, smooth, striate, spinulose, verruculose or
9. Ascomata immersed in thalli of terrestrial lichens, rarely tuberculate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
on algae or fungi, asexual morphs where known, 20. Ascomata pale to medium cream or pink orange,
Acremonium-like or Kutilakesa-like. . . . . . . . . . . . . 10 asci with dimorphic, 1(−2)-septate
9. Ascomata immersed in thalli of bryophytes, herbaceous spores . . . . . . . . . . . . . . . . .Ovicuculospora
tissue, bark or wood, rarely fungicolous, asexual morphs 20. Ascomata hyaline, pale yellow to orange, Asci with
where known, Acremonium-like . . . . . . . . . . . . . . . 11 monomorphic 1-septate spores. . . . . . . . . . . . . . . . 21
 Fungal Diversity

21 Ascomata immersed in an effused hyphal stroma, hairs, ascospores striate, rarely spinulose,
ascospores generally striate, less often smooth herbicolous. . . . . . . . . . . . . . . . . . . . . . . . . . . .Ijuhya
or tuberculate, asexual morph Acremonium-like,
usually on monocotyledonous plant
debris. . . . . . . . . . . . . . . . . . . Protocreopsis Key to the asexual morph genera of Bionectriaceae
21. Ascomata immersed in an effused hyphal stroma
or superficial with tan hyphae covering the 1. Conidiophores monomorphic, neither sporodochial nor
ascomatal wall, ascospores spinulose or verrucose, synnematous, conidia ellipsoidal, polysymmetrical,
asexual morphs synnematous (Stilbella-like) or sometimes slightly curved, on living
Acremonium-like, corticolous or sometimes on myxomycetes . . . . . . . . . . . . . . Rhopalocladium
other ascomycetes . . . . . . . . . . . . . Stilbocrea 1. Conidiophores sporodochial, synnematous or simple,
22. Ascomata white to pale yellow or orange, smooth, or conidia spherical, ellipsoidal, ovate or fusiform, not on
covered with white hyphae or flexuous hairs, asco- myxomycetes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
spores smooth to spinulose, rarely striate, fungicolous 2. Conidiophores simple, septate, unbranched, with termi-
or on liverworts and mosses. . . . . . . . . . . . . . . . . . 23 nal phialides. . . . . . . . . . . . . . . . . . . . . . . . . Spicellum
22. Ascomata white to orange, sometimes with white 2. Conidiophores sporodochial, synnematous . . . . . . . . . 3
warts, with or without hyphal or fasciculate hairs, as- 3. Synnemata, cylindrical-capitate . . . . . . . .Didymostilbe
cospores smooth, spinulose or striate, corticolous, less 3. Sporodochia cerebriform . . . . . . . . . . . . Stromatocrea
often fungicolous or herbicolous . . . . . . . . . . . . . . 25 (This key is a modification of the key of Rossman et al.
23. Ascomata on liverworts or mosses . . . . . .Bryocentria 1999)
23. Ascomata on myxomycetes, pyrenomycetes or
dematiaceous hyphomycetes, rarely on Boliniaceae Rick, Brotéria, sér. bot. 25: 65 (1931)
Aphyllophorales . . . . . . . . . . . . . . . . . . . . . . . . . . 24 Facesoffungi number: FoF 01129
24. Ascomata on Asterina, Meliola, Schnifferula, growing Saprobic on decorticated wood or decaying wood in terres-
on superficial, black hyphae covering living trial habitats. Sexual morph: Ascostromata absent or present,
leaves. . . . . . . . . . . . . . . . . . . . . . . . Dimerosporiella immersed, erumpent to superficial, clypeate to irregular in
24. Ascomata on other fungi, including myxomycetes, shape, light brown to black, coriaceous, membranaceous or
not on Meliola or similar fungi on living powdered (or furaceous), with or without brown, septate in-
leaves . . . . . . . . . . . . . . . . . . . Nectriopsis terwoven hyphae, if present surrounding the apex of the
25. Ascomata globose to subglobose, occasionally ascomata. Ascomata perithecial, solitary to gregarious, brown
doliiform, becoming cupulate upon drying, ascomatal to black, immersed, erumpent to superficial, globose, cylindri-
wall of globose, thin walled cells, ascospores often cal to obpyriform, coriaceous or membranaceous, smooth,
striate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26 papillate or papilla absent, if present with periphysate ostiole,
25. Ascomata globose to subglobose or ovoid, gener- stellate or not. Peridium thick (20–150 μm), outer layer hard,
ally not cupulate upon drying, ascomatal wall of composed of brown, thick-walled cells of textura angularis or
thick walled cells, ascospores smooth, spinulose textura intricata; inner layer soft, composed of hyaline, thin-
or striate. . . . . . . . . . . . . . . . . . . . . . . 27 walled cells of textura prismatica. Hamathecium composed of
26. Ascomata of three regions, with orange oil droplets in abundant, persistent, filamentous, tapering, septate, branched
the middle region of the wall . . . . . . . . Ochronectria paraphyses. Asci 8-spored, unitunicate, cylindrical to clavate,
26. Ascomata of two regions, without orange oil long pedicellate, some with an apical ring. Ascospores 2–3-
droplets. . . . . . . . . . . . . . . . . . . . . . Hydropisphaera seriate, hyaline to brown, versicolored or concolourous (hya-
27. Ascomata with a flattened apex, often with solitary or line or brown), ellipsoid to cylindrical, straight to slightly
fasciculate hairs forming an apical fringe, ascospores curved to suballantoid, 0–2-septate, smooth-walled, mostly
striate or spinulose . . . . . . . . . . . . . . . . . . . . . . . . . 28 with guttules. Asexual morph: Undetermined.
27. Ascomata without distinct hairs, smooth to warted or Type: Bolinia (Nitschke) Sacc
with short hyphal hairs, ascospores smooth, spinulose Notes: The family Boliniaceae was introduced by Rick (1931)
or rarely striate. . . . . . . . . . . . . . . . . . . Clonostachys based on black ascostromata and cylindrical asci with smooth
28. Ascomata yellow-brown to dark brown, globose to ascospores, and is typified by Camarops (Karsten 1873). Earlier,
subglobose, with solitary stiff or hyphal hairs, not this family was placed in Xylariales based on morphological
forming a distinct fringe, ascospores striate or spinu- similarity (Barr 1990; Romero and Samuels 1991). Andersson
lose, herbicolous or corticolous . . . . . . . Lasionectria et al. (1995) concluded that Boliniaceae was more closely related
28. Ascomata white to pale yellow, globose to ovoid, with a to Sordariales based on SSU rDNA sequence data. This family
flattened apical disk, often with a fringe of fasciculate was subsequently placed in Boliniales by Kirk et al. (2001).
Fungal Diversity

Based on phylogenetic and morphological data six genera were Apiorhynchostoma Petr., Annls mycol. 21(3/4): 185
accepted in Boliniaceae (Lumbsch and Huhndorf 2010). (1923)
Untereiner et al. (2013) added Apiorhynchostoma and Type species: Apiorhynchostoma apiculata (Sacc.) Petr.,
Pseudovalsaria based on the LSU rDNA sequence data of Annls mycol. 21(3/4): 185 (1923)
Apiorhynchostoma curreyi and Pseudovalsaria ferruginea. Camaropella Lar.N. Vassiljeva, Mikol. Fitopatol. 31(1): 6
(1997)
Bolinia (Nitschke) Sacc., Syll. fung. (Abellini) 1: 352 Type species: Camaropella pugillus (Schwein.) Lar.N.
(1882) Vassiljeva, Mikol. Fitopatol. 31(1): 6 (1997)
Facesoffungi number: FoF 01130, Fig. 20 Camarops P. Karst., Bidr. Känn. Finl. Nat. Folk 23: 6, 53
Saprobic on decorticated wood or decaying wood in terres- (1873)
trial habitats. Sexual morph: Ascostromata immersed, Type species: Camarops hypoxyloides P. Karst., Bidr.
erumpent to superficial, irregular in shape, black, coriaceous, Känn. Finl. Nat. Folk 23: 53 (1873)
membranaceous. Ascomata perithecial, gregarious, brown to Cornipulvina Huhndorf et al., Fungal Diversity 20: 61
dark brown, immersed, erumpent through bark of host sub- (2005)
strate at maturity, globose to subglobose, coriaceous to Type species: Cornipulvina ellipsoides Huhndorf et al.,
membranaceous, smooth, papillate, with periphysate ostiole. Fungal Diversity 20: 63 (2005)
Peridium thick, outer layer coriaceous, thick-walled, com- Endoxyla Fuckel, Jb. nassau. Ver. Naturk. 25–26: 321
posed of brown to dark brown cells of textura intricata; inner (1871)
layer membranaceous, composed of hyaline cells of textura Type species: Endoxyla operculata (Alb. & Schwein.)
prismatica. Hamathecium composed of abundant, persistent, Sacc., Syll. fung. (Abellini) 1: 181 (1882)
filamentous, septate, branched paraphyses. Asci 8-spored, Mollicamarops Lar.N. Vassiljeva, Mycotaxon 99: 160
unitunicate, cylindrical, slightly curved, blunt apex with bare- (2007)
ly visible apical ring, long pedicellate. Ascospores 2–3-seriate, Type species: Mollicamarops stellata Lar.N. Vassiljeva,
pale brown when young, dark brown at maturity, subglobose Mycotaxon 99: 160 (2007)
to ellipsoid, aseptate, smooth, with guttulate. Asexual morph: Neohypodiscus J.D. Rogers, Y.M. Ju & Læssøe,
Undetermined. Mycologia 86(5): 684 (1994)
Type species: Bolinia tubulina (Alb. & Schwein.) Sacc., Type species: Neohypodiscus rickii (Lloyd) J.D. Rogers,
Syll. fung. (Abellini) 1: 352 (1882) Y.M. Ju & Læssøe, Mycologia 86(5): 685 (1994)
Notes: The name Bolinia was first introduced as a subgenus Pseudovalsaria Spooner, Trans. Br. mycol. Soc. 86(3): 405
by Nitschke (1867), and later raised to generic rank by Saccardo (1986)
(1882). Karsten (1873) introduced the genus Camarops based Type species: Pseudovalsaria foedans (P. Karst.) Spooner,
on the single species C. hypoxyloides P. Karst. Rehm (1904) Trans. Br. mycol. Soc. 86(3): 405 (1986)
and Theissen (1999) considered Bolinia tubulina to be a syno-
nym of Camarops hypoxyloides, but Miller (1930) disagreed
based on a re-evaluation of specimens. Miller (1930) however, Key to genera of Boliniaceae
accepted C. polyspermum (Mont.) J.H. Mill as a synonym of
C. hypoxyloides. Nannfeldt (1972) reviewed the taxonomy of 1. Ascostroma immersed . . . . . . . . . . . . . . . . . . . . . . . . . . 2
these taxa and concluded that Camarops and Bolinia were two 1. Ascostroma erumpent or superficial . . . . . . . . . . . . . . 5
different genera, but having similar morphological characters. 2. Ascostroma poorly-developed . . . . . . . . . . . . . . . . . . 3
However, Lumbsch and Huhndorf (2010) excluded Bolinia and 2. Ascostroma well-developed . . . . . . . . . . . . . . . . . . . . 4
included it as the synonym of Camarops (Index Fungorum 3. Ascospores 2-septate, with a hyaline basal cell and two
2016). Bolinia tubulina (current name Camarops tubulina) is dark brown cells at the center, the shorter brown cell
the type of Bolinia (Index Fungorum 2016). In our opinion, with an apical germ pore at the end Apiorhynchostoma
Bolinia tubulina should be accepted as the type species and 3. Ascospores 1-septate, with a hyaline cell and a brown
Camarops should be regarded as an independent genus. cell with an apical germ pore at the end . . . . Endoxyla
4. Ascospores aseptate, brownish . . . . . . . . .Camaropella
4. Ascospores 1-septate, dark brown . . . . Pseudovalsaria
Other genera included 5. Ascospores hyaline . . . . . . . . . . . . . . . . Cornipulvina
5. Ascospores brown or composed of a brown cell and
Apiocamarops Samuels & J.D. Rogers, Mycotaxon 28(1): 54 hyaline cell . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
(1987) 6. Ascospores one colour 7
Type species: Apiocamarops alba Samuels & J.D. Rogers, 6. Ascospores 1-septate, composed of a large, porate brown
Mycotaxon 28(1): 54 (1987) cell and a small, hyaline cell. . . . . . . . . Apiocamarops
Fungal Diversity
 Fungal Diversity

ƒFig. 20 Bolinia tubulina (Material examined: SWEDEN, Bökeberg Maharachchikumbura et al. 2015). The combined LSU and
Forest in Femsjö, on the trunks, 18 October 1967, PRM PR647008) a ITS gene analysis also confirmed the phylogenetic placement
Material and herbarium packet b, c Stroma with immersed ascomata d
Stroma in cross section e Ascoma in cross section f Peridium g Periphyses
of Atrotorquata in Cainiaceae (Senanayake et al. 2015).
h Paraphyses i–k Asci l Apical ring of ascus m–p Ascospores. Notes: Another genus, Amphibambusa was introduced from bamboo
Figs e-p are stained in KOH. Scale bars: d = 1 mm, b. e = 500 μm, in Liu et al. (2015) and LSU gene analysis and morphology
c = 200 μm, g = 30 μm, f = 20 μm, h–k = 10 μm, l–p = 5 μm indicated that the genus belongs to Cainiaceae. Based on both
morphological and phylogenetic data, Senanayake et al.
7. Ascomata with stellate ostioles. . . . . . . Mollicamarops (2015) accepted the genera Amphibambusa, Arecophila,
7. Ascomata lacking stellate ostioles . . . . . . . . . . . . . . . 8 Atrotorquata, Cainia and Seynesia in Cainiaceae. The asexual
8. Perithecia monostichous. . . . . . . . . . . . . . . . Camarops morph of Cainiaceae from a pure culture of Cainia desmazieri
8. Perithecia polystichous. . . . . . . . . . . . . . . . . . . Bolinia C. Moreau & E. Müll. ex Krug has been observed and was
reported by Muller and Corbaz (1956) as Rhabdospora-like.
Cainiaceae J.C. Krug, Sydowia 30 (1–6):123 (1978) However, later attempts at obtaining the asexual morph from
[1977] Cainia desmazieri in culture have not been successful (Kang
Facesoffungi number: FoF 00687 et al. 1999a, b; Senanayake et al. 2015). The genera included
Saprobic on dead grasses, bamboo and other monocotyle- in this family are confused and may be wrongly placed. Their
dons, appearing as shiny black dots, slightly effuse from the placement is pending molecular data.
substrate. Sexual morph: Pseudostromata poorly developed
or lacking, or sometimes clypeate, scattered, superficial, Cainia Arx & E. Müll., Acta bot. neerl. 4 (1): 111 (1955)
dome-shaped or slightly effuse, dark brown to black. Facesoffungi number: FoF 00688; Fig. 21
Ascomata immersed, solitary or aggregated, globose to Saprobic or pathogenic on Poaceae and Cyperaceae,
subglobose, coriaceous, brown, ostiolate. Ostiolar papilla forming leaf or stem spots, appearing as shiny, black, scattered
short, internally lined with hyaline periphyses. Peridium one dots on host surface. Sexual morph: Pseudoclypeus indis-
or two-layered, outer layer comprising of thick-walled, brown tinct, scattered, dome-shaped, effuse, dark brown to black,
cells of textura angularis and inner layer comprising of hya- smooth, each containing one, or rarely several ascomata.
line thin-walled cells of textura angularis. Hamathecium Ascomata immersed, scattered, solitary or aggregated,
comprising abundant, filamentous paraphyses, slightly con- ampulliform, coriaceous, with ostiolate. Ostioles short, inter-
stricted at the septa. Asci 8-spored, unitunicate, cylindrical to nally lined with hyaline, filamentous periphyses. Peridium
broadly cylindrical, short pedicellate, with a complex J+, cy- comprising an outer layer of thick-walled, brown, textura
lindrical apical ring or series of rings. Ascospores uniseriate, angularis cells and inner layer comprising of thick-walled,
overlapping uniseriate to biseriate, hyaline when young and hyaline, textura angularis cells. Hamathecium comprising nu-
dark brown at maturity, sphaerical to ellipsoidal, unicellular to merous, septate paraphyses, slightly constricted at the septa,
1-septate, slightly constricted at the septum, wall ornamented Asci 8-spored, unitunicate, cylindrical to clavate, short pedi-
with longitudinal germ slits or germ pores, surrounded by a cellate, apex rounded, with a complex, J+, dome-shaped to
gelatinous sheath. Asexual morph: Coelomycetous. cylindrical series of rings. Ascospores overlapping uniseriate
Conidiomata pycnidial, scattered, immersed, globose to to biseriate, dark brown, ellipsoidal, 1- septate, constricted at
subglobose, black. Conidiophores hyaline, denticulate, the septum, wall with longitudinal striations, surrounded by a
sympodially proliferating. Conidiogenous cells with 1–3 gelatinous sheath. Asexual morph: Coelomycetous.
phialides, filiform, branched or simple, septate, hyaline. Conidiomata pycnidia, scattered, immersed, globose to
Conidia elongate fusiform, falcate to lunate, unicellular or subglobose, black. Conidiophores hyaline, denticulate.
septate, hyaline, with pointed ends. Conidiogenous cells with 1–3 phialides, filiform, simple or
Type: Cainia Arx & E. Müll. branched, septate, hyaline. Conidia elongate-fusiform to fili-
Notes: The family Cainiaceae was introduced by Krug form, falcate to lunate, with holoblastic conidiogenesis, uni-
(1977) to accommodate the species with unique apical ring cellular or septate, hyaline, sometimes with pointed ends
of the asci, which consisted of a series of rings, and ascospores (description of asexual morph is based on Krug 1977 and
with longitudinal germ slits. The genera Arecophila, Kang et al. 1999a, b).
Atrotorquata, Cainia, Ceriophora, Reticulosphaeria and Type species: Cainia graminis (Niessl) Arx & E. Müll.,
Ommatomyces have been placed in this family at various Acta bot. neerl. 4(1):112 (1955)
times (Kohlmeyer and Kohlmeyer 1993; Hyde et al. 1996). Notes: The genus was introduced by von Arx and Müller
Seynesia was included based on phylogenetic analysis by (1955). It was distinguished from Amphisphaeriaceae based
Maharachchikumbura et al. (2015). The placement of on the complex structure of the apical ring and presence of
Arecophila, Cainia and Seynesia in Cainiaceae was con- germ slits in the ascospores and thus, accommodated in
firmed based on molecular analysis (Smith et al. 2003; Cainiaceae. Senanayake et al. (2015) designated a reference
 Fungal Diversity

Fig. 21 Cainia graminis (Material examined: ITALY, Province of Forlì- of ascomata e Peridium f Paraphyses g-i Asci j J+ apical ring in Melzer’s
Cesena [FC], Santa Sofia, Corniolo, stem of Lolium temulentum L reagent k-q Ascospores r Sheath surrounding ascospore after treating
(Poaceae), 25 September 2013, Erio Camporesi, IT 1462, MFLU 15- with Indian ink. Scale bars: a = not to scale, c, g–j = 100 μm,
0724)) a Ascomata on host b Close up of ascomata c, d Vertical section d = 250 μm, e, f, o–r = 25 μm, k–n = 10 μm

specimen for the type Cainia graminis. Currently, five epithets Type species: Amphibambusa bambusicola D.Q. Dai &
have been listed for Cainia in Index Fungorum (2016). K.D. Hyde, Liu et al., Fungal Divers. 72: 7 (2015)
Arecophila K.D. Hyde, Nova Hedwigia 63(1–2):82 (1996)
Type species: Arecophila gulubiicola K.D. Hyde, Nova
Hedwigia 63(1–2):91 (1996)
Other genera included Atrotorquata Kohlm. & Volkm.-Kohlm., Syst. Ascom.
12(1–2):8 (1993)
Amphibambusa D.Q. Dai & K.D. Hyde, in Liu et al., Fungal Type species: Atrotorquata lineata Kohlm. & Volkm.-
Divers. 72: 7 (2015) Kohlm., Syst. Ascom. 12(1–2):8 (1993)
Fungal Diversity

Seynesia Sacc., Syll. fung. (Abellini) 2:668 (1883) subcylindrical, pigmented in the apical. Conidiogenous cells
Type species: Seynesia nobilis (Welw. & Curr.) Sacc., Syll. enteroblastic or holoblastic, phialidic, hyaline, with brown
fung. (Abellini) 2:668 (1883) apex, smooth-walled. Conidia aggregated, hyaline, aseptate
to multi-septate, cylindrical to allantoid, sometimes 4–5-radi-
ate and consisting of a main axis and three or four arcuate
Key to the genera of Cainiaceae branches inserted near its base, with or without guttules
(from Réblová et al. 2004).
1. Apical apparatus consisting of a wedge-shaped or conical, Type: Calosphaeria Tul. & C. Tul.
subapical ring . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 Notes: The family Calosphaeriaceae was introduced by
1. Apical apparatus consisting of several rings . . . . . . . . 4 Munk (1957) based on immersed, globose ascomata, clavate,
2. Ascospores bearing a single conical appendage emerg- pedicellate asci with hyaline allantoid ascospores,
ing from the outer sheath . . . . . . . . . . . . . . . . Seynesia enteroblastic, phialidic conidiogenesis, micronematous conid-
2. Ascospores not bearing an appendage as above . . . . . 3 iophores and hyaline, aseptate conidia, and is typified by
3.Ascospores fusiform to broad Calosphaeria. Many asexual morphs have been reported for
fusiform . . . . . . . . . . . . . . . .Amphibambusa this family (Hyde et al. 2011; Wijayawardene et al. 2012), but
3. Ascospores ellipsoidal . . . . . . . . . . . . . . . . Arecophila information is incomplete. Calosphaeria has been referred to
4. Apical apparatus wedge-shaped, ascospores having 5–7 various families based on the allantoid ascospores. The genera
longitudinal striations at each apex. . . . . .Atrotorquata Calosphaeria and Pleurostoma were placed in the
4. Apical apparatus dome-shaped to cylindrical or Diatrypaceae (Fuckel 1870; Saccardo 1882; Winter 1885;
appearing as plates, ascospores bearing a germ slit at Schroeter 1897); Allantosporae (Berlese 1900); Valsaceae
each end . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Cainia (Traverso 1906); and Allantosphaeriaceae (von Höhnel
1918). Petrak (1923, 1941) included Phragmocalosphaeria
Calosphaeriaceae Munk, Dansk bot. Ark. 17(no. 1): 278 and Calosphaeriopsis in this family because of similar mor-
(1957) phological characters. Wehmeyer (1975) accepted
Facesoffungi number: FoF 01133 Calosphaeriaceae in Allantosphaeriales, while Barr (1983)
Saprobic on wood or decorticated wood or leaves in fresh- recognized this family in Calosphaeriales. Lumbsch and
water and terrestrial habitats, sometimes fungicolous. Sexual Huhndorf (2010) included eight genera (Calosphaeria,
morph: Ascostromata scattered, erumpent through leaf epi- Conidiotheca, Jattaea, Kacosphaeria,
dermis, ellipsoid, long axis parallel to the leaf surface. Phragmocalosphaeria, Sulcatistroma, Togniniella,
Ascomata perithecial, scattered or gregarious, dark brown to Wegelina) in this family, while Maharachchikumbura et al.
black, superficial, erumpent or immersed, globose to (2015) accepted Calosphaeria (= Calosphaeriophora),
subglobose, coriaceous or membranous, tuberculate or J a t t a e a ( = P h r a g m o c a l o s p h a e r i a , = We g e l i n a ) ,
smooth or with brown, septate, hyphal coating, ostiolate. Kacosphaeria, Sulcatistroma, Togniniella (= Phaeocrella),
Neck long or short neck, periphysate. Peridium thick (10– and Tulipispora.
250 μm), outer layer coriaceous or membranous, composed
of dark brown, reddish brown to brown cells of textura Calosphaeria Tul. & C. Tul., Select. fung. carpol. (Paris) 2:
angularis or prismatica, or porrecta; inner layer membranous, 108 (1863)
composed of hyaline cells of textura prismatica. Facesoffungi number: FoF 01134, Figs 22, 23
Hamathecium composed of numerous, broad, septate, un- Saprobic on wood or decorticated wood or leaves in
branched, tapering, paraphyses. Ascogenous hyphae discrete, freshwater and terrestrial habitats. Sexual morph:
hyaline, smooth, branched, producing a sympodial sequence Ascomata perithecial, scattered or solitary, dark brown
of hyaline, ovoid to ellipsoidal cells, often with mucronate to black, immersed, globose to subglobose, coriaceous
apex, in dense clusters, each giving rise to an ascus. Asci 8- or membranous, tuberculate or smooth or with hyphal
spored, unitunicate, clavate, long or short pedicellate, mostly coating, ostiolate. Neck long or short neck, periphysate.
in fascicles, apical ring distinct or absent. Ascospores 2-seriate Peridium thick, outer layer coriaceous or membranous,
or overlapping, hyaline to light brown, allantoid to ellipsoid, composed of dark brown, reddish brown to brown cells
straight to slightly curved, aseptate to multi-septate, smooth- of textura angularis; inner layer membranous, composed
walled, sometimes with guttules. Asexual morph: of hyaline cells of textura prismatica. Hamathecium
Hyphomyce to us. Con id io phores micro nematous, composed of numerous, broad, septate, unbranched, ta-
mononematous or semimacronematous, brown or hyaline, pering, paraphyses. Ascogenous hyphae discrete, hya-
straight or flexuous, septate, branched or unbranched, with a line, smooth, branched, producing a sympodial sequence
terminal. Phialides terminal or lateral, aggregated, of hyaline, ovoid to ellipsoidal cells, often with mucro-
monophialidic, smooth, hyaline, elongate ampulliform to nate apex, in dense clusters, each giving rise to an
Fungal Diversity
 Fungal Diversity

ƒFig. 22 Calosphaeria princeps (Material examined: USA, Kansas, ampulliform to subcylindrical, pigmented in the apical.
Stockton, on inner bark of Prunus cerasus L. (Rosaceae), 5 October Conidiogenous cells enteroblastic or holoblastic,
1906, S-F263455). a Herbarium packet and material b Material c
Ascomata in mass d Ascoma e Ascoma in cross section f Peridium at
phialidic, hyaline, with brown apex, smooth-walled.
bottom of ascoma substructure g Peridium of neck with periphyses h Conidia aggregated, hyaline, aseptate, cylindrical to
Peridium in cross section i Paraphyses with asci j–k Asci l Ascospores allantoid, with or without guttules (From Réblová
in ascus m Ascospores. Notes: Figs i-m soaked in 3 % KOH; Figs k–l et al. 2004 and Révay et al. 2009).
stained in Melzer’s reagent. Scale bars: c = 1 mm, d = 200 μm,
e = 100 μm, f–i = 10 μm, j–m = 2 μm
Type species: Calosphaeria princeps Tul. & C. Tul.,
Select. fung. carpol. (Paris) 2: 109 (1863)
Notes: The genus Calosphaeria was introduced by Tulasne
ascus. Asci 8-spored, unitunicate, clavate, long or short and Tulasne (1863a). In the type species C. princeps, the sexual
pedicellate, mostly in fascicles, apical ring distinct or morph is characterized by immersed ascomata, tuberculate peri-
absent. Ascospores 2-seriate or overlapping, hyaline, thecia, clavate asci and hyaline ascospores. The asexual morph
allantoid to ellipsoid, straight to slightly curved, of Calosphaeria pulchella (Pers.) J. Schröt is
aseptate, smooth-walled, sometimes with guttules. Calosphaeriophora pulchella Réblová et al. The monotypic
Asexual morph: Hyphomycetous. Conidiophores genus, Calosphaeriophora, is characterized by micronematous
micronematous, hyaline, straight or flexuous, septate, conidiophores, and smooth aseptate conidia (Barr 1985;
branched, with a terminal. Phialides terminal or lateral, Réblová et al. 2004; Damm et al. 2008). In our opinion,
aggregated, monophialidic, smooth, hyaline, elongate Calosphaeriophora should be included in Calosphaeriaceae.

Fig. 23 Calosphaeria princeps a

Asci and ascospores. b-d
Calosphaeriophora pulchella
(redrawn from Réblová et al.
2004) b, c Conidiophores d
Conidia. Scale bars: a = 2 μm, b–
d = 5 μm
 Fungal Diversity

Other genera included scurfy, flat or slightly convex, composed of rather thin-walled,
reddish yellow cells of textura angularis, textura globosa and
Calosphaeriophora Réblová et al., Stud. Mycol. 50(2): 542 textura epidermoidea. Ascomata perithecial, deeply imbedded
(2004) in stromatic columns, globose, with a long, periphysate,
Type species: Calosphaeriophora pulchella Réblová et al., ostiolar neck. Peridium thick, composed of several layers of
Stud. Mycol. 50(2): 542 (2004) compressed, reddish brown cells of textura angularis.
Jattaea Berl., Icon. fung. (Abellini) 3(1–2): 6 (1900) Paraphyses hypha-like, numerous, tapering towards the apex,
Type species: Jattaea algeriensis Berl., Icon. fung. not embedded in a gelatinous matrix. Asci 8-spored,
(Abellini) 3(1–2): 6 (1900) unitunicate, broad cylindrical, short pedicellate, apically
Kacosphaeria Speg., Boln Acad. nac. Cienc. Córdoba rounded or truncate, with a J-, discoid, refractive, apical ring.
11(2): 214 (1888) Ascospores bi-seriate, hyaline, 1-celled, ellipsoidal to cylindri-
Type species: Kacosphaeria antarctica Speg., Boln Acad. cal, smooth-walled, lacking a mucilaginous sheath. Asexual
nac. Cienc. Córdoba 11(2): 214 (1888) morph: Undetermined.
Sulcatistroma A.W. Ramaley, Mycotaxon 93: 140 (2005) Type: Catabotrys Theiss. & Syd.
Type species: Sulcatistroma nolinae A.W. Ramaley, Notes: The monotypic family Catabotrydaceae was pro-
Mycotaxon 93: 140 (2005) posed by Petrak (1954) to accommodate the genus
Togniniella Réblová et al., Stud. Mycol. 50(2): 543 (2004) Catabotrys. Later this family was validated by Barr (1990).
Type species: Togniniella acerosa Réblová et al., Stud. According to the characteristics of hamathecium and asci,
Mycol. 50(2): 545 (2004) Catabotrydaceae has been placed in Sordariales (Barr
Tulipispora Révay & J. Gönczöl, in Révay et al., Nova 1990). Hyde et al. (2000) placed Catabotrydaceae in
Hedwigia 88(1–2): 42 (2009) Xylariales, while Kirk et al. (2001) suggested that it is mor-
Type species: Tulipispora ingoldii Révay & J. Gönczöl, phologically similar to some Boliniaceae and thus the order
Nova Hedwigia 88(1–2): 42 (2009) Boliniales was introduced to include both families. Huhndorf
et al. (2004a) showed that the relationship between
Catabotrydaceae and Boliniaceae is not supported by molec-
Key to genera of Calosphaeriaceae ular data, hence Catabotrydaceae was excluded from
Boliniales and placed in Sordariomycetidae family incertae
1. Hyphomycete . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 sedis. Although analysis of combined β-tubulin, LSU and
1. Ascomycete . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 RPB2 sequence data by Miller and Huhndorf (2005) indicated
2. Conidia multi-septate, branched . . . . . . . . . Tulipispora that C. decidua is a member of Diaporthales, however, recent
2. Conidia aseptate, unbranched . . . . .Calosphaeriophora molecular studies by Maharachchikumbura et al. (2015) con-
3. Ascomata immersed in ascostromata . . . Sulcatistroma firmed that Catabotrydaceae clusters with
3. Ascomata without ascostromata . . . . . . . . . . . . . . . . . 4 Amplistromataceae, hence it has been placed in
4. Ascospores light greyish brown . . . . . . . . . . . . Jattaea Amplistromatales.
4. Ascospores hyaline . . . . . . . . . . . . . . . . . . . . . . . . . . 5 Type species: Catabotrys deciduum (Berk. & Broome)
5. Peridium composed of brown to hyaline cells of textura Seaver & Waterston, Mycologia 38(2): 184 (1946)
prismatica . . . . . . . . . . . . . . . . . . . . . . . . .Togniniella
5. Peridium outer layer composed of brown cells of textura Cephalothecaceae Höhn., Annls mycol. 15(5): 362 (1917)
angularis; inner layer composed of hyaline cells of Facesoffungi number: FoF 01330
textura prismatica . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 Saprobic or epiphytic on rotting or dead plants or fungi,
6. Ascomata globose to subglobose, with hyphal coating, or causing systemic mycotic infection in humans with
asexual morphs Calosphaeriophora . . . . Calosphaeria burns. Sexual morph: Ascomata perithecial, solitary to
6. Ascomata pyriform, asexual morph gregarious, superficial, subglobose to globose, dark brown
Undetermined . . . . . . . . . . . . Kacosphaeria to black, glabrous, carbonaceous, covered by sulphureous
hyphae, ostiole central, without periphyses. Peridium
Catabotrydaceae Petr. ex M.E. Barr, Mycotaxon 39: 83 cephalothecoid, comprising 2–3 layers, outer layer com-
(1990) posed of dark brown cells of textura angularis to
Facesoffungi number: FoF 01372; Fig. 24 prismatica, inner layer composed of elongated, hyaline
Saprobic on dead leaves and stems of tropical monocoty- cells of textura prismatica. Ascogenous hyphae septate,
ledons. Sexual morph: Stromata irregularly scattered, occa- hyaline. Asci 8-spored, unitunicate, pyriform to subglobose
sionally coalescing, conspicuous, multi-loculate, superficial, to globose, evanescent, apedicellate, without an apical
with base slightly penetrating the epidermis at regular inter- ring. Ascospores irregularly arranged, brown, variously-
vals, discoid to pulvinate, reddish brown to black, surface shaped, unicellular, without germ pores, smooth-walled.
Fungal Diversity

Fig. 24 Catabotrys deciduum (Material examined: a-b, d-j. Sri Lanka, Stromata on host surface d–e Transverse section of stromata with
on unidentified dead palm, G.H.K. Thwaites 172 (ex herb. M.J. ascomata f Ostiole with periphyses g Stromatic tissue h Peridium i
Berkeley), November 1867, K (M) 35063, holotype; Hyde and Cannon Paraphyses j–k Immature and mature asci l ascospores. Scale bars:
(1999); c, i. Australia, Queensland, Cape Tribulation, J.E. Taylor, 15 d = 500 μm, e = 200 μm, f–h = 100 μm, i = 20 μm, j–l = 10 μm
April 1995, HKU(M) 4306) a Herbarium material with label b–c

Asexual morph: Hyphomycetous. Mycelium pale- Conidiophores long, cylindrical, stiffly upright, septate,
yellowish to brownish-yellow, branched septate. verticillate and whorled or not. Conidiogenous cells
 Fungal Diversity

phialidic, cylindrical, hyaline to brown, smooth-walled. family (Maharachchikumbura et al. 2015). The sexual morph
Conidia in chains, hyaline to brown, cylindrical, ovate or of these genera have carbonaceous, dark brown to black
obovate, with or without an apiculate or truncate base, 1- ascomata, with a central ostiole and a cephalothecoid peridi-
celled, smooth-walled. um (Suh and Blackwell 1999). Ascospores are small, brown,
Type: Cephalotheca Fuckel 1-celled, without germ pores, and smooth-walled (Chesters
Notes: The family Cephalothecaceae was introduced by 1935; Kirschstein 1936; Malloch and Cain 1970; Yaguchi
von Höhnel (1917) and is typified by Cephalotheca with et al. 2006). The genus Phialemonium was included in
C. sulfurea Fuckel as the type species (von Höhnel 1917). Cephalothecaceae as an asexual morph based on phylogenet-
This family is characterized by ascomata with a ic analysis (Perdomo et al. 2013b).
cephalothecoid peridium. The peridial cells form plate-like
complexes made up of radiating groups of cells, where each Cephalotheca Fuckel, Jb. nassau. Ver. Naturk. 25–26: 297
plate is separated by well-defined lines of dehiscence (1871)
(Malloch and Cain 1970). They can be small and simple, or Facesoffungi number: FoF 01331, Figs 25, 26
very large and complex (Malloch and Cain 1970; Suh and Saprobic in soil or on wood or mushrooms. Mycelium
Blackwell 1999). Four genera, Albertiniella, Cephalotheca, brown, branched. Sexual morph: Ascomata perithecial, soli-
Cryptendoxyla and Phialemonium are presently placed in the tary to gregarious, superficial, immature ascomata, disc-like,

Fig. 25 Cephalotheca sulfurea

(Material examined: GERMANY,
Oestrich, on rotting planks of
Oak, lying on damp, ammoniacal
ground, G00266459, holotype) a
Herbarium material b Ascomata
covered by sulphureous hyphae c
Close up of ascoma d Squash
mount of ascoma e Cross section
of ascoma f Peridium g–j Asci k
Ascogenous hyphae l–n
Ascospores. Scale bars: b–
c = 100 μm, d, k = 20 μm,
e = 50 μm, f = 10 μm, g-j, l–
n = 5 μm
Fungal Diversity

are saprobes inhabiting soil, wood, and mushrooms (Suh et al.

2006). Cephalotheca is a poorly known genus, which has both
sexual and asexual morphs. Fungal isolate SB5-1 was identi-
fied as a new strain of Cephalotheca sulfurea through molec-
ular and phylogenetic approaches. Culture filtrates (CFs) of
isolate SB5-1 were analyzed for the presence of gibberellins,
and it was found that all physiologically active gibberellins
were present. Gibberellins are well known for plant growth
promotion (Hamayun et al. 2012).

Other genera included

Albertiniella Kirschst., Annls mycol. 34(3): 183 (1936)

Type species: Albertiniella reticulata Kirschst., Annls
mycol. 34(3): 183 (1936)
Cryptendoxyla Malloch & Cain, Can. J. Bot. 48(10): 1816
(1970)
Type species: Cryptendoxyla hypophloia Malloch & Cain,
Can. J. Bot. 48(10): 1816 (1970)
Phialemonium W. Gams & McGinnis, Mycologia 75(6):
978 (1983)
Type species: Phialemonium obovatum W. Gams &
McGinnis, Mycologia 75(6): 978 (1983)

Key to genera of Cephalothecaceae

Fig. 26 Cephalotheca sulfurea (redrawn from Chesters 1935) a, b
Conidiophores with conidiogenous cells and conidia c, d
Conidiogenesis e Conidia. Scale bars: a–d = 10 μm, e = 5 μm 1. Asexual morph comprising of simple or branched, hyaline
conidiophores, phialidic pegs conidiogenous, hyaline or
pigmented, cylindrical to globose, 1-celled, arranged in
rather flat, with a central ostiole, with a thin and dark brown slimy heads or in long chains conidia . . . . Phialemonium
wall composed of the cells of textura prismatica to textura 1. Mostly known in sexual morph . . . . . . . . . . . . . . . . . 2
angularis; mature ascomata subglobose to globose, dark brown 2. Asci pyriform to subglobose . . . . . . . . Cryptendoxyla
to black, glabrous, carbonaceous, covered by sulphureous hy- 2. Asci globose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
phae. Peridium of mature ascomata cephalothecoid, comprising 3. Ascospore globose . . . . . . . . . . . . . . . . . . Albertiniella
two layers, outer layer composed of dark brown, rows of cells 3. Ascospore ovate to fusiform. . . . . . . . . . Cephalotheca
textura angularis, inner layer composed of elongated, hyaline
cells of textura prismatica. Ascogenous hyphae of septate, hy- Ceratocystidaceae Locq. ex Réblová et al., Stud. Mycol.
aline, woven ascogenous hyphae and asci irregularly disposed 68(1): 188 (2011)
on ascogenous hyphae. Asci 8-spored, unitunicate, subglobose Facesoffungi number: FoF 01248
to globose, lacking a pedicel, lacking an apical ring. Ascospores Saprobic or pathogenic on plant material. Sexual morph:
irregularly arranged, ovate to fusiform, reniform, brown to dark Ascomata globose to pyriform or ovoid, light brown or dark
brown, 1-celled, without germ pores, smooth-walled. Asexual brown to black, with spines or septa, unornamented or with
morph: Hyphomycetous. Mycelium pale-yellowish to brown, undifferentiated ornamental hyphae, ostiolate. Ostioles com-
branched septate. Conidiophores long, cylindrical, septate, with prising long, tapering, dark brown or dark grey to black necks.
apex expanded and branched. Conidiogenous cells phialidic, Peridium thin-walled, composed of polyhedral pseudoparen-
hyaline to brown, smooth-walled. Conidia in chains, hyaline chymatous cells of textura angularis, pale yellow to reddish
to brown, broad fusiform to obovate, 1-celled, smooth-walled. brown, appearing dark brown or black. Hamathecium com-
Type species: Cephalotheca sulfurea Fuckel, Jb. nassau. prising light brown to hyaline, aseptate, convergent or slightly
Ver. Naturk. 25–26: 297 (1871) Bot. 15(5): 742 (2002) divergent, paraphyses, tapered at the apex. Asci dehiscent.
Notes: Cephalotheca was established by Fuckel (1872) and Ascospores 0–1-septate, hyaline, varied in shape, hat-shaped,
includes about 20 epithets (Index Fungorum 2016). Species ellipsoidal, or elongate to slightly curved, with round ends, or
 Fungal Diversity

oblong cylindrical, or narrow fusiform to spindle-shaped, hy- authors placed Ceratocystis in Chadefaudiellaceae, while oth-
aline, often with eccentric wall thickening or surrounded by a er authors placed it in its own family, Ceratocystidaceae (as
sheath, accumulating as masses at apices of ascomatal necks. “Ceratocystaceae”), which was formerly proposed by
Asexual morph: Conidiophores phialidic, arising laterally Locquin (1972) but was not validly published. The family
from vegetative hyphae, lageniform, tubular, rectangular, cy- Ceratocystidaceae was subsequently validated by Réblová
lindrical to flask-shaped, sometimes with slightly flared et al. (2011) and placed in Microascales in the subclass
collarette, single to aggregated in sporodochia or synnemata, Hypocreomycetidae (Réblová et al. 2011; De Beer et al.
septate, tapering to the apex, hyaline, unbranched or sparingly 2013). Ceratocystidaceae forms a monophyletic group, in-
branched, conidiophores consisting of two types; primary co- cluding Ambrosiella, Ceratocystis and Thielaviopsis, which
nidiophores (Ceratocystis) and secondary conidiophores is distinct from Gondwanamycetaceae based on strong boot-
(Huntiella). Conidiogenous cells phialidic, oblong or cylindri- strap support (Réblová et al. 2011). Davidsoniella and
cal, hyaline, subhyaline or pale brown, tapering towards the Huntiella were introduced as new genera in
apex. Conidia unicellular, varied in shape, cylindrical to ob- Ceratocystidaceae by De Beer et al. (2014). Seven genera
long, barrel-shaped to subglobose, rectangular, single or were emended based on morphological characters, eco-
formed in chains or as terminal aleurioconidia, with rounded logical differences, as well as DNA-sequence data for
or truncate ends, hyaline to light brown or pale brown, or three gene regions (60S, LSU, MCM7), these include
hyaline becoming grey; some of genera with two types of Ambrosiella, Ceratocystis sensu stricto, Chalaropsis,
conidia; primary and secondary conidia for genera such as Davidsoniella, Endoconidiophora, Huntiella and
Ceratocystis, Huntiella, and Thielaviopsis. Aleurioconidia Thielaviopsis (De Beer et al. 2014). Cornuvesica was
globose to subglobose, ovoid to pyriform, singly or in chains, referred to Microascales, genera incertae sedis
pale brown to brown. (Maharachchikumbura et al. 2015).
Type: Ceratocystis Ellis & Halst.
Notes: The family level classification of Ceratocystis has Ceratocystis Ellis & Halst., New Jersey Agric. Coll. Exp.
been discussed since the genus was removed from the Sta. Bull. 76: 14 (1890)
Ophiostomatales (Barr 1990; Samuels 1993). Recently, some Facesoffungi number: FoF 01249; Figs 27, 28

Fig. 27 Ceratocystis fimbriata

(Material examined: USA, New
Jersey, Swedesboro, on leaves of
Ipomoea batatas, 12 April 1891,
B. D. Halsted, BPI 595863–
595866 neotype) a Herbarium
material b Ascomata on Ipomoea
batatas L. (Convolvulaceae) c
Enlarged view of ascomatal base
and ascomatal neck d e Neck
region Scale bars: b–d = 100 μm,
d = 50 μm
Fungal Diversity

Fig. 28 Ceratocystis fimbriata. a

Habitat on wood b Ascospores
being released from ostiolar
hyphae c Ascoma with spines or
septa, unornamented or with
undifferentiated ornamental
hyphae d Conidiophore e
Germinated spores f
Aleurioconidia (redrawn from
Halsted and Fairchild 1891)

Saprobic or pathogenic on stems or roots of vascular Secondary conidia barrel-shaped to subglobose, hyaline
plants, some species associated with bark beetles. Sexual to light brown. Aleurioconidia globose ovoid to pyri-
morph: Ascomata single to aggregated, superficial or im- form, single or in chains, pale-brown to brown (De
mersed in substrate, globose to subglobose, dark brown to Beer et al. 2014).
black, with ornamented or unornamented hyphae, with Type species: Ceratocystis fimbriata Ellis & Halst., Bull.
or without spines, spines if present dark brown to black. New York Agricultural Experimental Station 76: 14 (1890)
Ostiole with long necks, cylindrical, tapering to the Notes: The genus Ceratocystis with C. fimbriata as the type
apex, straight, dark-brown to black, hyaline at apex, was introduced by Ellis and Halsted (1890) for the pathogen
ostiolar hyphae divergent, aseptate, tapering to apex, that causes black rot of sweet potatoes in the USA.
hyaline to light brown. Peridium thin–walled, compris- Ceratocystis species are associated with bark beetles, which
ing pseudoparenchymatous cells of textura angularis in are symbiotically associated with insects as well as important
surface view. Hamathecium containing centrum causal agents of sap-stain in timber and many important path-
interascal tissue absent. Asci dehiscent. Ascospores hya- ogens of plants, including pineapples and palms (Hedgcock
line, 1-celled, variable in shape, fusoid, oblong, ob- 1906; Mitchell 1937; Seifert 1993; Alvarez et al. 2012;
ovoid, cylindrical, or hat–shaped, curved, lunate, with Mbenoun et al. 2014).
or without a gelatinous sheath, accumulating in cream- Ceratocystis was removed from the order Ophiostomatales
coloured masses at apices of the ascomatal necks. (Barr 1990; Samuels 1993) and the taxonomic confusion be-
Asexual morph: Conidiophores simple tubular, flask- tween Ceratocystis and Ophiostoma was subsequently re-
shaped, tapering at the apex, with phialidic solved using DNA sequence data, showing that these genera
conidiogenesis. Primary conidiophores phialidic, flask- are unrelated (Hausner et al. 1993a, b; Spatafora and
shaped. Secondary conidiophores flaring or wide- Blackwell 1994). Ophiostoma belongs to the order
mouthed. Primary conidia cylindrical, hyaline. Ophiostomatales (Sordariomycetidae) and Ceratocystis was
 Fungal Diversity

accommodated in the Ceratocystidaceae (Microascales, Type species: Ambrosiella xylebori Brader ex von Arx &
Hypocreomycetidae) (Réblová et al. 2011; De Beer et al. Hennebert, Mycopath. Mycol. appl. 25(7): 314 (1965)
2013). Recently, these differences have been substantially am- Chalaropsis Peyronel, Staz. Sper. Argar. Ital. 49: 595
plified by the discovery of many new and often cryptic spe- (1916)
cies, revealed through DNA-sequence comparisons Type species: Chalaropsis thielavioides Peyronel, Staz.
(Wingfield et al. 1996; Harrington and Wingfield et al. Sper. Argar. Ital. 49: 585 (1916)
1998; De Beer et al. 2014). Davidsoniella Z.W. de Beer et al., in de Beer et al., Stud.
Huntiella Z.W. de Beer et al., in de Beer et al., Stud. Mycol. 79: 210 (2014)
Mycol. 79: 211 (2014) Fig. 29 Type species: Davidsoniella virescens (R.W. Davidson)
Z.W. de Beer et al., in de Beer et al., Stud. Mycol. 79: 210
(2014)
Other genera included Endoconidiophora Münch, Naturwiss. Z. Forst–Landw. 5:
564 (1907)
Ambrosiella Brader ex von Arx & Hennebert, Mycopath. Type species: Endoconidiophora coerulescens Münch,
Mycol. appl. 25: 314 (1965) Naturwiss. Z. Forst–Landw. 5: 564 (1907)

Fig. 29 Huntiella chinaeucensis (Material examined: THAILAND, conical spines f Ascoma neck g Hat-shaped ascospores being released
Chiang Rai Province, Mae Suai District, Mae Lao garden, on stumps of from ostiolar hyphae h Globose ascoma with ascoma neck i Hat-shaped
Tectona grandis L. (Lamiaceae), 24 December 2012, M. Doilom, MFLU ascospores j Cylindrical conidia k Conidiophores with conidiogenous
15-3204). a Ascomata on Tectona grandis wood b, c Globose ascomata cells Scale bar b–c = 500 μm, d, h = 100 μm. e–g = 20 μm, i, j = 5 μm,
with long necks with subhyaline to cream masses at tips of ascomata k = 10 μm
necks on teak wood d Immature globose ascoma e Ascomata base with
Fungal Diversity

Type species: Huntiella moniliformis (Hedgc.) Z.W. de 4. Ascomatal globose to ovoid, surrounded by distinct bas-
Beer et al., Stud. Mycol. 79: 212 (2014) al spines, ascospores elongate to slightly curved with
Saprobic on wood. Sexual morph: Ascomata superficial, round ends or some species oblong cylindrical, and
globose to subglobose, ornamented with dark brown to black, surrounded by distinct translucent
conical spines; dark brown to black. Ostiolar neck long, black, sheath. . . . . . . . . . . . . . . . . . . . . . . Endoconidiophora
tapering to apex, with a disk-like base. Ostiole comprising
hyaline hyphae. Asci not observed. Ascospores hyaline, hat-
shaped in side view, aseptate, being released from ostiolar Key to the asexual genera of Ceratocystidaceae
hyphae, by accumulating in cream-coloured masses at apices
of the ostiolar neck. Asexual morph: Conidiophores long, Notes: The morphological characters of the asexual morph in
macronematous, straight or flexuous, tapering to apex, arising several genera are indistinguishable. Generic delineations are
from hyphae, septate, slightly constricted at septa, hyaline, better achieved by using DNA sequence data.
smooth. Conidiogenous cells phialidic, cylindrical, tapering 1. Sexual morph not known or not observed . . . . . . . . . 2
toward apex, hyaline. Conidia cylindrical to barrel-shaped, 1. Sexual morph known . . . . . . . . . . . . . . . . . . . . . . . . . 3
truncate at the ends, hyaline, single to catenate, aseptate. 2. Conidiophores single to aggregated in sporodochia; co-
Notes: The genus Huntiella was introduced by De Beer nidia globose, associated with ambrosia
et al. (2014), and placed in the family Ceratocystidaceae. beetles . . . . . . . . . . . . . . . . . . . . . . . . . . . Ambrosiella
They introduced 17 Huntiella species with another new com- 2. Conidiophores sympodially or irregularly branched; co-
bination of 17 species in Ceratocystis based on morphological nidia cylindrical . . . . . . . . . . . . . . . . . . . . Chalaropsis
and phylogenetic analyses of 60S ribosomal RNA, LSU and 3. Aleurioconidia not present . . . . . . . . . . . . . . . . . . . . . 4
MCM7 dataset (De Beer et al. 2014). The species included in 3. Aleurioconidia present . . . . . . . . . . . . . . . . . . . . . . . . 6
this study show 100 % similarity to Huntiella moniliformis 4. Conidiogenous cells cylindrical . . . . . . . . . . . . . . . . . 5
(Hedgc.) De Beer et al. (2014), based on nucleotide blast 4. Conidiogenous cells flask-shaped. . . . . . . . . Huntiella
results of ITS gene, as well as the distinctive features of mor- 5. Conidia cylindrical with flattened ends, barrel-
phology. Based on morphological features as well as DNA shaped . . . . . . . . . . . . . . . . . . . . . . . . . Davidsoniella
s e q ue n ce da t a , w e r ec o gn i z e d ou r co l l ec t i on a s 5. Conidia rectangular with two attachment
H. moniliformis. Thielaviopsis has been reported as asexual points. . . . . . . . . . . . . . . . . . . . . . Endoconidiophora
morph of Huntiella moniliformis (Chen et al. 2013). 6. Secondary conidia cylindrical to oblong; conidiophores
Thielaviopsis Went, Meded. Proefstn Suik Riet W. Java 5: lageniform . . . . . . . . . . . . . . . . . . . . . . . Thielaviopsis
4 (1893) 6. Secondary conidia barrel to subglobose; conidiophores
Type species: Thielaviopsis ethacetica Went [as flared or wide-mouthed . . . . . . . . . . . . . . Ceratocystis
‘ethaceticus’], Annuario Soc. Alpinisti Trident.: 4 (1893)
Ceratostomataceae G. Winter [as ‘Ceratostomeae’],
Rabenh. Krypt.–Fl., Edn 2 (Leipzig) 1.2: 247 (1885)
Facesoffungi number: FoF 01803
Key to the sexual genera of Ceratocystidaceae Saprobic or weakly parasitic, often found growing on other
fungi, commonly isolated from soil. Sexual morph:
1. Ascospores one-celled, hat-shaped . . . . . . . . . . . . . . . . . 2 Ascomata perithecial or cleistothecial, usually translucent, yel-
1. Ascospores one-celled, not hat-shaped (elongate or ob- low to pale brown, ostiolate or not, often with long-necks, with
long cylindrical, or narrow fusiform to spindle-shaped)3 smooth ostiolar setae. Interascal tissue absent. Peridium
2. Ascomatal bases globose to pyriform, with ornamented membranaceous, comprising pale yellow to pale yellowish-
hyphae, conical spines, disk-like bases of the ascomatal brown, cell of a textura angularis or textura globulosa. Asci
necks . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Huntiella 8-spored, unitunicate, thin-walled, clavate, without an apical
2. Ascomatal bases globose, with unornamented ring, deliquescing. Ascospores biseriate, brown to dark brown,
hyphae . . . . . . . . . . . . . . . . . . . . . . . . . . Ceratocystis ellipsoidal to citriform, occasionally discoid or fusiform, usual-
3. Ascomatal necks long, brown to dark brown or black, ly 2-germ pored at each end, aseptate, smooth to strongly
with non-digitate or non-stellate appendages on the ornamented, sheath absent. Asexual morph:
ascomatal. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 Hyphomycetous. Conidiophores simple, semi-macronematous,
3. Ascomatal necks long, dark grey, with digitate or stellate mononematous, some genera (Acrospeira) branched towards
appendages on the asco . . . . . . . . . . . . . Thielaviopsis the apex, some genera (Gonatobotrys, Erythrocarpon) conidi-
4. Ascomatal globose, surrounded by ornamental hyphae, ophore with several roughened swellings along its length,
ascospores elongate, narrow fusiform to spindle shaped, reddish-brown to dark. Conidiogenous cells integrated, termi-
slightly curved, with hyaline sheath . . . . Davidsoniella nal, monoblastic, some genera (Pteridiosperma) with phialides,
 Fungal Diversity

singly on aerial hyphae or rarely on conidiophores, lageniform,

hyaline. Conidia globose, spindle-shaped, ovate to pyriform,
some genera (Pteridiosperma) are aggregated in small globose
heads at the apices of phialides, one-celled, some genera
(Erythrocarpon, Acrospeira) containing with 2–3-transverse
septa, hyaline, brown and verrucose.
Type: Melanospora Corda
Notes: The family Ceratostomataceae was established by
Winter (1885). Barr (1990), Hawksworth et al. (1995), and
Samuels and Blackwell (2001) placed Melanospora and its
allies in Ceratostomataceae (synonymy: Melanosporaceae),
based on the ascospore colour and ornamentation and ribo-
somal DNA sequences. The family was placed in Sordariales
based on the characters of the ascospores, with dark with germ
pores, which are the most conspicuous similarities (Huang
1976; Uecker 1976). Zhang and Blackwell (2002) placed
Fig. 30 Melanospora zamiae (holotype). a Ascomata b Ascospores c
Ceratostomataceae in the order Melanosporales based on
Ascomata and ascospores mass (redrawn from Corda 1837)
phylogenetic analyses, while emphasizing the morphological
characters, Sordaria-type centrum composed of paraphyses in
addition to pseudoparenchyma, which differs from the cen- (Gonatobotrys) produced in grape-like clusters around each
trum of Melanospora. Most of the species of swelling, some genera (Pteridiosperma) produced from single
Ceratostomataceae are saprobic on plant material. Certain phialides on aerial hyphae or rarely on conidiophores.
species have a widespread host range, and are closely Type species: Melanospora zamiae Corda, Icon. fung.
associated with other fungi, on soil, or rotting vegetation and (Prague) 1: 24 (1837)
some species are considered as potential biocontrol agents as Notes: The genus Melanospora was introduced by Corda
they can grow on other plant pathogens. Recently, (1837) with the type species, M. zamiae, and subsequently
Maharachchikumbura et al. (2015) provided an updated out- referred to the family Ceratostomataceae (Winter 1885) and
line of the family Ceratostomataceae and included ten genera. Sordariales (Hawksworth and Eriksson 1986). Melanospora
was traditionally placed within the Ceratostomataceae,
Melanospora Corda, Icon. fung. (Prague) 1: 24 (1837) Sordariales based on morphological similarities with the
Facesoffungi number: FoF 01804; Fig. 30 Chaetomiaceae (Hawksworth et al. 1995). Later molecular
Saprobic or weakly parasitic on plant materials, commonly phylogenetic studies indicated that Melanospora belonged in
isolated from soil and closely associated with other fungi. Hypocreales (Rehner and Samuels 1995; Jones and Blackwell
Sexual morph: Ascomata superficial to immersed, globose 1998). Zhang and Blackwell (2002) and Chaudhary et al.
to ovoid, gregarious, inconspicuously ostiolate, setose, pale yel- (2006) considered the Melanospora clade to be a sister group
low, translucent, appearing dark brown to black, due to the of Hypocreales and found that sequences derived from other
massed ascospores, with short neck or sometimes absent, with genera of Ceratostomataceae (synonymy: Melanosporaceae)
a ring of rigid, hyaline, smooth and thick-walled setae around clustered within the main Hypocreales clade. Zhang et al.
the ostiole, setae right to slightly curved or sinuous, pale yellow, (2006) concluded that the family belongs to
aseptate, thick-walled. Peridium membranaceous, thick-walled, Hypocreomycetidae but that it was a sister group of the
pale yellow to pale yellowish-brown, cell of a textura globulosa Coronophorales and introduced a new order Melanosporales.
or textura angularis. Paraphyses absent. Asci 8-spored, The order was published formally by Hibbett et al. (2007). A
unitunicate, broadly clavate to obovate, fasciculate, apex round- similar placement was proposed by Schoch et al. (2007) in a
ed, without apical ring, short stipitate, broadly rounded apex, study of marine fungal lineages within the Hypocreomycetidae.
evanescent walled. Ascospores irregularly biseriate, at first hy- However, the phylogenetic inconsistencies of the order
aline and guttulate, brown to dark brown, large ellipsoidal, one- Melanosporales have not been addressed.
celled, smooth and thick-walled, with two terminal germ pores, Most species in Melanospora are parasitic and associated on
surrounded by a dark ring-like structure. Asexual morph: wide host range such as basidiomycetes, sexual and asexual
Hyphomycetous. Conidiophore simple, erect, brown or dark ascomycetes as well as with other fungi and some species ob-
brown. Conidiogenous cell phialides, single on aerial hyphae tain nutrients by fusing with the host protoplasts, an interaction
or rarely on conidiophores, lageniform, hyaline, some genera called fusion biotrophism (Jeffries and Young 1994). Harveson
(Acrospeira) integrated, terminal, monoblastic. Conidia glo- (1999) reported Melanospora species as potential biocontrol
b o s e , s pi nd l e- s ha p e d, 1– ( −4 ) c e l l , s o m e g e n er a agents as they can grow on other plant pathogenic fungi.
Fungal Diversity

Asexual morphs of the genus belong in a wide range of Type species: Acrospeira mirabilis Berk. & Broome,
genera, including Gonatobotrys, Harzia, Papulaspora and in Berkeley, Intr crypt. Bot. (London): 305, fig. 69a
Proteophiala (Hyde et al. 2011). (1857)
Arxiomyces P.F. Cannon & D. Hawksw., Trans Br Mycol
Melanospora collipora Stchigel & Guarro, in Stchigel Soc 81(3): 644 (1983)
et al., Mycol. Res. 101(4): 446 (1997); Fig. 31. Type species: Arxiomyces vitis (Fuckel) P.F. Cannon & D.
Notes: Melanospora collipora was described by Stchigel et al. Hawksw, Trans. Br. Mycol. Soc. 81(3): 644 (1983)
(1996), isolated from a soil sample from India. This species was Erythrocarpon Zukal, Verh zool-bot Ges Wien 35: 337
introduced to Melanospora based on its large ellipsoidal, 1-celled (1886)
ascospores, with two terminal germ pores, surrounded by ring- Type species: Erythrocarpon microstomum Zukal, Verh
like structure (Fog. 31). Melanospora collipora differs from other zool-bot Ges Wien 35: 337 (1886)
species in conidial size and character of germ pores, and hyaline Gonatobotrys Corda, Pracht-Fl. Eur. Schimmelbild.: 9
rim around the germ pores (Stchigel et al. 1996). (1839)
Type species: Gonatobotrys simplex Corda, Pracht - Flora.
Europaeischer Schimmel-Bildungen: 9 (1839)
Pteridiosperma J.C. Krug & Jeng, Mycotaxon 10(1): 44
Other genera included (1979)
Type species: Pteridiosperma foveolatum (Udagawa & Y.
Acrospeira Berk. & Broome, in Berk., Intr. crypt bot Horie) J.C. Krug & Jeng, Mycotaxon 10(1): 45 (1979)
(London): 305 (1857) Pustulipora P.F. Cannon, Mycotaxon 15: 526 (1982)

Fig. 31 Melanospora collipora

(holotype). a Ascoma b Ascus c,
d Ascospores e Detail of germ
pores f Chlamydospores. Scale
bars: a = 25 μm, b, c, f = 10 μm,
d, e = 1 μm. (redrawn from
Stchigel et al. 1996)
 Fungal Diversity

Type species: Pustulipora corticola P.F. Cannon, Key to the sexual genera of Ceratostomataceae
Mycotaxon 15: 526 (1982); Fig. 32
Notes: Pustulipora was established by Cannon and 1. Ascomata with necks . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
Hawksworth (1982), based on P. corticola, as the type species 1. Ascomata lacking necks. . . . . . . . . . . . . . . . . . . . . . . 3
isolated from bark (unidentified) in UK. Pustulipora was 2. Peridium coriaceous to carbonaceous, dark brown to
placed in Ceratostomataceae based on its ascospores, which black; ascospores with a large sunken germ pore and
are muriform-ellipsoidal with two simple terminal germ pores small basal appendage . . . . . . . . . . . . . . . .Arxiomyces
(Fig. 32) and other characters which confirmed the close rela- 2. Peridium membranous brownish or pale yellow to red-
tionship of this genus to Ceratostomataceae (Cannon and dish brown, and translucent; ascospores with two apical
Hawksworth 1982). germ pores. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
Rhytidospora Jeng & Cain, Mycotaxon 5(1): 278 (1977) 3. Ascomata with ostiole . . . . . . . . . . . . . . . . . . . . . . . . 5
Type species: Rhytidospora tetraspora Jeng & Cain, 3. Ascomata without ostiole . . . . . . . . . . . . . . . . . . . . . . 6
Mycotaxon 5(1): 279 (1977) 4. Ascospores with a thick hyaline ridge running full
Setiferotheca Matsush., Matsush. Mycol Mem 8: 34 (1995) length of the ascospore between the germ
Type species: Setiferotheca nipponica Matsush., Matsush. pores . . . . . . . . . . . . . . . . . . . . Vittatispora
Mycol. Mem. 8: 35 (1995) 4. Ascospores without a thick hyaline ridge running full
Vittatispora P. Chaudhary et al., Mycologia 98(3): 461 (2006) length of the ascospore between the germ
Type species: Vittatispora coorgii P. Chaudhary et al., pores . . . . . . . . . . . . . . . . . . . . . . . . . . . Melanospora
Mycologia 98(3): 461 (2006) 5. Ascospores with two apical germ pores . . . . . . . . . . . 7

Fig. 32 Pustulipora corticola

(UK, Nottinghamshire,
Nottingham, Clumber park, on
unidentified bark, 7 September
1956, W.D. Graddon., IMI
284968, holotype). a, b
Herbarium specimen c Ascomata
superficial on host tissue d
Section through ascoma e
Peridium f Ascospores attached to
the inner wall of ascoma g–j
Ascospores. Scale bars:
d = 100 μm, e–j = 20 μm
Fungal Diversity

5. Ascospores with a single germ pore at one and asci which are catenate and irregularly disposed in the
end………………………..……..Setiferotheca centrum at maturity (Udagawa and Furuya 1973). However,
6. Ascospores black, with wing-like ridges at convex sur- the classification of Faurelina has been problematic. Despite
faces…………..……Pteridiosperma the similarities with Chadefaudiella noted by Locquin-Linard
6. Ascospores brown to dark brown, without wing-like (1975), Parguey-Leduc and Locquin-Linard (1976) concluded
ridges at convex surfaces . . . . . . . . . . . . Rhytidospora that Faurelina should be placed in the Loculoascomycetes
7. Asci cylindrical, ephemeral, 8-spored. . Erythrocarpon (now in Dothideomycetes; Hyde et al. 2013). Faurelina was
7. Asci clavate to obovoid, 4-spored . . . . . . .Pustulipora later transferred by von Arx (1978) to the family
Microascaceae. Recently both genera were placed in
Chadefaudiellaceae, Microascales (Cannon and Kirk 2007;
Key to the asexual genera of Ceratostomataceae Maharachchikumbura et al. 2015). However, phylogenetic
re-evaluation of Microascales by Réblová et al. (2011) con-
1. Conidia 1-celled . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 cluded that Chadefaudiella is morphologically slightly differ-
1. Conidia 3- or 4-cell . . . . . . . . . . . . . . . . . . . . . . . . . . 3 ent from Faurelina and further molecular analysis may lead to
2. Conidia produced in grape-like clusters around each a re-establishment of the family Chadefaudiellaceae in the
swelling on hyphae . . . . . . . . . . . . . . . . Gonatobotrys order Microascales, but with the exclusion of Faurelina.
2. Conidia produced from single phialides on The family is poorly understood and needs recollecting.
hyphae . . . . . . . . . . . . . . . . . . . . . . . . Pteridiosperma
3. Conidia globose, with 2 transverse septa . . Acrospeira Chadefaudiella Faurel & Schotter, Revue Mycol., Paris
3. Conidia spindle-shaped, with 3 transverse 30: 339 (1959)
septa. . . . . . . . . . . . . . . . . . . . . . . . . . Erythrocarpon Facesoffungi number: FoF 02103; Fig. 33
Saprobic on mammalian dung. Sexual morph: Ascomata
Chadefaudiellaceae Faurel & Schotter ex Benny & relatively large (500–1500 μm high, 150–250 μm wide), light
Kimbr., Mycotaxon 12(1): 46 (1980) yellow, elongate or hemisphaerical, immersed at the base, pale
Facesoffungi number: FoF 01666 yellow to brown, basal perithecial envelope black, globular,
Saprobic on mammalian dung. Sexual morph: Ascomata carbonaceous, foot cylindrical, light yellow to brown, translu-
perithecial, elongate or hemisphaerical, immersed at the base, cent, striated, upper part with brown, anastomosing setae or
pale yellow to brown, basal perithecial envelope black, glob- setae lacking, carbonaceous, lacking ostioles. Peridium com-
ular, carbonaceous, foot cylindrical, light yellow to brown, posed of pseudoparenchymatous cells forming a textura
translucent, striated, upper part with brown, anastomosing se- angularis embedded in the substrate and an aerial
tae or setae lacking, lacking ostioles. Peridium composed of “capillitium”. Asci 8-spored, globose, evanescent.
pseudoparenchymatous cells forming a textura angularis em- Ascospores 1-celled, fusiform, thick, striate, not turn into
bedded in the substrate and an aerial “capillitium”. Asci over- reddish-brown upon application of Melzer’s reagent (non-
lapping, 8-spored, globose or clavate, catenulate, evanescent. dextrinoid). Asexual morph: Undetermined.
Ascospores 1-celled, hyaline to pale brown, ellipsoidal or fu- Type species: Chadefaudiella quezelii Faurel & Schotter,
siform, without germ pores, striate or striations lacking, turn- Comptes rendus hebdomadaires des séances de l’Académie
ing reddish brown in Melzer’s reagent (dextrinoid) or non- des sciences, Paris 249(1): 152 (1959)
dextrinoid formed in a mazaedial mass. Asexual morph: Notes: The genus Chadefaudiella was described for a
Hyphomycetous, forming arthrospores. single species, Chadefaudiella quezelii. A later study by
Type: Chadefaudiella Faurel & Schotter Faurel and Locquin (1972) introduced another species,
Notes: Chadefaudiellaceae was introduced by Benny and Chadefaudiella thomasii Faurel & Locq. isolated from
Kimbrough (1980) to accommodate the genus dung. Benny and Kimbrough (1980) proposed a new
Chadefaudiella. Cannon and Kirk (2007) added a second ge- family Chadefaudiellaceae in the order Microascales to
nus to the family, Faurelina (Locquin-Linard 1975). Parguey- include the genus Chadefaudiella and this has been
Leduc (1977) placed Chadefaudiella in the order followed by Cannon and Kirk (2007), Réblová et al.
Microascales because of its perithecial ascomata, catenate as- (2011) and Maharachchikumbura et al. (2015). The ge-
ci, and characteristic centrum structures, i.e. “asci arising from nus is poorly understood and needs recollecting.
a fertile layer lining the bottom of the cavity, ascogenous hy-
phae ramifying upwards, asci extricated without croziers and
liberated by basal dissolution to float free in the centrum” Other genus included
(Benny and Kimbrough 1980). Faurelina was included in this
f a m i l y be c au s e it h as ch a r a ct er s r em i ni s c en t o f Faurelina Locq.-Lin., Revue Mycologique, Paris 39(2): 127
Chadefaudiella, such as a cephalothecoid ascomatal wall (1975)
 Fungal Diversity

and humans. Sexual morph: Ascomata perithecial or

cleistothecial, colourless to light brown, egg yellow, grey-
olivaceous to (greenish) grey, to grey, golden brown, brown
to black, solitary to scattered to gregarious, superficial or im-
mersed to semi-immersed, subglobose to obpyriform, globose;
ellipsoidal; ovoid to pyriform, cylindrical; covered with
hair/setae or glabrous, hairs maybe simple or branched and of
one to two types, ostiole or non-ostiole. Peridium with thick
wall, comprising several layers of hyaline or brown cells of
textura epidermoidea to textura intricata. Hamathecium septate
with hyaline paraphyses. Asci 4- or 8-spored, unitunicate, cla-
vate, or clavate to cylindrical, obovate or ellipsoidal, pedicel-
late, without apical structures or with an indistinct thickened
ring, evanescent, with wall dissolving at maturity. Paraphyses
absent or greatly reduced. Ascospores irregularly arranged, at
first colourless and dextrinoid, (translucent, greenish, dark),
brown to black, and opaque when mature, ellipsoidal, globose,
subglobose, oval, fusiform or triangular, with single or some-
times two germ pores, aseptate, with thick, smooth walls.
Asexual morph: Hyphomycetous. Colonies brown or orange
or white to (black, green) grey brown or grey, woolly, border
even, reverse uncolored or buff-coloured to brown, aerial my-
celium cottony or felty, with funiculose setae and hyphae.
Mycelium abundant, intramatrical and aerial, composed of sep-
tate, hyaline or dark brown hyphae. Conidiophores absent or
simple, vegetative hyphae, hyaline, branched, septate, smooth-
walled or slightly rough. Conidiogenous cells phialidic, verti-
cillate or solitary, hyaline, consisting of a lageniform or ellip-
soid swollen basal portion, or reduced or cylindrical. Conidia
dimorphic, first kind holoblastic, hyaline to brown, smooth,
globose to obclavate, 1-celled, formed on hyphae or cylindrical
conidiogenous cells, single or in racemose clusters; second kind
produced form phialides, hyaline to brown, lutescens, or dark
Fig. 33 Chadefaudiella quezelii a Ascoma b Longitudinal section of brown, subglobose to globose or ellipsoidal, occasionally cy-
ascoma c Peridium. Scale bars: a–b = 200 μm, c = 100 μm (redrawn
lindrical to pyriform or fusiform, formed single or in chains, 1-
from Faurel and Schotter 1959)
celled, with thick, smooth-walled.
Type species: Faurelina fimigena Locq.-Lin. [as Type: Chaetomium Kunze
‘fimigenes’], Revue Mycol., Paris 39(2): 127 (1975) Notes: The family Chaetomiaceae was introduced by Winter
(as Chaetomiea) in 1885, and is typified by Chaetomium
(Grunow et al. 1887). Earlier, the family was placed in the order
Chaetomiales by Ames (1961), Alexopoulos (1962) and
Key to genera of Chadefaudiellaceae Mukerji (1968). Barr (1976b), Müller and von Arx (1973) trans-
ferred the family Chaetomiaceae to the order Sphaeriales, while
1. Ascomata with apical anastomosing setae; ascospores non Hawksworth and Wells (1973) and Mehrotra and Aneja (1990)
dextrinoid. . . . . . . . . . . . . . . . . . . . . . . . . . . Chadefaudiella placed it in Sordariales. Later molecular data showed
1. Ascomata without apical anastomosing setae; ascospores Chaetomiaceae belonged to the order Sordariales (Huhndorf
dextrinoid. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Faurelina et al. 2004a; Kirk et al. 2008; Lumbsch and Huhndorf 2010;
Liu 2011; Maharachchikumbura et al. 2015). Twenty genera are
Chaetomiaceae G. Winter [as ‘Chaetomieae’], Rabenh. presently placed in the family (Maharachchikumbura et al.
Krypt.-Fl., Edn 2 (Leipzig) 1.2: 153 (1885) 2015). However, Diplogelasinospora and Zopfiella need to be
Facesoffungi number: FoF 01900 transferred to Lasiosphaeriaceae, which was shown by Cai et al.
Saprobic or parasitic on plant debris, straw, seeds, dung, (2006b, c) and Morgenstern et al. (2012) based on gene se-
feathers of birds, soil, paper, textiles, air, mushrooms, rabbit quence data. Members of this family are ubiquitous fungi,
Fungal Diversity

commonly residing in soil, on lignin and cellulosic or similar Chaetomium Kunze, in Kunze and Schmidt,
materials such as paper, cotton, fabrics, straw, manure. Species Mykologische Hefte (Leipzig) 1: 15 (1817)
have also been found as human opportunistic pathogens (von Facesoffungi number: FoF 01901; Fig. 34
Arx et al. 1986; Mukerji and Manoharachary 2010; Ahmed Saprobic or parasitic on plant debris, straw, seeds, dung,
et al. 2015). feathers of birds, soil, paper, textiles and humans. Sexual

Fig. 34 Chaetomium globosum (Material examined: THAILAND, d, e Ascomatal hairs f Squash mount showing surface of peridium g, h
Phayao Province, Muang District, on dead moist twig of Tectona Immature asci i Mature asci j Immature ascospores k-m Mature
grandis L. (Lamiaceae), M. Doilom, 12 March 2012, MFLU 15-3206, ascospores n Germinating ascospore. Scale bars: a = 1000 μm,
living culture, MFLUCC 12-0290) a Ascomata on PDA after 7 days b b = 500 μm, c = 200 μm, d, e, g, = 20 μm, f, h–k, n = 10 μm, l, m = 5 μm
Close-up ascomata on PDA c Ascomata with yellowish ascomatal hairs
 Fungal Diversity

morph: Ascomata dark, olive brown to greyish yellow, grey or Bommerella Marchal, Bull. Soc. R. Bot. Belg. 24(2): 164
green, superficial, globose, subglobose to ovoid, pyriform or (1885)
ampulliform, numerous, with broadly rounded base and narrow Type species: Bommerella trigonospora Marchal, Bull.
ostiole. Peridium with thick wall, composed of several layers of Soc. R. Bot. Belg. 24(2): 164 (1885)
pale brown cells of textura intricata or textura epidermoidea. Boothiella Lodhi & Mirza, Mycologia 54(2): 217 (1962)
Ascomatal hairs numerous, usually unbranched, brown, flexu- Type species: Boothiella tetraspora Lodhi & J.H. Mirza,
ous, undulate or coiled, often tapering, septate, broad at the Mycologia 54(2): 217 (1962)
base. Asci 8-spored, rarely 4-spored, unitunicate, clavate, ob- Botryotrichum Sacc. & Marchal, in Marchal, Bull. Soc. R.
ovate, fusiform, or nearly cylindrical, pedicellate, apically Bot. Belg. 24(1): 66 (1885)
rounded, evanescent. Ascospores overlapping biseriate to Type species: Botryotrichum piluliferum Sacc. & Marchal,
crowded, hyaline when young, later brown, variable in shape, in Marchal, Bull. Soc. R. Bot. Belg. 24(1): 66 (1885)
most are ellipsoidal, broadly fusiform or limoniform, aseptate, Chaetomidium (Zopf) Sacc., Syll. fung. (Abellini) 1: 39
thick and smooth-walled, containing several small guttules, (1882)
with an apical germ pore. Asexual morph: Hyphomycetous. Type species: Chaetomidium fimeti (Fuckel) Zopf, Syll.
Colonies brown or orange or white, woolly, border even, re- fung. (Abellini) 1: 39 (1882)
verse buff-colored to brown, aerial mycelium cottony or felty, Corynascella Arx & Hodges, in von Arx, Stud. Mycol. 8:
with funiculose setae and hyphae. Mycelium abundant, 23 (1975)
intramatrical and aerial, composed of hyaline or dark brown, Type species: Corynascella humicola Arx & Hodges, Stud.
septate, hyphae. Conidiophores reduced to conidiogenous cells Mycol. 8: 23 (1975)
or simple, vegetative hyphae, branched, hyaline, septate, with Chaetomiopsis Moustafa & Abdul-Wahid, Mycologia
smooth or slightly rough wall. Conidiogenous cells phialidic, 82(1): 129 (1990)
hyaline, lateral or intercalary on aerial mycelium, or immersed Type species: Chaetomiopsis dinae Moustafa & Abdul-
in agar or on conidiophore. Conidia hyaline to brown, Wahid, Mycologia 82(1): 129 (1990)
subglobose to globose, or oblong, occasionally cylindrical to Crassicarpon Y. Marín et al., Mycologia 130(3): 629
pyriform, formed single or in chains, aseptate, smooth-walled, (2015)
wall thick with age. Type species: Crassicarpon thermophilum (Fergus &
Type species: Chaetomium globosum Kunze, in Kunze Sinden) Y. Marín et al., Mycologia 130(3): 629 (2015)
and Schmidt, Mykologische Hefte (Leipzig) 1: 16 (1817) Emilmuelleria Arx, Sydowia 38: 6 (1986) [1985]
Notes: Chaetomium is a cosmopolitan genus with more Type species: Emilmuelleria spirotricha (R.K. Benj.) Arx,
than 150 species (Asgari and Zare 2011a; Zhang et al. Sydowia 38: 6 (1986) [1985]
2012a, b). The genus belongs to Sordariomycetes, family Farrowia D. Hawksw., Persoonia 8(2): 173 (1975)
Chaetomiaceae with C. globosum as the type species Type species: Farrowia longicollea (Krzemien. & Badura)
(Zhang et al. 2012a, b). Special characteristics of D. Hawksw. [as ‘Farrowia longicollis’], (1975) Persoonia
Chaetomium are ascomata covered with hairs or setae 8(2): 173 (1975)
(Hawksworth and Wells 1973), clavate or fusiform or some- Guanomyces M.C. González, Hanlin & Ulloa, Mycologia
times cylindrical thin-walled asci, lacking paraphyses before 92(6): 1139 (2000)
ascomata mature (von Arx et al. 1986) and 1-celled, brown or Type species: Guanomyces polythrix M.C. González et al.,
gray-brown ascospores with germ pores. A comprehensive Mycologia 92(6): 1141 (2000)
classification of Chaetomium was provide by Dreyfuss Humicola Traaen, Nytt Mag. Natur. 52: 31 (1914)
(1976) based on morphological characters and some physio- Type species: Humicola fuscoatra Traaen, Nytt Mag.
logical traits (temperature and nutritional requirements, Natur. 52: 33 (1914)
growth and fruiting rate, and compatibility). Chaetomium spe- Madurella Brumpt, Compt.-Rend. Séances Mém. Soc.
cies can be potential biological control agents (Soytong et al. Biol. 58: 999 (1905)
2001), can produce bioactive metabolites (Wang et al. 2012; Type species: Madurella mycetomi (Laveran) Brumpt [as
Li et al. 2014), others produce mycotoxins and infect skin and ‘Madurella mycetomatis’], (1905)
nails in humans (Pieckova 2003). Myceliophthora Costantin, C. r. hebd. Séanc. Acad. Sci.,
Paris 114: 849 (1892)
Type species: Myceliophthora lutea Costantin, C. r. hebd.
Other genera included Séanc. Acad. Sci., Paris 114: 2 (1892)
Staphylotrichum J. Mey. & Nicot, Bull. trimest. Soc.
Achaetomium J.N. Rai et al., Can. J. Bot. 42(6): 693 (1964) mycol. Fr. 72: 322 (1957) [1956]
Type species: Achaetomium globosum J.N. Rai & J.P. Type species: Staphylotrichum coccosporum J.A. Mey. &
Tewari, in Rai et al., Can. J. Bot. 42(6): 693 (1964) Nicot, Bull. trimest. Soc. mycol. Fr. 72(4): 323 (1957) [1956]
Fungal Diversity

Subramaniula Arx, Proc. Indian Acad. Sci., Pl. Sci. 94(2– 13. Asci 4-6-spored . . . . . . . . . . . . . . . . . . Crassicarpon
3): 344 (1985) 13. Asci 8-scospored . . . . . . . . . . . . . . . . . . . . . . . . . . 14
Type species: Subramaniula thielavioides (Arx et al.) Arx, 14. Asci cylindrical, clavate or obovate . . . .Corynascella
Proc. Indian Acad. Sci., Pl. Sci. 94(2–3): 344 (1985) 14. Asci globose to broadly ellipsoidal. . . . . .Corynascus
Taifanglania Z.Q. Liang et al., Fungal Diversity 34: 72 15. Conidiogenous blastic . . . . . . . . . . . . . . . . . . . . . . 16
(2009) 15. Conidiogenous phialidic, conidia amerospores, basipe-
Type species: Taifanglania hechuanensis Z.Q. Liang tal chains with or without connectives . . Taifanglania
et al., Fungal Diversity 34: 72 (2009) 16. Conidiophore absent . . . . . . . . . . . . . . . . . . . . . . . 17
Thielavia Zopf, Verh. bot. Ver. Prov. Brandenb. 18: 101 16. Conidiophore present . . . . . . . . . . . . . . . . . . . . . . . 18
(1876) 17. Conidia brown . . . . . . . . . . . . . . . . . . . . . . Humicola
Type species: Thielavia basicola Zopf, Sber. bot. Ver. Prov. 17. Conidia hyaline . . . . . . . . . . . . . . . . . . . . .Madurella
Brandenb. 18: 105 (1871) 18. Conidiophore branched . . . . . . . . . . . . . . . . . . . . . 19
18. Conidiophore reduced, unbranched, conidiogenus
monoblastic or polyblastic, conidia hyaline,
Key to genera of Chaetomiaceae ovoid. . . . . . . . . . . . . . . . . . . . . . . . Myceliophthora
19. Conidiophore apically branched, brown, conidia pale
1. Sexual morph present . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 brown . . . . . . . . . . . . . . . . . . . . . . . Staphylotrichum
1. Asexual morph present. . . . . . . . . . . . . . . . . . . . . . . 15 19. Conidiophore branched from base, hyaline, conidia
2. Ascospores with one germ pore . . . . . . . . . . . . . . . . . 3 hyaline . . . . . . . . . . . . . . . . . . . . . . . Botryotrichum
2. Ascospores with two germ pores . . . . . . . . . . . . . . . 12
3. Ascomata perithecioid . . . . . . . . . . . . . . . . . . . . . . . . 4 Chaetosphaerellaceae Huhndorf et al., in Huhndorf et al.,
3. Ascomata cleistothecioid . . . . . . . . . . . . . . . . . . . . . . 8 Mycol. Res. 108(12): 1387 (2004)
4. Ascospores triangular; ascomata bearing ampulliform Facesoffungi number: FoF 01114
setae . . . . . . . . . . . . . . . . . . . . . . . . . . . . Bommerella Saprobic on woody substrates in terrestrial habitats. Sexual
4. Ascospores limoniform, ovoid, globose or fusoid; morph: Ascomata perithecial, dark brown to black, scattered
ascoma ornamentation different from ampulliform or densely gregarious, superficial, sitting on or in a subiculum
setae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 or absent, pyriform, obpyriform or ovoid, coriaceous, turbinate
5. Ascomata glabrous, urn-shaped . . . . . Subranmaniula or tuberculate or smooth, with or without, brown, branched or
5. Ascomata bearing hyphae and/or appendages, not urn- unbranched setae, papilla at the apex present or lacking,
shaped . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 collabent or not collapsing, ostiolate. Subiculum thin or thick,
6. Ascomata attached to substrate by basal tuft of hyphae; brown to dark brown, septate, branched or unbranched with
appendages not ornamented . . . . . . . . . . . . . Farrowia spiny hyphae. Peridium thick (10–80 μm), munk pores present,
6. Ascomata not attached to substrate by basal tuft of hy- outer layer composed of dark brown to brown cells of textura
phae; appendages ornamented . . . . . . . . . . . . . . . . . . 7 angularis; inner layer composed of hyaline cells of textura
7. Peridium dark and wide beneath prismatica. Hamathecium composed of paraphyses, often pres-
vestiture . . . . . . . . . . . . . . . .Achaetomium ent only in young ascomata. Asci 8-spored, unitunicate, clavate,
7. Peridium brown or pallid and narrow beneath long or short pedicellate, apical ring distinct, indistinct or ab-
vestiture . . . . . . . . . . . . . . . . . . . . . . . . .Chaetomium sent, evanescent. Ascospores 1–3-seriate or overlapping, with
8. Ascomata with pallid or light brown, translucent brown or brown median cells and hyaline end cells, oblong
peridium . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 cylindrical, ellipsoid or fusiform, 1–3-septate. Asexual morph:
8. Ascomata with dark peridium . . . . . . . . . . . . . . . . . 10 Hyphomycetous. Conidiophores mononematous, brown, sep-
9. Germ pore not protuberant . . . . . . . . . . . . . . Thielavia tate, branched, percurrent, with a terminal ampulla.
9. Germ pore protuberant . . . . . . . . . . . . . . . . .Boothiella Conidiogenous cells enteroblastic, phialidic, holoblastic or
10. Ascomata bearing appendages . . . . . . . . . . . . . . . . 11 polyblastic, integrated, terminal, brown. Conidia brown, oval
10. Ascomata glabrous. . . . . . . . . . . . Thermothelomyces or elliptical, solitary or catenate, 1–3-septate.
11. Ascomata bearing smooth, coiled Type: Chaetosphaerella E. Müll. & C. Booth
appendages. . . . . . . . . . . . . . . . . . . .Emilmuelleria
11. Ascomata bearing verrucoses and/or smooth, straight or Notes: The family Chaetosphaerellaceae was introduced by
undulate appendages, sometimes with circinate Huhndorf et al. (2004a) based on its superficial, ostiolate
tips . . . . . . . . . . . . . . . . . . . . . . . . . . . Chaetomidium ascomata on a subiculum, clavate or cylindrical asci, with
12. Ascomata perithecioid, bearing setae or branched hy- pigmented ellipsoid ascospores, and enteroblastic, phialidic or
phae or circinate appendages . . . . . . . . .Chaetomium holoblastic conidiogenesis, and is typified by Chaetosphaerella
12. Ascomata cleistothecioid . . . . . . . . . . . . . . . . . . . . 13 (Müller and Booth 1972). Sivanesan (1976) synonymized
 Fungal Diversity

Chaetosphaeria fusispora P. Larsen as Chaetosphaerella only in young ascomata. Asci 8-spored, unitunicate, clavate,
fusispora Sivan. In a re-evaluation of Chaetosphaeria species, long or short pedicellate, apical ring distinct or indistinct, ev-
Réblová (1999a, b, c, d) accepted only Chaetosphaerella anescent. Ascospores 1–3-seriate or overlapping, with brown
phaeostroma (Durieu & Mont.) E. Müll. & C. Boot and median cells and hyaline end cells, cylindrical to oblong,
C. fusca (Fuckel) E. Müll. & C. Booth in Chaetosphaerella slightly curved, 3-septate, slightly constricted or not constrict-
and placed this genus in the family Helminthosphaeriaceae ed in the center, middle cells may become slightly inflated,
based on the characteristic ostiolate ascomata without a smooth-walled. Asexual morph: Hyphomycetous.
quellkörper. The new genera Tengiomyces and Crassochaeta Conidiophores mononematous, brown, septate, branched,
with versicolorous ascospores were placed in percurrent, with a terminal, ampulla. Conidiogenous cells ho-
Helminthosphaeriaceae (Tengiomyces indicus (Varghese & loblastic or enteroblastic, polyblastic, integrated, terminal,
V.G. Rao) Réblová) and Trichosphaeriaceae (Crassochaeta smooth-walled. Conidia brown, elliptical, solitary, 3-septate,
nigrita (Sacc.) Réblová and C. fusispora (Sivan.) Réblová) slightly constricted in the middle, the central cells longer and
(Réblová 1999a, d). Réblová (1999d) transferred darker than the end cells, sometimes with guttules.
Chaetosphaerella fusispora to Crassochaeta fusispora (Sivan.) Type species: Chaetosphaerella phaeostroma (Durieu &
Réblová based on different asexual morphs; Chaetosphaerella is Mont.) E. Müll. & C. Booth, Trans. Br. mycol. Soc. 58(1):
associated with Oedemium and Veramycina asexual morphs, 77 (1972)
while Crassochaeta has Undetermined asexual morphs with Notes: The genus Chaetosphaerella was introduced by
Arthrinium-like conidia. Réblová (1999a) reported that Müller and Booth (1972) and included two species
Chaetosphaerella was similar to the family Nitschkiaceae based (C. phaeostroma and C. fusca (Fuckel) E. Müll. & C.
on their characteristic peridium. Booth). The type species C. phaeostroma, is characterized
LSU rDNA sequence data analysis (Huhndorf et al. by black ascomata surrounded by a subiculum, with tubercu-
2004b), showed that Chaetosphaerella and Crassochaeta late or roughened perithecia, clavate asci and pigmented
clustered in the same clade i.e. Chaetosphaerellaceae. ascospores. Varghese and Rao (1979) and Sivanesan (1976)
Mugambi and Huhndorf (2010) introduced included C. fusispora and C. indica Varghese & V.G. Rao in
Spinulosphaeria in Sordariomycetes genera incertae Chaetosphaerella based on the morphological characters,
sedis, based on morphology and analysis of LSU se- however, Réblová (1999a, d) excluded these two from the
quence data of S. nuda Mugambi and Huhndorf. In genus as they had different asexual morphs.
Maharachchikumbura et al. (2015), Oedemium was in-
cluded in Chaetosphaerellaceae based on the type spe-
cies Chaetosphaerella phaeostroma linked with
Other genera included
Oedemium minus (Link) S. Hughes. The type species
o f O e d e m i u m ( O . a t ru m L i n k ) i s l i n k e d w i t h
Crassochaeta Réblová, Mycotaxon 71: 46 (1999)
Chaetosphaerella fusca (Müller and Booth 1972;
Type species: Crassochaeta nigrita (Sacc.) Réblová,
Réblová 1999a, b, c, d). In a combined gene dataset
Mycotaxon 71: 48 (1999)
including LSU, SSU, TEF and RPB2 sequences,
Oedemium Link, in Willdenow, Sp. pl., Edn 4 6(1): 42
Spinulosphaeria nuda formed a sister group to
(1824); Fig. 36
Chaetosphaerella species with high support in the fam-
Type species: Oedemium atrum Link, in Willdenow, Sp.
ily Chaetosphaerellaceae (Mugambi and Huhndorf
pl., Edn 4 6(1): 42 (1824)
2010; Maharachchikumbura et al. 2015).
Spinulosphaeria Sivan., Trans. Br. mycol. Soc. 62(1): 5
Chaetosphaerella E. Müll. & C. Booth, Trans. Br. mycol.
(1974)
Soc. 58(1): 76 (1972)
Type species: Spinulosphaeria thaxteri (Pat.) Sivan.,
Facesoffungi number: FoF 01115; Fig. 35
Trans. Br. mycol. Soc. 62(1): 36 (1974)
Saprobic on woody substrates in terrestrial habitats. Sexual
morph: Ascomata perithecial, dark brown or black, densely
gregarious, superficial, sitting on a subiculum, obpyriform,
coriaceous, tuberculate or slightly roughened, with brown se- Key to genera of Chaetosphaerellaceae
tae, with a small papilla at the apex, collabent when dry,
ostiolate. Subiculum thin, brown to dark brown, septate, 1. Asexual morph with brown, septate conidia, produced in
branched with spiny hyphae. Peridium thick (20–70 μm), short acropetal chains, conidiogenesis monoblastic or sympo-
munk pores present, outer layer composed of dark brown to dial simple, brown conidiophores Oedemium
brown cells of textura angularis; inner layer composed of 1. Asexual morph differs and/or sexual morph present . . 2
none pigmented cells of textura prismatica. Hamathecium 2. Ascomata scattered, ascospores 1-septate when
composed of numerous, filiform, paraphyses, often present mature . . . . . . . . . . . . . . . . . . . . . . . Spinulosphaeria
Fungal Diversity

Fig. 35 Chaetosphaerella
phaeostroma (Material
examined: FRANCE, Paris, on
dead wood, Montagne C, PC
PC0167641) a Herbarium packet
b Herbarium material c Ascomata
on host d Ascoma cross section e,
f Peridium g Septate and spiny
hypha h–j Immature and mature
asci k–m Ascospores. Notes: d-m
soaked in 3 % KOH. Scale bars:
c = 200 μm, d = 100 μm,
e = 50 μm, f = 30 μm, g–
j = 20 μm, k–m = 10 μm

2. Ascomata densely gregarious, ascospores 3-septate Facesoffungi number: FoF 01139

when mature,. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 Saprobic on wood in terrestrial habitats, some fungicolous.
3. Ascomata collapsing or not, tuberculose, asexual morphs Sexual morph: Ascomata perithecial, dark brown to black,
Oedemium . . . . . . . . . . . . . . . . . . . . Chaetosphaerella gregarious or scattered, solitary, superficial or basally im-
3. Ascomata not collapsing, with spiny setae on the surface, mersed on a thin stroma or on a thin subiculum or the
asexual morphs Arthrinium-like . . . . . . . Crassochaeta subiculum absent, ovoid, globose to subglobose, carbona-
ceous, coriaceous or membranaceous, rough, smooth or with
Chaetosphaeriaceae Réblová et al., Sydowia 51(1): 56 short setae, papillate, the apex collapsing when dry. Ostiole
(1999) lined with hyaline periphyses or absent. Subiculum thin,
 Fungal Diversity

Fig. 36 Oedemium minus (Link)

S. Hughes (Asexual morph of
Chaetosphaerella phaeostroma,
Silva 2015) a Conidia b, c
Conidiogenous cells and conidia
d Setae. Scale bars = 20 μm

scanty, brown to dark brown, septate, of unbranched hyphae. excipulum, smooth, thick-walled, multi-septate.
Peridium thin-walled (10–40 μm), composed of two layers, Conidiophores lining the basal stroma in a dense layer or
outer layer comprising brown cells of textura epidermoidea or arising from conidiomatal cavity, brown, 4–6-septate, un-
angularis, carbonaceous; inner layer comprising hyaline cells branched, cylindrical, thin-walled, smooth. Conidiogenous
of textura prismatica, thin, membranaceous. Hamathecium cells integrated, determinate, holoblastic or enteroblastic,
with numerous, septate, unbranched, tapering, filiform or cy- phialidic with conspicuous periclinal thickening at an attenu-
lindrical paraphyses. Asci 8-spored, unitunicate, thin-walled, ated apex, brown, smooth, subcylindrical to lageniform.
clavate to cylindrical, long or short pedicellate, with J-, refrac- Conidia hyaline to brown, aseptate, thin-walled, smooth, fu-
tive apical ring. Ascospores 2–3-seriate, hyaline or brown or siform to allantoid, curved or straight, obtuse to subobtusely
becoming dark colored in part, fusiform, cylindrical to ellip- rounded at apex, truncate at base, eguttulate or guttulate, with
soid, sometimes curved, 0–3-septate, with or without a single, cellular, unbranched, flexuous, with tubular append-
fragmenting, smooth or striate, with guttules, sheath or ap- age at each end, separated by a septum, with basal asymmet-
pendages Asexual morph: Hyphomycetous. Conidiophores rically located appendage.
macronematous, mononematous, scattered or gregarious, dark Type: Chaetosphaeria Tul. & C. Tul.
brown or hyaline, straight or flexuous, septate, branched or Notes: The family Chaetosphaeriaceae was introduced by
unbranched, with short encircling collar hyphae. Réblová et al. (1999) based on Chaetosphaeria (Tulasne and
Conidiogenous cells monophialidic or polyphialidic, holo- Tulasne 1863a). Müller and von Arx (1962) included
blastic or enteroblastic, proliferating percurrently or sympodi- Chaetosphaeria in the Sphaeriaceae. Réblová et al. (1999)
al, hyaline, with a distinct funnel-shaped collarette, smooth- suggested Chaetosphaeriaceae belonged in the order
walled. Conidia aggregated, continuous or mucilaginous, hy- Sordariales based on morphological characters. Huhndorf
aline to brown, aseptate to multi-septate, flexuous, fusiform, (2004b), placed this family in Chaetosphaeriales, based on
cylindrical to allantoid, curved or straight, with obtuse to LSU nrDNA sequence data. Zignoëlla was distinguished from
subobtusely rounded apex, truncate at base, eguttulate or Chaetosphaeria based on the color of the ascospores
guttulate. Coelomycetous. Conidiomata stromatic, scattered (Saccardo 1883; Munk 1953), but Booth (1957) regarded
or aggregated, superficial, cupulate or globose, unilocular, se- Zignoëlla as a synonym of Chaetosphaeria. Based on mor-
tose, comprising black to dark brown cells of textura phological characters, Chaetosphaeria did not appear to be-
angularis or intricata. Setae numerous, black to brown, sep- long to Helminthosphaeriaceae or Trichosphaeriaceae
tate, ovoid to cylindrical or arising from the outer elements of (Samuels et al. 1997a; Réblová et al. 1999). This family is
Fungal Diversity

morphologically diverse with the genera Catenularia, comprising of hyaline cells of textura prismatica, thin,
Cylindrotrichum, Chalara, Chloridium, Custingophora, membranaceous. Hamathecium with abundant, cylindri-
Dictyochaeta, Menispora, Phialophora and Zanclospora re- cal, septate branched paraphyses. Asci 8-spored,
ported as the asexual morphs (Réblová et al. 1999). unitunicate, clavate to cylindrical, short pedicellate, with
Infundibulomyces was included in Chaetosphaeriaceae based small, distinct, refractive, J- apical ring. Ascospores 2–
on SSU and LSU gene data by Somrithipol et al. (2008). 3-seriate, hyaline, allantoid or ellipsoid, curved, septate,
Lumbsch and Huhndorf (2010) listed ten sexual genera in smooth-walled, with guttules. Asexual morph:
the family, while Maharachchikumbura et al. (2015) listed Hyphomycetous. Conidiophores macronematous,
35 sexual and asexual genera. Hashimoto et al. (2015) intro- mononematous, scattered or gregarious, dark brown or
duced Neopseudolachnella and Pseudodinemasporium to the hyaline, straight or flexuous, septate, branched or un-
family and key to 37 genera are showing here. The genera of branched, with short encircling collar hyphae.
the family are confused and a monograph is needed with mo- Conidiogenous cells monophialidic or polyphialidic, ho-
lecular support for accepted genera. loblastic to enteroblastic, proliferating percurrently or
sympodially, hyaline, with a distinct funnel-shaped
Chaetosphaeria Tul. & C. Tul., Select. fung. carpol. (Paris) collarette, smooth-walled. Conidia aggregated, continu-
2: 252 (1863) ous or mucilaginous, hyaline to brown, aseptate to
Facesoffungi number: FoF 01140, Fig. 37 multi-septate, flexuous, fusiform, cylindrical to allantoid,
Saprobic on wood in terrestrial habitats. Sexual curved or straight, obtuse to subobtusely rounded at
morph: Ascomata perithecial, black, gregarious, superfi- apex, truncate at base, with or without appendage,
cial or basally immersed in a thin basal stroma, ovoid, eguttulate or guttulate.
globose to subglobose, carbonaceous or membranaceous, Type species: Chaetosphaeria innumera Berk. & Broome
the apex collapsing when dry, smooth or covered by ex Tul. & C. Tul., Select. fung. carpol. (Paris) 2: 252 (1863)
setae, papillate, ostiole lined with hyaline periphyses,. Notes: The genus Chaetosphaeria was introduced by
Peridium thick (10–20 μm), composed of two layers, Tulanse and Tulanse (1863a) and is typified by C. innumera.
outer layer comprising brown cells of textura Saccardo (1883) included the genus in the “Sphaeriaceae,
e p i d e r m o i d e a , t h i n , c a r b o n a c e o u s ; i n n e r la y e r Phaeophragmiae” which included several species with dark

Fig. 37 Chaetosphaeria innumera (redrawn from Discover Life, http://www.discoverlife.org/) a Ascoma with setae. b Vertical section of ascoma c
Conidiophores and conidia (asexual morph: Dictyochaeta fuegiana) d. Asci and paraphyses. e. Ascospores. Scale bars: a–b = 100 μm, c–e = 10 μm
 Fungal Diversity

ascospores, along with the type species. Huhndorf et al. Type species: Dinemasporium graminum (Lib.) Lév.,
(2004b) referred it to the Chaetosphaeriaceae based on the Annls Sci. Nat., Bot., sér. 3 5: 274 (1846)
LSU nrDNA analyses. The sexual morph is characterized by Dictyochaeta Speg., Physis, Rev. Soc. Arg. Cienc. Nat. 7:
superficial perithecia, cylindrical asci and brown ascospores, 18 (1923); Fig. 38
while the asexual morph is characterized by enteroblastic Type species: Dictyochaeta fuegiana Speg., Physis, Rev.
conidiogenous cells, and aggregated conidia (Réblová Soc. Arg. Cienc. Nat. 7: 18 (1923)
1999a, b, c, d). Dictyochaetopsis Aramb. & Cabello, Mycotaxon 38: 12
Fernández et al. (2006) concluded that two kinds of pat- (1990)
terns are observed in the asexual morph of Chaetosphaeria. Type species: Dictyochaetopsis apicalis (Berk. & M.A.
1. Broadly integrated conidiogenous cells and endogenous Curtis) Aramb. & Cabello, Mycotaxon 38: 12 (1990)
conidiogenous loci with aseptate to multi-septate conidia Exserticlava S. Hughes, N.Z. J Bot. 16(3): 332 (1978)
(Exserticlava S. Hughes 1978); 2. Narrow conidiogenous Type species: Exserticlava vasiformis (Matsush.) S.
locus in a discrete or integrated conidiogenous cells with Hughes, N.Z. J Bot. 16(3): 332 (1978)
aseptate to uni-septate conidia (Chloridium Link 1809). Gonytrichum Nees & T. Nees, Nova Acta Phys.-Med.
Acad. Caes. Leop.-Carol. Nat. Cur. 9: 244 (1818)
Type species: Gonytrichum caesium Nees, Nova Acta
Acad. Leop. Carol. Ac. Naturf. Fo. 9: 244 (1818)
Other genera included Hemicorynespora M.B. Ellis, Mycol. Pap. 131: 19 (1972)
Type species: Hemicorynespora deightonii M.B. Ellis,
Ascochalara Réblová, Sydowia 51(2): 212 (1999) Mycol. Pap. 131: 20 (1972)
Type species: Ascochalara gabretae Réblová, Sydowia Infundibulomyces Plaingam, Somrith. & E.B.G. Jones
51(2): 212 (1999) 2003
Brunneodinemasporium Crous & R.F. Castañeda, in Type species: Infundibulomyces cupulata Plaingam et al.,
Crous et al., Persoonia, Mol. Phyl. Evol. Fungi 28: 128 (2012) Can. J. Bot. 81(7): 733 (2003)
Type species: Brunneodinemasporium brasiliense Crous Kionochaeta P.M. Kirk & B. Sutton, Trans. Br. mycol.
& R.F. Castañeda, in Crous et al., Persoonia, Mol. Phyl. Evol. Soc. 85(4): 712 (1986) [1985]
Fungi 28: 129 (2012) Type species: Kionochaeta ramifera (Matsush.) P.M. Kirk
Catenularia Grove, in Saccardo, Syll. fung. (Abellini) 4: & B. Sutton, Trans. Br. mycol. Soc. 85(4): 715 (1986) [1985]
303 (1886) Lecythothecium Réblová & Winka, Mycologia 93(3): 481
Type species: Catenularia simplex Grove, in Saccardo, (2001)
Syll. fung. (Abellini) 4: 303 (1886) Type species: Lecythothecium duriligni Réblová & Winka,
Chloridium Link, Mag. Gesell. naturf. Freunde, Berlin Mycologia 93(3): 482 (2001)
3(1–2): 13 (1809) Melanopsammella Höhn., Annls mycol. 17(2/6): 121
Type species: Chloridium viride Link, Mag. Gesell. naturf. (1920) [1919]
Freunde, Berlin 1: 13 (1805) Type species: Melanopsammella inaequalis (Grove)
Codinaea Maire, Publ. Inst. Bot. 3(4): 15 (1937) Höhn., Annls mycol. 17(2/6): 121 (1920) [1919]
Type species: Codinaea aristata Maire, Publ. Inst. Bot. Menispora Pers., Mycol. eur. (Erlanga) 1: 32 (1822)
3(4): 15 (1937) Type species: Menispora glauca (Link) Pers., Mycol. eur.
Codinaeopsis Morgan-Jones, Mycotaxon 4(1): 166 (1976) (Erlanga) 1: 32 (1822)
Type species: Codinaeopsis gonytrichoides (Shearer & J.L. Miyoshiella Kawam., Jap. J. Bot. 4: 295 (1929)
Crane) Morgan-Jones, Mycotaxon 4(1): 167 (1976) Type species: Miyoshiella fusispora (Kawam.) Kawam.,
Craspedodidymum Hol.-Jech., Česká Mykol. 26(2): 70 Jap. J. Bot. 4: 295 (1929)
(1972) Neopseudolachnella A. Hashim et al., Mycologia 107(2):
Type species: Craspedodidymum elatum Hol.-Jech., Česká 385 (2015)
Mykol. 26(2): 70 (1972) Type species: Neopseudolachnella acutispora A. Hashim.
Cryptophiale Piroz., Can. J. Bot. 46: 1123 (1968) et al., in Hashimoto et al., Mycologia 107(2): 385 (2015)
Type species: Cryptophiale kakombensis Piroz., Can. J. Phaeostalagmus W. Gams, in Gams & Holubová-Jechová,
Bot. 46: 1124 (1968) Stud. Mycol. 13: 90 (1976)
Dendrophoma Sacc., Michelia 2(no. 6): 4 (1880) Type species: Phaeostalagmus cyclosporus (Grove) W.
Type species: Dendrophoma cytisporoides Sacc., Michelia Gams, in Gams & Holubová-Jechova, Stud. Mycol. 13: 91
2(no. 6): 4 (1880) (1976)
Dinemasporium Lév., Annls Sci. Nat., Bot., sér. 3 5: 274 Phialogeniculata Matsush., in Kobayasi, Bull. natn. Sci.
(1846) Mus., Tokyo, N.S. 14: 471 (1971)
Fungal Diversity

Fig. 38 Dictyochaeta sp.

(Asexual morph) a–c Colonies on
drying veneer in surface view d–f
Conidiophores with conidia g–n
Conidia. Scale bars: b = 500 μm,
c = 200 μm, e–f = 20 μm, d, g–
n = 10 μm

Type species: Phialogeniculata guadalcanalensis Type species: Rattania setulifera Prabhug. & Bhat,
Matsush., in Kabayasi et al., Bull. natn. Sci. Mus., Tokyo Mycotaxon 108: 220 (2009)
14(3): 472 (1971) Sporoschisma Berk. & Broome, in Berkeley, Gard.
Pseudobotrytis Krzemien. & Badura, Acta Soc. Bot. Pol. Chron., London: 540 (1847)
23: 761 (1954) Type species: Sporoschisma mirabile Berk. &
Type species: Pseudobotrytis fusca Krzemien. & Badura, Broome, in Berkeley, Gard. Chron., London: 540
Acta Soc. Bot. Pol. 23: 762 (1954) (footnote) (1847)
Pseudodinemasporium A. Hashim et al., in Hashimoto Striatosphaeria Samuels & E. Müll., Sydowia 31(1–6):
et al., Mycologia 107(2): 390 (2015) 131 (1979) [1978]; Fig. 39
Type species: Pseudodinemasporium fabiforme A. Type species: Striatosphaeria codinaeaphora Samuels &
Hashim. et al., in Hashimoto et al., Mycologia 107(2): 390 E. Müll., Sydowia 31(1–6): 132 (1979) [1978]
(2015) Tainosphaeria F.A. Fernández & Huhndorf, Fungal
Pseudolachnea Ranoj., Annls mycol. 8(3): 393 (1910) Diversity 18: 44 (2005)
Type species: Pseudolachnea insignis Velen., Monogr. Type species: Tainosphaeria crassiparies F.A. Fernández
Discom. Bohem. (Prague): 314 (1934) & Huhndorf, in van der Walt & Scott, Fungal Diversity 18: 44
Pyrigemmula D. Magyar & Shoemaker, in Magyar et al., (2005)
Mycol. Progr. 10(3): 310 (2011) Thozetella Kuntze, Revis. gen. pl. (Leipzig) 2: 873 (1891)
Type species: Pyrigemmula aurantiaca D. Magyar & Type species: Thozetella nivea (Berk.) Kuntze, Revis. gen.
Shoemaker, in Magyar et al., Mycol. Progr. 10(3): 309 (2011) pl. (Leipzig) 2: 873 (1891)
Rattania Prabhug. & Bhat, Mycotaxon 108: 218 (2009) Umbrinosphaeria Réblová, Mycotaxon 71: 17 (1999)
 Fungal Diversity

Fig. 39 Striatosphaeria
codinaeaphora (Material
examined: BRAZIL, Roraima. ca
219 km North of Boa Vista, on the
Boa Vista Sta. Elena Venezuela
Rd. (BR 174), on dead log,
Dumont KP, Hosford DR,
Samuels GJ, Buck WR, Araujo I,
Souza MA, Bernardi JC; 1
December 1977, NY 01050494,
isotype) a Herbarium material b
Ascomata c, d Ascomata cross
section e Peridium f Paraphyses g,
h Asci i-n Ascospores Scale bars:
b–c = 200 μm, d–e = 100 μm, g–
h = 50 μm, f = 20 μm, i–n = 5 μm

Type species: Umbrinosphaeria caesariata (Clinton & Key to sexual genera of Chaetosphaeriaceae
Peck) Réblová, Mycotaxon 71: 18 (1999)
Zanclospora S. Hughes & W.B. Kendr., N.Z. J Bot. 3: 151 1. Ascomata immersed or semi-immersed. . . . . . . . . . . . . . 2
(1965) 1. Ascomata superficial . . . . . . . . . . . . . . . . . . . . . . . . . 3
Type species: Zanclospora novae-zelandiae S. Hughes & 2. Ascomata globose to subglobose; ascospores 1–3-sep-
W.B. Kendr., N.Z. Jl Bot. 3: 152 (1965) tate, hyali ne, n ot frag m e n t i n g , a l l a n t o i d o r
Zignoëlla Sacc., Michelia 1(no. 3): 346 (1878) ellipsoid. . . . . . . . . . . . . . . . . . . . . . . Chaetosphaeria
Type species: Zignoëlla pulviscula (Curr.) Sacc., Michelia 2. Ascomata ovoid; ascospores multi-septate, versicolored,
1(no. 3): 346 (1878) fusiform, straight or inequilateral . . . . Lecythothecium
Fungal Diversity

3. Ascomata carbonaceous . . . . . . . . . . . . . . . . . . . . . . . 4 12. Conidia aseptate, brown, ellipsoid to

3. Ascomata membranous . . . . . . . . . . . . . . . . . . . . . . . 7 obovoid. . . . . . . . . . . . . . . . . . . Craspedodidymum
4. Ascospores fragmenting at maturity . . . . . Ascochalara 12. Conidia 1-septate, obclavate,
4. Ascospores not fragmenting . . . . . . . . . . . . . . . . . . . . 5 guttulate . . . . . . . . . . . . . . . . . . Phialogeniculata
5. Ascospores fusiform, straight or inequilateral . . . . . . . 6 13. Conidia thick-walled, capitate hyphae present . . . . 14
5. Ascospores ellipsoid, striate, germ pore in the median 13. Conidia thin-walled, capitate hyphae absent . . . . . . 15
septum. . . . . . . . . . . . . . . . . . . . . . . . . Striatosphaeria 14. Conidia obovoid, aseptate . . . . . . . . . . . Catenularia
6. Ascospores hyaline, concolourous, 3- 14. Conidia ellipsoid, 1–3-septate. . . . . . . . . Exserticlava
septate . . . . . . . . . . . . . . . . . . . Miyoshiella 15. Conidia inverted T-shaped or L-shaped, straight or
6. Ascospores with middle cells brown and the end cells obpyriform . . . . . . . . . . . . . . . . . . . . . . . . Thozetella
hyaline, versicolored, multi-septate . Umbrinosphaeria 15. Conidia slightly curved or not curved. . . . . . . . . . . 16
7. Ascomata smooth or setose . . . . . . . . . . . . . . . . . . . . 8 16. Conidia curved. . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
7. Ascomata rough. . . . . . . . . . . . . . . . . . .Tainosphaeria 16. Conidia not curved. . . . . . . . . . . . . . . . . . . . . . . . . 21
8. Ascospores 1-septate, fragmenting at maturity, 17. Conidia lunate . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
hyaline . . . . . . . . . . . . . . . . . . . . . Melanopsammella 17. Conidia clavate, fusiform or cylindrical . . . . . . . . . 19
8. Ascospores 0–3-septate, not fragmenting, 18. Conidia with appendages at the ends . . . . . Codinaea
greenish . . . . . . . . . . . . . . . . . . . . . . . . . . . Zignoëlla 18. Conidia lacking appendages . . . . . . . . . . Zanclospora
19. Conidia cylindrical, hyaline to yellow-green, strongly
recurved. . . . . . . . . . . . . . . . . . . . . . . . . . Menispora
Key to asexual genera of Chaetosphaeriaceae 19. Conidia clavate or fusiform . . . . . . . . . . . . . . . . . . 20
20. Conidia aseptate, rounded at the apex. . . Kionochaeta
1. Coelomycetous taxon . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 20. Conidia 0–multi-septate, with appendages at the
1. Hyphomycetous taxon . . . . . . . . . . . . . . . . . . . . . . . . 6 ends . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Rattania
2. Conidiomata comprising black to brown cells of textura 21. Conidia aseptate. . . . . . . . . . . . . . . . . . . . . . . . . . . 22
intricata . . . . . . . . . . . . . . . . . . . . . . . . Dendrophoma 21. Conidia septate. . . . . . . . . . . . . . . . . . . . . . . . . . . . 23
2. Conidiomata comprising black to brown cells of textura 22. Conidia subglobose to oval, hyaline to light
angularis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 brown. . . . . . . . . . . . . . . . . . . . . . . . . Gonytrichum
3. Conidia fusiform with obtuse ends, 1-septate, hyaline, 22. Conidia cylindrical, ovoid, reniform . . . . . Chloridium
smooth macroconidia with appendages, intermixed with 23. Conidia hyaline . . . . . . . . . . . . . . . . . . . . . . . . . . . 24
aseptate microconidia. . . . . . . . . . . . . . Pseudolachnea 23. Conidia grey, brown to reddish brown . . . . . . . . . . 25
3. Conidia oblong to fusiform, curved or straight, hyaline to 24. Conidia globose to subglobose. . . . . . . .Cryptophiale
brown, aseptate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 24. Conidia ellipsoid or cylindrical with rounded
4. Conidiomata setae absent . . . . . . . . .Infundibulomyces ends . . . . . . . . . . . . . . . . . . . . . . . . Phaeostalagmus
4. Conidiomata setae present . . . . . . . . . . . . . . . . . . . . . 5 25. Conidia 1-septate, ellipsoid to clavate with apiculate
5. Conidiophores brown, unbranched apex. . . . . . . . . . . . . . . . . . . . . . . . . . Pseudobotrytis
...………………………… Brunneodinemasporium 25. Conidia multi-septate, oblong, obovoid to
5. Conidiophores apex hyaline, base brown . . . . . . . . . . 6 obclavate . . . . . . . . . . . . . . . . . . . . . . 26
6. Conidia cylindrical to subcylindrical 26. Conidia acrogenous, obovoid to obclavate, constricted
……………........................…… Neopseudolachnella at the basal septum. . . . . . . . . . . . . Hemicorynespora
6. Conidia ovoid to ellipsoidal . . . Pseudodinemasporium 26. Conidia not acrogenous, not constricted, long
6. Conidia allantoid to lunate . . . . . . . . . Dinemasporium oblong . . . . . . . . . . . . . . . . . . . . . . . . Pyrigemmula
7. Conidia collarette present . . . . . . . . . . . . . . . . . . . . . . 8 Clavicipitaceae Earle, in Mohr, Contr. U.S. natnl. Herb. 6:
7. Conidia collarette absent . . . . . . . . . . . . . . . . . . . . . 13 170 (1901)
8. Conidia multi-septate . . . . . . . . . . . . . . . . . . . . . . . . . 9 Facesoffungi number: FoF 01313
8. Conidia 0–3-septate . . . . . . . . . . . . . . . . . . . . . . . . . 10 Obligate saprotrophic, parasites and symbiotic with in-
9. Conidia short, cylindrical . . . . . . . . . . . . Sporoschisma sects, fungi, grasses, rushes or sedges. Sexual morph:
9. Conidia lunate, with or without a long appendages at the Stromata or subiculum darkly or brightly coloured, fleshy or
ends . . . . . . . . . . . . . . . . . . . . . . . . . .Dictyochaetopsis tough. Perithecia superficial to completely immersed, ordinal
10. Conidia with appendages . . . . . . . . . . . . . . . . . . . . 11 or oblique in arrangement. Asci cylindrical with thickened
10. Conidia with papillate at the ends. . . . . . . . . . . . . . 12 ascus apex. Ascospores usually cylindrical and multi-septate,
11. Conidia aseptate, hyaline, cylindrical . . Codinaeopsis disarticulating into part-spores or non-disarticulating. Asexual
11. Conidia 0–3-septate, fusiform to ellipsoid morph: Clavicipitaceae can be isolated from these genera:
……..........…......................................... Dictyochaeta Aschersonia, Metarrhizium, Nomuraea, Pochonia,
 Fungal Diversity

Paecilomyces, Rotiferophthora, Tolypocladium and Claviceps Tul., Annls Sci. Nat., Bot., sér. 3 20: 43 (1853)
Verticillium. Facesoffungi number: FoF 01316; Fig. 40
Type: Claviceps Tul. Parasitic of the ovaries of grasses and a few species of
Notes: The name Clavicipitaceae was introduced by Earle rushes and sedges. Unfertilized ovaries are especially
(1901). Nannfeldt (1932) accepted the name, which was used susceptible to infection. Sexual morph: Stromata
as variously classified by Diehl (1950), Luttrell (1951), stipitate, spherical, pale brown to orange-brown when
Rogerson (1970), Eriksson (1982) Eriksson and fresh. Stroma surface dotted with mid to dark brown,
Hawksworth (1985), Spatafora and Blackwell (1993) and weakly to strongly papillate ostioles. Ascomata numer-
Rehner and Samuels (1994). Sung et al. (2007) divided the ous, 150–250 per stroma, ovoid with a somewhat elon-
family Clavicipitaceae into three monophyletic groups gate neck region. Perithecia are at least partially embed-
(C la v i c i p i t a c e a e, Ophioco rdyc i pita ceae a nd ded within the stromata and are distributed over the
Cordycipitaceae) based on phylogenetic analyses. surface, resulting in a punctate appearance. Asci, nar-
Rogerson (1970) list 13 genera under Clavicipitaceae. rowly cylindrical, thin-walled and not fissitunicate, with
W h i t e e t a l . ( 2 00 0 ) c h a r ac t e r i z e d s i x g en e r a i n a conspicuously thickened capitate apex that is penetrat-
Clavicipitaceae based on morphological and phylogenetic ed by a narrow channel, 8-spored. Ascospores arranged
analyses. Nigrocornus was introduced to accommodate a spe- in a fascicle, not helically coiled, thin, filiform-shaped.
cies of Balansia, which were significantly different from that of Ascospores that land on stigmas of a susceptible host
the type and all other Balansia species (Ryley 2003). Sung et al. germinate and produce infection hyphae that grow down
(2007) list 22 genera under Clavicipitaceae. They also listed the element to infect the base of the ovary. Within sev-
Berkelella, Cavimalum, Dussiella, Epicrea, Helminthascus, eral days of infection, a sphacelium producing large
Konradia, Moelleriella, Mycomalus, Neobarya, numbers of conidia develops. A sugary syrup commonly
Neocordyceps, Podocrella, Romanoa, Sphaerocordyceps, and referred to as the honeydew stage is replaced by a scle-
Stereocrea as uncertain genera in the family. Kirk et al. (2008) rotium, generally 1–4 times larger than the host seed.
recorded 43 genera in this family within the order Hypocreales. Asexual morph: Undetermined.
Chaverri et al. (2008) introduced Samuelsia under Type species: Claviceps purpurea (Fr.) Tul., Annls Sci.
Clavicipitaceae based on morphology and phylogenetic Nat., Bot., sér. 3 20: 45 (1853)
analyses. Ustilaginoidea was introduced by Brefeld (1895) Notes: This genus was described by Tulasne (1853). There
and revised by Tanaka et al. (2008) as a genus of were some doubts as to the taxonomic status of the genus in
Clavicipitaceae. Conoideocrella and Orbiocrella have been in- the 1950s, which has been corrected in recent years based on
troduced by Johnson (2009). Chamaeleomyces was introduced molecular sequence data and acceptance of Clavicipitaceae
as a new monotypic genus isolated from the liver of Chamaeleo (Pazoutova and Parbery 1999; Yokoyama et al. 2006; Sung
calyptratus based on morphological and phylogenetic analyses et al. 2007; Tanaka et al. 2008). This genus occur on a wide
(Sigler et al. 2010). Periglandula was introduced with range of host plants and is pathogenic on some economic
P. ipomoeae U. Steiner as the type species, which was isolated plants (Hulvova et al. 2013). The genus produces secondary
from Ipomoea asarifolia in Ecuador (Steiner et al. 2011). metabolites with toxicity and pharmaceutical properties
Lumbsch and Huhndorf (2010) list 32 genera under (Hulvova et al. 2013). There are 36 species of Claviceps,
Clavicipitaceae. According to phylogenetic analyses, Kepler which can infect about 600 species of monocotyledonous
et al. (2012) placed Tyrannicordyceps under Clavicipitaceae, plants (Hulvova et al. 2013).
which live on sclerotia of Claviceps and Collarina was intro-
duced by Crous et al. (2014b). Maharachchikumbura et al.
(2015) list 48 genera under this family including Other genera included
Aciculosporium, Amphichorda, Atkinsonella, Balansia,
Cavimalum, Chamaeleomyces, Claviceps, Conoideocrella, Aciculosporium I. Miyake, Bot. Mag., Tokyo 22: (307)
Corallocytostroma, Diploöspora, Dussiella, Ephelis, Epichloë, (1908)
Epicrea, Helminthascus, Heteroepichloë, Hypocrella, Konradia, Type species: Aciculosporium take I. Miyake, Bot. Mag.,
Loculistroma, Metacordyceps, Metarhiziopsis, Metarhizium, Tokyo 22: (307) (1908)
Metapochonia, Moelleriella, Mycomalus, Myriogenospora, Amphichorda Fr., Syst. orb. veg. (Lundae) 1: 170 (1825)
Neobarya, Neoclaviceps, Neocordyceps, Neotyphodium, Type species: Amphichorda felina (DC.) Fr., Syst. orb.
Nigrocornus, Nomuraea, Orbiocrella, Parepichloë, veg. (Lundae) 1: 170 (1825)
Periglandula, Pochonia, Pseudogibellula, Pseudomeria, Atkinsonella Diehl, Agriculture Monogr., US Dept Agric.
Regiocrella, Romanoa, Rotiferophthora, Samuelsia, 4: 48 (1950)
Shimizuomyces, Sphacelia, Sphaerocordyceps, Stereocrea, Type species: Atkinsonella hypoxylon (Peck) Diehl, Agric.
Tyrannicordyceps and Ustilaginoidea. Monogr. U.S.D.A. 4: 49 (1950)
Fungal Diversity

Fig. 40 Claviceps purpurea

(Material examined: CHINA.
Province of Sichuan, on Phalaria
arundinacea L. (Poaceae). 19
September 1989, HMAS 65629,
holotype) a, b Overview of
sclerotium and host c Sclerotium
with stromata d Part of stromata e
Cross section of ascomata f
Ascomata g, h Asci i, j
Ascospores. Scale bars: e = 1 mm,
f = 50 μm, g, h = 20 μm, i,
j = 20 μm

Balansia Speg., Anal. Soc. cient. argent. 19(1): 45 (1885) Corallocytostroma Y.N. Yu & Z.Y. Zhang, Acta Microbiol.
Type species: Balansia claviceps Speg., Anal. Soc. cient. Sin. 20(3): 232 (1980)
argent. 19(1): 45 (1885) Type species: Corallocytostroma oryzae Y.N. Yu & Z.Y.
Cavimalum Y Oshim. Doi, Dargan& K.S. Thind, Bull. Zhang, Actamicrobiol. Sin. 20(3): 234 (1980)
natn. Sci. Mus., Tokyo, B 3(1): 23 (1977) Diploöspora Grove, J. Bot., Lond. 54: 220 (1916)
Type species: Cavimalum indicum Yoshim. Doi et al., Type species: Diploöspora rosea Grove, J. Bot., Lond. 54:
Bull. natn. Sci. Mus., Tokyo, B 3(1): 24 (1977) 220 (1916)
Chamaeleomyces Sigler, J. Clin. Microbiol. 48(9): 3186 Dussiella Pat., Bull. Soc. mycol. Fr. 6: 107 (1890)
(2010) Type species: Dussiella tuberiformis (Berk. & Ravenel)
Type species: Chamaeleomyces granulomatis Sigler, J. Pat., Bull. Soc. mycol. Fr. 4(3): 106 (1888)
Clin. Microbiol. 48(9): 3188 (2010) Ephelis Fr., Summa veg. Scand., Section Post.
Collarina Giraldo et al., in Crous et al., Persoonia, Mol. (Stockholm): 370 (1849)
Phyl. Evol. Fungi 33: 271 (2014) Type species: Ephelis mexicana Fr., Summa veg. Scand.,
Type species: Collarina aurantiaca Giraldo et al., in Crous Section Post. (Stockholm): 370 (1849)
et al., Persoonia, Mol. Phyl. Evol. Fungi 33: 271 (2014) Epichloë (Fr.) Tul. & C. Tul., Select. fung. carpol. (Paris) 3:
Conoideocrella D. Johnson et al., Mycol. Res. 113(3): 286 24 (1865)
(2009) Type species: Epichloë typhina (Pers.) Tul. & C. Tul.,
Type species: Conoideocrella luteorostrata (Zimm.) D. Select. fung. carpol. (Paris) 3: 24 (1865)
Johnson et al., Mycol. Res. 113(3): 286 (2009) Epicrea Petr., Sydowia 4(1–6): 325 (1950)
 Fungal Diversity

Type species: Epicrea insignis Petr., Sydowia 4(1–6): 325 Type species: Neobarya parasitica (Fuckel) Lowen,
(1950) in Eriksson & Hawksworth, Syst. Ascom. 5(1): 121
Helminthascus Tranzschel, Trudy S. Petersb. Obschch. (1986)
Est. Otd. Bot. 28: 331 (1898) Neoclaviceps J.F. White et al., Mycologia 93(1): 91 (2001)
Type species: Helminthascus arachnophthorus Type species: Neoclaviceps monostipa J.F. White et al., in
Tranzschel [as ‘arachnophthora’], Trudy S. Petersb. Sullivan et al., Mycologia 93(1): 92 (2001)
Obschch. Est. Otd. Bot. 28: 331 (1898) Neocordyceps Kobayasi, J. Jap. Bot. 59(6): 187 (1984)
Heteroepichloë E. Tanaka, C. Tanaka, Gafur & Tsuda, Type species: Neocordyceps kohyasanensis Kobayasi, J.
Mycoscience 43(2): 92 (2002) Jap. Bot. 59(6): 187 (1984)
Type species: Heteroepichloë bambusae (Pat.) E. Tanaka Neotyphodium Glenn, C.W. Bacon & Hanlin, Mycologia
et al., Mycoscience 43(2): 92 (2002) 88(3): 377 (1996)
Hypocrella Sacc., Michelia 1(no. 3): 322 (1878) Type species: Neotyphodium coenophialum (Morgan-
Type species: Hypocrella discoidea (Berk. & Broome) Jones & W. Gams) Glenn et al., in Glenn et al., Mycologia
Sacc., Michelia 1(no. 3): 322 (1878) 88(3): 377 (1996)
Konradia Racib., Parasit. Alg. Pilze Java’s (Jakarta) 2: 15 Nigrocornus Ryley & Langdon, in Ryley, Mycology
(1900) Series (New York) 19: 266 (2003)
Type species: Konradia bambusina Racib., Parasit. Alg. Type species: Nigrocornus scleroticus (Pat.) Ryley,
Pilze Java’s (Jakarta) 2: 15 (1900) Mycology Series (New York) 19: 267 (2003)
Loculistroma F. Patt. & Charles, Bull. Bureau Plant Nomuraea Maubl., Bull. Soc. mycol. Fr. 19(3): 295 (1903)
Industry U.S. Dep. Agric. 171: 11 (1910) Type species: Nomuraea prasina Maubl., Bull. Soc.
Type species: Loculistroma bambusae F. Patt. et al., Bull. mycol. Fr. 19(3): 296 (1903)
U.S. Department of Agriculture, Bureau Plant Industry 171: Orbiocrella D. Johnson et al., Mycol. Res. 113(3): 286
11 (1910) (2009)
Metacordyceps G.H. Sung et al., in Sung et al., Stud. Type species: Orbiocrella petchii (Hywel-Jones) D.
Mycol. 57: 27 (2007) Johnson et al., Mycol. Res. 113(3): 287 (2009)
Type species: Metacordyceps taii Z.Q. Liang & A.Y. Liu, Parepichloë J.F. White & P.V. Reddy, Mycologia 90(2):
in Liang et al., Acta Mycol. Sin. 10(4): 257 (1991) 231 (1998)
Metarhiziopsis D.W. Li et al., Mycologia 100(3): 462 Type species: Parepichloë cinerea (Berk. & Broome) J.F.
(2008) White & P.V. Reddy, Mycologia 90(2): 231 (1998)
Type species: Metarhiziopsis microspora D.W. Li et al., Periglandula U. Steiner et al., in Steiner et al., Mycologia
Mycologia 100(3): 462 (2008) 103(5): 1137 (2011)
Metarhizium Sorokīn, Veg. Parasitenk. Mensch Tieren 2: Type species: Periglandula ipomoeae U. Steiner et al., in
268 (1879) et al., Mycologia 103(5): 1140 (2011)
Type species: Metarhizium anisopliae (Metschn.) Sorokīn, Pochonia Bat. & O.M. Fonseca, Publicações Inst. Micol.
Rastitel’nye parazity cheloveka i zhivotnykh kak prichina Recife 462: 4 (1965)
zarazhykh bolezneĭ (Petersburg) 2: 268 (1883) Type species: Pochonia humicola Bat. & O.M. Fonseca,
Metapochonia Kepler et al., in Kepler et al., Mycologia Publicações Inst. Micol. Recife 462: 5 (1965)
106(4): 820 (2014) Pseudogibellula Samson & H.C. Evans, Acta bot. neerl.
Type species: Metapochonia suchlasporia (W. Gams & 22(5): 524 (1973)
Dackman) Kepler et al., in Kepler et al., Mycologia 106(4): Type species: Pseudogibellula formicarum (Mains)
820 (2014) Samson & H.C. Evans, Acta bot. neerl. 22(5): 524 (1973)
Moelleriella Bres., Boll. Soc. bot. ital. 44: 292 (1897) Pseudomeria G.L. Barron, Can. J. Bot. 58(4): 443 (1980)
Type species: Moelleriella sulphurea (Bres.) Bres., in Type species: Pseudomeria mucosa G.L. Barron, Can. J.
Saccardo & Sydow, Syll. fung. (Abellini) 14(1): 626 (1899) Bot. 58(4): 443 (1980)
Mycomalus Möller, Bot. Mitt. Trop. 9: 300 (1901) Regiocrella P. Chaverri & K.T. Hodge, Mycologia 97(6):
Type species: Mycomalus bambusinus Möller, Bot. Mitt. 1232 (2006) [2005]
Trop. 9: 300 (1901) Type species: Regiocrella camerunensis P. Chaverri &
Myriogenospora G.F. Atk., Bull. Torrey bot. Club 21(5): H.C. Evans, in Chaverri et al., Mycologia 97(6): 1232
225 (1894) (2006) [2005]
Type species: Myriogenospora paspali G.F. Atk., Bull. Romanoa Thirum., R.C. Ist. Sup. Sanitä, (Rome) 17: 1326
Torrey bot. Club 21(5): 225 (1894) (1954)
Neobarya Lowen, in Eriksson & Hawksworth, Syst. Type species: Romanoa terricola Thirum., R.C. Ist. Sup.
Ascom. 5(1): 121 (1986) Sanitä, (Rome) 17(12): 1326 (1954)
Fungal Diversity

Rotiferophthora G.L. Barron, Can. J. Bot. 69(3): 495 (1991) 5. Ascospores, septa indistinct or lacking . . . . . . . . . . . . 9
Type species: Rotiferophthora globospora G.L. Barron, 6. Ascospores disarticulating . . . . . . . . . . . . . . . Balansia
Can. J. Bot. 69(3): 495 (1991) 6. Ascospores no disarticulating . . . . . . . . . . . . . . . . . . . 7
Samuelsia P. Chaverri & K.T. Hodge, in Chaverri et al., 7. Asci with 2–6 ascospores . . . . . . . . . . .Shimizuomyces
Stud. Mycol. 60: 59 (2008) 7. Asci with 8 ascospores. . . . . . . . . . . . . . . . . . . . . . . . 8
Type species: Samuelsia rufobrunnea P. Chaverri & K.T. 8. Perithecia superficial . . . . . . . . . . . . Tyrannicordyceps
Hodge, in Chaverri et al., Stud. Mycol. 60: 62 (2008) 8. Perithecia immersed . . . . . . . . . . . . . . . . Neocordyceps
Shimizuomyces Kobayasi, Bull. natn. Sci. Mus., Tokyo, B 9. Stipe pale brown to orange brown . . . . . . . . Claviceps
7(1): 1 (1981) 9. Stipe reddish-brown . . . . . . . . . . . . . . . . .Neoclaviceps
Type species: Shimizuomyces paradoxus Kobayasi [as 10. Host bamboo . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
‘paradoxa’], Bull. natn. Sci. Mus., Tokyo, B 7(1): 1 (1981) 10. Host grasses . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
Sphacelia Lév., Mém. Soc. Linn. Paris 5: 578 (1827) 11. Ascospores hyaline. . . . . . . . . . . . . . . . . . . . . . . . . 12
Type species: Sphacelia segetum Lév. Mém. Soc. Linn. 11. Ascospores some other colour . . . . . . . . . . . . . . . . 16
Paris 5: 578 (1827) 12. Ascospores multi-septate . . . . . . . . . . . . . . . . . . . . 13
Sphaerocordyceps Kobayasi, Bull. natn. Sci. Mus., Tokyo, 12. Ascospores irregularly septate . . . . . . . . .Cavimalum
B 7(1): 2 (1981) 13. Perithecia immersed, oval, pyriform or obpyriform . . 14
Type species: Sphaerocordyceps palustris (Berk. & 13. Perithecia superficial, cylindrical . . . . . . . . .Dussiella
Broome) Kobayasi, Bull. Natn. Sci. Mus., Tokyo, B 7(1): 2 14. Stromata on woody culm, light brown or dark
(1981) brown . . . . . . . . . . . . . . . . . . . . . . . . 15
Stereocrea Syd. & P. Syd., Annls mycol. 15(3/4): 216 (1917) 14. Stromata half encircling the leaves,
Type species: Stereocrea schizostachyi Syd. & P. Syd., black . . . . . . . . . . . . . . . . . Heteroepichloë
Annls mycol. 15(3/4): 216 (1917) 15. Ascospores disarticulating into part
Tyrannicordyceps Kepler & Spatafora, Index Fungorum spores . . . . . . . . . . . . . . . . . . Mycomalus
12: 1 (2012) 15. Ascospores lacking part spores. . . . . Aciculosporium
Type species: Tyrannicordyceps fratricida (Tanda & 16. Ascospores filiform, yellowish, part spores, brown to
Kobayasi) Kepler & Spatafora, in Kepler et al., Index black . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Konradia
Fungorum 12: 1 (2012) 16. Ascospores fusiform, olivaceous, 3 septate, lacking part
Ustilaginoidea Bref., Unters. Gesammtgeb. Mykol. spores. . . . . . . . . . . . . . . . . . . . . . . . . . Loculistroma
(Liepzig) 12: 194 (1895) 17. Stromata developing on inflorescences of grasses. . 18
Type species: Ustilaginoidea oryzae (Pat.) Bref., Unters. 17. Stromata developing on leaves, tillers or culms of
Gesammtgeb. Mykol. (Liepzig) 12: 194 (1895) grasses. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
Accepted name in key 18. Part-spores unicellular, curved, needle-shaped,
Albomyces I. Miyake ex I. Hino, Trans. Mycol. Soc. Japan narrower at one end . . . . . . . . . . . . . . . Atkinsonella
3: 113 (1962) 18. Part-spores 1-septate, not curved, elongate
Type species: Albomyces take I. Hino, Trans. Mycol. Soc. cylindrical . . . . . . . . . . . . . . . . . . . . . . . . . Balansia
Japan 3: 112 (1962) 19. Ascospores breaking into part spores . . . . . . . . . . . 20
Aschersonia Mont., Annls Sci. Nat., Bot., sér. 3 10: 121 19. Ascospores not breaking into part spores . . . . . . . . 21
(1848) 20. Part spores 1-septate. . . . . . . . . . . . . . . . . . . Balansia
Type species: Aschersonia tahitensis Mont., Annls Sci. 20. Part spores aseptate . . . . . . . . . . . . . Myriogenospora
Nat., Bot., sér. 3 10: 122 (1848) 21. Perithecia flask or obpyriform . . . . . . . . Nigrocornus
22. With conidiomata, with conidia . . . . . . . . . . . . . . . 23
22. With synnemata, lacking conidia . . . . . .Periglandula
Key to genera of Clavicipitaceae 23. Conidia holoblastic . . . . . . . . . . . . . . . . . . . . . . . . 24
23. Conidia not holoblastic. . . . . . . . . . . . . . . . . . . . . . 25
1. Host is a plant . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 24. Conidiogenous cells, arising as integrated or discrete,
1. Host is not a plant . . . . . . . . . . . . . . . . . . . . . . . . . . 29 cylindrical to narrowly obpyriform Corallocytostroma
2. Sexual morph produced . . . . . . . . . . . . . . . . . . . . . . . 3 24. Conidiogenous cells arising as a palisade layer, slender,
2. Asexual morph produced . . . . . . . . . . . . . . . . . . . . . 22 filiform. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ephelis
3. Stromata stalked. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 25. Conidia 0–1-septate . . . . . . . . . . . . . . . . . . . . . . . . 26
3. Stromata lacking stalks. . . . . . . . . . . . . . . . . . . . . . . 10 25. Conidia 2-septate . . . . . . . . . . . . . . . . . . . Albomyces
4. Ascospores yellow to green . . . . . . . . . . . . Stereocrea 26. Conidia 1-septate, later swelling. . . . . . . . . .Dussiella
4. Ascospores hyaline or some other colour . . . . . . . . . . 5 26. Conidia 1-celled, not swelling . . . . . . . . . . . . . . . . 27
5. Ascospores with distinct septa . . . . . . . . . . . . . . . . . . 6 27. Conidia verruculose . . . . . . . . . . . . . . Ustilaginoidea
 Fungal Diversity

27. Conidia smooth-walled . . . . . . . . . . . . . . . . . . . . . 28 47. Conidiogenous cells or conidia without slime. . . . . 50

28. Conidia catenulate . . . . . . . . . . . . . . . Metarhiziopsis 48. Conidia fusiform or fusoid, generally with acute
28. Conidia discrete . . . . . . . . . . . . . . . . . Neotyphodium ends . . . . . . . . . . . . . . . . . . . . . . . . . 49
29. Host insects, nematodes, rotifers, protozoans or 48. Conidia ovoid to ellipsoid, can be reniform or
soil. . . . . . . . . . . . . . . . . . . . . . . . . . 30 apiculate . . . . . . . . . . . . . . . . . . Sphacelia
29. Host Chameleon or other fungi . . . . . . . . . . . . . . . 52 49. Conidiogenous cells flask-shaped, formed in a thick
30. Sexual morph produced . . . . . . . . . . . . . . . . . . . . . 31 compact palisade . . . . . . . . . . . . . . . . . . . Hypocrella
30. Asexual morph produced . . . . . . . . . . . . . . . . . . . . 40 49. Conidiogenous cells filiform, can branch at acute an-
31. Stroma with stalk . . . . . . . . . . . . . . . . . . . . . . . . . . 32 gles, not formed in a thick compact
31. Stroma lacking stalk. . . . . . . . . . . . . . . . . . . . . . . . 34 palisade . . . . . . . . . . . . . . . . . . . . . . . . . Aschersonia
32. Asci cylindrical, no appendant . . . . . . . . . . . . . . . . 33 50. Phialides flask-shaped . . . . . . . . . . . . . . . . . . . . . . 51
32. Asci flexuous, filled with moniliform 50. Phialides cylindrical . . . . . . . . . . . . . . . . . . . . . . . . 52
strings . . . . . . . . . . . . . . . . . . . . .Sphaerocordyceps 51. Conidiophores irregularly branched . . . . . Regiocrella
33. Perithecia superficial on the stroma or clustered at the 51. Conidiophores regularly branched . . . . . Moelleriella
apex. . . . . . . . . . . . . . . . . . . . . . . . Tyrannicordyceps 52. Phialides with a brownish funnel-shaped
33. Perithecia completely or half immersed in the collarette . . . . . . . . . . . . . . . . . . . . . . . . . Collarina
stroma. . . . . . . . . . . . . . . . . . . . . . . . Metacordyceps 52. Phialides without a brownish funnel-shaped
34. Perithecia flask-shaped or conical. . . . . . . . . . . . . . 35 collarette. . . . . . . . . . . . . . . . . . . . . . . . . .Samuelsia
34. Perithecia obpyriform, subglobose or 53. Host fungus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 54
subcylindrical . . . . . . . . . . . . . . . . . . . 36 53. Host chameleons. . . . . . . . . . . . . . . Chamaeleomyces
35. Stromata superficial from the host, discrete, variously 54. Host Claviceps . . . . . . . . . . . . . . . . . . . . . . . .Epicrea
coloured. . . . . . . . . . . . . . . . . . . . . . . Conoideocrella 54. Host Hypocrella chusqueae . . . . . . . . . . . . Neobarya
35. Stromata surrounding host, crowd, ring-like,
ochraceous . . . . . . . . . . . . . . . . . . . . . . . Orbiocrella Clypeosphaeriaceae G. Winter [as ‘Clypeosphaerieae’],
36. Perithecia obpyriform or subglobose . . . . . . . . . . . 37 Rabenh. Krypt.-Fl., Edn 2 (Leipzig) 1.2: 554 (1886)
36. Perithecia subcylindrical. . . . . . . . . . . . . . . .Dussiella Facesoffungi number: FoF 01776
37. Stromata tissue changing colour in 3 % KOH . . . . 38 Hemibiotrophic or saprobic on woody or herbaceous
37. Stromata tissue not changing in 3 % plants. Sexual morph: Pseudoclypeus comprising both
KOH . . . . . . . . . . . . . . . . . . . Moelleriella host and fungal tissues, black. Ascomata immersed to
38. Stromata tissue changing colour in 3 % KOH to brown, erumpent, rarely superficial, solitary or aggregated, glo-
reddish or dark reddish . . . . . . . . . . . . . . . . . . . . . . 39 bose to subglobose, coriaceous, brown to black,
38. Stromata tissue becoming purple in 3 % ostiolate, papillate. Papilla short, narrow, internally
KOH . . . . . . . . . . . . . . . . . . . . . . . . . . . Regiocrella lined with hyaline, filamentous periphyses. Peridium
39. Stroma surface minutely pulverulent or comprising dark brown to light brown, thick-walled
pruinose. . . . . . . . . . . . . . . . . . . . . . . . . Hypocrella cells of textura angularis, inwardly hyaline.
39. Stroma surface smooth or slightly pruinose . .Samuelsia Hamathecium comprising numerous, hypha-like, septate,
40. Conidia green in mass . . . . . . . . . . . . . .Metarhizium flexuose, paraphyses, embedded in a gelatinous matrix.
40. Conidia some other colour in mass. . . . . . . . . . . . . 41 Asci 8-spored, unitunicate, cylindrical to broadly cylin-
41. Conidia with adhesive hapteron . . . . . . .Pseudomeria drical, pedicellate, with a wedge-shaped, J-, or J+ apical
41. Conidia without adhesive hapteron. . . . . . . . . . . . . 42 ring. Ascospores uniseriate to biseriate, hyaline to
42. Conidia 1-multi-septate . . . . . . . . . . . . . . . . . . . . . 43 brown, ellipsoidal to fusiform, sometimes oval, straight
42. Conidia 1-celled. . . . . . . . . . . . . . . . . . . . . . . . . . . 45 or curved, unicellular or septate, wall smooth or
43. Conidia, 1-septate, not dictyochlamydospores. . Dussiella ornamented or striate, sometimes with sheaths, append-
43. Conidia dictyochlamydospores. . . . . . . . . . . . . . . . 44 ages, rarely with germ slits or germ pores. Asexual
44. Phialides smooth . . . . . . . . . . . . . . . . Metapochonia morph: Undetermined.
44. Phialides aculeate. . . . . . . . . . . . . . . . . . . . Pochonia Type: Clypeosphaeria Fuckel
45. Conidia adhering in chains . . . . . . . . . . . . . . . . . . . 46 Notes: The family Clypeosphaeriaceae was introduced by
45. Conidia not as above . . . . . . . . . . . . . . . . . . . . . . . 47 Winter (1887) to include Anthostomella, Clypeosphaeria,
46. Conidiogenous cells flask-shaped, with short, narrow Hypospila, Linospora and Trabutia. Barr (1989) revived the
neck. . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nomuraea family Clypeosphaeriaceae and included Apiorhynchostoma,
46. Conidiogenous cells cylindrical to clavate, without a Clypeosphaeria, Endoxyla, Melomastia, Pseudovalsaria,
neck. . . . . . . . . . . . . . . . . . . . . . . . . . . .Metarhizium Saccardoella and Urosporella, which are related and
47. Conidiogenous cells or conidia with slime . . . . . . . 48 morphologically similar to the family Amphisphaeriaceae.
Fungal Diversity

However, Barr (1989) revisited the family and excluded all Brunneiapiospora K.D. Hyde et al., Sydowia 50(1): 40
genera and retained Clypeosphaeria as the type of this (1998)
monotypic family. Hawksworth et al. (1995) treated this fam- Type species: Brunneiapiospora javensis K.D. Hyde et al.,
ily with Apiorhynchostoma, Ceratostomella, Clypeosphaeria, Sydowia 50(1): 50 (1998)
Crassoascus, Duradens, Frondicola, Jobellisia, Melomastia Crassoascus Checa et al., Mycotaxon 46: 300 (1993)
and Pseudovalsaria (Kang et al. 1999c; Smith et al. 2003). Type species: Crassoascus fusisporus Checa et al.,
Lumbsch and Huhndorf (2010) accepted only Mycotaxon 46: 301 (1993)
Apiorhynchostoma, Aquasphaeria, Brunneiapiospora, Palmomyces K.D. Hyde et al. Sydowia 50(1): 59 (1998)
Clypeosphaeria, Crassoascus, Duradens, Palmomyces and Type species: Palmomyces montaneus K.D. Hyde et al.,
Pseudovalsaria in the family. Maharachchikumbura et al. Sydowia 50(1): 60 (1998)
(2015) and Senanayake et al. (2015) accepted Apioclypea,
Aquasphaeria, Brunneiapiospora, Clypeosphaeria,
Crassoascus and Palmomyces as genera of Key to genera of Clypeosphaeriaceae
Clypeosphaeriaceae. In this paper we exclude Aquasphaeria
which we place in Sordariomycetes genera incertae sedis. 1. Ascomata erumpent . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
1. Ascomata immersed, clypeus, epiphyllous . . . . . . . . . 2
Clypeosphaeria Fuckel, Jb. nassau. Ver. Naturk. 23–24: 2. Globose or sphaeroid, J+, fusoid asymmetric, clear
117 (1870) [1869–70] brown ascospore, 5-celled, gattulate, apiculate
Facesoffungi number: FoF 02104; Fig. 41 ends . . . . . . . . . . . . . . . . . . . . . . . . . . . . Crassoascus
Saprobic bark of dead wood. Sexual morph: Clypeus 2. Globose or subglobose, overlapping uniseriate,
dome-shaped, well-developed, black. Ascomata 350– apiosporous, with a mucilaginous sheath. . . . . . . . . . 3
400 μm high, 380–450 μm wide, immersed under the clypeus, 3. Clypeus, J+ or J-, hyaline ascospore. . . . . . Apioclypea
solitary, scattered, globose to subglobose, coriaceous, black. 3. Subglobose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
Ostiole central, papillate, papilla short, narrow, internally lined 4. Subglobose or ovoid, clypeate, J+, ellipsoidal, brown
with hyaline, filamentous periphyses. Peridium 20–35 μm ascospore, unicellular, 3–5 pseudosepta, rarely
wide, comprising outer, brown, elongated, flat, cells of textura apiosporous, lacking a mucilaginous
angularis and inner, hyaline, elongated, flat cells of textura sheath . . . . . . . . . . . . . . . . . . . . . . . . .Clypeosphaeria
angularis. Hamathecium comprising numerous, hypha-like, 4. Subglobose; Epiphyllous or Clypeus . . . . . . . . . . . . . 5
aseptate, unbranched, flexuose, paraphyses, tapering towards 5. Clypeus, J+ or J-, hyaline to light brown asco-
the apex. Asci 8-spored, unitunicate, cylindrical, short pedicel- spore….………........ Brunneiapiospora
late, apex rounded, with a J+ apical ring. Ascospores 5. Epiphyllous, J-, biseriate, hyaline ascospore, obcalvate,
uniseriate, dark brown, ellipsoidal to fusiform, unicellular, 1-septate . . . . . . . . . . . . . . . . . . . . . . . . . Palmomyces
straight or slightly curved, not constricted at the septa,
smooth-walled. Asexual morph: Undetermined. Coniocessiaceae Asgari & Zare, Mycol. Progr. 10(2): 195
Type species: Clypeosphaeria notarisii Fuckel, Jb. nassau. (2011)
Ver. Naturk. 23–24: 117 (1870) Facesoffungi number: FoF 01296
Notes: Clypeosphaeria notarisii was introduced by Fuckel Saprobic on grasses, soil and dung. Sexual morph:
and is discussed by Kang et al. (1999c). Barr (1989) men- Ascomata small, less than 200 μm diam, superficial,
tioned that Clypeosphaeria has been synonymized under sev- subglobose to pyriform, glabrous or pilose, ostiolate, com-
eral genera. Hyde et al. (1998) treated Clypeosphaeria as a monly with hypha-like ostiolar projections. Peridium thin,
monotypic genus, excluding all other species. Currently 48 membranaceous, translucent or sometimes opaque, outer layer
species have been listed under the genus Clypeosphaeria in comprising pale-brown to dark brown cells of textura
Index Fungorum (2016). Clypeosphaeria is characterized by intricata. Hamathecium comprising septate, tapering, hy-
0–5-pseudoseptate ascospores, without germ pores (Kang phae-like, thin-walled, filamentous, branched. paraphyses,
et al. 1999c). abundant or few. Asci 4–spored, unitunicate, cylindrical to
subcylindrical, pedicellate, without an apical ring.
Ascospores uniseriate, dark brown to black, ellipsoidal, with
narrowly rounded ends, smooth–walled, one-celled, with
Other genera included germ-slit, full length, straight. Asexual morph:
Hyphomycetous. nodulisporium-like: Conidiophores
Apioclypea K.D. Hyde, J. Linn. Soc., Bot. 116(4): 316 (1994) micronematous to macronematous, simple or branched,
Type species: Apioclypea livistonae K.D. Hyde, J. Linn. smooth-walled or verrucose, hyaline. Conidiogenous cells in-
Soc., Bot. 116(4): 317 (1994) tegrated, terminal, discrete, elongating sympodially, with
 Fungal Diversity

Fig. 41 Clypeosphaeria
notarisii (Material examined:
GERMANY, Forest of Oestriche,
December 1823, Fuckel,
G00127177, holotype) a
Herbarium package b Herbarium
material c Ascomata on the
surface of host d Section of
ascoma e–g Asci with ascospores
h Peridium i, j Ascus apex with
J+, subapical ring k–n
Ascospores. Scale bars:
c = 500 μm, d = 50 μm, e–
g = 20 μm, h, i–n = 5 μm

persistent, conspicuous denticles, hyaline. Conidia globose, family is unique in the order Xylariales in having a hyaline
subglobose to pyriform, smooth-walled or verruculose, with asexual morph with polyblastic conidiogenesis
rounded apex, hyaline, attenuated and truncated base and a (nodulisporium-like), contrasting with the generally
more or less distinct projection at the point of attachment to pigmented nodulisporium-like asexual morphs in Xylariales
the conidiogenous cells. (Asgari and Zare 2011b). Members of Coniocessiaceae can be
Type: Coniocessia Dania García et al. distinguished from Xylariaceae in possessing astromatic
Notes: The family Coniocessiaceae was introduced by ascomata, and asci without apical structures (Asgari and
Asgari and Zare (2011b) to accommodate the genus Zare 2011b). Asgari and Zare (2011b) introduced four new
Coniocessia with a nodulisporium-like asexual morph. The species to Coniocessia and provided molecular data indicating
Fungal Diversity

this was a distinct lineage of Xylariales, but closely related to subulate or indistinguishable from a normal hyphal cell,
the family Diatrypaceae according to their phylogenetic anal- collarettes present, but usually hard to distinguish, only occa-
ysis. Thus they introduced the new family, which confirms its sionally somewhat flared. Conidia accumulating near the
distinctness as a family and place it in the order Xylariales. point of formation (in chains), hyaline, orange or pink in mass,
elliptical to oblong-elliptical to reniform, 1-celled, smooth-
Coniocessia Dania García et al., in García et al., Mycol. walled.
Res. 110(11): 1284 (2006) Type: Coniochaeta (Sacc.) Cooke
Facesoffungi number: FoF 01297; Figs 42, 43 Notes: The family Coniochaetaceae was introduced by
Type species: Coniocessia nodulisporioides (D. Hawksw.) Malloch and Cain (1971) to accommodate Coniochaeta and
Dania García et al., in García et al., Mycol. Res. 110(11): 1285 Coniochaetidium, and is typified by Coniochaeta.
(2006) Coniochaetaceae is a single family within the order
Notes: García et al. (2006) introduced the genus Coniochaetales in the subclass Sordariomycetidae (Huhndorf
Coniocessia to accommodate the type species, et al. 2004a; García et al. 2006). Two genera, Coniochaeta and
C. nodulisporioides Dania García et al., by synonymising Barrina are presently placed in the family based on morpho-
Coniochaeta nodulisporioides D. Hawksw., which was placed logical and phylogenetic analyses (Ramaley 1997; Huhndorf
in Xylariales genera incertae sedis, based on morphological et al. 2004a; García et al. 2006; Kirk et al. 2008; Lumbsch
and sequences data of SSU and LSU rDNA genes (García and Huhndorf 2010; Wijayawardene et al. 2012; Khan et al.
et al. 2006). 2013; Miller et al. 2014; Maharachchikumbura et al. 2015).
Recently, Asgari and Zare (2011b) introduced four new spe- Coniochaetaceae is characterized by 4- to 8-, or multi-spored
cies (C. anandra Asgari & Zare, C. cruciformis Asgari & Zare, asci (Asgari and Zare 2011b), ascospores with longitudinal
C. maxima Asgari & Zare and C. minima Asgari & Zare) in the germ slit (Asgari et al. 2007; Ivanová and Bernadovičová
genus Coniocessia based on morphological and molecular data 2013), and an asexual morph with phialidic or polyblastic
of ITS and LSU rDNA. All species of Coniocessia clustered conidiogenous cells (Asgari and Zare 2006; Asgari et al.
into a single monophyletic clade (Asgari and Zare 2011b). 2007). This family occurs on various substrates and media:
such as plants (wood, bark, leaves, leaf litter), animal dung, soil
Coniochaetaceae Malloch & Cain, Can. J. Bot. 49: 878 and strongly acidic water with high heavy metal concentrations
(1971) and humans (Damm et al. 2010; Ivanová and Bernadovičová
Facesoffungi number: FoF 01332 2013; Khan et al. 2013).
Saprobic on dung, plants litter or in soil, water, pathogenic
on plants or pathogens of immune-compromised humans. Coniochaeta (Sacc.) Cooke, Grevillea 16 (no. 77): 16
Sexual morph: Ascomata perithecial or cleistotheciod, soli- (1887)
tary to gregarious, superficial, semi-immersed or immersed, Facesoffungi number: FoF 01333, Figs 44, 45
subglobose to globose, or pyriform, dark brown to black, gla- Saprobic on plants litter, dung or in soil, water, pathogenic
brous or hairy brown to black, ostiolate or lacking ostioles. on plants or pathogens of immunocompromised humans.
Ostioles periphysate, with or without setae. Peridium Sexual morph: Ascomata perithecial or cleistotheciod, soli-
membranaceous to pseudoparenchymatous, rarely coriaceous; tary to gregarious, superficial or semi-immersed, pyriform and
composed of several layers of cells of textura angularis or ostiolate or globose and non-ostiolate, setose, hairy or gla-
textura intricata, or less frequently cephalothecoid. brous, dark brown to black or slightly pigmented. Peridium
Hamathecium comprising numerous, filiform, simple, septate, membranaceous to pseudoparenchymatous, rarely coriaceous;
evanescent paraphyses. Asci (4-), 8- to multi-spored, composed of several layers with the cells of textura angularis,
unitunicate, cylindrical to fusoid or clavate, globose to textura intricata, or less frequently cephalothecoid.
subglobose, thin-walled, evanescent, short pedicellate, with a Hamathecium paraphysate or absent. Paraphyses when pres-
truncate to rounded apex, with a non-amyloid apical ring. ent numerous, hyaline, filiform, simple, septate, and evanes-
Ascospores 1-seriate or irregularly arranged, hyaline, brown cent. Asci 4-, 8- to multi-spored, unitunicate, cylindrical, cla-
to dark brown, olive-greenish to dark olivaceous or black, vate, sub-globose or globose, evanescent, short pedicellate,
ellipsoid to fusiform, broadly ellipsoidal to globose, lenticular with a truncate to rounded apex, usually with a conspicuous
or cruciform, with rounded to apiculate ends, flattened on one to indistinct, non-amyloid, apical ring, thin-walled.
or both sides; 1-celled, with or without a germ slit, smooth Ascospores 1-seriate or irregularly arranged, narrowly ellip-
walled or pitted. Asexual morph: Hyphomycetous. Colonies soid to fusoid, broadly ellipsoidal to globose, lenticular or
are frequently characterized by pink or orange and a yeast-like cruciform, with rounded to apiculate ends, flattened on one
appearance. Conidiophores hyphae variable, frequently close- or both sides; hyaline, brown to dark brown, olive-greenish to
ly septate. Conidiogenous cells phialidic, polyblastic, of vari- dark olivaceous or black, 1-celled, with a cleft germ, smooth-
ous size and morphology, phialides somewhat ampulliform, walled or pitted. Asexual morph: Hyphomycetous.
 Fungal Diversity

Fig. 42 Coniocessia spp. (Material examined: a-e IRAN, West IRAN, Parsabad, Ardabil, on wheat seeds, B. Asgari. IRAN 14376F,
Azerbaijan, Miandoab, on wheat seeds, B. Asgari. IRAN 14375F, holotype of C. minima; u–y JORDAN, from soil, A. H. Moubasher,
holotype of C. anandra; f–g IRAN, Ardabil Bilesavar, on wheat straw, 1976, IMI 204247, holotype of C. nodulisporioides) a, f, k, p, u
B. Asgari, IRAN 14374F, holotype of C. cruciformis; k–o SPAIN, Ascomata b, g, l, q, v Peridium c, h, m, r, w Paraphyses d, I, n, s, x
Caceras, on goat dung, P. Blaser, May 1971. Received from CBS as Asci e, j, o, t, y Ascus. Scale bars: a, f, k, p, u = 50 μm, b–e, g–j, l–o, q–t,
C. nodulisporioides. IRAN 14377F, holotype of C. maxima; p–t v–y = 10 μm
Fungal Diversity

Fig. 43 Asexual morph of

Coniocessia spp. (Material
examined: a-c IRAN, Ardabil
Bilesavar, on wheat straw, B.
Asgari, IRAN 14374F, holotype
of C. cruciformis; d–f SPAIN,
Caceras, on goat dung, P. Blaser,
May 1971. Received from CBS
as C. nodulisporioides. IRAN
14377F, holotype of C. maxima;
g–i IRAN, Parsabad, Ardabil, on
wheat seeds, B. Asgari. IRAN
14376F, holotype of C. minima;
j–l JORDAN, from soil, A. H.
Moubasher, 1976, IMI 204247,
holotype of C. nodulisporioides).
a, d, g, j Asexual morph b, e, h, k
Conidiophores and
conidiogenous cells c, f, i, l
conidia. Scale bars: a, d,
g = 50 μm, b–c, e-f, h-i, k–
l = 10 μm, j = 20 μm

Belonging to Lecythophora, Verticillium, Paecilomyces, and Notes: The genus Coniochaeta (Sacc.) is typified by
Cladobotryum. Colonies are frequently characterized by pink C. ligniaria, introduced by Greville (1823–1824) as
or orange, colony surface more or less smooth, slimy, usually Sphaeria ligniaria. Saccardo (1882) treated Coniochaeta as
without aerial hyphae, occasionally with short tufts of white a subgenus under Rosellinia and it was raised to generic rank
hyphae in advancing zone. Conidiophores hyphae variable, by Cooke in 1887 (Greville 1823–1824; Saccardo 1882;
frequently closely septate. Conidiogenous cells phialidic, Asgari et al. 2007; Asgari and Zare 2011b). Asexual morphs
phialides somewhat ampulliform, subulate or indistinguish- of Coniochaeta are reported to belong to cladobotryum-like,
able from a normal hyphal cell, terminal or lateral on hyphae, Lecythophora, nodulisporium-like and Paecilomyces, which
or intercalary hyphal cells producing phialidic openings di- have phialidic and polyblastic conidiogenesis (Barr 1990;
rectly or on a short extension; collarettes present but usually Asgari and Zare 2006; Asgari and Zare 2011b). Coniochaeta
hard to distinguish, only occasionally somewhat flared. has 86 epithets listed in Index Fungorum (2016). Coniomela,
Conidia elliptical to oblong-elliptical to reniform, hyaline, or- Cucurbitariella, Cucurbitula, Germslitospora, Pleosporopsis,
ange or pinky in masse, 1-celled, smooth-walled. Sphaerodermatella, Sphaerodermella, Sphaeropyxis,
Type species: Coniochaeta ligniaria (Grev.) Cooke, Coniochaetidium, Ephemeroascus and Poroconichaeta are
Grevillea 16(no. 77): 16 (1887) listed as synonyms of Coniochaeta (Huhndorf et al. 2004a;
 Fungal Diversity

Fig. 44 Coniochaeta ligniaria (Material examined: SWEDEN, material b Host (hare dung) c Ascomata on the host d Vertical section of
Småland, Döderhult par., 2 km ENE of Bohult, on fire-place in ascoma e Ostioles with setae f–g Blunt setae h–k Asci l Asci and
coniferous forest, on hare dung (Lepus) in moist chamber, Stockholm, paraphyses m–o Ascospores. Scale bars: c = 100 μm, d = 30 μm, e–
leg Nils Lundqvist, 30 May 1985, S F139191 (lgt 15438-g) a Herbarium n = 10 μm
Fungal Diversity

Fig. 45 Coniochaeta ligniaria (redrawn from Weber and Begerow 2002) a–c Conidiophores d Conidiogenous cell e Conidia. Scale bars: a–e = 10 μm

García et al. 2006; Kirk et al. 2008; Lumbsch and Huhndorf 1. Ascomata superficial or semi-immersed, ascospores
2010; Wijayawardene et al. 2012; Khan et al. 2013; Miller coloured . . . . . . . . . . . . . . . . . . . . . . . . . Coniochaeta
et al. 2014; Maharachchikumbura et al. 2015). Previous phy-
logenetic studies have linked the type species Coniochaeta Cordanaceae Nann., Repert. mic. uomo: 498 (1934)
ligniaria and Lecythophora lignicola ITS, LSU, Actin and Facesoffungi number: FoF 01673
Beta-tubulin molecular data, as well as previous phylogenetic Saprobic or pathogenic on wood and leaves of various
studies (Weber et al. 2002; Huhndorf et al. 2004a; García et al. shrubs, deciduous and evergreen trees, conifers, bamboo and
2006; Perdomo et al. 2013a, b). Khan et al. (2013) resolved grasses in terrestrial and rarely in freshwater habitats. Sexual
the phylogenetic placement of Lecythophora species and syn- morph: Ascomata perithecial, solitary or gregarious, superfi-
onymized the genus to Coniochaeta. cial, globose to ovoid, sometimes with basal stroma, setose or
glabrous, ostiolate, papillate, with periphyses. Peridium com-
posed of 3–5 layers of cells of textura prismatica.
Hamathecium comprising simple, unbranched, septate, hya-
Other genus included
line paraphyses. Asci 8-spored, unitunicate, cylindrical, short
pedicellate, with or without an inconspicuous, non-amyloid,
Barrina A.W. Ramaley, Mycologia 89(6): 962 (1997)
apical ring. Ascospores uniseriate, ellipsoid to fusiform, 1-
Type species: Barrina polyspora A.W. Ramaley,
septate, light brown to brown, sometimes with pores at the
Mycologia 89(6): 963 (1997)
ends. Asexual morph: Hyphomycetous. Colonies effuse,
dark brown to black. Conidiophores erect, mononematous,
macronematous, brown, branched or unbranched, with a fer-
Key to genera of Coniochaetaceae tile apex. Conidiogenous cells polyblastic, denticulate.
Conidia brown to dark brown, obovate or ellipsoidal, 1-sep-
1. Ascomata immersed, ascospores hyaline. . . . . . . .Barrina tate, smooth, with or without germ pores at the end of the
 Fungal Diversity

conidia (description modified from Hughes (1955), Müller Notes: The genus Cordana was introduced by Preuss
and Samuels (1982) and Hernández-Restrepo et al. (2014)). (1851), while describing the species Cordana polyseptata
Type: Cordana Preuss Preuss, C. pauciseptata Preuss and C. pedunculata Preuss.
Notes: The family Cordanaceae was established by Later a fourth species C. parvispora Preuss was also described
Nannfeldt with Cordana as the type genus. Based on (Preuss 1852), but in both publications, a type was not desig-
molecular data, Hernández-Restrepo et al. (2015a) intro- nated. Saccardo (1877) retained only C. pauciseptata in
duced the order Cordanales to accommodate this family, Cordana selecting it as the lectotype and classifying the
which is easily distinguished from the species of its other three Cordana species under Acrothecium. Kuntze
sister order Coniochaetales (Coniochaetaceae), in hav- (1891) disagreed with this classification and renamed the ge-
ing a basal stroma, ascospores lacking germ slits and nus as Preussiaster in honour of its first author. Cordana
polyblastic asexual morphs (Hernández-Restrepo et al. pauciseptata was reclassified as Preussiaster pauciseptatus
2015a). The family contains a single genus thus only (Preuss) O. Kuntze and the three species placed by Saccardo
the family description is given. (1877) under Acrothecium were also added (Kuntze 1891).
However, Hughes (1955) stated that any three of the speci-
Cordana Preuss, Linnaea 24: 129 (1851) mens originally described by Preuss (1851) should be the
Facesoffungi number: FoF 01674; Fig. 46 lectotype. After studying the type material of Cordana

Fig. 46 Cordana pauciseptata (redrawn from Hughes 1955) a, b the first species reported to have a Cordana asexual morph (redrawn from
Conidiophores with developing conidia on MEA (DAOM 33990) c Müller and Samuels 1982) g Vertical section of ascoma h Asci with
Conidiophore with proliferated conidia on PDA (DAOM 28664) d, e ascospores i Non amyloid apical ring of asci with a mature ascospore j
Hypha bearing chlamydospores on MEA (DAOM 33990) f Mature Ascospores. Scale bars: a, b, e–j = 10 μm, c, d = 20 μm
conidia (DAOM 33990). Sexual morph Porosphaerella cordanophora,
Fungal Diversity

pauciseptata, Hughes (1955) re-described and illustrated it as eleven genera (including Ascopolyporus, Cordyceps,
Brachysporium polyseptatum (= Brachysporium bloxami), Hyperdermium and Torrubiella Beauveria, Engyodontium,
which resulted in C. pauciseptata being an older epithet for Isaria, Lecanicillium, mariannaea-like, Microhilum and
the species. However, this type material was reported to be Simplicillium) to this family, while currently Index
lost and an epitype was designated by Hernández-Restrepo Fungorum (2016) lists 14 genera.
et al. (2014). Akanthomyces was first introduced by Lebert (1858) and
Cordana species are characterized by brown, septate co- later revised by Mains (1950). Most of the known
nidiophores with swollen conidiogenous zones, terminal and Akanthomyces species are parasites of insects or spiders, ex-
intercalary conidiogenous cells, and pale brown to brown, 1- cept Akanthomyces johnsonii (Massee) Vincent et al., which
septate or aseptate conidia (Hernández-Restrepo et al. 2015a). was isolated from leaves and stems (Hsieh et al. 1997; Huang
Müller and Samuels (1982) linked Porosphaerella et al. 1999). Gibellula was established by Cavara (1894) be-
cordanophora to Cordana pauciseptata, the type species of cause of its unusual conidiogenous cells. Since then, a number
Cordana using culture studies. Romero and Samuels (1991) of species have been described from spiders (Petch 1932a;
published a second species in Porosphaerella (P. setosa A.I. Mains 1950; Samson and Evans 1973, 1977; Humber and
Romero & Samuels), but no asexual morph was reported. Rombach 1987; Hsieh et al. 1997). Pseudogibellula was sep-
Fernández and Huhndorf (2004) established that arated from Gibellula as conidia were produced singly on
Porosphaerella borinquensis was linked to Pseudobotrytis sympodial conidiogenous cells and is common on a wide
terrestris as the asexual morph formed in vitro. From its asex- range of arthropod hosts and grows freely on agar media
ual morph the genus is known to be cosmopolitan. Molecular (Samson and Evans 1973). The asexual morph genera
studies have shown that Cordanaceae is closely related to Pseudogibellula, Gibellula and Granulomanus are exclusive-
Coniochaetales (Réblová and Winka 2000; Huhndorf et al. ly associated with Torrubiella (Johnson et al. 2009).
2004a; Réblová and Seifert 2007; Hernández-Restrepo et al. Rotiferophthora was described by Barron (1991) based on
2014; Zelski et al. 2014; Maharachchikumbura et al. 2015). distinct morphological differences from Diheterospora and
Hernández-Restrepo et al. (2014) accepted 19 species in this Verticillium, and this was confirmed by Zare et al. (2001)
genus and provided a key for their identification. Cordana and Sung et al. (2007).
species have been recorded from various temperate and trop- The asexual morph of this family is linked to Beauveria,
ical regions in the world, including Africa, South America, Isaria, Lecanicillium and Torrubiella (Sung et al. 2007;
South East Asia and New Zealand. Kepler et al. 2013). The genus Lecanicillium is characterized
Type species: Cordana pauciseptata Preuss, Linnaea 24: by slender aculeate phialides that are produced singly or in
129 (1851) whorls and usually arising from prostrate aerial hyphae; sin-
gle-celled, globose, hyaline to subhyaline conidia are pro-
Cordycipitaceae Kreisel ex G.H. Sung et al., in Sung et al., duced mostly at the tip of phialides and remain attached in
Stud. Mycol. 57: 48 (2007) heads of fascicles as glioid masses (Zare and Gams 2001; Zare
Facesoffungi number: FoF 01314 et al. 2001). Some Lecanicillium species are known to be
Parasites or pathogens of scale insects or mosses, or asexual morph forms of Cordyceps and Torrubiella (Petch
saprobes in leaf litter and upper soil layers. Sexual morph: 1932a; Evans and Samson 1982; Zare and Gams 2001; Zare
Stromata or subiculum, fleshy, pallid, or brightly coloured. et al. 2001). Some asexual morphs of Akanthomyces,
Perithecia superficial to completely immersed, oriented at Gibellula, Hirsutella, Isaria, and Simplicillium have been
right angles to the surface of the stroma. Asci mostly 8–spored, linked to Torrubiella (Samson et al. 1988). Some species of
cylindrical, with thickened ascus apex. Ascospores usually Cordyceps produce asexual morphs that are linked to
cylindrical, multi-septate, disarticulating into part-spores or Beauveria, which produce basally-inflated conidiogenous
remaining intact at maturity. Asexual morph: See notes. cells that produce conidia sympodially on divergent denticles
Type: Cordyceps Fr. (de Hoog 1972). Some species from Isaria are the asexual
Notes: The family name Cordycipitaceae was first used by morphs of Cordyceps and Torrubiella (Sung et al. 2007). In
K r e i s e l ( 1 9 6 9 ) , w h i l e We h m e y e r ( 1 9 7 6 ) u s e d this genus the usually hyaline conidia are produced in dry
Cordycipitoideae as a sub-family for Clavicipitaceae based chains from short flask-shaped phialides (Bischoff and
on the type genus Cordyceps. Cordycipitaceae was validly White 2003).
segregated from Clavicipitaceae by Sung et al. (2007), based
on morphology and multi-gene phylogenetic analyses. Most Cordyceps Fr., Observ. mycol. (Havniae) 2: 316
of the species in the family are entomogenous and produce (cancellans) (1818)
superficial to partially immersed to completely immersed peri- Facesoffungi number: FoF 01317; Figs 47, 48
thecia, on a fleshy stroma or subicula, that are pallid or bright- Parasitic on arthropods, worldwide. Sexual morph:
ly coloured (Sung et al. 2007). Sung et al. (2007) assigned Stromata or subiculum pallid or brightly pigmented, fleshy,
Fungal Diversity
 Fungal Diversity

ƒFig. 47 Cordyceps militaris – sexual morph (Material examined: Beauveria Vuill., Bull. Soc. bot. Fr. 59: 40 (1912)
CHINA, Province of Liao-Ling, on dead larva. 18 June 2014, Ting Chi Type species: Beauveria bassiana (Bals.-Criv.) Vuill.,
Wen TL 2014091004, MFU 15-3202) a Overview of stromata and host b
Yellow, half superficial ascomata on stroma c Host d Cross section
Bull. Soc. bot. Fr. 12: 40 (1912)
showing the stroma and ascomata e, f Ascomata g–i Asci j Cap of Coremiopsis Sizova & Suprun, Vestn. Moskov. Univ., Ser.
ascus k Part spores l Asci with ostioles. Scale bars: b = 1000 μm, biol. 12(2): 55 (1957)
d = 400 μm, e, g, h = 1000 μm, f, i, l = 500 μm, j = 50 μm, k = 2 μm Type species: Coremiopsis rosea Sizova & Suprun, Vestn.
Moskov. Univ., Ser. biol. 2: 55 (1957)
Engyodontium de Hoog, Persoonia 10(1): 53 (1978)
stromata growing from the heads of adult Lepidoptera, Type species: Engyodontium parvisporum (Petch) de
usually simple, rarely branched, Stipe flexuous, white. Hoog, Persoonia 10(1): 53 (1978)
Fertile head rounded, differentiated from stipe. Ascomata Gibellula Cavara, Atti Ist. bot. R. Univ. Pavia, 2 Sér. 3: 347
perithecial, superficial to completely immersed, ordinal in (1894)
arrangement, elongate or ampulliform, with the ostioles Type species: Gibellula pulchra Cavara [as ‘pulcra’], Atti
opening on the surface of the head. Peridium comprising Ist. bot. R. Univ. Pavia, 2 Sér. 3: 347 (1894)
three layers. Asci hyaline, cylindrical, with thickened apex. Granulomanus de Hoog & Samson, Persoonia 10(1): 70
A s c o s p o re s h y al i n e , c y l i n d r i c a l, m u lt i- se p t at e , (1978)
disarticulating into part-spores or non-disarticulating, rarely Type species: Granulomanus aranearum (Petch) de Hoog
possessing a thread-like structure connecting the fusiform & Samson, in de Hoog, Persoonia 10(1): 70 (1978)
ends. Asexual morph: Hyphomycetous. Colonies positive Hyperdermium J.F. White et al., Mycologia 92(5): 910 (2000)
pale, reverse yellowish. Hyphae hyaline, narrow, septate, Type species: Hyperdermium bertonii (Speg.) J.F. White
branched. Conidiophores verticillately or singly branched. et al., in Sullivan et al., Mycologia 92(5): 910 (2000)
Conidiogenous cells phialidic, solitary, verticillate, hyaline, Isaria Pers., Neues Mag. Bot. 1: 121 (1794)
inflated at the base, rarely cylindrical, tapering to a narrow Type species: Isaria farinosa (Holmsk.) Fr., Syst. mycol.
tip, and lacking a collarette. Conidia hyaline, smooth, (Lundae) 3(2): 271 (1832)
subglobose to ovoid. Lecanicillium W. Gams & Zare, Nova Hedwigia 72(3–4):
Type species: Cordyceps militaris (L.) Fr., Observ. mycol. 332 (2001)
(Havniae) 2: 317 (cancellans) (1818) Type species: Lecanicillium lecanii (Zimm.) Zare & W.
Notes: The genus name Cordyceps was validly published Gams, in Gams & Zare, Nova Hedwigia 72(3–4): 333 (2001)
by Link (1833) and first given as the group name by Fries in Microhilum H.Y. Yip & A.C. Rath, J. Invert. Path. 53(3): 361
1818 as Cordylia (Rogers 1954). Cordyceps includes about (1989)
280 species (Index Fungorum 2016) and was formally moved Type species: Microhilum oncoperae H.Y. Yip & A.C.
to Cordycipitoideae in 2007 (Sung et al. 2007). Sung et al. Rath, J. Invert. Path. 53(3): 362 (1989)
(2007) transferred some species from Cordyceps into several Pseudogibellula Samson & H.C. Evans, Acta bot. neerl.
genera including Ophiocordyceps (Ophiocordycipitaceae), 22(5): 524 (1973)
Tolypocladium (Ophiocordycipitaceae) as Elaphocordyceps, Type species: Pseudogibellula formicarum (Mains)
Metacordyc eps (C la v ici p i t a c e a e) , Claviceps Samson & H.C. Evans, Acta bot. neerl. 22(5): 524
(Clavicipitaceae), Tyrannicordyceps (Clavicipitaceae), (1973)
Epichloë (Clavicipitaceae), Podostroma (Hypocreaceae) and Rotiferophthora G.L. Barron, Can. J. Bot. 69(3): 495 (1991)
Podocrea (Hypocreaceae). Type species: Rotiferophthora globospora G.L. Barron,
Can. J. Bot. 69(3): 495 (1991)
Simplicillium W. Gams & Zare, Nova Hedwigia 73(1–2): 38
Other genera included (2001)
Type species: Simplicillium lanosiniveum (J.F.H. Beyma)
Akanthomyces Lebert, Z. Wiss. Zool. 9: 449 (1858) Zare & W. Gams [as ‘Simplicillium lanosoniveum’], (2001)
Type species: Akanthomyces aculeatus Lebert, Z. Wiss. Syspastospora P.F. Cannon & D. Hawksw., J. Linn. Soc., Bot.
Zool. 9: 449 (1858) 84(2): 152 (1982)
Ascopolyporus Möller, Bot. Mitt. Trop. 9: 300 (1901) Type species: Syspastospora parasitica (Tul.) P.F.
Type species: Ascopolyporus polychrous Möller, Bot. Mitt. Cannon & D. Hawksw., J. Linn. Soc., Bot. 84(2): 152
Trop. 9: 300 (1901) (1982)
Beejasamuha Subram. & Chandrash., Can. J. Bot. 55(3): Torrubiella Boud., Revue mycol., Toulouse 7: 226
247 (1977) (1885)
Type species: Beejasamuha samala Subram. & Type species: Torrubiella aranicida Boud., Revue mycol.,
Chandrash., Can. J. Bot. 55(3): 248 (1977) Toulouse 7: 226 (1885)
 Fungal Diversity

Fig. 48 Cordyceps militaris -

asexual morph (Material
examined: CHINA, Province of
Liao-Ling, on dead larva. 28
February 2015, Yuan Pin Xiao
MFLU) a, b Conidiophores and
developing conidia c Conidia d
Culture from above on PDA
medium after 7 days e Culture
from below on PDA medium after
7 days f Culture from above on
PDA medium after 40 days g
Culture from below on PDA
medium after 40 days. Scale bars:
a = 20 μm, b, c = 5 μm

Key to genera of Cordycipitaceae 4. Stromata with 1–3 perithecia per stroma

……………………………………...……Hyperdermium
1. Sexual morph . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 4. Stromata crowded . . . . . . . . . . . . . . . . Ascopolyporus
1. Asexual morph . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 5. Always with synnemata, only on insects . . . . . . . . . . 6
2. On insects . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 5. Synnemata not present; on insects, nematodes, other
2. On insects hidden within plants or on plants . . . . . . . 4 micro fauna, or other fungi or mycetozoans in soil . . . 11
3. Stromata with stipe . . . . . . . . . . . . . . . . . . .Cordyceps 6. Vesiculate conidiophores arising from synnematous
3. Stromata without stipe . . . . . . . . . . . . . . . . Torrubiella conidiomata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
Fungal Diversity

6. Conidiogenous cells lacking vesicles . . . . . . . . . . . . . 8 ascospores, unitunicate, thin-walled, clavate to cylindrical, long
7. Conidiogenous cells phialidic, on spiders . . .Gibellula pedicellate, apex blunt, without an apical ring. Ascospores
7. Conidiogenous cells sympodial, with short denticles; on crowded, hyaline, cylindrical to allantoid, slightly curved,
insects . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudogibellula aseptate, smooth-walled, mostly with guttules. Asexual morph:
8. Conidiogenous cells monophialidic . . . . . . . . . . . . . . 9 Undetermined.
8. Conidiogenous cells polyphialidic . . . . . . . . Beauveria
9. Synnemata determinate, with a terminal capitulum of Coronophora Fuckel, Fungi rhenani exsic., fasc. 10: no. 961
conidiophores, not on spiders . . . . . . . . . . . . . . . . . . . 10 (1864)
9. Synnemata indeterminate, with conidiophores emerging Facesoffungi number: FoF 01118, Fig. 49
from an extended conidiogenous layer, on spiders Type species: Coronophora gregaria Fuckel, Jb. nassau.
………………………………....................Akanthomyces Ver. Naturk. 23–24: 229 (1870) [1869–70]
10. Conidia produced in dry chains; phialidic; on insects Notes: The monotypic family Coronophoraceae was intro-
..........................................................................................Isaria duced by von Höhnel (1907). The family was placed in
10. Conidia produced in slime masses; denticulate, on ar- Sordariales based on lack of ostioles, while other characteris-
thropods or in soil . . . . . . . . . . . . . . . . . . Engyodontium tics were similar to Lasiosphaeriaceae, e.g. Lasiosphaeria
11. No resting structures produced, not parasites of delloid noonaedaniae (Carroll and Munk 1964). Coronophoraceae
rotifers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 was reported as a synonym of Nitschkiaceae and has been
11. Resting structures formed, parasites of delloid rotifers variously referred to Coronophorales or placed in the order
Rotiferophthora Sordariales (Huhndorf et al. 2004b). Mugambi and Huhndorf
12. Conidiogenous cells monoblastic or monophialidic, (2010), used TEF and RPB2 sequence dataset, which included
with a single conidiogenous locus . . . . . . . . . . . . . . . . 13 the type species, C. gregaria, to show that Coronophoraceae
12. Conidiogenous cells polyblastic, producing conidia is a distinct family from Nitschkiaceae. The genus
from more than one conidiogenous locus . . . . . . . . . . 15 Coronophora was introduced by Fuckel (1864) and includes
13. Conidiophores with a single verticil; phialides pointed 26 epithets (Index Fungorum 2016). Coronophora is charac-
………………………………………………………...14 terized by immersed, erumpent ascomata, clavate asci and
13. Conidiophores with several layers of verticils, phialides allantoid ascospores. Although the family Coronophoraceae
flask-shaped . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Isaria was reported as lacking ostioles (Carroll and Munk 1964), an
14. Phialides single, at right angles to subtending hyphae ostiole is present in C. consobrina P. Karst. (Fuckel 1870).
……………………………………………......Simplicillium
14. Phialides usually in whorls of 2 or more, not at right Cryphonectriaceae Gryzenh. & M.J. Wingf., in Gryzenhout
angles to subtending hyphae Lecanicillium et al., Mycologia 98(2): 246 (2006)
15. Rachis with an acropetal, zigzag sequence of slender Facesoffungi number: FoF 01382
denticles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Beauveria Saprobic or pathogenic in forest trees and economic crops.
15. Rachis with more or less swollen portions of the Sexual morph: Ascostromata scattered, immersed or
conidiogenous cells . . . . . . . . . . . . . . . . . . . . . Microhilum erumpent, aggregated, oval to circular from above, comprising
two layers, outer layer of yellowish orange to light brown cells,
Coronophoraceae Höhn., Sber. Akad. Wiss. Wien, Math.- purpling in KOH and inner layer of hyaline cells, mixed with
naturw. Kl., Abt. 1 116: 624 (1907) plant cells. Ascomata immersed, aggregated, several in one stro-
Facesoffungi number: FoF 01117 mata, globose to subglobose, brown, with long neck, or ostiolar
Saprobic on plant stems or wood in terrestrial habitats, or canal sometimes immersed in stromatic tissues, or superficial,
parasitic on other fungi. Sexual morph: Ascomata perithecial, necks covered in umber stromatic tissue of textura porrecta,
gregarious or solitary, superficial, black, sub-immersed or inner wall of the necks or ostiolar canal with hyaline, filamen-
erumpent through bark of host, ovoid to subglobose, carbona- tous periphyses. Peridium comprising inner layer of small, hy-
ceous, coriaceous or membranaceous, smooth, with tubercula or aline cells of textura angularis and outer layer of small, brown
hairs, laterally collapsing when dry, short neck present or absent, cells of textura angularis. Hamathecium comprising a few cel-
with or without ostioles. Peridium thick (30–80 μm), composed lular paraphyses and parenchymatous cells, attached at the base
of three layers, outer layer comprising dark tissues, thin (10– and asci dissolving at maturity. Asci 8-spored, unitunicate,
30 μm), carbonaceous; middle layer comprising dark brown to cylindrical-fusoid to clavate, pedicellate, with distinct, J- refrac-
brown cells of textura angularis, thick (20–50 μm), tive ring. Ascospores overlapping uniseriate to biseriate, oval to
membranaceous and the inner layer comprising hyaline cells of fusiform, 1-median-septate, rarely 2-septate, not constricted at
textura prismatica, thin (5–10 μm), membranaceous. the septa, hyaline, smooth-walled. Asexual morph:
Hamathecium composed of numerous, hyaline, septate, un- Coelomycetous. Conidiomata occurring as a part of ascomata
branched, filamentous paraphyses. Asci with numerous as conidial locules or solitary structures, uni- to multi-loculate,
Fungal Diversity
 Fungal Diversity

ƒFig. 49 Coronophora gregaria (Material examined: BELGIUM, on globose to subglobose, 4–6 aggregated in a single stroma,
bark of Prunus cerasus L. and Sorbus aucuparia L., Autumn, BR no. with black to brown ostiole, ostiolar canal slender, covered
5020094489111) a Material label b Host substrate c Ascomata erumpent
through bark of host d Ascomata without bark of host e Peridium f
with orange to fuscous-black stromatic tissue. Hamathecium
Paraphyses with asci g-j Asci. k-o Ascospores. Notes: Figs d-f, h-n are aparaphysate, comprising parenchymatous tissues. Asci 8-
strained in 3 % KOH. Scale bars: d = 2 mm, c = 1 mm, e–f = 50 μm, g– spored, unitunicate, fusiform to cylindrical, apex blunt, base
j = 20 μm, – = 5 μm with small pedicel. Ascospores overlapping uniseriate or
biseriate, hyaline, ellipsoid to fusiform, 1-septate. Asexual
morph: Coelomycetous. Conidiomata eustromatic, erumpent,
pyriform, subglobose to pulvinate, necks absent or present, if pyriform to pulvinate, orange to fuscous black, occurring in
present, with one to several attenuated necks, superficial or the same stroma as ascomata. Conidiophores cylindrical to
semi-immersed, orange to fuscous-black. Conidiophores cylin- bottle-shaped, unbranched, hyaline. Conidiogenous cells
drical, aseptate, hyaline, sometimes reduced to conidiogenous phialidic, simple or branched. Conidia hyaline, minute, gen-
cells. Conidiogenous cells lining the inner cavity of the erally ovoid to cylindrical, aseptate.
conidioma, phialidic, sometimes with inflated bases, Type species: Cryphonectria parasitica (Murrill) M.E.
ampulliform, inconspicuous, with attenuated or truncate apices, Barr, Mycologia Memoirs 7: 143 (1978)
hyaline, smooth. Conidia minute, sometimes both micro and Notes: Cryphonectria is typified by C. parasitica the causal
macro conidia present, broadly ellipsoid to fusoid, obovoid- agent of chestnut blight (Anagnostakis 1987; Heiniger and
cylindrical to allantoid, aseptate, hyaline. Rigling 1994). This genus consists of 17 species (Index
Type: Cryphonectria (Sacc.) Sacc. & D. Sacc. Fungorum 2016). Castlebury et al. (2002) showed some
Notes: The family Cryphonectriaceae was introduced to Endothia species to be the asexual morph of Cryphonectria.
accommodate the Cryphonectria-Endothia complex and other Cryphonectria was synonymized under Endothia (Kobayashi
allied genera (Gryzenhout et al. 2006). This family is typified 1970), however Barr (1978) separated these two genera based
by Cryphonectria parasitica (Murrill) M.E. Barr, which is a on the variation of ascospore septation and stromatal morphol-
serious pathogen causing chestnut blight. Members of this family ogy and transferred many Endothia species to Cryphonectria.
can be distinguished from other families of Diaporthales by Cryphonectria parasitica, C. cubensis (Bruner) Hodges and
having orange stromatic tissues, which are purple in KOH and C. eucalypti M. Venter & M.J. Wingf. are serious pathogens of
yellowing in lactic acid. Most members of this family are plant American chestnut and Eucalyptus. However, other taxa are
pathogens and form cankers, blights and dieback in economically saprobes (Roane et al. 1986).
important plants and forest trees. Castlebury et al. (2002), in a
study based on LSU sequence data of a large number of genera in
the Diaporthales, recognized six major lineages within the order,
namely, Gnomoniaceae sensu-stricto, Melanconidaceae sensu- Other genera included
stricto, Schizoparme complex including the asexual genera
Coniella and Pilidiella, Cryphonectria-Endothia complex (a pre- Amphilogia Gryzenh.et al., Taxon 54(4): 1017 (2005)
cursor to the Cryphonectriaceae), Valsaceae sensu-stricto, and Type species: Amphilogia gyrosa (Berk. & Broome)
Diaporthaceae sensu-stricto. Cryphonectriaceae was formally Gryzenh. et al., in Gryzenhout et al., Taxon 54(4): 1017
established by Gryzenhout et al. (2006) when analyzing both (2005)
LSU sequence data of fungal taxa in Diaporthales. Initially Aurantiosacculus Dyko & B. Sutton, in Dyko et al.,
Chrysoporthe, Rostraureum, Cryphonectria, Endothia and Mycologia 71(5): 922 (1979)
Amphilogia were placed in the family. Aurantiosacculus, Type species: Aurantiosacculus eucalypti (Cooke &
Aurapex, Aurifilum, Celoporthe, Chrysocrypta, Chrysoporthe, Massee) Dyko & B. Sutton, in Dyko et al., Mycologia
Chrysoporthella, Cryptometrion, Diversimorbus, Endothiella, 71(5): 922 (1979)
Foliocryphia, Holocryphia, Immersiporthe, Latruncellus, Aurapex Gryzenh. & M.J. Wingf., Mycologia 98(1): 112
Luteocirrhus, Mastigosporella, Microthia, Prosopidicola and (2006)
Ursicollum have since been added to the family. Type species: Aurapex penicillata Gryzenh. & M.J.
Wingf., in Gryzenhout et al., Mycologia 98(1): 112 (2006)
Cryphonectria (Sacc.) Sacc. & D. Sacc., Syll. fung. (Abellini) Aurifilum Begoude et al., in Begoude et al., Antonie van
17: 783 (1905) Leeuwenhoek 98(3): 273 (2010)
Facesoffungi number: FoF 02105; Fig. 50 Type species: Aurifilum marmelostoma Begoude et al., in
Saprobic or pathogenic on forest trees and economic crops. Begoude et al., Antonie van Leeuwenhoek 98(3): 273 (2010)
Sexual morph: Ascostromata comprising erumpent to super- Celoporthe Nakab. et al., Stud. Mycol. 55: 261 (2006)
ficial, orange epistromatic portion and immersed, hyaline, par- Type species: Celoporthe dispersa Nakab.et al., in Nakab.
enchymatous portion. Ascomata perithecial, immersed, et al., Stud. Mycol. 55: 261 (2006)
Fungal Diversity
 Fungal Diversity

ƒFig. 50 Cryphonectria parasitica (Material examined: USA, New York. Type species: Lasmenia balansae Speg. [as ‘balanse’],
Bronx Co., Bronx, North of Botanical Museum, Bronx Park, on Castanea Anal. Soc. cient. argent. 22(2): 152 (1886)
dentata (Marsh.) Borkh. (Fagaceae), 26 November 1905, W.A. Murrill,
01293321, 01293322 (NY, type of Diaporthe parasitica). a, b Herbarium
Latruncellus M. Verm. et al., in Vermeulen et al., Mycologia
packet c Herbarium specimen d Ascostromata on substrate and horizontal 103(3): 562 (2011)
section of ascostroma e Cross section of ascoma f Peridium g–j Asci k–n Type species: Latruncellus aurorae M. Verm., et al., in
Ascospores o–p Conidiostromata q Cross section of conidioma r-v Vermeulen et al., Mycologia 103(3): 562 (2011)
Conidia attached to the conidiophores w Conidia. Scale bars: d = 1 cm,
e–f = 100 μm, g–n = 10 μm, o = 200 μm, p = 500 μm, q = 100 μm, r–
Luteocirrhus C. Crane & T.I. Burgess IMA Fungus 4(1): 115
w = 10 μm (2013)
Type species: Luteocirrhus shearii C. Crane & T.I.
Burgess IMA Fungus 4(1): 115 (2013)
Chromendothia Lar.N. Vassiljeva, Mikol. Fitopatol. 27(4): 5 Mastigosporella Höhn. Sber. Akad. Wiss. Wien, Math.-
(1993) naturw. Kl., Abt. 1 123: 135 (1914)
Type species: Chromendothia appendiculata Lar.N. Type species: Mastigosporella hyalina (Ellis & Everh.)
Vassiljeva, Mikol. Fitopatol. 27(4): 5 (1993) Höhn., Sber. Akad. Wiss. Wien, Math.-naturw. Kl., Abt. 1
Chrysocrypta P.W. Crous & B.A. Summerell, in Crous et al., 123: 135 (1914)
Persoonia, Mol. Phyl. Evol. Fungi 28: 165 (2012) Microthia Gryzenh. & M.J. Wingf., Stud. Mycol. 55: 44
Type species: Chrysocrypta corymbiae P.W. Crous & B.A. (2006)
Summerell, in Crous et al., Persoonia, Mol. Phyl. Evol. Fungi Type species: Microthia havanensis (Bruner) Gryzenh.
28: 165 (2012) & M.J. Wingf., in Gryzenhout, et al., Stud. Mycol. 55:
Chrysofolia Crous & M.J. Wingf., Persoonia, Mol. Phyl. 44 (2006)
Evol. Fungi 34: 207 (2015) Prosopidicola Crous & C.L. Lennox, Stud. Mycol. 50(1):
Type species: Chrysofolia colombiana Crous, Rodas & ‘187’ [191] (2004)
M.J. Wingf., Persoonia, Mol. Phyl. Evol. Fungi 34: 207 Type species: Prosopidicola mexicana Crous & C.L.
(2015) Lennox, in Lennox et al., Stud. Mycol. 50(1): ‘187’ [191]
Chrysoporthe Gryzenh. & M.J. Wingf., Stud. Mycol. 50(1): (2004)
129 (2004) Rostraureum Gryzenh. & M.J. Wingf., Mycol. Res. 109(9):
Type species: Chrysoporthe cubensis (Bruner) Gryzenh. & 1039 (2005)
M.J. Wingf., in Gryzenhout et al., Stud. Mycol. 50(1): 130 Type species: Rostraureum tropicale Gryzenh. & M.J.
(2004) Wingf., Mycol. Res. 109(9): 1040 (2005)
Cryptometrion Gryzenh. & M.J. Wingf., Australas. Pl. Path. Ursicollum Gryzenh. & M.J. Wingf., Stud. Mycol. 55: 44
39(2): 166 (2010) (2006)
Type species: Cryptometrion aestuescens Gryzenh. & M.J. Type species: Ursicollum fallax Gryzenh. & M.J. Wingf.,
Wingf., Australas. Pl. Path. 39(2): 166 (2010) in Gryzenhout et al., Stud. Mycol. 55: 46 (2006).
Diversimorbus S.F. Chen et al., Fungal Biol. 117: 300 (2014)
Type species: Diversimorbus metrosiderotis S.F. Chen
et al., Fungal Biol. 117: 301 (2014) Key to genera of Cryphonectriaceae
Endothia Fr., Summa veg. Scand., Section Post. (Stockholm):
385 (1849) 1. Conidiomata orange………….……….………………… 2
Type species: Endothia gyrosa (Schwein.) Berk. [as 1. Conidiomata uniformly brown to black, with or without
‘gyrosum’], Outl. Brit. Fung. (London): 384 (1860) orange necks……………….……….………..…….…. 14
Foliocryphia Cheew. & Crous, Persoonia 23: 65 (2009) 2. Sexual morph known………………………....…........ 3
Type species: Foliocryphia eucalypti Cheew. & Crous, 2. Sexual morph not known……….….....................….. 17
Persoonia 23: 65 (2009) 3. Ascospores brown….….…….....……. Chromendothia
Holocryphia Gryzenh. & M.J. Wingf., in Gryzenhout et al., 3. Ascospores hyaline……………………...............…… 4
Stud. Mycol. 55: 48 (2006) 4. Conidiomata pulvinate to globose, ascospores septate or
Type species: Holocryphia eucalypti (M. Venter & M.J. aseptate…....…......………..........................................….. 5
Wingf.) Gryzenh. & M.J. Wingf., in Gryzenhout et al., Stud. 4. Conidiomata conical, rostrate, pyriform or convex, with
Mycol. 55: 48 (2006) or without a neck, ascospores septate……….…..…..... 10
Immersiporthe S.F. Chen et al., in Chen et al., Pl. Path. 62: 5. Ascospores septate………..................……………...... 6
674 (2013) 5. Ascospores aseptate…….………………………....…. 9
Type species: Immersiporthe knoxdaviesiana S.F. Chen 6. Ascostromata superficial, conidiomata paraphysate..... 7
et al., in Chen et al., Pl. Path. 62: 674 (2013) 6. Ascostromata erumpnt, conidiomata aparaphysate ..... 8
Lasmenia Speg., Anal. Soc. cient. argent. 22(4): 199 (1886) 7. Perithecia valsoid……............................ Diversimorbus
 Fungal Diversity

7. Perithecia diatrypoid….............……..……… Microthia 21Conidia ellipsoid, straight to allantoids….... Chrysofolia

8. Conidiomata usually larger than 350 μm, uni- to multi- 22. Conidiophores brown………....……… Prosopidicola
loculate, ascospores with median septum........ 22 Conidiophores hyaline………...................………… 23
Cryphonectria 23. Conidiomata with one to four attenuated
8. Conidiomata usually smaller than 350 μm diam, uniloc- necks………...….…………………...… Chrysoporthella
ular, ascospores with median to submedian sep- 23. Conidiomata with or without necks……....………. 24
tum……..….........…..…..…..…..…....….. Cryptometrion 24. Conidiophores irregularly branched……. Foliocryphia
9. Stromata strongly developed, large, erumpent, mostly 24. Conidiophores unbranched……….......................… 25
superficial, numerous conidial locules, no paraphyses 25. Conidia cylindrical or slightly
…………….......…………………………........ Endothia allantoid……………………….………....... Luteocirrhus
9. Stromata small to medium, semi-immersed, few co- 25. Conidia narrowly ellipsoid to fusi-
nidial locules or one convoluted locule, paraphyses form….……………….…....…………... Mastigosporella
present……..................................….............………
Holocryphia Diaporthaceae Höhn. ex Wehm., Am. J. Bot. 13: 638 (1926)
10. Conidiomata with necks, …………………..……… 11 Facesoffungi number: FoF 01383
10. Conidiomata without necks, ….………............…… 13 Pathogenic, endophytic or saprobic on terrestrial and rarely
11. Conidiomata with prominent, delimited neck…...... 12 submerged plants. Sexual morph: Stromata present or absent.
11. Conidiomata with neck continuous with base, rostrate, If present, pulvinate, erumpent, flat or slightly convex, orbic-
white sheath of tissue surrounding perithecial ular, circular or somewhat irregular, sclerotioid, subhyaline,
necks.....……….……….……….………… Rostraureum stromatic disk coriaceous, whitish to brownish black, with or
12. Conidiomata rostrate to pyriform with large base, neck without black zone or a crust consisting of fungus tissue, sol-
attenuated or not, sexual morph Undetermined itary or containing up to 10 ascomata in a stroma. Ascomata
.......................................................................... Ursicollum perithecial, immersed to erumpent, solitary or aggregated in a
12. Conidiomata conical with constricted, fattened neck, valsoid configuration, globose or compressed, coriaceous,
shaped like a chess pawn…….................…. Latruncellus black, ostiolate, papillate. Papilla short or long, erumpent,
13. Conidiomata conical, uniformly orange…. Amphilogia convergent, cylindrical to conical, black, internal wall covered
13. Conidiomata convex, with blackened ostiolar open- by hyaline periphyses, composed of vertically arranged par-
ings………….….….….….…....….….………. Aurifilum enchymatous tissues. Peridium comprising outer layer of flat-
14. Conidiomata uniformly black when mature….….... 15 tened, thick-walled, dark brown cells of textura angularis and
14. Conidiomata black with orange neck, teleomorph inner layer of hyaline, thin-walled cells of textura angularis.
Undetermined……...….…………………….…. Aurapex Hamathecium comprising septate, unbranched, cylindrical pa-
15. Sexual morph reported…………..………..……….. 16 raphyses, tapering upwards. Asci 8-spored, unitunicate, cla-
15. Sexual morph not reported……….…….…….……. 22 vate, oblong-clavate to broadly fusoid, sessile, with a distinct
16. Conidiomata base tissue of textura globulosa when apical ring. Ascospores biseriate to partially biseriate, ellip-
sectioned longitudinally, perithecial necks long and cov- soid, oblong to fusoid, unicellular or 1-septate, constricted at
ered with dark tissue, emerging from orange stro- septum, with cap-like appendages at both ends or without
ma….…...…….….…....…….….….......… Chrysoporthe appendages, hyaline, dark to blackish brown, sometimes nar-
16. Conidiomata base tissue prosenchymatous, apices of rowly rounded ends and multi-guttulate, smooth-walled.
conidiomata can be orange to scarlet when young, perithe- Asexual morph: Conidiomata acervular or pycnidial, glo-
cial necks short, orange to umber stroma.…… Celoporthe bose, initially immersed, erumpent at maturity, solitary,
17. Chlamydospores present……........……..…. Lasmenia scattered, coriaceous, black, elongated ostiolar neck, often
17. Chlamydospores absent…………………………… 18 with yellowish, conidial mass extruding from ostiole.
18. Conidiomata pulvinate…………………….……… 19 Peridium comprising 3–4 layers of light brown cells of textura
18. Conidiomata globose to subglobose…….….…..…. 20 angularis. Conidiophores arising from the inner layer of the
19. Conidiomata uniloculate with necks.….… Endothiella locule, ampulliform, cylindrical, straight to sinuous, septate,
19. Conidiomata m ulti-l oculate without necks palisades, simple, branched, hyaline to pale brown, smooth,
……………………………………...........Immersiporthe bearing conidia acrogenously. Conidiogenous cells
20. Conidiogenous cells subcylindrical to enteroblastic, phialidic, cylindrical, terminal, slightly tapering
lageniform…….….….….….….….….….. towards the apex, sometime with a small collarette, determi-
Aurantiosacculus nate. Alpha conidia abundant, ovate, fusoid to ellipsoidal,
20. Conidiogenous cells ampulliform…………….....… 21 aseptate, hyaline, smooth, base sub-truncate, straight to
21Conidia fusoid-ellipsoid, apex acutely round- curved, occasionally slightly sigmoid, pale brown, aseptate,
ed……………………………………...….. Chrysocrypta smooth, with many guttules, hyaline, sometimes with short
Fungal Diversity

appendages at both ends. Beta conidia fusiform to hooked, (Wehmeyer 1933). Diaporthe has a worldwide distribu-
aseptate, hyaline, smooth, base sub-truncate. tion and the species are saprobes, pathogens and endo-
Type: Diaporthe Nitschke. phytes. Phomopsis was previously known as the asexual
Notes: The family Diaporthaceae was established by von morph and as these genera were linked Udayanga et al.
Höhnel (1917) and was accommodated in the order 2012a, b; 2014). Rossman et al. (2015) based on prin-
Diaporthales. Wehmeyer (1975) confined this family to include ciple of priority gave Diaporthe over Phomopsis, to
Diaporthe and Mazzantia. Later, Diaporthaceae was synony- resolve nomenclatural problems.
mized under Valsaceae (Barr 1978). However, analysis of LSU
sequence data of diaporthalean taxa showed the distinct place-
ment of Diaporthaceae in Diaporthales where it formed a well- Other genera included
supported clade. This family previously only accommodated
Diaporthe (Phomopsis) and Mazzantia (Castlebury et al. Allantoporthe Petr., Hedwigia 62: 289 (1921)
2002). Pustulomyces, isolated from decaying bamboo culms, Type species: Allantoporthe tessella (Pers.) Petr., Hedwigia
was placed in Diaporthaceae by Dai et al. (2014b), based on 62: 289 (1921)
combined analysis of LSU, SSU and TEF sequence data. The Apioporthella Petr., Annls mycol. 27(5/6): 401 (1929)
phylogenetic placement of Phaeodiaporthe has been confirmed Type species: Apioporthella bavarica Petr., Annls mycol.
within Diaporthaceae by Voglmayr and Jaklitsch (2014) based 27(5/6): 401 (1929)
on analysis of LSU sequence data. Maharachchikumbura et al. Clypeoporthella Petr., Annls mycol. 22(1/2): 149 (1924)
(2015) listed Allantoporthe, Apioporthella, Clypeoporthella, Type species: Clypeoporthella brencklei Petr., Annls
Diaporthe, Diaporthella, Leucodiaporthe, Mazzantia, mycol. 22(1/2): 149 (1924)
Mazzantiella, Ophiodiaporthe and Pustulomyces as genera of Diaporthella Petr., Annls mycol. 22(1/2): 30 (1924)
Diaporthaceae. Type species: Diaporthella aristata (Fr.) Petr., Annls
mycol. 22(1/2): 30 (1924)
Diaporthe Nitschke, Pyrenomyc. Germ. 2: 240 (1870); Leucodiaporthe M.E. Barr & Lar.N. Vassiljeva, in Vasilyeva,
Facesoffungi number: FoF 02106 Fig. 51 Rossman & Farr, Mycologia 99(6): 917 (2008) [2007]
Pathogenic, endophytic or saprobic on plants. Sexual Type species: Leucodiaporthe acerina M.E. Barr & Lar.N.
morph: Ascomata perithecial, deeply immersed, aggregated, Vassiljeva, Mycologia 99(6): 919 (2008) [2007]
globose, subglobose or irregular, coriaceous, black, papillate, Mazzantia Mont., Bull. Soc. bot. Fr. 2: 525 (1855)
ostiolate. Papilla tapering towards the host surface, internally Type species: Mazzantia galii (Fr.) Mont., Syll. gen. sp.
covered by hyaline, periphyses. Peridium comprising brown, crypt. (Paris): 246 (1856)
thick-walled, cells of textura angularis. Hamathecium with- Ophiodiaporthe Y.M. Ju et el., Mycologia 105(4): 868
out paraphyses. Asci unitunicate, 8-spored, elongate to cla- (2013)
vate, apex with distinct, J- apical ring, sessile. Ascospores Type species: Ophiodiaporthe cyatheae Y.M. Ju et al.,
overlapping uniseriate, elongated to elliptical, hyaline, 1-sep- Mycologia 105(4): 868 (2013)
tate, often 4 guttulate, with larger guttules at center and smaller Phaeodiaporthe Petr., Annls mycol. 17(2/6): 99 (1920)
ones at the ends, smooth-walled. Asexual morph: [1919]
Conidiomata pycnidia, globose, initially immersed, erumpent Type species: Phaeodiaporthe keissleri Petr., Annls mycol.
at maturity, black, elongated neck, often with yellowish co- 17(2/6): 99 (1920) [1919]
nidial mass extruding from ostiole. Peridium comprising 3–4 Pustulomyces D.Q. Dai et al., in Dai et al., Cryptog. Mycol.
layers of light brown cells of textura angularis. 35(1): 64 (2014)
Conidiophores ampulliform, straight to sinuous, unbranched, Type species: Pustulomyces bambusicola D.Q. Dai et al.,
hyaline, smooth. Conidiogenous cells phialidic, cylindrical, in Dai et al., Cryptog. Mycol. 35(1): 64 (2014)
terminal, slightly tapering towards the apex. Alpha conidia Stenocarpella Syd. & P. Syd., Annls mycol. 15(3/4): 258
abundant, ovate to ellipsoidal, aseptate, mostly biguttulate, (1917)
hyaline, smooth, base sub-truncate. Beta conidia fusiform, Type species: Stenocarpella zeae Syd. & P. Syd., Annls
aseptate, hyaline, smooth, base sub-truncate. mycol. 15(3/4): 258 (1917)
Type species: Diaporthe eres Nitschke, Pyrenomyc. Germ.
2: 245 (1870)
Notes: The genus Diaporthe was introduced by Key to genera of Diaporthaceae
Nitschke (1867) to include taxa in the Sphaeriales with
stromata often with blackened zones in the substrate, 1. Coelomycetes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
ellipsoid to fusiform ascospores and enclosed, unilocular 1. Ascomycetes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
pycnidia that contain spermatia, stylospores and conidia 2. Conidia elongate fusiform to sigmoid. . . Pustulomyces
 Fungal Diversity

Fig. 51 Diaporthe eres (Material.: B 70 0009145 lectotype specimen (a- Ascospores i Conidiomata j Conidia attached to the conidiogenous cells
h) see Udayanga et al. 2014 for details (i–l) Ar5193, ex-epitype) a k Alpha conidia l Beta conidia. Scale bars: a = 500 μm, b = 100 μm, c–e,
Ascomata on substrate b Cross section of ascoma c–e Asci f–h i–l = 20 μm, f–h = 10 μm

2. Conidia subcylindrical to narrowly ellipsoid 3. Ascomata aggregated……......................................….. 4

…………………………………....……..... Stenocarpella 4. Stromata without blackened zone in substrate and
3. Ascomata solitary…………....…………. Clypeoporthe brownish stromatic disk...................… ...........Diaporthella
Fungal Diversity

4. Stromata with blackened zone in substrate and brown to and blunt apex, hyaline. (Pollack and Uecker 1974; Hyde and
blackish stromatic disk.….............................................… 5 Jones 1992; Klaysuban et al. 2014; Senanayake et al. 2015)
5. Ascospores brown…….................….... Phaeodiaporthe Type: Diatrype Fr., Summa veg. Scand.
5. Ascospores hyaline…...................................………… 6 Notes: The family Diatrypaceae was typified by Diatrype.
6. Septa submedian……………….............. Apioporthella It is characterized by perithecial ascomata embedded in a
6. Septa median………….......…………….....………… 7 black stroma, long stalked asci and allantoid ascospores
7. Ascomata more than 20 in a stromata……..............… 8 (Glawe and Rogers 1984; Rappaz 1987). It comprised 15
7. Ascomata 1 to very few(less than 20) in a stroma- genera: Anthostoma, Cryptosphaeria, Cryptovalsa, Diatrype,
ta……………………..……............…………........…… 9 Diatrypella, Diatrypasimilis, Echinomyces, Eutypa,
8. Ascospores overlapping uniseriate, elongate to elliptical, Eutypella, Leptoperidia, Monosporascus, Pedumispora,
often with 4-guttules, larger guttules at the center and P e ro n e u t y p a , P h a e o i s a r i a a n d Q u a t e r n a r i a i n
smaller ones at the ends…….....................…… Diaporthe Maharachchikumbura et al. (2015). Detailed taxonomic re-
8. Ascospores biseriate, fusoid, without views for the family were provided by Trouillas et al. (2011)
guttules….................................................… Allantoporthe and Mehrabi et al. (2015) and sequence data is lacking for
9. Ascospores bi-celled…….............…… Ophiodiaporthe only Echinomyces, Leptoperidia, Peroneutypa and
9. Ascospores unicellular……….........……………….. 10 Quaternaria (Senanayake et al. 2015). Recent papers with
10. Asci oblong-clavate, ascospores elliptic to fusoid sequence data providing backbones trees are Carmaran et al.
…………………….……............................................... 11 (2006) and Trouillas et al. (2010). The placement of species in
10. Ascospores ovoid…...................…… Leucodiaporthe genera is very confused with many genera being polyphyletic
11. Perithecia with long neck…......……. Clypeoporthella and the family is in need of a thorough monographic revision
11. Perithecia with very short or almost lacking neck based on molecular data and morphology. These fungal taxa
………………..……………….............................. Mazzantia produce extracellular ligninolytic enzymes, cellulose and ca-
talyse the hydrolysis of cellulose, break down of lignin in the
Diatrypaceae Nitschke [as ‘Diatrypeae’], Verh. naturh. Ver. cell walls of plants. Therefore, some species in the family have
preuss. Rheinl. 26: 73 (1869) the physiological capacity in decay wood (Trouillas et al.
Facesoffungi number: FoF 00679 2011; Mehrabi et al. 2015).
Saprobic or pathogenic on woody plants in terrestrial and
aquatic habitats. Sexual morph: Stromata eustromatic or Diatrype Fr., Summa veg. Scand., Section Post. (Stockholm)
pseudostromatic, well-developed, immersed to erumpent, 384 (1849)
rarely superficial, mostly black or dark brown, with somewhat Facesoffungi number: FoF 00702 Figs 52, 53
carbonaceous outer layer, inner layer pale, loosely packed, Saprobic on bark of woody hosts. Sexual morph:
parenchymatous. Ascomata perithecial, immersed in stromatic Stromata scattered or aggregated on host, sometime spread
tissues, mostly brown to black, globose to sub-globose, with on large area of host, erumpent to superficial, orbicular, disc-
ostiolar necks. Ostioles sulcate, inner layer covered with hya- like plane or convex, sometime spread on host evenly, arising
line, periphyses. Peridium consists of two layers, an inner through the cracks in bark or epidermis or spread beneath the
hyaline layer and an outer layer of brown to black cells of epidermis, edges of cracks remaining as pointed, angular
textura angularis. Hamathecium comprising long, wide, parts, with numerous perithecia immersed in one stroma, os-
branched, septate, paraphyses. Asci 8-spored or polysporous, tiole opening through host bark and appearing as black spots,
rarely 1-spored or 2-spored, unitunicate, cylindrical, clavate to composed of an outer layer of dark brown, small, tightly
pyriform, fusiform, with a very long stalk, with a more or less packed, thin parenchymatous cells and an inner layer of yel-
truncate apex, with a J- or J+ apical ring. Ascospores crowded, lowish white, large, loosely packed, parenchymatous cells.
most hyaline to light brown, rarely jet-black, allantoid, ellip- Ascomata perithecia, immersed in stromatic tissues, aggregat-
soidal, globose or filiform. Asexual morph: Coelomycetous, ed, pale brown, globose to sub-globose, narrowing towards
non stromatic, occurring on host as acervuli. Conidiomata the apex and very narrow at the base of ostiolar canal, thin
acervuli sub cortical, erumpent, yellow to red, with branched walled, ostiolate. Ostiolar canal, periphysate, ostiolar opening
conidiophores and in culture as pycnidia, superficial, solitary covered with carbonaceous, black cells, apex wider than base.
or aggregated, yellow, dark brown to black, subconical, glo- Periphyses hyaline, filamentous, short, bent towards the cav-
bose to subglobose, and thick peridium, comprising brown, ity. Peridium thin, comprising an outer layer of brown, thick
thick-walled cells of textura angularis with branched conidi- walled cells of textura angularis and a thin, inner strata of
ophores, arising from pseudoparenchymatous cells or inter- hyaline, thick walled cells of textura angularis. Paraphyses
woven hyphae. Conidiogenous cells in dense palisades, cylin- arising from base of perithecia, composed of long, wide, glo-
drical, straight or curved, apically distorted or annulated. bose to ovoid thin-walled cells, branched, septate, and slightly
Conidia filiform, curved, or rarely straight with flattened base constricted at septa, narrowing and tapering towards the apex,
 Fungal Diversity

Fig. 52 Diatrype disciformis

(Material examined: USA,
Washington, Snohomish Co.
Marysville. J.M. Grant, January
1928, NY 00305625) a
Herbarium material b Stromata
on substrate c Ostioles appear on
stroma d Ascomata formed in a
stroma e Peridium f Cross section
of ascoma g Paraphyses h–j Asci
k–l Ascospores. Scale bars: b–
c = 1000 μm, d = 200 μm, e–
f = 50 μm, g–j = 15 μm, k–
l = 5 μm

apex blunt. Asci unitunicate, 8-spored, with very long, brown to black, globose to subglobose, shiny, smooth
narrow, thin walled stalk, with cylindrical, thick walled, surface. Peridium thick, consisting of brown, thick-
swollen upper portion, apex flat, with J-, cylindrical, walled, textura angularis cells. Conidiophores branched,
conspicuous apical ring, attached to the base by stalk. arising from pseudoparenchymatous cells or interwoven
Ascospores biseriate, hyaline, allantoid, unicellular, thin hyphae. Conidiogenous cells dense palisades, cylindri-
walled, with small, fat globules at each end, smooth cal, straight or curved, apically distorted or annulated.
walled. Asexual morph: Conidiomata appears as Conidia filiform, curved or rarely straight with flattened
brownish yellow, watery, bubble-like, conidial mass base and blunt apex, hyaline. (Senanayake et al. 2015)
rounded from white, mycelia clumps, pycnidial, superfi- Type species: Diatrype disciformis (Hoffm.) Fr., Summa
cial, solitary or aggregated, subconical, yellow, dark veg. Scand., Section Post. (Stockholm): 385 (1849)
Fungal Diversity

Type species: Cryptosphaeria millepunctata Grev., Fl.

Edin.: 360 (1824)
Cryptovalsa Ces. & De Not. ex Fuckel, Jb. nassau. Ver.
Naturk. 23–24: 212 (1870) [1869–70]
Type species: Cryptovalsa protracta (Pers.) De Not., Sfer.
Ital.: 40 (1863)
Diatrypella (Ces. & De Not.) De Not. Sfer. Ital.: 29 (1863)
Type species: Diatrypella verruciformis (Ehrh.) Nitschke
1867
Diatrypasimilis J.J. Zhou & Kohlm.et al. Mycologia 102(2):
432 (2010)
Type species: Diatrypasimilis australiensis J.J. Zhou &
Kohlm., et al., Mycologia 102(2): 432 (2010)
Echinomyces Rappaz, Mycol. helv. 2(549): 547 (1987)
Type species: Echinomyces obesa (Syd.) Rappaz, Mycol.
helv. 2(3): 548 (1987)
Eutypa Tul. & C. Tul., Select. fung. carpol. (Paris) 2: 52 (1863)
Type species: Eutypa lata (Pers.) Tul. & C. Tul., Select.
fung. carpol. (Paris) 2: 56 (1863)
Eutypella (Nitschke) Sacc., Atti Soc. Veneto-Trent. Sci. Nat.,
Padova, Sér. 4 4: 80 (1875)
Type species: Eutypella cerviculata (Fr.) Sacc., Syll. fung.
(Abellini) 1: 146 (1882)
Leptoperidia Rappaz, Mycol. helv. 2(547): 544 (1987)
Type species: Leptoperidia macropunctata (Rehm)
Rappaz, Mycol. helv. 2(3): 545 (1987)
Monosporascus Pollack & Uecker, Mycologia 66(2): 348
(1974)
Type species: Monosporascus cannonballus Pollack &
Uecker, Mycologia 66(2): 348 (1974)
Fig. 53 Asexual morph of Diatrype (a, b, d Libertella betulina, taken
from Sutton 1980; c Diatrype subaffixa, taken from Chlebicki and Pedumispora K.D. Hyde & E.B.G. Jones, Mycol. Res. 96(1):
Krzyzanowska 1995; e Diatrype disciformis, taken from Rappaz 1987) 78 (1992)
a, b conidiophore c–e conidia. Scale bars: a–e = 10 μm Type species: Pedumispora rhizophorae K.D. Hyde &
E.B.G. Jones, Mycol. Res. 96(1): 78 (1992)
Peroneutypa Berl., Icon. fung. (Abellini) 3(3–4): 80 (1902)
Notes: Diatrype was established by Fries (1849) with Type species: Peroneutypa bellula (Desm.) Berl., Icon.
Diatrype disciformis (Hoffm.) Fr. as the type species. It com- fung. (Abellini) 3(3–4): 81 (1902)
monly inhabits decaying wood. A few species are reported as Phaeoisaria Höhn., Sber. Akad. Wiss. Wien, Math.-naturw.
pathogens that form cankers on forest trees (Senanayake et al. Kl., Abt. 1 118: 330 [56 repr.] (1909)
2015). Species in this genus are quite resistant to harsh con- Type species: Phaeoisaria bambusae Höhn., Sber. Akad.
ditions. The asexual morph of Diatrype is reported as Wiss. Wien, Math.-naturw. Kl., Abt. 1 118: 329 [55 repr.] (1909)
Libertella-like and Dumortieria-like (Kirk et al. 2008; Quaternaria Tul. & C. Tul., Select. fung. carpol. (Paris) 2:
Wijayawardene et al. 2012; Senanayake et al. 2015). 104 (1863)
Type species: Quaternaria persoonii Tul. & C. Tul., Select.
fung. carpol. (Paris) 2: 105, tab. 12, fig. 16–25 (1863).

Other genera included

Key to genera of Diatrypaceae
Anthostoma Nitschke, Pyrenomyc. Germ. 1: 110 (1867)
Type species: Anthostoma decipiens (DC.) Nitschke, 1. Sexual morph . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
Pyrenomyc. Germ. 1: 111 (1867) 1. Asexual morph: hyphomycetous forming synnemata
Cryptosphaeria Ces. & De Not., Comm. Soc. crittog. Ital. with hyaline or coloured, ovoid to ellipsoidal, aseptate co-
1(4): 231 (1863) nidia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Phaeoisaria
 Fungal Diversity

2. Ascospores globose, elongate fusiform, or oblong to Saprobic on intertidal wood and bark in marine habitats.
ellipsoidal . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 Sexual morph: Ascomata subglobose to globose or pyriform,
2. Ascospores allantoid . . . . . . . . . . . . . . . . . . . . . . . . 5 light brown to dark brown or black, immersed, oblique or
3. Ascospores elongate fusiform, septate Pedumispora vertical to the host surface, clypeate, coriaceous, ostiolate,
3. Ascospores aseptate . . . . . . . . . . . . . . . . . . . . . . . . . 4 periphysate, papillate. Peridium comprising several layers of
4. Ascospores globose, with 1–2 spores in each ascus brown to dark brown cell layers of textura angularis.
Monosporascus Paraphyses numerous, hyaline, mostly unbranched, attached
4. Ascospores oblong to ellipsoidal, with 8 spores in each to the apex of the ascomatal cavity, in a gelatinous matrix. Asci
ascus . . . . . . . . . . . . . . . . . . . . . . . . . . . Diatrypasimilis 8-spored, unitunicate, thin-walled, cylindrical to oblong, ped-
5. Asci with more than 8 spores . . . . . . . . . . . . . . . . . . 6 icellate, J-, persistent. Ascospores 1–2-seriate, hyaline, ellip-
5. Asci with 8 spores . . . . . . . . . . . . . . . . . . . . . . . . . . 7 soidal, 1-many septate, constricted at the septa, with a filamen-
6. Ascostromata erumpent through host surface, dis- tous appendage at one or both ends. Appendages bristle-like,
coid………………………………………….. Diatrypella origin undetermined. Asexual morph: Undetermined.
6. Ascostromata immersed, erumpent through host surface Type: Etheirophora Kohlm. & Volkm.-Kohlm.
by sulcate ostioles, eutypoid. . . . . . . . . . . . . Cryptovalsa Notes: This family was introduced to accommodate the gen-
7. Ascostromata semi-immersed to erumpent through the era Etheirophora (E. bijubata Kohlm. & Volkm.-Kohlm.,
host periderm (ectostromatic) . . . . . . . . . . . . . . . . . . . . 8 E. blepharospora Kohlm. and Volkm.-Kohlm., E. unijubata
7. Ascostromata deeply immersed in the host periderm Kohlm. & Volkm.-Kohlm.) and Swampomyces Kohlm. &
(entostromatic) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 Volkm.-Kohlm. (S. armeniacus Kohlm. & Volkm.-Kohlm.,
8. Ascomata erumpent through the ectostromata, with long S. triseptatus K.D. Hyde & Nakagiri), which grouped together
cylindrical, prominent ostioles in the center Anthostoma in a well-supported clade in an analysis of LSU and SSU se-
8. Ascomata immersed in pustulate, effuse, acervuloid, or quence data (Jones et al. 2014). However, the genera
discoid ectostromata. . . . . . . . . . . . . . . . . . . . . . . . . . . 9 Etheirophora and Swampomyces are not congeneric. They form
9. Ectostromata composed of carbonaceous cells, a sister clade with Falcocladium species (Falcocladiaceae) in an
acervuloid . . . . . . . . . . . . . . . . . . . . . . . . . Echinomyces unsupported clade in Hypocreomycetidae, order incertae sedis.
9. Ectostromata composed of pseudoparenchymatous cells, This was also shown by Maharachchikumbura et al. (2015) in
discoid . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Diatrype an enlarged data set. Subsequently, Jones et al. (2015) intro-
10. Ascomata clustered, forming valsoid configuration, duced the order Torpedosporales to accommodate the families
breaking through entostroma by short to long necks. . 11 Etheirophoraceae, Juncigenaceae and Torpedosporaceae. The
10. Ascomata scattered, arranged in linear entostroma, with order Torpedosporales forms a sister clade to the orders
short to long necks. . . . . . . . . . . . . . . . . . . . . . . . . . . 12 Falcocladiales, Coronophorales and Melanosporales with high
11. Entostromata slightly raised on the host, with long cy- statistical support.
lindrical, packed necks . . . . . . . . . . . . . . . . . . . Eutypella
11. Entostromata immersed in the host, with, individually, Etheirophora Kohlm. & Volkm.-Kohlm., Mycol. Res. 92(4):
protruding necks at the center. . . . . . . . . . . . Quaternaria 414 (1989)
12. Ascomata forming very long necks, through the host Facesoffungi number: FoF 02107; Fig. 54
surface . . . . . . . . . . . . . . . . . . . . . . . . . . . . Peroneutypa Saprobic on dead mangrove wood and bark. Sexual
12. Ascomata forming short papilla protruding the host morph: Ascomata immersed in wood, solitary, light coloured
surface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13 or dark brown to black, subglobose to globose or pyriform,
13. Peridium thin-walled, composed of a single layer of elongate, clypeate, coriaceous, ostiolate, papillate,
melanized cells, difficult to separate from the periphysate. Peridium composed of several layers of brown
entostroma…………………………………. Leptoperidia to dark brown cells of textura angularis. Paraphyses numer-
13. Peridium thick-walled, compose of two distinct layers, ous, septate, rarely branched in a gelatinous matrix, hyaline.
separating from entostroma . . . . . . . . . . . . . . . . . . . . 14 Asci 8-spored, unitunicate, thin-walled, persistent, cylindrical
14. Asci generally spindle-shaped, with sub-olivaceous to to oblong, pedicellate, J−, sometimes apically thickened.
brown ascospores. . . . . . . . . . . . . . . . . . Cryptosphaeria Ascospores 1–2-seriate, clavate, ellipsoidal, 1–3-septate, hya-
14. Asci cylindric-clavate, with pale yellow asco- line, some slightly constricted at the septa, with or without
spores………………………………………......... Eutypa appendages. Asexual morph: Undetermined.
Type species: Etheirophora bijubata Kohlm. & Volkm.-
Etheirophoraceae Rungjind. et al., in Jones et al., Cryptog. Kohlm., Mycol. Res. 92(4): 414 (1989)
Mycol. 35(2): 134 (2014) Notes: Kohlmeyer and Volkmann-Kohlmeyer (1989) intro-
Facesoffungi number: FoF 01281 duced the genus Etheirophora (type species E. bijubata) to
Fungal Diversity

Fig. 54 Etheirophora bijubata (Material examined: USA, Hawaii, 01315470, microslide from holotype) a Herbarium material label b–d
Haena Beach, Hanalei, Kauai (Pacific Ocean), 22° 13′ 18″N, 159° 34′ Ascomata cross sections e Section of apical region of ascoma f
16″W (22.222, −159.571), intertidal wood embedded in concrete, Paraphyses g–h Asci i–m Ascospores with apical appendages. Scale
collected by J. Kohlmeyer 5098 with B. Kohlmeyer; 8 May 1987, NY bars: b–d = 100 μm, e = 50 μm, f–h = 20 μm, i–m = 10 μm

accommodate three marine species from tropical locations, blepharospora Kohlm. & E. Kohlm. The genus was assigned
including a species previously referred to as Keissleriella to the order Sphaeriales by Kohlmeyer and Volkmann-
 Fungal Diversity

Kohlmeyer (1989) and to the Halosphaeriales by molecular study, by comparing sequences of F. thailandicum
Hawksworth et al. (1995) and Kirk et al. (2001). Schoch with sequences in GenBank, Crous et al. (2007) suggested an
et al. (2007), based on molecular and morphological data, affinity with the order Hypocreales. Jones et al. (2014)
referred it to the TBM clade comprising Bertia, showed that the genus Falcocladium is monophyletic and is
M e l a n o s p o r a a n d To r p e d o s p o r a , w i t h i n well-separated from the order Hypocreales. Jones et al. (2014)
Hypocreomycetidae, with affinities to the Coronophorales. introduced the family Falcocladiaceae based on SSU and
Etheirophora grouped with a range of unresolved taxa, LSU sequence data to accommodate the genus
Juncigena, Swampomyces and Torpedospora and the asexual Falcocladium and suggested further taxon sampling to deter-
genera Glomerulispora and Moheitospora, in the TBM clade mine its ordinal position. The Falcocladiaceae group in the
with high bootstrap value (Abdel-Wahab et al. 2010). Jones Hypocreomycetidae with the family Etheirophoraceae as a
et al. (2014) demonstrated that the genus Etheirophora and sister subclade, could not be accommodated in any order
two Swampomyces species formed a distinct clade in the (Jones et al. 2014), but subsequently Maharachchikumbura
Hypocreomycetidae and introduced the family et al. (2015) introduced the order Falcocladiales to accommo-
Etheirophoraceae. date the family Falcocladiaceae.

Falcocladium S.F. Silveira et al., Mycotaxon 50: 447 (1994)

Other genera included Facesoffungi number: FoF 02108; Fig. 55
Saprobic on leaves, and leaf litter, associated with tropical
Swampomyces Kohlm. & Volkm.-Kohlm., Bot. Mar. 30(3): forests. Sexual morph: Undetermined. Asexual morph:
198 (1987) Conidiomata sporodochial or synnematal or penicilliate when
Type species: Swampomyces armeniacus Kohlm. & formed on aerial mycelium, single to aggregate, arising direct-
Volkm.-Kohlm., Bot. Mar. 30(3): 200 (1987) ly from the mycelium or from a stroma of thick walled, red-
brown chlamydospores, with hyaline, aseptate, thick walled,
stipe extensions, arising from any position on a conidiomatal
Key to the genera of Etheirophoraceae branch, or in the position of phialide, sometimes more than
one occurring in the same penicilliate conidioma.
1. Ascospores with polar appendages Etheirophora Conidiophores up to three series of branches, branches hya-
1. Ascospores lacking appendages Swampomyces line, aseptate to multi-septate, subcylindrical. Conidiogenous
Falcocladiaceae Somrithipol et al., in Jones et al., cells in groups of 2–6, ampulliform, with inconspicuous
Cryptog. Mycol. 35(2): 134 (2014) collarettes. Conidia hyaline, aseptate to 1-septate, falcate, with
Facesoffungi number: FoF 01288 acute, short apical and basal appendages.
Saprobic on leaf litter and leaves. Sexual morph: Type species: Falcocladium multivesiculatum S.F. Silveira
Undetermined. Asexual morph: Hyphomycetous. et al., Mycotaxon 50: 448 (1994).
Conidiomata sporodochial or synnematal or penicilliate, di- Notes: Falcocladium species are found on a wide range of
rectly arising from the mycelium or a stroma or from host substrates including Eucalyptus grandis,
microsclerotia, thick-walled, with aseptate, stipe extensions, E. camaldulensis leaves and leaf litter associated with tropical
that terminates in hyaline, thin-walled sphaeropendunculate forests (Somrithipol et al. 2007). Unique characters of the
vesicles. Conidiophores up to three series of branches per genus Falcocladium include white sporodochia, thick-walled,
conidiomata, branches hyaline, aseptate to multi-septate, aseptate stipe extensions, that terminate in thin-walled vesicles
smooth-walled, subcylindrical. Conidiogenous cells in and conidia that are hyaline, aseptate to 1-septate, falcate and
whorls, ampulliform with elongated necks and minute with short apical and basal appendages (Crous et al. 1994;
collarettes, phialidic. Conidia hyaline, aseptate to 1-septate, Somrithipol et al. 2007). Species are delineated by the mor-
smooth-walled, falcate, with short apical and basal phology of the terminal vesicle, conidial measurements and
appendages. septation (Somrithipol et al. 2007). A multi-gene molecular
Type: Falcocladium S.F. Silveira et al. study established the monophyly of Falcolcadium species,
Notes: Although the morphology of the genus and their inclusion in a new family Falcocladiaceae (Jones
Falcocladium has been well-studied and documented (Crous et al. 2014) and ordinal placement in the Falcocladiales
et al. 1994, 1997, 2007; Somrithipol et al. 2007), studies at the (Maharachchikumbura et al. 2015). The order forms a sister
molecular level are few. Crous et al. (1994) described the clade to the Coronophorales and Melanosporales with high
genus Falcocladium with F. multivesiculatum S.F. Silveira statistical support (Maharachchikumbura et al. 2015).
et al. as the type species and subsequently introduced further
new species in F. sphaeropedunculatum Crous & Alfenas and Flammocladiaceae Crous et al., in Crous et al., Sydowia 67:
F. thailandicum Crous & Himaman. In a preliminary 103 (2015)
Fungal Diversity

Fig. 55 Falcocladium
multivesiculatum (Material
examined: BRAZIL. Espirito
Santo, Aracruz, from Eucalyptus
grandis W. Hill ex Maiden
(Myrtaceae) leaf litter, Silvaldo F.
Silveira, January 1993, PREM
51541, holotype) a Herbarium
material of F. multivesiculatum
(dried culture on MEA) b, c Stipe
extensions d Conidiomata on
mycelium e Conidia. Scale bars:
b, c = 100 μm, d, e = 5 μm

Facesoffungi number: FoF 01902; Fig. 56 at septa. Asexual morph: Coelomycetous. Conidiomata
Saprobic on twigs of Acer platanoides Sexual sporodochial, determinate, hyaline, becoming orange.
morph: Ascomata perithecial, pale luteous to yellow-or- Conidiophores subcylindrical, septate, branched.
ange, aggregated in clusters, linked by a stromatic base, Conidiogenous cells subcylindrical, terminal and interca-
covered in a dirty white crustose layer, not discolouring lary, hyaline, smooth, proliferating sympodially at apex.
in 3 % KOH, turning pale luteous to dirty white, with a Conidia slimy, solitary, hyaline, smooth, granular to
characteristic papillate, periphysate ostiolar area; wall of guttulate, straight to gently curved, subcylindrical to
smooth, 3–4 layers of subhyaline textura angularis. Asci narrowly obclavate (Crous et al. 2015b).
8-spored, fusoid-ellipsoidal to subclavate, unitunicate. Type: Flammocladiella Crous et al. in Crous et al.,
Ascospores fasciculate, hyaline, fusoid-ellipsoidal with Sydowia 67: 103 (2015)
obtuse ends, septate, warty, not or slightly constricted Type species: Flammocladiella aceris Crous et al.
 Fungal Diversity

Fig. 56 Flammocladiella aceris (CBS 138906, ex-type) (redrawn from Crous et al. 2015b). a Ascomata arranged in rosette on host tissue b Asci c
Ascospores d, e Conidiophores with conidiogenous cells f Conidia. Scale bars: a = 250 μm, b–e = 10 μm

Notes: Based on LSU sequence data Crous et al. (2015b) to conspicuous. Conidia unicellular, hyaline, cylindrical, cla-
introduced Flammocladiaceae in the Hypocreales. vate, falcate, forming appressoria when germinating.
Flammocladiella has phylogenetic affinities with members Type: Colletotrichum Corda
of the Clavicipitaceae and Ophiocordycipitaceae. However Notes: Glomerellaceae is a monotypic family comprising
it is easily distinguished from other members of Hypocreales mainly pathogens. Chadefaud (1960) introduced the ordinal
based on its yellowish ascomata that aggregate in clusters on a name Glomerellales including Colletotrichum (= Glomerella)
single stromatic base, covered by a crustose layer, having a and three other genera in a non-ranked group “Eu-
papillate, periphysate ostiolar region, and a sporodochial asex- Glomérellales”, but this was not validly published.
ual morph forming flame-like conidial masses (Crous et al. Previously, Colletotrichum was placed in family
2015b). Phyllachoraceae, but has long been considered to be an out-
lier due to its non-stromatic nature (Cannon 1991). The family
Glomerellaceae Locq. ex Seifert & W. Gams, in Zhang et al. Glomerellaceae was introduced by Locquin (1984), but was
Mycologia 98(6): 1083 (2007) invalidly published. Uecker (1994), based on preliminary
Facesoffungi number: FoF 01100 sequence-based studies, along with ontogenetic studies, con-
Parasitic, endophytic and saprobic on plant leaves, stems firmed that Colletotrichum does not belong in the same order
and fruits. Sexual morph: Ascomata solitary or gregarious, as Phyllachora. Kirk et al. (2001) placed Glomerellaceae with
globose to subglobose, dark brown to black, ostiole an uncertain position in Sordariomycetidae. Zhang et al.
periphysate. Peridium composed of pale to medium brown, (2006) validated the family Glomerellaceae with a Latin de-
flattened cells of textura angularis. Hamathecium composed scription, while placing it within the Hypocreomycetidae. Kirk
of paraphyses. Asci 8-spored, unitunicate, cylindrical to et al. (2008) placed this family in an uncertain position in the
subfusoid, short pedicellate, with a refractive, J-, apical ring. subclass Hypocreomycetidae. Réblová et al. (2011) further
Ascospores uniseriate to overlapping biseriate, hyaline, uni- elucidated the phylogenetic position of Glomerellaceae by
cellular, oval, fusiform or rhomboid. Asexual morph: analysis of combined ITS, LSU, SSU and RPB2 sequence
Coelomycetous. Conidiomata acervular, conidiophores and data. The order Glomerellales was validated by Réblová
setae formed on cushions of pale to medium brown, roundish et al. (2011), who provided a Latin diagnosis. Two new fam-
to angular cells, comprising thick-walled hyphae. Setae ilies, Australiascaceae and Reticulascaceae occupied a com-
straight, constricted and slightly wavy. Conidiophores hyaline mon clade with Glomerellaceae (Réblová et al. 2011) in
to pale brown. Conidiogenous cells enteroblastic, hyaline to Glomerellales.
pale brown, cylindrical to ellipsoidal, doliform to The family Glomerellaceae was established based on the
ampulliform, collarette distinct, periclinal thickening visible genus Glomerella (Zhang et al. 2006), which had been
Fungal Diversity

synonymized under its asexual morph Colletotrichum Peridium composed of pale to medium brown flattened cells
(Maharachchikumbura et al. 2015). of textura angularis. Hamathecium composed of hyaline, sep-
tate paraphyses, branched at the base, rounded at the tips. Asci
Colletotrichum Corda, in Sturm, Deutschl. Fl., 3 Abt. (Pilze 8-spored, unitunicate, cylindrical to subfusoid, short pedicel-
Deutschl.) 3(12): 41 (1831) late, with an inamyloid, refractive ring at the apex. Ascospores
Facesoffungi number: FoF 01094; Figs 57, 58 uni- to biseriate, aseptate, hyaline, oval, fusiform or rhomboid,
Parasitic, endophytic and saprobic on plant leaves, stems, one end ± acute and one ended round or both ends rounded,
fruits. Sexual morph: Ascomata solitary or gregarious, glo- sometimes slightly curved, smooth-walled. Asexual morph:
bose to subglobose, dark brown to black, ostiole periphysate. Coelomycetous. Conidiomata acervular, conidiophores and

Fig. 57 Colletotrichum ti (Material examined: NEW ZEALAND, setae d Base of the setae e Conidiophores and conidiogenous cells f
Taupo, Orakei Korako on Cordyline australis (Forst. f.) Hook. f. Conidiophores and conidiogenous cells stained with Congo Red reagent
(Asparagaceae), July 1965, R.F.R. McNabb, PDD 30206, holotype). a, g Hyaline conidia with broadly rounded ends h Appressoria i Ex-
b, e Holotype c, d, f, g, h, i Ex-holotype culture a Herbarium details b holotype culture. Scale bars: d = 20 μm, c, e–g = 10 μm, h = 5 μm
Lesions of the dried herbarium specimen with black conidiomata c Young
 Fungal Diversity

Fig. 58 Sexual morph of

Colletotrichum karstii (Holotype
redrawn from Yang et al. 2011) a
Immature asci b Mature ascus c
Hyaline ascospores. Scale bars:
a–c = 10 μm

setae formed on cushions of pale to medium brown, roundish within a molecular phylogenetic system was carried out with
to angular cells. Setae may or may not be present, if present the use of 18S rDNA sequence data (Illingworth et al. 1991;
usually emerging from darkened hyphae, straight, constricted Berbee and Taylor 1992). Winka and Eriksson (2000) consid-
and slightly wavy, hyaline, pale brown to medium brown, ered Colletotrichum to be more closely related to
dark chestnut to black, basal cell often paler, hyaline towards Hypocreomycetidae. The study of Wanderlei-Silva et al.
the tip, smooth-walled or verruculose, towards the tip often (2003) showed Colletotrichum to be a sister group to
verruculose, 1–6-septate, often septate only at the base, base Hypocreales. Zhang et al. (2006) confirmed the phylogenetic
cylindrical, conical or slightly inflated, tip ± rounded to ± position of Colletotrichum within the Hypocreomycetidae.
acute. Conidiophores hyaline to pale brown, simple or septate, Maharachchikumbura et al. (2015) also confirmed the posi-
branched or unbranched, smooth-walled, Conidiogenous cells tion of Colletotrichum in the family Glomerellaceae.
enteroblastic, hyaline to pale brown, smooth-walled, cylindri- Colletotrichum species are phytopathogens causing anthrac-
cal, ellipsoidal, doliform or ampulliform, collarette distinct, nose disease of many crops and fruits worldwide (Hyde et al.
periclinal thickening visible to conspicuous, Conidia hyaline, 2009a, b; Cannon et al. 2012b; Hyde et al. 2014). Species of
smooth-walled, aseptate, cylindrical, clavate, falcate, straight Colletotrichum are also important as endophytes of living plant
or slightly curved, apex acute to rounded, sometimes reduced tissues (Manamgoda et al. 2013; Hyde et al. 2014). Species of
in to filiform appendage, base rounded to truncate, forms ap- Colletotrichum have been studied extensively as model organ-
pressoria when germinating. isms in genetic research (Cannon et al. 2012b). Some species of
Type species: Colletotrichum lineola Corda, in Sturm, Colletotrichum have been defined using ITS sequences data, but
Deutschl. Fl., 3 Abt. (Pilze Deutschl.) 3(12): 41 (1831) ITS alone is insufficient for resolving Colletotrichum species
Notes: The genus Colletotrichum was introduced by Corda well. Multi-marker phylogenetic analysis, epitypification and
(1831) for C. lineola Corda (Damm et al. 2009; Cannon et al. knowledge of species complexes have contributed to a better
2012b). Colletotrichum being an asexual fungal genus was understanding of the genus (Cai et al. 2009; Hyde et al. 2014).
included in morphological classifications of the Ascomycota
due to its sexual genus Glomerella (Cannon et al. 2012b). Gnomoniaceae G. Winter [as ‘Gnomonieae’], Rabenh.
Ainsworth (1971) listed Colletotrichum as a member of the Krypt.-Fl., Edn 2 (Leipzig) 1.2: 570 (1886)
family Phyllachoraceae. However, Barr (1976a) included Facesoffungi number: FoF 01903
Colletotrichum in the family Melogrammataceae, but Saprobic on bark and leaves of overwintered plants.
Hawksworth et al. (1983) placed Colletotrichum in its tradi- Sexual morph: Ascomata immersed to erumpent, solitary or
tional position in the family Phyllachoraceae, which was aggregated, globose to subglobose, black, coriaceous, thin-
adopted by Hawksworth et al. (1995). However, due to its walled, with one or more long, central or eccentric necks with
astromatic nature Colletotrichum had been considered to be hyaline periphyses. Peridium comprising few layers of brown,
an outlier within the family Phyllachoraceae (Cannon et al. thick-walled cells of textura angularis. Hamathecium com-
2012b). Preliminary studies together with ontogenetic re- prising hyaline, septate, cellular paraphyses. Asci 8-spored,
search confirmed that Colletotrichum and Phyllachora do unitunicate, oval, fusiform to almost filiform, short stalked,
not belong in the same family (Uecker 1994). Kirk et al. with a distinct, J- apical ring. Ascospores biseriate, overlap-
(2001, 2008) placed Colletotrichum in the family ping uniseriate to fasciculate, oval, fusiform, ovoid to subu-
Glomerellaceae. The first attempt to place Colletotrichum late, small, unicellular to 1-septate, rarely multi-septate, ends
Fungal Diversity

mostly rounded, rarely pointed, appendages absent or subu- species reported as pathogens and endophytes in leaves of
late, navicular or whip-shaped, smooth. Asexual morph: herbaceous or woody trees (Rossman et al. 2007). Mejía
Conidiomata formed on the bark together with perithecia, et al. (2011) reported Betulaceae, Fagaceae, and Salicaceae
acervuli, subcuticular, flat. Conidiophores simple, hyaline, as common host families. Gnomoniaceae groups with
annellations not clearly visible. Conidiogenous cells phialidic. Melanconidaceae as a sister clade in Diaporthales, and com-
Conidia one-celled, acrogenous, filiform or fusiform, curved, prises 32 genera (Maharachchikumbura et al. 2015).
hyaline, thin walled, obtusely crescent.
Type: Gnomonia Ces. & De Not. Gnomonia Ces. & De Not., Comm. Soc. crittog. Ital. 1(4):
Notes: Gnomoniaceae was introduced by Winter in 1886 231 (1863)
with Gnomonia gnomon (Tode) J. Schröt., as the type. This Facesoffungi number: FoF 02109; Fig. 59
family is characterized by immersed, rarely erumpent or su- Saprobic on overwintered, fallen or attached leaves of
perficial ascomata, without a stroma, or aggregated with a shrubs, usually epiphyllous or on petioles, rarely hypophyllous.
rudimentary stroma. Gnomoniaceae comprises microfungal Sexual morph: Ascomata perithecial, solitary, without stroma

Fig. 59 Gnomonia gnomon (Material examined: FINLAND, Helsinki, Ascomata on substrate e Rehydrated ascoma in water f Cross section of
Helsinki University Botanical Garden, overwintered fallen leaves of ascoma g–h Peridium i-l Asci m-p Ascospores. Scale bars: c = 1 mm,
Corylus avellana L. (Betulaceae), 2004 April 19, Shchigel, Dmitry S., d = 200 μm, e = 1 mm, f = 20 μm, g, h = 10 μm, i–l = 20 μm, m–
BPI 844273, epitype) a Packet of herbarium b Herbarium specimen c, d p = 10 μm
 Fungal Diversity

or sometime very poorly developed stromatic tissues, black, Clypeoporthe Höhn., Sber. Akad. Wiss. Wien, Math. -naturw.
initially immersed, later become erumpent, rarely partly Kl., Abt. 1 128: 584 (1919)
erumpent, sometimes wide opening with pinkish white powdery Type species: Clypeoporthe monocarpa Höhn., Sber.
collar around the neck, globose to subglobose when moist, Akad. Wiss. Wien, Math. -naturw. Kl., Abt. 1 128: 584 (1919)
bowl-shaped when dry, with 1–3 necks Necks central to margin- Cryptosporella Sacc., Michelia 1(no. 1): 30 (1877)
al, never truly lateral, slightly curved, longer, sometimes almost Type species: Cryptosporella hypodermia (Fr.) Sacc.,
absent. Asci 8-spored, unitunicate, oval to fusiform, short (1877)
stalked, with J- apical ring. Ascospores overlapping uniseriate Cylindrosporella Höhn., Sber. Akad. Wiss. Wien, Math. -
or irregularly multi-seriate, one septate, fusiform to acerose, ends naturw. Kl., Abt. 1 125(1–2): 96 (1916)
narrowly rounded, appendages ovoid, subulate or acicular with Type species: Cylindrosporella carpini (Lib.) Höhn., Sber.
diffuse ends or rarely absent. Asexual morph: Undetermined. Akad. Wiss. Wien, Math.-naturw. Kl., Abt. 1 125(1–2): 96
Type species: Gnomonia gnomon (Tode) J. Schröt., in (1916)
Cohn, Krypt.-Fl. Schlesien (Breslau) 3.2(4): 390 (1897) [1908] Depazea Fr., Observ. mycol. (Havniae) 2: 365 (1818)
Notes: Gnomonia was introduced by Cesati and De Notaris Type species: Depazea frondicola Fr., Observ. mycol.
(1863) and typified by Gnomonia gnomon. This genus is char- (Havniae) 2: 365 (1818)
acterized by having non-stromatic solitary, thin-walled, im- Diplacella Syd., Annls mycol. 28(1/2): 101 (1930)
mersed perithecia with long necks and ascospores with one Type species: Diplacella paulliniae (Gonz. Frag. & Cif.)
median septum. Species of Gnomonia occur on overwintered Syd., Annls mycol. 28(1/2): 101 (1930)
leaves and plant twigs. Gnomonia comprises 273 species Ditopella De Not., Sfer. Ital.: 42 (1863)
(Index Fungorum 2016). Type species: Ditopella fusispora De Not., Sfer. Ital.: 48
(1863)
Ditopellopsis J. Reid & C. Booth, Can. J. Bot. 45(9): 1479
Other genera included (1967)
Type species: Ditopellopsis clethrae J. Reid & C. Booth,
Alnecium Voglmayr & Jaklitsch, Persoonia, Mol. Phyl. Evol. Can. J. Bot. 45(9): 1479 (1967)
Fungi 33: 76 (2014) Gloeosporidina Petr., Annls mycol. 19(3–4): 214 (1921)
Type species: Alnecium auctum (Berk. & Broome) Type species: Gloeosporidina moravica Petr., Annls
Voglmayr & Jaklitsch, Persoonia, Mol. Phyl. Evol. Fungi mycol. 19(3–4): 214 (1921)
33: 76 (2014) Gnomoniella Sacc., Michelia 2(no. 7): 312 (1881)
Ambarignomonia Sogonov, Stud. Mycol. 62: 35 (2008) Type species: Gnomoniella tubiformis (Tode) Sacc. [as
Type species: Ambarignomonia petiolorum (Schwein.) ‘Gnomoniella tubaeformis’], (1882)
Sogonov, Stud. Mycol. 62: 36 (2008) Gnomoniopsis Berl., Icon. fung. (Abellini) 1(3): 93 (1893)
Amphiporthe Petr., Sydowia 24(1–6): 257 (1971) [1970] Type species: Gnomoniopsis chamaemori (Fr.) Berl., Icon.
Type species: Amphiporthe hranicensis (Petr.) Petr., fung. (Abellini) 1(3): 93 (1893)
Sydowia 24(1–6): 257 (1971) [1970] Mamiania Ces. & De Not., Comm. Soc. crittog. Ital. 1(4): 210
Anisomyces Theiss. & Syd., Annls mycol. 12(3): 270 (1863)
(1914) Type species: Mamiania fimbriata (Pers.) Ces. & De Not.,
Type species: Anisomyces papilloideoseptatus (Henn.) Comm. Soc. crittog. Ital. 1(4): 211 (1863)
Theiss. & Syd., Annls mycol. 12(3): 270 (1914) Millerburtonia Cif., Mycopath. Mycol. appl. 6(1): 26
Apiognomonia Höhn., Ber. dt. bot. Ges. 35: 635, 637 (1917) (1951)
Type species: Apiognomonia veneta (Sacc. & Speg.) Type species: Millerburtonia oyedaeae Cif., Mycopath.
Höhn., Hedwigia 62: 47 (1920) Mycol. appl. 6(1): 27 (1951)
Apioplagiostoma M.E. Barr, Mycol. Mem. 7: 101 (1978) Occultocarpon L.C. Mejía & Zhu L. Yang, in Mejía et al.,
Type species: Apioplagiostoma populi (E.K. Cash & Fungal Divers 52(1): 101 (2012)
Waterman) M.E. Barr, Mycol. Mem. 7: 102 (1978) Type species: Occultocarpon ailaoshanense L.C. Mejía &
Asteroma DC., in de Candolle & Lamarck, Fl. franç., Edn 3 Zhu L. Yang, in Mejía et al. Fungal Divers 52(1): 101 (2012)
(Paris) 5/6: 162 (1815) Ophiognomonia (Sacc.) Sacc., Syll. fung. (Abellini) 14(1):
Type species: Asteroma phyteumae DC., in de Candolle & 613 (1899)
Lamarck, Fl. franç., Edn 3 (Paris) 6: 162 (1815) Type species: Ophiognomonia melanostyla (DC.) Sacc.,
Bagcheea E. Müll. & R. Menon, Phytopath. Z. 22(4): 417 Icon. fung. (Abellini) 2: 146 (1899)
(1954) Phragmoporthe Petr., Annls mycol. 32(5/6): 354 (1934)
Type species: Bagcheea castaneae E. Müll. & R. Menon, Type species: Phragmoporthe ploettneriana (Henn.) Petr.,
Phytopath. Z. 22(4): 418 (1954) Annls mycol. 32(5/6): 354 (1934)
Fungal Diversity

Phylloporthe Syd., Annls mycol. 23(3/6): 348 (1925) 7. Ascomata forming under clypeus….........…. Clypeoporthe
Type species: Phylloporthe evernoniae Syd., Annls mycol. 7. Ascomata do not form under clypeus…....................…… 8
23(3/6): 349 (1925) 8. Ascospores slightly isthmoid with a median septum, often
Plagiostoma Fuckel, Jb. nassau. Ver. Naturk. 23–24: 118 readily separating as part spores…..............… Pleuroceras
(1870) [1869–70] 8. Ascospores non-isthmoid, not separating into part
Type species: Plagiostoma euphorbiae (Fuckel) Fuckel, spores….......…...................................……..…..………. 9
Jb. nassau. Ver. Naturk. 23–24: 118 (1870) [1869–70] 9. Appendages generally present….................................… 10
Pleuroceras Riess, Hedwigia 1(6): 25 (1854) 9. Appendages generally absent…………........................... 11
Type species: Pleuroceras ciliatum Riess, Hedwigia 1(6): 10. Perithecia mostly epiphyllous…………… 14
25 (1854) 10. Perithecia mostly hypophyllous……....................... 15
Skottsbergiella Petr., in Skottsberg, Nat. Hist. Juan Fernandez 11. Ascospores with mucilaginous sheath……….. Diplacella
Easter Isl. 2: 481 (1927) 11. Ascospores without mucilaginous sheath…..............… 12
Type species: Skottsbergiella diaporthoides Petr., in 12.Perithecia occurring on both sides of the
Skottsberg, Nat. Hist. Juan Fernandez Easter Isl. 2: 481 (1927) leaf…….............………..…........................ Gnomoniella
Sirococcus Preuss, Linnaea 26: 716 (1855) 12. Perithecia occurring on only upper or lower side of
Type species: Sirococcuss trobilinus Preuss, Linnaea 26: leaf……......……………..……...............................…. 13
716 (1853) 13. Asci inoperculate without conspicuous apical
Spataporthe Bronson et al., Int. J. Pl. Sci. 174(3): 278–292 ring…………...................................…........ Spataporthe
(2013) 13. Asci with characteristic apical ring………...............… 16
Type species: Spataporthe taylorii Bronson et al., Int. J. Pl. 14. Appendages ovoid to sabulate……. Ambarignomonia
Sci. 174(3): 278–292 (2013) 14. Appendages cuneiform with diffuse ends or ovoid, subu-
Uniseta Ciccar., Nuovo G. bot. ital. 54: 711 (1948) late acicular………....................................….. Gnomonia
Type species: Uniseta flagellifera (Ellis & Everh.) Ciccar., 15. Ascospores fusiform; arranged irregularly fasciculate or,
Nuovo G. bot. ital. 54: 15 (1947) obliquely in one longitudinal row….....…. Apiognomonia
Xenotypa Petr., Sydowia 9(1–6): 499 (1955) 15.Ascospores oval to filiform; arranged mostly unevenly par-
Type species: Xenotypa aterrima (Fr.) Petr., Sydowia 9(1– allel, also irregularly multi-seriate or obliquely uniseriate,
6): 499 (1955) occasionally parallel….......................…. Ophiognomonia
Zythia Fr., Summa veg. Scand., Section Post. (Stockholm): 16. Necks present…..................................................…..…. 17
407 (1849) 16. Necks absent…….................................… Gloeosporidina
Type species: Zythia resinae (Ehrenb.) P. Karst., Meddn 17. Necks lateral ………………........….....… Gnomoniopsis
Soc. Fauna Flora fenn. 16: 104 (1890) [1889] 17. Necks central ………………....................................… 18
18.Ectostromatic disc present; dark brown to
black………………………………...……….. Ditopella
Key to genera of Gnomoniaceae 18. Ectostromatic disc absent……….........................……. 19
19. Ascospores not apiosporous…......……....… Plagiostoma
1. Sexual morph known . . . . . . . . . . . . . . . . . . . . . . . . . . 2 19. Ascospores apiosporous…............................... Mamiania
2. Sexual morph not known; asexual morph common in na- 20. Stromata plectenchymatous ……................ Phylloporthe
ture…………………....................……….........………. 29 20. Stromata pseudoparenchymatous…............................... 21
3. Life mode parasitic……...............……...... Millerburtonia 21. Ectostromatic disc present; rectangular, hexagonal to ir-
2. Life mode saprobic, or endophytic……………....…..…. 3 regularly-shaped..………........................… Ditopellopsis
3. Pseudostromata / stromata absent; perithecia immersed in 21. Ectostromatic disc absent………………….................. 22
host tissues…………………........................… 4 22.Mycelial clumps at the base of perithecia ……
4. Pseudostromata / stromata present; perithecia immersed in ………………………..........................… Occultocarpon
stromatic tissues……...................……….........................20 22.Mycelial clumps not at the base of perithecia ...……… 23
4. Pseudostromata / stromata present; perithecia immersed in 23.Cytoplasm of ascospores granular and divided into two
stromatic tissues…………...………...………...………... 5 parts with wide vacuous space forming diplastic polari-
4. Perithecia aggregated; necks parallel to substrate and not ty…………………………………….......….... Bagcheea
fused . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cryptosporella 23.Cytoplasm of ascospores not divided into two parts…..24
5. Infected lesions distinct and colourful . . . . . . . . . . . . . . 6 24. Ascospores unicellular…….….….….….….…..….…. 25
5. Infected lesions indistinct……..…………….................... 7 24. Ascospores multi-cellular.…..…..…..…..…..…..….… 26
6. Infected leaves with dark purple to brown pigmenta- 25.Ascospores elongate allantoid to cylindri-
tion………….......................................... Apioplagiostoma cal….…..…..…..…..…..…..…..…..….......…. Xenotypa
6. Infected leaves with tan to grey pigmentation…..… Zythia 25.Ascospores oval to short allantoid….......… Amphiporthe
 Fungal Diversity

26. Ascospores not constricted at the septa……….......…. 27 conidiogenous loci are located at the base of the shallow
26. Ascospores slightly constricted at the septa…….....…. 28 collarette (Kolařík and Hulcr 2008). Réblová et al. (2011)
27. Ascospores 1-septate………................…. Skottsbergiella placed this family in the order Microascales based on
27. Ascospores 3-septate………..........……. Phragmoporthe analysis of SSU and a combined dataset of LSU, SSU
28. Ascospores not apiosporous; ….………......…. Alnecium and RPB2 sequence data and this was followed by
28. Ascospores apiosporous; ………….…....…. Anisomyces Maharachchikumbura et al. (2015).
29. Conidia hyaline………….............................................. 30
29. Conidia brown……..…....................................… Uniseta Gondwanamyces G.J. Marais & M.J. Wingf., Mycologia
30. Conidiomata pycnidia………........................................ 31 90(1): 139 (1998)
30. Conidiomata acervuli……………............................… 32 Facesoffungi numbers: FoF 02228; Figs 60, 61
31. Conidia released as chains ……………............. Depazea Pathogenic on various Protea species, occurring in the
31. Conidia release one at a time ……………....... Asteroma infructescences and parasitic on bark beetles. Sexual
32. Conidia spindle-shaped, 1-septate…........….. Sirococcus morph: Ascomata globose to subglobose, black, with long
32.Conidia filiform to fusiform, unicellu- neck, tapered towards the apex, terminating in short and
lar……………………………….…… Cylindrosporella divergent, 5–11 ostiolar hyphae. Asci evanescent, hyaline.
Ascospores hyaline, aseptate, fusiform, with or without a
Gondwanamycetaceae Réblová et al., Stud. Mycol. 68(1): sheath, sometimes gelatinous sheath giving a lunate to
188 (2011) falcate appearance to the ascospores. Asexual morph:
Facesoffungi number: FoF 01282 Conidiophores macronematous, mononematous, oliva-
Pathogenic on plants or parasitic on beetles. Sexual ceous-brown, septate, arising from well-developed rhizoids;
morph: Ascomata perithecial, black, necks relatively long, stipe erect, simple, inflated at the apex. Conidiogenous
tapered towards the apex, terminating in ostiolar hyphae. cells (phialides) produced terminally on conidiophores, dis-
Peridium fragile, thin-walled, interascal tissue absent. Asci 8 crete, ovoid, olivaceous-brown, producing conidia at the
to multi-spored, evanescent. Ascospores hyaline, aseptate, fu- apex and leaving minute collarettes. Conidia holoblastic,
siform to lunate or falcate, with or without a gelatinous sheath. hyaline, one-celled, smooth-walled, cylindrical to allantoid,
Asexual morph: Hyphomycetous. Conidiophores mono- rounded at the apex and truncate at the base, produced in
verticillate or penicillate, brown. Conidiogenous cells mucoid masses at the apex of conidiophores.
phialidic. Conidia aseptate, slimy. Type species: Gondwanamyces proteae Wingf. et al., in
Type: Gondwanamyces G.J. Marais & M.J. Wingf. Marais et al., Mycologia 90(1): 139 (1998)
Notes: The family Gondwanamycetaceae, a strongly Notes: Based on RFPL analyses, cycloheximide sensitivity,
supported monophyletic sister clade to Ceratocystidaceae, cell wall saccharides and morphology of asexual morphs, the
was introduced for the genus Gondwanamyces and its genus Gondwanamyces, was introduced to accommodate
asexual morph Custingophora by Réblová et al. (2011). Ceratocystiopsis proteae M.J. Wingf. et al. and Ophiostoma
Studies of Viljoen et al. (1999) and Kolařík and Hulcr capense M.J. Wingf. & P.S. van Wyk. These fungi were de-
(2008) also documented the phylogenetic relationship of scribed as having asexual morphs in the genus Knoxdaviesia
the asexual genera Knoxdaviesia and Custingophora to (Wingfield et al. 1988; Wingfield and van Wyk 1993), which
the sexual morphs of this family. The morphological char- was synonymized under Custingophora by Kolařík and Hulcr
acters of this clade include the apparent absence of (2008). Gondwanamyces species are characterized by
interascal filaments in the ascomatal centrum and hyaline, ascomata, that are similar to those of species of Ceratocystis
allantoid ascospores, with a hyaline sheath, giving the and Ophiostoma, with globose ascomatal bases and long
spore a falcate to lunate appearance. Sexual morphs of necks, bearing ascospores in slimy masses (Marais et al.
this family have been reported from infructescences of 1998). Phylogenetic studies of Marais et al. (1998) and
Protea (Wingfield et al. 1988; Marais et al. 1998) and Zhang et al. (2006) have shown that these fungi reside in the
from sapwood associated with Scolytidae (bark beetles) order Microascales and are closely related to, but distinct,
(Bright and Torres 2006; Kolařík and Hulcr 2008), which from species of Ceratocystis. Gondwanamyces was first ob-
produce dark, globose ascomata with a long, filiform neck, served in infructescences of Protea spp. infested by insects
evanescent asci, and hyaline, fusiform ascospores, with or (Wingfield et al. 1988; Marais et al. 1998), whereas, some
without a gelatinous sheath. Distinctive morphological recently described species of Gondwanamyces are associated
characters of asexual morphs of Gondwanamyces includes, with Scolytidae (bark beetles) (Bright and Torres 2006;
conidiophores that are erect, darkly pigmented, and paler Kolařík and Hulcr 2008). The genera Gondwanamyces and
towards the apex, which are either monoverticillate, some- Custingophora are currently placed in the family
times with a terminal vesicle or divergently penicillate, Gondwanamycetaceae (Réblová et al. 2011;
with whorls of phialides producing hyaline conidia. The Maharachchikumbura et al. 2015).
Fungal Diversity

Fig. 60 Gondwanamyces
proteae (Material examined:
SOUTH AFRICA, Cape
Province, Stellenbosch, from
flower within inflorescence,
infested by insects, L.J. Strauss, 7
October 1985, PREM 48924,
holotype) a Herbarium specimen
b-d Ascomata on host e Long
neck of ascomata f, g Asci h
Ascospores. Scale bars: b, c,
d = 100 μm. e = 50 μm, f–
h = 5 μm

Other genus included 1. Asexual morph with monoverticillate or penicillate co-

nidiophores, bearing cylindrical to allantoid conidia in mucoid
Custingophora Stolk et al., Persoonia 5(2): 195 (1968) masses Custingophora
Type species: Custingophora olivacea Stolk et al., in Stolk
& Hennebert, Persoonia 5(2): 197 (1968) Graphiaceae Z.W. de Beer, Seifert & M. J. Wingf., CBS
fungal biodiversity series 12: 1–19 (2013)
Facesoffungi number: FoF 01099
Key to genera of Gondwanamycetaceae Saprobic on wood; sometimes causing wounds on trees
bark or beetles. Sexual morph: Undetermined. Asexual
1. Sexual morph with globose ascomatal bases and long morph: Hyphomycetous. Conidiomata macronematous,
necks, bearing fusiform ascospores in slimy masses synnematous, determinate, with dematiaceous, compact,
Gondwanamyces stipes flared at the fertile tip; hyphae of stipe pigmented,
 Fungal Diversity

swollen apex. Conidiogenous cells in several whorls, cylindrical

to subulate, straight or sometimes slightly curved, phialidic,
sometimes with percurrent proliferations. Conidia cylindrical
to ovoid, with rounded apices and subtruncate to truncate bases,
aseptate, hyaline, smooth-walled.
Type species: Graphium penicillioides Corda, Icon. fung.
(Prague) 1: 18 (1837)
Notes: Corda (1837) introduced Graphium with
G. penicillioides as the type species. It is a hyphomycetous
genus, including species with dark synnemata, terminating in
a slimy head of aseptate conidia. Synnematous species with
dry conidia were also introduced to this genus by Saccardo
(1886). Later, more than 120 synnematous fungal species
were added (Seifert and Okada 1993; Index Fungorum
2016). Goidanich (1935) re-described Graphium more or less
in its classical concept, and many of the synnematous asexual
morphs of Ophiostoma were classified under Graphium by
Fig. 61 Knoxdaviesia proteae a Conidiophores b Conidiogenous cells Hedgcock (1906) and Seifert and Okada (1993). Okada
and conidia (re-drawn from Wingfield et al. <CitationRef et al. (2000) designated an epitype for G. penicillioides,
CitationID="CR687>1988). Scale bars; b = 10 μm, a = 50 μm
representing Graphium sensu stricto and the eleven
Graphium species related to Ophiostoma were transferred to
simple, septate, branched. Conidiophores penicillately Pestum (Okada et al. 1998, 2000; Harrington et al. 2001).
branched, with branching in two or three levels, forming Based on available SSU and ITS sequences, Cruywagen
metulae at the tip. Conidiogenous cells in whorls of two to et al. (2010) included eight described and seven undescribed
six, phialidic, with percurrent, annellidic extensions; species together with G. penicillioides in Graphium sensu
conidiogenesis enteroblastic. Conidia produced in a transpar- stricto. These asexual morphs were isolated from stained sap-
ent, slimy droplet, aseptate, cylindrical to obovoid, sometimes wood, wounds on trees, or from bark or ambrosia beetles
slightly curved with age, truncate at base, often with a distinct (Mouton et al. 1994; Okada et al. 1998, 2000; Jacobs et al.
basal frill, hyaline. Rarely synanamorphic with obovoid, 2003; Geldenhuis et al. 2004; Hulcr et al. 2007; Massoumi
pigmented conidia. Alamouti et al. 2007; Cruywagen et al. 2010; Paciura et al.
Type: Graphium Corda 2010). De Beer et al. (2013) provided a description of
Notes: Based on the phylogenetic distance between Graphium sensu stricto based on G. penicillioides and the
Graphium and other families of Microascales, the family seven species that comprised a monophyletic lineage.
Graphiaceae was introduced by de Beer et al. (2013). Separation of Graphium sensu stricto from Microascus and
Graphiaceae is monophyletic (Lackner et al. 2014) and com- its allied genera was supported by multi-gene analyses (Zhang
prises a single genus Graphium, which is a synnematous hy- et al. 2006; Spatafora et al. 2006; Schoch et al. 2009). De Beer
phomycete described by Corda (1837), with G. penicillioides et al. (2013) also confirmed that the distinctive Graphium
as the type species. Maharachchikumbura et al. (2015) also lineage was distinct from Microascus and introduced the
placed Graphium species under the family Graphiaceae, in new family, Graphiaceae.
the order Microascales. Halosphaeriaceae E. Müll. & Arx, ex Kohlm., Can. J. Bot.
50: 1951 (1972)
Graphium Corda, Icon. fung. (Prague) 1: 18 (1837) Facesoffungi number: FoF 01294
Facesoffungi numbers: FoF 02143; Fig. 62 Saprobic on algae, immersed or submersed on phanero-
Saprobic on sapwood; sometimes causing wounds on trees, gams, wood, bark, leaves, and other cellulosic plant remains,
or from bark or ambrosia beetles. Sexual morph: grains of sand, or in calcareous shell fragments, rarely para-
Undetermined. Asexual morph: Hyphomycetous. Colonies sitic or symbiotic, found in marine and freshwater habitats
scattered, black, upright, with abundant, mostly single, some- (oceans and estuaries). Sexual morph: Ascomata subglobose,
times in pairs or triplets, erect synnemata, sometimes with aerial cylindrical or pyriform, hyaline or dark; sometimes subiculate,
mycelium on host surface, with cylindrical, dark brown to black rarely stromatic; superficial or immersed. Ostioles papillate to
stipes and divergent light brown to grey capitula, surrounded by long cylindrical; ostiolar canal with periphyses or pseudopa-
watery conidial masses, at first colourless, then white, but quick- renchyma; rarely without ostioles. Peridium soft,
ly becoming olive brown to almost black. Conidiophores subcarbonaceous or carbonaceous, composed of flattened,
branched, generally biverticillate, with whorls of two to six, with thick- or thin-walled cells. Centre of immature ascomata
Fungal Diversity

Fig. 62 Graphium penicillioides

(Material examined: CZECH
REPUBLIC, České Budějovice,
isolated from wood core of
Populus nigra L. (Salicaceae), 3
September 1998, T. Kirisits, PRM
842988, epitype) a, b Herbarium
material c Colony on the surface
of the substrate d Synnema on the
host surface e Synnema after
spore dispersal f Conidiogenous
cells g Polyblastic conidia. Scale
bars: c, d = 500 μm, e = 20 μm, f,
g = 10 μm

consisting of polygonal, thin-walled, pseudoparenchymatic seriate, hyaline or light brown, 1- multi-celled, mostly with
cells, sometimes with pits, at maturation separating to form characteristic ornamentations, appendages or gelatinous
catenophyses or compressed by the asci and dissolving. sheaths, or both. Asexual morph: Varied types.
Paraphyses absent. Asci fusiform, clavate or rarely Conidiophores present or lacking, septate or lacking septa,
subglobose, with or without apical structures, thin-walled, 1- hyaline to pale brown, conidiogenous cells monoblastic, ter-
layered, persistent or swelling and deliquescing at or before minal, conidia apical, helicoid; or conidiophores
ascospore maturity. Hymenial layer at base of venter, flat or pleurogenous on the mycelium, conidiogenous cells holoblas-
convex. Mature ascospores filling the venter of the ascoma, tic, terminal, sympodial or irregular, denticulate with
released singly through the ostiole or rarely within the ascus, schizolytic secession, conidia aleuriospores C to U-shaped,
which swells after dispersal. Ascospores overlapping 2–3 rarely sigmoid, solitary, septate, hyaline, or conidiophores
 Fungal Diversity

monoblastic, ellipsoidal cylindrical or clavate, septate, hyaline referred to a new family Lulworthiaceae (Campbell et al.
to pale brown, conidia unicellular, thick-walled, pale brown to 2003; Pang et al. 2003; Koch et al. 2007). Hibbett et al.
dark brown, catenulate, or conidiogenous cells monoblastic to (2007) and Schoch et al. (2007) referred Halosphaeriaceae
polyblastic, conidia terminal, clavate, 1–4 septate, thick- to the order Microascales, while others continued to include
walled pale brown to dark brown, or conidia terminal, hyaline, it in the order Halosphaeriales (Zhang et al. 2006; Jones et al.
branched, septate, with main axis give rise to 2–3 side 2009a). Maharachchikumbura et al. (2015) and Jones et al.
branches, typically tetraradiate. (2015) accept the placement of the Halosphaeriaceae as one
Type: Halosphaeria Linder of the families in Microascales. However, the higher level
Notes: The family Halosphaeriaceae, which is the most classification of this family remains controversial.
diverse group of marine ascomycetes, was introduced by Halosphaeria Linder, Farlowia 1: 412 (1944)
Müller and von Arx (1962) and Eriksson (1984) with Facesoffungi numbers: FoF 02142; Figs 63, 64, 65
Halosphaeria as the type genus (Barghoorn and Linder Saprobic on wood in marine environments. Sexual
1944). Members of the Halosphaeriaceae are amongst the morph: Ascomata globose, subglobose, obpyriform, ellip-
most intensively studied of marine ascomycetes at the mor- soidal, immersed to superficial, ostiolate, papillate, coria-
phological, ultrastructural and molecular level, with 161 spe- ceous, pale brown to black, necks variable in length,
cies in 62 genera species (Pang 2002; Jones et al. 2009a, 2015; periphysate, centrum breaking up into deliquescing
Maharachchikumbura et al. 2015). Morphological characters catenophyses. Asci 8-spored, unitunicate, thin-walled, cla-
unique to the family Halosphaeriaceae include: perithecial vate, pedicellate, lacking an apical ring, deliquescing early.
ascomata, necks (usually with periphyses), presence of Ascospores biseriate, hyaline, broad-ellipsoidal, 1-septate,
catenophyses that easily deliquesce, unitunicate, thin-walled not constricted at the septa, with 3–4 spoon-shaped equa-
asci that deliquesce early, asci with or lacking an apical ring torial appendages and one at each end of the spore.
and appendaged ascospores (Jones 1995). Based on scanning Asexual morph: Undetermined.
and transmission electron microscopic studies of ascospore Type species: Halosphaeria appendiculata Linder,
appendage morphology and ontogeny, a number of genera Farlowia 1: 412 (1944)
were found to be polyphyletic and new genera introduced to Notes: The genus Halosphaeria is monotypic with
accommodate them, e.g. Ondiniella, and Marinospora, while H. appendiculata Linder as the type species. Other species
others were reinstated, e.g. Antennospora, Arenariomyces and assigned to the genus by Kohlmeyer (1972) have been trans-
Halosphaeriopsis (Jones et al. 1983, 1984; Jones 1995). ferred to other genera as the result of ultrastructural and mo-
Subsequent molecular studies have supported these changes. lecular studies: Antennospora, Halosphaeriopsis,
The family Halosphaeriaceae was shown to be polyphyletic Lautisporopsis, Okeanomyces, Ondinella, and Remispora
and the genera Kohlmeyeriella, Lulworthia and Lindra (Jones et al. 2009a, 2015).

Fig. 63 Halosphaeria appendiculata (Material examined: TAIWAN, layers of elongated cells with large lumina c Spore passing through ostiole
New Taipei City, Tungpeichiao, rocky shore, on drift wood, 13 April d Clavate ascus e Ellipsoidal ascospore with spoon-shaped appendages at
2012, K. L. Pang) a Immersed ascoma b Peridium composed of a few polar and equatorial positions Scale bar: a, d = 30 μm, b–c, e = 10 μm
Fungal Diversity

Fig. 64 Halosphaeriaceae
selected genera a Aniptodera
longispora (Material examined:
MALAYSIA, Tioman island,
Mangrove, on dead mangrove
wood, 12 July 2010, K. L. Pang)
b Ceriosporopsis halima
(Material examined: TAIWAN,
Keelung, National Taiwan Ocean
University, rocky shore, on drift
wood, 2 March 2012, K. L. Pang)
c Arenariomyces trifurcatus
(Material examined: TAIWAN,
New Taipei City, Hemei, rocky
shore, on drift wood, 22 February
2011, K. L. Pang) d Marinospora
calyptrata (Material examined:
GREAT BRITAIN, Langstone
Harbour, Rocky shore, on drift
wood, 10 August 2010, K. L.
Pang) e Cucullosporella
mangrovei (Material examined:
MALAYSIA, Tioman island,
Mangrove, on dead mangrove
wood, 12 July 2010, K. L. Pang) f
Appendichordella amicta. Scale
bars: a–f = 15 μm

Other genera included Ascosacculus J. Campbell, J.L. Anderson & Shearer,

Mycologia 95 (3): 545 (2003)
Alisea J. Dupont & E.B.G. Jones, Mycol. Res. 113(12): 1358 Type species: Ascosacculus aquaticus (K.D. Hyde) J.
(2009) Campb. et al., Mycologia 95 (3): 545 (2003)
Type species: Alisea longicolla J. Dupont & E.B.G. Jones, Bathyascus Kohlm., Revue Mycol. 41(2): 190 (1977)
Mycol. Res. 113(12): 1358 (2009) Type species: Bathyascus vermisporus Kohlm., Revue
Aniptodera Shearer & M.A. Mill., Mycologia 69(5): 893 (1977) Mycol., Paris 41(2): 191 (1977)
Type species: Aniptodera chesapeakensis Shearer & M.A. Carbosphaerella I. Schmidt, Feddes Repert. 80(2–3): 108
Mill., Mycologia 69(5): 894 (1977) (1969)
Anisostagma K.R.L. Petersen & Jørg. Koch, Mycol. Res. Type species: Carbosphaerella pleosporoides I. Schmidt,
100: 209 (1996) Feddes Repert. 80: 108 (1969)
Type species: Anisostagma rotundatum K.R.L. Petersen & Ceriosporopsis Linder, Farlowia 1: 408 (1944)
Jørg. Koch, Mycol. Res. 100(2): 211 (1996) Type species: Ceriosporopsis halima Linder, Farlowia
Antennospora Meyers, Mycologia 49: 501 (1957) 1(3): 409 (1944)
Type species: Antennospora quadricornuta (Cribb & J.W. Chadefaudia Feldm.-Maz., Revue Générale de Botanique 64:
Cribb) T.W. Johnson, J. Elisha Mitchell Scient. Soc. 74: 46 (1958) 150 (1957)
Appendichordella R.G. Johnson et al., Can. J. Bot. 65(5): 941 Type species: Chadefaudia marina Feldm.-Maz., Rev.
(1987) gén. Bot. 64: 150 (1957)
Type species: Appendichordella amicta (Kohlm.) R.G. Corallicola Volkm.-Kohlm. & Kohlm., Mycotaxon 44(2):
Johnson et al., Can. J. Bot. 65(5): 941 (1987) 418 (1992)
Arenariomyces Höhnk, Veröff. Inst. Meeresf. Bremerhaven 3: Type species: Corallicola nana Volkm.-Kohlm. & Kohlm.,
28 (1954) Mycotaxon 44(2): 418 (1992)
Type species: Arenariomyces trifurcatus Höhnk, Veröff. Corollospora Werderm., Notizbl. Bot. Gart. Berlin-Dahlem:
Inst. Meeresf. Bremerhaven 3: 30 (1954) 248 (1922)
 Fungal Diversity

Fig. 65 Halosarpheia fibrosa (Material examined: BERMUDA, St. 00966760, holotype) a, b Herbarium material of Halosarpheia fibrosa
George Island, Coot Pond, intertidal wood of Avicennia germinans, c Ascomata on host surface d Section through ascomata e Peridium f-i
with teredinids and algae, 14 May 1976, J. J. Kohlmeyer, NY Ascospores. Scale bars: c = 500 μm, d, e = 100 μm. f–i = 20 μm

Type species: Corollospora maritima Werderm., Notizbl. Haiyanga K.L. Pang & E.B.G. Jones, Raffles Bull. Zool.,
Königl. bot. Gart. Museum Berlin 8: 248 (1922) Suppl. 19: 8 (2008)
Cucullosporella K.D. Hyde & E.B.G. Jones, Mycotaxon 37: Type species: Haiyanga salina (Meyers) K.L. Pang &
200 (1990) E.B.G. Jones, Raffles Bull. Zool., Suppl. 19: 8 (2008)
Type species: Cucullosporella mangrovei (K.D. Hyde & Haligena Kohlm., Nova Hedwigia 3: 87 (1961)
E.B.G. Jones) K.D. Hyde & E.B.G. Jones, Mycotaxon 37: Type species: Haligena elaterophora Kohlm., Nova
200 (1990) Hedwigia 3: 87 (1961)
Ebullia K.L. Pang, Mycoscience 56: 40 (2015) Halosarpheia sensu stricto Kohlm. & E. Kohlm., Trans. Br.
Type species: Ebullia octonae (Kohlm.) K.L. Pang, Mycol. Soc. 68(2): 208 (1977)
Mycoscience 56: 40 (2015) Type species: Halosarpheia fibrosa Kohlm. & E. Kohlm.,
Fluviatispora K.D. Hyde, Mycol. Res. 98: 720 (1994) Trans. Br. Mycol. Soc. 68(2): 208 (1977)
Type species: Fluviatispora tunicata K.D. Hyde, Mycol. Halosphaeriopsis T.W. Johnson, J. Elisha Mitchell Scient.
Res. 98: 722 (1994) Soc. 74: 44 (1958)
Gesasha Abdel-Wahab & Nagahama, Nova Hedwigia 92(3– Type species: Halosphaeriopsis mediosetigera (Cribb & J.W.
4): 501 (2011) Cribb) T.W. Johnson, J. Elisha Mitchell Scient. Soc. 74: 44 (1958)
Type species: Gesasha mangrovei Abdel-Wahab & Havispora K.L. Pang & Vrijmoed, Mycologia 100(2): 293
Nagah., Nova Hedwigia 92(3–4): 507 (2011) (2008)
Fungal Diversity

Type species: Havispora longyearbyenensis K.L. Pang & Type species: Neptunella longirostris (Cribb & J.W. Cribb)
Vrijmoed, Mycologia 100(2): 293 (2008) K.L. Pang & E.B.G. Jones, Mycol. Progr. 2(1): 35 (2003)
Iwilsoniella E.B.G. Jones, Syst. Ascomyc. 10(1): 8 (1991) Nereiospora E.B.G. Jones et al., J. Linn. Soc. Bot. 87(2): 204
Type species: Iwilsoniella rotunda E.B.G. Jones, Syst. (1983)
Ascomyc., 10(1): 8 (1991) Type species: Nereiospora comata (Kohlm.) E.B.G. Jones
Kitesporella Jheng & K.L. Pang, Bot. Mar. 55: 462 (2012) et al., J. Linn. Soc. Bot. 87(2): 206 (1983)
Type species: Kitesporella keelungensis J.S. Jheng & K.L. Nimbospora J. Koch, Nordic J. Bot. 2(2): 166 (1982)
Pang, Bot. Mar. 55(5): 462 (2012) Type species: Nimbospora effusa Jørg. Koch, Nordic
Kochiella Sakay. et al., Fungal Divers. 46: 96 (2011) Journal of Botany 2 (2): 166 (1982)
Type species: Kochiella crispa (Kohlm.) Sakay. et al. Nohea Kohlm. & Volkm.-Kohlm., Syst. Ascomyc. 10: 121 (1991)
Jones, Fungal Divers. 46: 96 (2011) Type species: Nohea umiumi Kohlm. & Volkm.-Kohlm.,
Lautisporopsis E.B.G. Jones et al., Mycotaxon 67: 1 Syst. Ascomyc. 10: 122 (1991)
(1998) Oceanitis Kohlm., Revue Mycol. 41(2): 193 (1977)
Type species: Lautisporopsis circumvestita (Kohlm.) Type species: Oceanitis scuticella Kohlm., Revue de
E.B.G. Jones et al., Can. J. Bot. 72(10): 1558 (1994) Mycologie 41 (2): 194 (1977)
Lignincola Höhnk, Veröff. Inst. Meeresf. Bremerhaven 3: 216 Ocostaspora E.B.G. Jones et al., Bot. Mar. 26: 353 (1983)
(1955) Type species: Ocostaspora apilongissima E.B.G. Jones
Type species: Lignincola laevis Höhnk, Veröff. Inst. et al., Bot. Mar. 26(7): 354 (1983)
Meeresf. Bremerhaven 3: 216 (1955) Okeanomyces K.L. Pang & E.B.G. Jones, J. Linn. Soc. Bot.
Limacospora Jørg. Koch & E.B.G. Jones, Can. J. Bot. 73(7): 146(2): 228 (2004)
1011 (1995) Type species: Okeanomyces cucullatus (Kohlm.) K.L.
Type species: Limacospora sundica (Jørg. Koch & E.B.G. Pang & E.B.G. Jones, J. Linn. Soc. Bot. 146(2): 228 (2004)
Jones) Jørg. Koch & E.B.G. Jones, Can. J. Bot. 73(7): 1013 Ondiniella E.B.G. Jones et al., Bot. Mar. 27: 136 (1984)
(1995) Type species: Ondiniella torquata (Kohlm.) E.B.G. Jones
Luttrellia Shearer, Mycologia 70(3): 692 (1978) et al., Bot. Mar. 27(3): 136 (1984)
Type species: Luttrellia estuarina Shearer, Mycologia Ophiodeira Kohlm. & Volkm.-Kohlm., Can. J. Bot. 66 (10):
70(3): 693 (1978) 2062 (1988)
Magnisphaera J. Campbell et al., Mycologia 95(3): 546 Type species: Ophiodeira monosemeia Kohlm. & Volkm.-
(2003) Kohlm., Can. J. Bot. 66(10): 2062 (1988)
Type species: Magnisphaera spartinae (E.B.G. Jones) J. Phaeonectriella R.A. Eaton & E.B.G. Jones, Nova Hedwigia
Campb. et al., Mycologia 95(3): 547 (2003) 19(3–4): 779 (1971) [1970]
Marinospora A.R. Caval., Nova Hedwigia 11: 548 (1966) Type species: Phaeonectriella lignicola R.A. Eaton &
Type species: Marinospora calyptrata (Kohlm.) A.R. E.B.G. Jones, Nova Hedwigia 19(3–4): 779 (1971) [1970]
Caval., Nova Hedwigia 11: 548 (1966) Praelongicaulis E.B.G. Jones et al., gen. nov. Fungal Divers.
Moana Kohlm. & Volkm.-Kohlm., Mycol. Res. 92 (4): 418 73: 54 (2015)
(1989) Type species: Praelongicaulis kandeliae (Abdel-Wahab &
Type species: Moana turbinulata Kohlm. & Volkm.- E.B.G. Jones) E.B.G. Jones et al., Fungal Divers. 73: 54 (2015)
Kohlm., Mycol. Res. 92(4): 418 (1989) Panorbis J. Campb. et al., Mycologia 95(3): 544 (2003)
Morakotiella Sakay., Mycologia 97(4): 806 (2005) Type species: Panorbis viscosus (I. Schmidt) J. Campb.
Type species: Morakotiella salina (C.A. Farrant & E.B.G. et al., Mycologia 95(3): 544 (2003)
Jones) Sakay., Mycologia 97(4): 806 (2005) Pileomyces K.L. Pang & Jheng, Bot. Stud. 53: 536 (2012)
Nais Kohlm., Nova Hedwigia 4: 409 (1962) Type species: Pileomyces formosanus K.L. Pang & J.S.
Type species: Naïs inornata Kohlm., Nova Hedwigia 4: Jheng, Bot. Stud. 53: 536 (2012)
409 (1962) Pseudolignincola Chatmala & E.B.G. Jones, Nova Hedwigia
Natantispora J. Campbell et al., Mycologia 95(3): 543 (2003) 83(1–2): 225 (2006)
Type species: Natantispora retorquens (Shearer & J.L. Type species: Pseudolignincola siamensis Chatmala &
Crane) J. Campbell et al., Mycologia 95(3): 543 (2003) E.B.G. Jones, in Jones, Chatmala & Pang, Nova Hedwigia
Nautosphaeria E.B.G. Jones, Trans. Br. Mycol. Soc. 47(1): 83(1–2): 226 (2006)
97 (1964) Remispora Linder, Farlowia 1(3): 409 (1944)
Type species: Nautosphaeria cristaminuta E.B.G. Jones, Type species: Remispora maritima Linder, Farlowia 1: 410
Trans. Br. Mycol. Soc. 47(1): 97 (1964) (1944)
Neptunella K.L. Pang & E.B.G. Jones, Mycol. Progr. 2(1): 35 Saagaromyces K.L. Pang & E.B.G. Jones, Mycol. Progr.
(2003) 2(1): 35 (2003)
 Fungal Diversity

Type species: Saagaromyces ratnagiriensis (S.D. Patil & 5. Ascospores ellipsoidal or spherical to round. . . . . . . 6
Borse) K.L. Pang & E.B.G. Jones, Mycological Progress 2 5. Ascospores rhomboid. . . . . . . . . . . . . . . . Kitesporella
(1): 35 (2003) 6. Ascospores ellipsoidal . . . . . . . . . . . . . . Chadefaudia
Sablicola E.B.G. Jones et al., Can. J. Bot. 82(4): 486 (2004) 6. Ascospores spherical to round . . . . . . . . . . . . . . . . . 7
Type species: Sablecola chinensis E.B.G. Jones et al., Can. 7. Asci with an apical pore, ascospores globose to
J. Bot. 82(4): 486 (2004) subglobose . . . . . . . . . . . . . . . . . . . . . Thalassogena
Thalassogena Kohlm. & Volkm.-Kohlm., Syst. Ascomyc. 6: 7. Asci lacking an apical pore . . . . . . . . . . . . . . . . . . . 8
223 (1987) 8. Ascomata cream-coloured, with long necks, ascospores
Type species: Thalassogena sphaerica Kohlm. & Volkm.- globose to ellipsoidal . . . . . . . . . . . . . . . Anisostagma
Kohlm., Syst. Ascomyc. 6(2): 225 (1987) 8. Ascomata brown, necks short, ascospores spheri-
Thalespora Chatmala & E.B.G. Jones, Nova Hedwigia 83(1– cal……………………………………...... Iwilsoniella
2): 228 (2006) 9. Ascospores 1-septate . . . . . . . . . . . . . . . . . . . . . . . 10
Type species: Thalespora appendiculata Chatmala & E.B.G. 9. Ascospores 1 to multi-septate. . . . . . . . . . . . . . . . . 50
Jones, in Jones, et al., Nova Hedwigia 83(1–2): 229 (2006) 10. Ascospores with no appendages . . . . . . . . . . . . . . 11
Tinhaudeus K.L. Pang et al., Fungal Divers. 75:160 (2015) 10. Ascospores with polar and/or equatorial appendages or
Type species: Tinhaudeus formosanus K.L. Pang et al., with sheaths . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
Fungal Divers. 75:164 (2015) 11. Asci deliquescing early. . . . . . . . . . . . . . . . . . . . Nais
Tirispora E.B.G. Jones & Vrijmoed, Can. J. Bot. 72(9): 1373 11. Asci persistent . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
(1994) 12. Asci persistent, no retraction of the plasmalemma 13
Type species: Tirispora unicaudata E.B.G. Jones & 12. Asci with plasmalemma retracted . . . . . . . . . . . . . 14
Vrijmoed, Can. J. Bot. 72(9): 1373 (1994) 13. Ascus tip thimble-shaped, or slightly thick-
Toriella Sakay., et al., Fungal Divers. 46(1): 99 (2011) ened…………………………………….. Lignincola
Type species: Toriella tubulifera (Kohlm.) Sakay. et al., 13. Ascus tip lacking thimble-shaped apical thicken-
Fungal Divers. 46(1): 100 (2011) ing…........……………………………………. Alisea
Trailia G.K. Sutherl., Trans. Br. Mycol. Soc. 5: 149 (1915) 14. Ascospores without unfurling bipolar append-
Type species: Trailia ascophylli G.K. Sutherl., Trans. Br. ages……………… Aniptodera
Mycol. Soc. 5(1): 149 (1915) 14. Ascospores with a faint sheath . . . . . . . . . Neptunella
Trichomaris Hibbits et al., Can. J. Bot. 59(11): 2123 (1981) 15. Ascospores with a single polar appendage . . . . . . 16
Type species: Trichomaris invadens Hibbits et al., Can. J. 15. Ascospores with bipolar appendages, or polar and
Bot. 59(11): 2123 (1981) equatorial appendages . . . . . . . . . . . . . . . . . . . . . 20
Tubakiella Sakay., et al., Fungal Divers. 46: 97 (2011) 16. Ascospores with a hamate polar appendage . . . . . 17
Type species: Tubakiella galerita (Tubaki) Sakay. et al., 16. Ascospores with an ephemeral drop of polar mucilage,
Fungal Divers. 46: 99 (2011) becoming 2-4-septate on germination…Okeanomyces
Tunicatispora K.D. Hyde, Aust. Syst. Bot. 3: 712 (1990) 17. Ascomata formed beneath a stroma, ascospores 6–21×
Type species: Tunicatispora australiensis K.D. Hyde, 6–8 μm . . . . . . . . . . . . . . . . . . . . . . . . . Ophiodeira
Aust. Syst. Bot. 3(4): 712 (1990) 17. Ascomata not stromatic . . . . . . . . . . . . . . . . . . . . 18
18. Ascospores ellipsoidal, 25–36 × 8–12 μm. . . . . . . 19
18. Ascospores filiform, 60–80 × 4–6 μm ascomata thick-
Key to genera of Halosphaeriaceae walled . . . . . . . . . . . . . . . . . . . . . . . . . . . . Oceanitis
19. Appendages gradually detach from the ascospore wall
1. Ascospores aseptate . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 to form an ellipsoidal sheet in water . . . . . Pileomyces
1. Ascospores septate . . . . . . . . . . . . . . . . . . . . . . . . . . 9 19. Appendages unfurling into fine thread in wa-
2. Ascospores appendaged . . . . . . . . . . . . . . . . . . . . . . 3 ter.……………………………………….... Tirispora
2. Ascospores lacking appendages . . . . . . . . . . . . . . . . 4 20. Ascospores with polar unfurling appendages 21
3. Ascospores with polar and 4 groups of equatorial hair- 20. Ascospore appendages with a different morphology 26
like appendages . . . . . . . . . . . . . . . . . . Nautosphaeria 21. Polar appendages emerging from a hood-like struc-
3. Ascospores with a single, uncoiling polar append- ture……………………………….... Cucullosporella
age…………………………………………..... Moana 21. Polar appendages not formed through a hood. . . . 22
3. Ascospores with a prominent sheath …… Fluviatispora 22. Ascospores longer than 35 μm and wider than
4. Ascospores longer than 50 μm, filiform, range 50– 20 μm…………………………………………..... 24
300 × 4–15 μm . . . . . . . . . . . . . . . . . . . . Bathyascus 22. Ascospores shorter and narrower than 35 μm and
4. Ascospores shorter than 50 μm, spherical, ellipsoidal or 20 μm, respectively . . . . . . . . . . . . . . . . . . . . . . . 23
rhomboid . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 23. Ascospores wider than 14 μm . . . . . . Saagaromyces
Fungal Diversity

23. Ascospores 12–14 μm wide . . . . . . . . . . Aniptodera 38. Ascospores with 5–7 appendages . . . . . . . Corallicola
24. Ascospores wider than 30 μm…………………........ 39. Equatorial appendage ring- or annulus-like. . . . . . 40
Halosarpheia sensu stricto 39. Equatorial appendages distinct . . . . . . . . . . . . . . . 42
24. Ascospores narrower than 30 μm . . . . . . . . . . . . . 25 40. Chamber-like polar appendage from which mucilage is
25. Ascospores 21–31 × 8–11 μm. . . . . . . . . . . Panorbis released . . . . . . . . . . . . . . Ceriosporopsis tubulifera
25. Ascospores 20–34 × 7–11 μm. . . . . . . . Natantispora 40. Polar appendages do not release mucilage . . . . . . 41
25. Ascospores 26-(33) –38 × 10– (12) 14 μm .... Tinhaudeus 41. Equatorial appendage annulus-like . . . . . . Ondiniella
25. Ascospores 40–60 × 10–15 μm, polyguttulate.......... 41. Equatorial appendage ring-like . . . . . . Lautisporopsis
Ascosacculus 42. Equatorial appendages lunate, with a cup-like polar
25. Ascospores 10–32 × 4–13 μm Halosarpheia sensu lato appendage . . . . . . . . . . . . . . . . . . . Halosphaeriopsis
26. Ascospores with two types of appendages . . . . . . 27 42. Appendages spoon-like or obclavate . . . . . . . . . . 43
26. Ascospores with only one type of appendage . . . . 30 43. Appendages spoon-like, do not fragment . Halosphaeria
27. Ascospores with a sheath and polar and lateral or sub- 43. Appendages become fibrillar at maturity . . . . . . . 44
polar appendages . . . . . . . . . . . . . . . . . . . . . . . . . 28 44. Polar appendage longer than equatorial appendages
27. Ascospores with no sheath . . . . . . . . . . . . . . . . . . 29 .................................................................. Ocostaspora
28. Ascospores with polar hair-like and lateral sheath-like 44. Appendages equal in length . . . . . . . . . . . . Sablecola
appendage . . . . . . . . . . . . . . . . . . . . . . . Nimbospora 45. Appendages with a cup-like exosporic fragments at
28. Ascospores with a fragmenting sheath, and polar their tips . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46
unfurling appendages . . . . . . . . . . . . . Tunicatispora 45. Appendages lacking cup-like fragments at their tips . . 47
29. Exosporium folds to form an annulus-like equatorial 46. Equatorial appendages two pairs, radiating, at the cen-
appendage, polar appendages form inside and end tral septum . . . . . . . . . . . . . . . . . . . . . . Marinospora
chamber consist of two electron-dense layers.. Toriella 46. Equatorial appendages one pair not radiating ...... Toriella
29. Sub-polar hair-like appendages arise from a pad, and on 47. Exosporic sheath highly fibrillar (mucilaginous), ap-
the opposite side long, sticky appendages that uncoil in pendage slug-like. . . . . . . . . . . . . . . . . Limacospora
water . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nohea 47. Exosporic sheath compact, not fibrillar, appendages not
30. Ascospores with polar or subpolar appendages. . . 31 slug-like . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48
30. Ascospores with polar and/or lateral appendages...... 39 48. Ascospores thick walled at apices. . . . . . . Tubakiella
30. Ascospores with an exosporic sheath . . . . . . . . . . 45 48. Ascospores wall evenly distributed. . . . . . . . . . . . 49
31. Ascospores with polar appendages . . . . . . . . . . . . 32 49. Apical appendages swells in water, equatorial ap-
31. Ascospores with subpolar appendages . . . . . . . . . 36 pendages subulate, evenly distributed around the
32. Ascospore appendages ephemeral . . . . . . . . Gesasha septum . . . . . . . . . . . . . . . . . . . . . Ebullia
32. Ascospore appendages permanent . . . . . . . . . . . . 33 49. Apical appendages tubular, 3–4 lateral appendages
33. Ascospore appendages formed by fragmentation of a subcylindrical with small caps on their tips
sheath . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34 .............................................................. Ceriosporopsis
33. Appendages coiled around the ascospores, uncoiling in 50. Ascospores with no appendages . . . . . . . . . . . . . . 47
water . . . . . . . . . . . . . . . . . . . . . . . . . . Morakotiella 50. Ascospores with appendages . . . . . . . . . . . . . . . . 49
34. Ascospores wing-like or radiating appendages 51. Ascospores filamentous, broad at one end, tapering at
.................................................................... Remispora the other. . . . . . . . . . . . . . . . . . . . . . . . . . . . . Trailia
34. Ascospores ellipsoidal without radiating appendages 51. Ascospores not tapering at one end . . . . . . . . . . . 48
....................................................................... Kochiella 52. Ascospores 5-septate (rarely 9), asci with 4 asco-
35. Appendages initially wrapped around the ascospore wall, spores . . . . . . . . . . . . . . . . . . . . Luttrellia
separating in water to form long filaments that are spoon- 52. Ascospores 3-septate, asci with 8 ascospores
shaped at the place of attachment . . . . . . . Morakotiella ........................................................ Pseudolignincola
35. Appendages initially hamate that deliquesce in water to 53. Ascospores hyaline or brown with polar and equatorial
form thin flat sheets . . . . . . . . . . . . . Praelongicaulis appendages. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 50
36. Ascospore appendages with a spade-like tip 53. Ascospores hyaline, with only polar appendages 52
................................................................... Arenariomyces 53. Ascospores with sheath, lacking polar and equatorial
36 Ascospores appendages lacking a spade-like tip 37 appendages . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 58
37. Ascospores with 2 sub-polar appendages 54. Appendages hair-like tufts, one polar and four equato-
............................................................. Antennospora rial . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51
37. Ascospores with more than 2 sub-polar appendages 38 54. Polar appendages spine-like, equatorial appendages
38. Ascospores with 3–4 sub-polar spoon-shaped append- formed by fragmentation of an exosporic sheath
ages . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Haiyanga Corollospora
 Fungal Diversity

55. Catenophyses present, ascospores hyaline, appendages apical appendage present or absent, if present elongate.
string-like, lacking an equatorial pad …..…Havispora Microconidia hyaline, oval to ellipsoid, aseptate, smooth-walled.
55. Catenophyses lacking, ascospores with hyaline Type: Harknessia Cooke
end cells and brown central cells, equatorial pad Notes: The family Harknessiaceae is typified by
present . . . . . . . . . . . . . . . . . Nereiospora H. eucalypti Cooke and has distinct morphological characters
56. Ascospores with a single polar appendage . . . . . . 53 with a wuestneia-like sexual morph. LSU analysis places the
56. Ascospores with bipolar appendages . . . . . . . . . . 54 family in Diaporthales (Crous et al. 2012). Crous et al. (2012)
57. Appendage an ephemeral drop of mucilage introduced six novel species of Harknessia on Eucalyptus and
.............................................................. Okeanomyces a multi-gene analysis (ITS, CAL and TUB) was provided for
57. Tetra radiate appendages formed after release from the these species. Most members in the family are associated with
ascoma. . . . . . . . . . . . . . . . . . . . . . . . . . Thalespora leaf spots and they are suspected to be pathogens. However,
58. Ascospores appendages hamate, unfurling in water ..... 55 many species have been isolated from asymptomatic plant
58. Ascospore appendages not hamate . . . . . . . . . . . . 57 tissues and are assumed to be saprobes.
59. Ascospores narrower than 7 μm. . . . . . . . . Oceanitis Harknessia Cooke, in Cooke & Harkness, Grevillea 9(no.
59. Ascospores wider than 7 μm . . . . . . . . . . . . . . . . 56 51): 85 (1881)
60. Ascospores verrucose, wider than 40 μm .. Magnisphaera Facesoffungi numbers: FoF 02144; Figs 66, 67
60. Ascospores not verrucose, narrower than 40 μm Saprobic or pathogenic, mostly on leaf tissues. Sexual
Halosarpheia sensu lato morph: Ascomata perithecial, single or aggregated, immersed
61. Ascospores appendages sub-polar, spine-like brown. Neck emergent to depressed, wall of 3–5 layers of brown
Arenariomyces cells of textura angularis. Paraphyses hyaline, septate, dispersed
61. Ascospores appendages broad strap-like. . . Haligena between asci. Asci 8-spored, unitunicate, cylindrical to clavate,
61. Ascospores appendages round . . . . . . . . Trichomaris short pedicellate, with J- apical ring. Ascospores uni- to biseriate,
62. Ascospores 12–20 μm wide, central cells dark, append- hyaline, ellipsoid to fusoid, aseptate, thick-walled, guttulate,
ages a fragmenting sheath net-like ..... Carbosphaerella smooth-walled. Asexual morph: Coelomycetous, appearing as
62. Ascospores hyaline, 7–11 μm wide ... Appendichordella nearly circular, black distinct spots. Conidiomata erumpent,
Harknessiaceae Crous, in Crous et al., Persoonia, Mol. scattered, pycnidial, unilocular, globose to subglobose, brown.
Phyl. Evol. Fungi 28: 55 (2012) Peridium comprising 3–4 layers of brown-walled cells of textura
Facesoffungi number: FoF 01387 angularis. Conidiophores short, cylindrical, almost globose,
Saprobic or pathogenic, associated with leaf spots. Sexual branched, hyaline, mixed with peridium cells. Conidiogenesis
morph: Ascomata perithecial, solitary or aggregated, immersed, cells hyaline to brown, holoblastic, bottle-shaped, cylindrical.
globose, coriaceous, brown, papillate. Papilla emergent to de- Conidia hyaline when young, brown at maturity, globose to
pressed, wall comprising 3–5 layers of brown-walled cells of ovoid, with a truncate apiculate apex and an obtuse to blunt base,
textura angularis. Hamathecium comprising hyaline, septate pa- smooth-walled, with longitudinal striations along the length of
raphyses. Asci 8-spored, unitunicate, cylindrical to clavate, short some conidia. Basal appendages hyaline, tubular, smooth, thin-
pedicellate, with J-, apical ring. Ascospores uniseriate to biseriate, walled, often collapsing.
hyaline, ellipsoid to fusoid, aseptate, thick-walled, guttulate, Type species: Harknessia eucalypti Cooke, Grevillea 9(no.
smooth-walled. Asexual morph: Coelomycetous. Conidiomata 51): 85 (1881)
eustromatic, pycnidial, scattered or aggregated, immersed, glo- Notes: Harknessia species are found in a large variety of
bose, coriaceous, with single or several locules, dark brown to ecosystems and are commonly associated with leaf spots, leaves
black. Peridium comprising thin-walled, almost hyaline to brown with tip dieback or leaf scorch and stem cankers (Crous et al.
cells of textura angularis. Ostiole wide, central, surrounded by 1989; Wijayawardene et al. 2016). Some species have been iso-
brown cells. Conidiophores lining the inner cavity or reduced to lated from leaf and twig litter (Marincowitz et al. 2008).
basal layer sometimes reduced to conidiogenous cells, sometimes However, whether Harknessia species are important pathogens
septate, branched. Conidiogenous cells holoblastic, discrete, or saprobes is not resolved. Harknessia species with hyaline
lageniform, subcylindrical to cylindrical, hyaline to pale yellow, conidia and apical appendages were placed in Mastigosporella
smooth, producing macroconidia and sometimes microconidia by von Höhnel (1915), and species with brown conidia with
from same conidiogenous cell, proliferating Sympodially, one apical and basal appendages were moved to Apoharknessia
or several times. Macroconidia with a basal appendage, hyaline (Lee et al. 2004), while species with very thick conidial walls
when young, brown at maturity, unicellular, although basal ap- and longitudinal slits were included in Dwiroopa (Farr and
pendage separated by a septum thick-walled, smooth-walled, Rossman 2003). However, there is molecular support to place
with or without light and dark longitudinal bands, sometimes Harknessia in a distinct lineage in the family Harknessiaceae.
longitudinally striate, guttulate, basal appendage cellular, cylindri- Helminthosphaeriaceae Samuels et al., Mycologia 89(1):
cal to subcylindrical, hyaline, thin-walled, devoid of contents, 144 (1997)
Fungal Diversity

Fig. 66 Sexual morph of Harknessia eucalyptorum (CPC 12697). a Leaf spot symptoms on Eucalyptus sp. b Ascomata with short neck c Paraphyses
d asci e Ascospores. Scale bars = 10 μm (redrawn from Crous et al. 2012)

Facesoffungi number: FoF 01142 Notes: The family Helminthosphaeriaceae was introduced by
Saprobic or parasitic on wood, stems or decorticated Samuels et al. (1997b) based on black, setose ascomata and cylin-
branch in terrestrial habitats, some fungicolous. Sexual morph: drical asci with brown to hyaline ascospores and only included
Ascomata perithecial, gregarious or scattered, dark brown to Helminthosphaeria (Fuckel 1870). Tengiomyces was added to the
black, solitary, superficial or immersed, ovoid, globose to family by Réblová (1999a) based on morphological data.
subglobose, carbonaceous or membranaceous, rough, tubercu- Echinosphaeria, Hilberina, Ruzenia and Synaptospora were in-
late, smooth or with setae, papillate or ostiole indistinct or ab- cluded based on analysis of LSU sequence data and morphological
sent, the apex collapsing when dry, hyaline periphyses lining characters (Miller and Huhndorf 2004; Miller et al. 2014).
the ostiole or absent. Peridium thick (30–60 μm), composed of Endophragmiella was accepted as the asexual morph of
two layers, outer layer comprising brown cells of textura Echinosphaeria. The family Helminthosphaeriaceae forms a
angularis or prismatica, carbonaceous or membranaceous; in- well-supported clade in the order Chaetosphaeriales, with
ner layer comprising of hyaline cells of textura prismatica, thin, Chaetosphaeriaceae as a sister clade (Maharachchikumbura
membranaceous. Hamathecium with numerous, septate, persis- et al. 2015). Currently seven genera are accepted in the family
tent or deliquescing, swollen, filiform or cylindrical based on morphological and phylogenic analyses of LSU, SSU,
pseudoparaphyses. Asci 8-spored, unitunicate, thin or thick- TEF and RPB2 sequence data (Maharachchikumbura et al. 2015).
walled, cylindrical to clavate, pedicellate, apex truncate, not
amyloid, apical ring refractive or indistinct. Ascospores 2-seri- Helminthosphaeria Fuckel, Jb. nassau. Ver. Naturk. 23–24:
ate, hyaline or brown or becoming dark colored in part, 166 (1870) [1869–70]
allantoid, clavate, cylindrical to ellipsoid, 0–3-septate, Facesoffungi number: FoF 01143, Fig. 68
smooth-walled, with or without guttules. Asexual morph: Saprobic on wood or decorticated branch in terrestrial hab-
Hyphomycetous. Conidiophores macronematous, itats, some fungicolous. Sexual morph: Ascomata perithecial,
mononematous, scattered or gregarious, brown, straight, sep- gregarious or scattered, dark brown to black, solitary, superfi-
tate, unbranched, smooth-walled. Conidiogenous cells cial or immersed, globose to subglobose, carbonaceous or
monoblastic, terminal, integrated, percurrent. Conidia solitary, membranaceous, rough, smooth or with setae, with or without
acrogenous, brown, septate, obclavate, fusiform to cylindrical, ostioles, hyaline periphyses lining ostiole or absent, the apex
straight, subtruncate to obtuse at the apex, truncate or swollen at collapsing when dry. Peridium thick (30–60 μm), composed
the base, eguttulate or guttulate. of two layers, outer layer comprising of brown cells of textura
Type: Helminthosphaeria Fuckel angularis, carbonaceous or membranaceous; inner layer
 Fungal Diversity

Fig. 67 Harknessia eucalypti (Material examined: USA, California, on Conidiomata on host substrate d Cross section of conidioma e Peridium
leaves of Eucalyptus globulus Labill. (Myrtaceae), Harkness 1280, f, g Conidia attached to conidiophores h–m Conidia. Scale bars:
isotype K (M) 195744). a, b Herbarium packet and specimen c d = 100 μm, e–g = 20 μm, h–m = 10 μm

comprising hyaline cells of textura prismatica, thin, without guttules. Asexual morph: Hyphomycetous.
membranaceous. Hamathecium with numerous, septate, un- Conidiophores mononematous, scattered to gregarious,
branched or branched, swollen, filiform or cylindrical paraph- brown, straight, septate, branched, smooth-walled.
yses. Asci 8-spored, unitunicate, cylindrical to clavate, pedi- Conidiogenous cells integrated, intercalary or percurrent.
cellate, apex truncate, not amyloid, apical ring refractive or Conidia solitary, hyaline to brown or multi-coloured, aseptate
indistinct,. Ascospores 2-seriate, hyaline or brown or greyish, to multi-septate, obclavate to cylindrical, eguttulate or
clavate to ellipsoid, 0–3-septate, smooth-walled, with or guttulate.
Fungal Diversity

Fig. 68 Helminthosphaeria clavariarum (from Samuels et al. 1997b) a Diplococcium sp.) d Ascospores (pores in ascospores are indicted by
Ascoma with setae and vertical section of ascoma b Asci and arrowhead) e Conidia (asexual morph Diplococcium sp.). Scale bars: a,
pseudoparaphyses c Conidiophore and conidia (asexual morph c = 50 μm, b = 20 μm, d–e = 10 μm

Type species: Helminthosphaeria clavariarum (Desm.) Type species: Echinosphaeria canescens (Pers.) A.N.
Fuckel, Jb. nassau. Ver. Naturk. 23–24: 166 (1870) [1869–70] Mill. & Huhndorf, Mycol. Res. 108(1): 31 (2004)
Notes: The genus Helminthosphaeria was introduced by Endophragmiella B. Sutton, Mycol. Pap. 132: 58 (1973)
Fuckel (1870), with H. clavariarum as the type species. This genus Type species: Endophragmiella pallescens B. Sutton,
was included in Melanosporaceae, in the order Sordariales based Mycol. Pap. 132: 62 (1973)
on its 1-septate ascospores with apical pores (Parguey-Leduc Hilberina Huhndorf & A.N. Mill., Mycol. Res. 108(1): 31
1960). Lundqvist (1972) excluded Helminthosphaeria from (2004)
Melanosporaceae and thought that it should be used as a subfam- Type species: Hilberina caudata (Fuckel) Huhndorf & A.N.
ily or family due to its distinct filamentous pseudoparaphyses and Mill., in Miller & Huhndorf, Mycol. Res. 108(1): 31 (2004)
asci lacking sheaths. Barr (1990) reevaluated the family Ruzenia O. Hilber, The Genus Lasiosphaeria and Allied Taxa
Sordariaceae and included Helminthosphaeria. Eriksson and (Kelheim): 7 (2002)
Hawksworth (1993) placed Helminthosphaeria in the order Type species: Ruzenia spermoides (Hoffm.) O. Hilber, in
Sordariales. Samuels et al. (1997b) monographed the genus, in- Hilber & Hilber, The Genus Lasiosphaeria and Allied Taxa
cluding the asexual morph Diplococcium, and introduced the fam- (Kelheim): 7 (2002) = Sphaeria spermoides Hoffm., Veg.
ily Helminthosphaeriaceae. The sexual morph is characterized by Crypt. 2: 12 (1790)
a fungicolous or lignicolous habit, superficial or immersed Synaptospora Cain, Beih. Sydowia 1: 4 (1957) [1956]
ascomata, cylindrical asci and hyaline to brown ascospores, and Type species: Synaptospora petrakii Cain, Beih. Sydowia
an asexual morph characterized by smooth or verrucose, septate 1: 5 (1957)
conidia (Samuels et al. 1997b; Goh and Hyde 1998; Hernández- Tengiomyces Réblová, Mycotaxon 70: 408 (1999)
Restrepo et al. 2012). Type species: Tengiomyces indicus (Varghese & V.G. Rao)
Réblová, Mycotaxon 70: 408 (1999)

Other genera included Key to genera of Helminthosphaeriaceae

Echinosphaeria A.N. Mill. & Huhndorf, Mycol. Res. 108(1): 1. Hyphomycete with acrogenous, cylindrical hyaline conidia
29 (2004) Endophragmiella
 Fungal Diversity

1. Ascomycete 2 of Hispidicarpomycetaceae. Maharachchikumbura et al. (2015)

2. Ascospores hyaline at maturity Ruzenia also accepted the taxonomic placement of the family
2. Ascospores brown or greyish-brown or versicolor at ma- Hispidicarpomycetaceae within the order Spathulosporales.
turity 3 Further collections, isolation and sequencing are necessary to
3. Ascospores versicolored Tengiomyces resolve the taxonomic position of these families.
3. Ascospores concolourous 4
4. Ascospores curved 5 Hispidicarpomyces Nakagiri, Mycologia 85(4): 639
4. Ascospores straight 6 (1993)
5. Ascospores L-shape Hilberina Facesoffungi number: FoF 02110, Fig. 69
5. Ascospores allantoid Echinosphaeria Parasitic on the red alga Galaxaura falcata. Sexual
6. Ascospores clavate to cylindrical, greyish-brown, not morph: Ascomata hemisphaerical, erumpent, solitary to gre-
contracted at the septa Helminthosphaeria garious, ostiolate, epapillate, black. Peridium composed of
6. Ascospores oval to ellipsoid, brown, constricted at the several cell layers with short hyphal projections at the outer
septa.. Synaptospora surface. Paraphyses present. Asci in a hymenium lining whole
Hispidicarpomycetaceae Nakagiri, Mycologia 85(4): 649 ascocarp venter, 8-spored, unitunicate, thin-walled, clavate to
(1993) pyriform, early deliquescing. Ascospores hyaline, ellipsoidal
Facesoffungi number: FoF 01098 to fusiform, unicellular, lacking appendages or a sheath.
Parasitic on marine alga. Sexual morph: Ascomata super- Reproductive state: Spermodochia composed of
ficial, solitary to gregarious, ostiolate, apapillate. Peridium penicilliate-like spermatiophores (antheridia), brown to dark
composed of three-layers: outer layer with thick-walled, short brown. Spermatia cylindrical, one-celled, hyaline.
hyphal projections, brown to dark brown, middle layer of light Trichogynes cylindrical, septate, simple, with thin-walled
brown to dark brown, thick-walled hyphal, flattened cells of and hyaline apical cell.
textura epidermoidea, and an inner layer of pale brown, thin- Type species: Hispidicarpomyces galaxauricola Nakagiri,
walled, flattened cells of textura epidermoidea, lining the Mycologia 85(4): 639 (1993)
whole ascomatal venter. Paraphyses septate branched. Asci Notes: A monotypic marine genus introduced by Nakagiri
8-spored, unitunicate, thin-walled, clavate to pyriform, deli- (1993) in the family Hispidicarpomycetaceae that needs fur-
quescing early and lacking an apical ring. Ascospores hyaline, ther taxon sampling to determine its phylogenetic position
ellipsoid to elliptic-fusiform, unicellular, lacking appendages within Ascomycota (Jones et al. 2009b, 2015). Nakagiri
or a sheath, overlapping. Reproductive state: of brown hy- (1993) described similar morphological features of
phae, Spermodochia composed of a mass spermatiospores. Hispidicarpomyces and Spathulospora, such as production
Spermatiospores, verticillate, asymmetrical penicilliate-like, of trichogynes and spermatiophores in a spermodochium.
brown at the base, paler and hyaline towards the apex. Stipes However, Hispidicarpomyces species possess different mor-
septate, branched, brown at the base. Spermatia cylindrical, phological features to Spathulospora, including a hyphoid
unicellular, hyaline. Trichogynes septate, cylindrical, brown. thallus, spermodochia formation, alternate development of
Type: Hispidicarpomyces Nakagiri spermodochium within the ascocarp, a peridium with a hispid
Notes: The family Hispidicarpomycetaceae was introduced to surface, a hymenium extending along the entire inside of the
accommodate a marine ascomycete, Hispidicarpomyces ascocarp venter, the hamathecium (paraphyses), ascospores
galaxauricola Nakagiri inhabiting the alga Galaxaura falcata without appendages and the host alga.
(Rhodophyta), which was found along the Japanese coast by
Nakagiri (1993). This fungus produces trichogynes and Hypocreaceae De Not. [as ‘Hypocreacei’], G. bot. ital. 2(1):
spermatiophores composed of spermodochia which resemble 48 (1844)
Spathulospora species, another ascomycete parasitic on a red = Trichodermataceae Fr. [as ‘Trichodermacei’], Syst. orb.
alga (Ballia spp.), but differs from the latter genus in possessing veg. (Lundae) 1: 144 (1825)
hyphoid thalli, in spermodochia formation, and ascomata struc- Facesoffungi number: FoF 01904
ture (Nakagiri 1993). The genera Spathulospora, Retrostium
(Spathulosporaceae) and Hispidicarpomyces
(Hispidicarpomycetaceae) were referred to the order Fig. 69 Hispidicarpomyces galaxauricola (Material examined: JAPAN,„
Spathulosporales by Nakagiri (1993). However, molecular data O-ura, Shimoda, 2 June 1990, IFO H-12135, Galaxaura falcata Kjellman
placed the genus Spathulospora (S. adelpha and S. antartica) in (Chaetangiaceae, Rhodophyta) holotype) a Herbarium material of
the order Lulworthiales (Inderbitzin et al. 2004), although the H. galaxauricola b Slides of H. galaxauricola c Ascomata on host
surface d Section through ascoma e Peridium f Short hyphal projections
type species (S. phyophila) has not been sequenced. arising from outer layer of peridium g-j Ascospores k–l Branching
Consequently, the affinity of Spathulospora within the spermatiophores m Penicilliate phialides of spermatiophores. Scale
Lulworthiales raises the question as to the phylogenetic position bars: c = 500 μm, d = 50 μm, e, f, k = 20 μm, l–m = 10 μm, g–j = 5 μm
Fungal Diversity
 Fungal Diversity

Biotrophic, hemibiotrophic, saprobic or hypersaprobic on major genera encompassing the majority of species. Currently,
various plants, other fungi, and myxomycetes, terrestrial and Maharachchikumbura et al. (2015) accepted 18 genera in the
aquatic habitats. Sexual morph: Stromatic tissue present or family.
lacking, when present soft and fleshy, pallid or brightly Trichoderma Pers., Neues Mag. Bot. 1: 92 (1794)
pigmented to light brownish, immersed erumpent, effuse, tuber- Facesoffungi number: FoF 02111, Figs 70, 71
culate or pulvinate, occasionally stipitate, then fertile region Biotrophic, hemibiotrophic, saprobic or hypersaprobic on
clavate, or subiculum of interwoven pallid or brightly various plants, on other members of the fungi including mem-
pigmented hyphae. Ascomata solitary or often arranged in bers of Basidiomycota, or on perennial bracket fungi in varying
groups, immersed in or erumpent to superficial on substrate, stages of decay, and less commonly on herbaceous substrates.
pallid, brightly pigmented, or shades of light brown or blue to Sexual morph: Stromata discrete to effused, composed of pseu-
violet (appearing black), rarely brown, globose, ovoid, doparenchyma cells or highly compacted hyphae, with
obpyriform or spheroid, collabent at times, apex papillate, with ascomatal elevations conspicuous or not, stromatal surface var-
periphysate ostiole, surface glabrous or warted or bearing hya- iously wrinkled, tuberculate or creased, margins of stromata free
line or pallid hyphal appendages or rarely thick-walled setae. from or adherent to the substrate, hyaline, white, yellow, reddish-
Peridium externally composed of pseudoparenchymatous cells, brown, dark brown to black. Ascomata immersed in the stroma,
sometimes with thick, sclerotial walls, internally composed of ascomatal wall and stromatal tissues KOH+ or KOH−. Asci
compressed rows of cells, pallid to brightly pigmented or unitunicate, 8-spored, cylindrical. Ascospores 1-septate,
brown, blue or violet. Paraphyses apical (periphysoids), usually disarticulating early in development into two equal or unequal,
deliquescent, occasionally visible at maturity, as cellular rem- subglobose, globose, ovoid, oblong or wedge-shaped part asco-
nants among asci or as remnants apical fringe. Asci basal to spores, hyaline or green, typically spinulose or warted, rarely
peripheral, mostly 8-spored, occasionally polysporous or less smooth. Asexual morph: Hyphomycetous. Conidiophores are
than eight, cylindric, oblong or inflated, apical ring often lack- mostly formed in pustules, conspicuously curved to sinuous or
ing, when present shallow, refractive, non-amyloid. Ascospores conspicuously curved, without or with weakly developed or well
uniseriate, biseriate or in fascicle, hyaline, yellowish, pinkish to developed main axis, branched, branching irregular, verticillate
greenish or occasionally brown, 1-celled or one to several sep- or sometimes tree-like, hyaline, smooth, septate. Phialides soli-
tate, occasionally with longitudinal septa, disarticulating into tary or forming in whorls, sometimes percurrently proliferating,
part-spores at times or budding to form conidia within ascus, straight, cylindrical or somewhat swollen at or below the middle,
ellipsoid, fusoid, allantoid, elongate or globose, with smooth, curved to sinuous or straight, hyaline, smooth. Conidia unicel-
verruculose or longitudinally striate cell wall. Asexual morph: lular, ovoid, ellipsoidal, oblong, globose to subglobose, green,
Hyphomycetous, grouped on sporodochia or synnemata, rarely rarely hyaline, guttulate, smooth, verruculose, spinulose or wart-
coelomycetous, conidiogenous cells enteroblastic phialidic, ed. Chlamydospores globose to subglobose and terminal at the
thick-walled structures present at times. ends of hyphae or intercalary within hyphal cells.
Type: Hypocrea Fr. (= Trichoderma Pers.) Type species: Trichoderma viride Pers., Neues Mag. Bot.
Notes: The family Hypocreaceae was recognized within 1: 92 (1794)
Hypocreales and divided into six subfamilies by Lindau ≡ Hypocrea rufa (Pers.) Fr., Summa veg. Scand.,
(1897). Seaver (1909a, b, 1910a, b, 1911) divided the order Section Post. (Stockholm): 383 (1849)
Hypocreales into two families ( Nectriaceae and Notes: Hypocrea was described by Fries in 1825, based on
Hypocreaceae) depending on their perithecial and stromatic Sphaeria rufa Pers., a species with hyaline ascospores.
characters. Petch (1938) also accepted Nectriaceae and Currently, the type species of the genus is represented by
Hypocreaceae as distinct families within the Hypocreales, while Hypocrea rufa (Pers.) Fr. The genus Hypocrea is characterized
Munk (1957) and Dennis (1960) placed them in the Sphaeriales. by typically hyaline or green, eight, 1-septate ascospores, that
Later, Miller (1949), Bessy (1950), Luttrell (1951), Dingley disarticulate at the septum when they are young, producing 16
(1951a, b, 1952a, b, 1953, 1954)), von Arx and Müller (1954), part-spores in each ascus. Hypocrea (with sexual morph) and
Müller and von Arx (1962), Gäumann (1964), Rogerson (1970) Trichoderma (with asexual morph) are linked and Rossman
and Barr (1990) placed Nectriaceae under Hypocreaceae as
one family. However, Kreisel (1969) accepted Hypocreaceae
Fig. 70 Trichoderma viride (Material examined: CZECH REPUBLIC,„
and Nectriaceae in the order Hypocreales, along with another South Bohemia, Frymburk, on partly decorticated logs of Pinus sylvestris
five families based on their bright coloured, fleshy ascomata, L. (Pinaceae), leg. W. Jaklitsch, W.J. 2753, 3 October 2004. WU 24013,
presence of apical paraphyses and production of phialides on epitype) a Herbarium label b, c Stromata on host substrate d Transverse
free conidiophores. Rossman et al. (1999) believed that section through stromata with perithecia e Ostiolar opening in transverse
section f Hair on surface of stroma g Outer layer of stromata h Peridium. i,
Nectriaceae and Hypocreaceae could be two separate families j Asci k Ascus in Melzer’s reagent l Ascospores m Ascospores in cotton
within the Hypocreales. In their work they included 12 genera blue. Scale bars: c = 1 mm, d = 200 μm, e = 50 μm, f = 10 μm, g–
in Hypocreaceae, including Hypocrea and Hypomyces as two i = 20 μm, j = 50 μm, k = 20 μm, l, m = 10 μm
Fungal Diversity
 Fungal Diversity

Fig. 71 Trichoderma viride (Material examined: Dry culture, WU 24013a, epitype) a Herbarium material and label b, c Sporulation on CMD d, e
Conidiophores f, g Conidia. Scale bars: b = 2 mm, c = 500 μm, d, e = 20 μm, f, g =10 μm

et al. (2013) proposed the use of Trichoderma over Hypocrea, Escovopsioides H.C. Evans & J.O. Augustin, PLoS ONE
and this was followed by Maharachchikumbura et al. (2015). 7(12): e51392, 6 (2012)
Type species: Escovopsioides nivea H.C. Evans & J.O.
Augustin, PLoS ONE 7(12): e51392, 6 (2012)
Other genera included Hypocreopsis P. Karst., Bidr. Känn. Finl. Nat. Folk 23: 251
(1873)
Aphysiostroma Barrasa et al., Can. J. Bot. 63(12): 2439 Type species: Hypocreopsis riccioidea (Bolton) P. Karst.,
(1986) [1985] Bidr. Känn. Finl. Nat. Folk 19: 221 (1871)
Type species: Aphysiostroma stercorarium Barrasa et al., Hypomyces (Fr.) Tul. & C. Tul., Annls Sci. Nat., Bot., sér. 4
Can. J. Bot. 63(12): 2441 (1986) [1985] 13: 11 (1860)
Arachnocrea Z. Moravec, Bull. trimest. Soc. mycol. Fr. Type species: Hypomyces lactifluorum (Schwein.) Tul. &
72: 161 (1956) C. Tul., Annls Sci. Nat., Bot., sér. 4 13: 11 (1860)
Type species: Arachnocrea stipata (Lib. ex Fuckel) Z. Lichenobarya Etayo et al., in Lawrey et al., Bryologist
Moravec, Bull. trimest. Soc. mycol. Fr. 72: 161 (1956) 118: 88 (2015)
Dialhypocrea Speg., Boln Acad. nac. Cienc. Córdoba 23(3 Type species: Lichenobarya usneae (Etayo) Etayo et al., in
−4): 475 (1919) [1918] Lawrey et al., Bryologist 118: 88 (2015)
Type species: Dialhypocrea puiggariana Speg., Boln Mycogone Link, Mag. Gesell. naturf. Freunde, Berlin 3(1–
Acad. nac. Cienc. Córdoba 23(3−4): 475 [no. 320, reprint 2): 18 (1809)
page 113] (1919) [1918] Type species: Mycogone rosea Link, Mag. Gesell. naturf.
Escovopsis J.J. Muchovej & Della Lucia, Mycotaxon 37: Freunde, Berlin 3(1–2): 18 (1809)
192 (1990) Protocrea Petch, J. Bot., Lond. 75: 219 (1937)
Type species: Escovopsis weberi J.J. Muchovej & Della Type species: Protocrea farinosa (Berk. & Broome) Petch,
Lucia [as ‘weberii’], Mycotaxon 37: 192 (1990) J. Bot., Lond. 75: 219 (1937)
Fungal Diversity

Pseudohypocrea Yoshim. Doi, Bull. natn. Sci. Mus., 5. Stromata thin, arachnoid to loosely arranged, cottony,
Tokyo, N.S. 15: 655 (1972) white to pink, ascospores hyaline to green…….....…..7
Type species: Pseudohypocrea citrinella (Ellis) Yoshim. 5. Stromata discrete or effused, pulvinate, hyaline, white,
Doi, Bull. natn. Sci. Mus., Tokyo 15(4): 655 (1972) pale yellow to yellow orange, reddish brown, dark brown
Rogersonia Samuels & Lodge, Sydowia 48(2): 250 (1996) to black, ascospores hyaline or green……....…….....…8
Type species: Rogersonia striolata Samuels & Lodge, 6. Part ascospores irregularly globose, septum sub-median,
Sydowia 48(2): 251 (1996) disarticulating into dimorphic part ascospores, asexual
Sepedonium Link, Mag. Gesell. naturf. Freunde, Berlin morph verticillium-like…..….................... Arachnocrea
3(1–2): 18 (1809) 6. Part ascospores conical, septum median, disarticulating
Type species: Sepedonium mycophilum (Pers.) Nees, Syst. into monomorphic-part ascospores, asexual morph
Pilze (Würzburg): 44 (1816) [1816–17] gliocladium-like………......……………...… Protocrea
Sibirina G.R.W. Arnold, Nova Hedwigia 19(1–2): 299 7. Stromata pale yellow to yellow orange, ascospores hya-
(1970) [1969] line…………………….………..........................… 9
Type species: Sibirina fungicola G.R.W. Arnold, Nova 7. Stromata hyaline, white, reddish brown, dark brown to
Hedwigia 18(1−2): 300 (1970) [1969] black, ascospores hyaline or green………… Trichoderma
Sphaerostilbella (Henn.) Sacc. & D. Sacc., Syll. fung. 8. Stromata prosenchymatous, ascospores smooth,
(Abellini) 17: 778 (1905) part ascospores conical to lemon-shaped, mono-
= Gliocladium Corda, Icon. fung. (Prague) 4: 30 (1840) morphic ……………………… Pseudohypocrea
Type species: Sphaerostilbella lutea (Henn.) Sacc. & D. 8. Stromata pseudoparenchymatous, ascospores spinulose,
Sacc., Syll. fung. (Abellini) 17: 778 (1905) part ascospores dimorphic………………. Dialhypocrea
Sporophagomyces K. Põldmaa & Samuels, in Pöldmaa 9. Ascospores typically fusiform, apiculate, often
et al., Can. J. Bot. 77(12): 1765 (1999) coarsely warted, rarely smooth or spinulose, ascomata
Type species: Sporophagomyces chrysostomus (Berk. & typically partly or completely immersed in densely cot-
Broome) K. Põldmaa & Samuels, in K. Põldmaa et al., Can. tony or highly compacted subiculum, hyaline or in
J. Bot. 77(12): 1765 (1999) shades of yellow, orange tan or green……… Hypomyces
Stephanoma Wallr., Fl. crypt. Germ. (Norimbergae) 2: 269 9. Ascospores ellipsoid to naviculate, non-apiculate,
(1833) ascomata immersed or superficial on stomata, purple, reddish
Type species: Stephanoma strigosum Wallr., Fl. crypt. brown to grey or in shades of yellow….…. 10
Germ. (Norimbergae) 2: 269 (1833) 10. Ascomata immersed in the well-developed stroma, red-
dish brown to grey…Hypocreopsis
10. Ascomata superficial on stromata, dark red to purple, or
in shades of yellow…….. Sphaerostilbella
Key to the genera of Hypocreaceae with sexual
morph

1. Coprophilous, asexual morph verticillium-like Key to the genera of Hypocreaceae with asexual
Aphysiostroma morph
1. Fungicolous or lignicolous rarely on herbaceous sub-
strates, asexual morph Acremonium-, Gliocladium-, 1. In leaf-cutting ant nests, conidiophore branched,
Stilbella-, Trichoderma- or verticillium-like 2 vesiculate…………………………………………...…..2
2. Ascospores transversely seriate, aseptate Rogersonia 1. On diverse fungi, soil, litter and wood, conidiophores
2. Ascospores smooth, spinulose, longitudinally seriate, or simple, branched, trichodermoid, penicillate or
ornamented but not transversely seriate. uniseptate or rarely verticillate…………………………………….......….3
multi-septate or aseptate 3 2. Form lageniform phialides on terminal and intercalary
3. Ascospores 0−1-septate, rarely 3-septate, not globose vesicles, hyaline smooth-walled conidia in long
disarticulating within the ascus 4 chains….......…………………………. Escovopsioides
3. Ascospores 1-septate, disarticulating while in the ascus 5 2. Form lageniform to subulate terminal and intercalary
4. Ascospores fusiform to naviculate or lanceolate, smooth- globose, clavate to cylindrical vesicles, conidia hyaline
walled, sometimes finely verrucose, mature ascospores at first becoming pigmented with an ornamented or mu-
disarticulating into part ascospores after discharge.…… cilaginous brown outer coat or sheath, arrange in short
Sporophagomyces basipetal chains………………....….............Escovopsis
4. Ascospores ellipsoid, lanceolate or fusiform, smooth, 3. Produce phragmo conidia (primarily three-septate), rest-
spinulose to tuberculate …....…….…….…….…........6 ing spores lacking……………........ Sporophagomyces
 Fungal Diversity

3. Produce amero, didymo, phragmo or stauro conidia, genera were included in the family by Hawksworth et al. (1995).
resting spores present………………..……..….……. 4 However, Hyde et al. (1998) excluded most of the genera from
4. Conidiomata postulate or absent, with trichodermoid the family, while Palmomyces and Charonectria were added
branching conidiophore, conidia green or hyaline (Hyde et al. 1998; Rossman et al. 1999). Hyponectriaceae com-
…………………………………….…… Trichoderma prises the genera Apiothyrium, Arecomyces, Arwidssonia,
4. Conidiomata absent, conidiophore unbranched, sparing- Cesatiella, Chamaeascus, Charonectria, Discosphaerina,
ly branched or penicillately or verticillately branched, Exarmidium, Frondicola, Hyponectria, Micronectria,
conidia hyaline or pigmented…..………………..........5 Papilionovela, Pellucida, Physalospora, Phragmitensis,
5. Conidiogenous cell terminal and intercalary in conidio- Rachidicola and Xenothecium (Sivanesan and Shivas 2002;
phore, produce stauro-conidia....................Stephanoma Maharachchikumbura et al. 2015). This is a problematic family
5. Conidiogenous cell phialidic, produce amero, didymo, and it is not clear if it belongs in Xylariales. The type species
phragmo or stauro conidia…………………...………6 Hyponectria buxi needs recollecting and sequencing as the se-
6. Produce two types of conidiophores and quence data in GenBank may be problematic (Jaklitsch and
conidia……….……………………….….....……….7 Voglmayr 2012). The family is almost certainly polyphyletic
6. Produce only one type of conidiophores and and has been used as a “waste bin” for genera that cannot be
conidia…………….………………………....……….8 assigned to other families (Jaklitsch et al. 2016). Consequently,
7. Conidiophores unbranched, sparingly branched or verti- other genera in this family should also be recollected and se-
cillate, produce hyaline or pigmented amero co- quenced to establish their phylogenetic affinities. In this paper
nidia……………………………….……. Sepedonium we include 17 genera that are listed in Maharachchikumbura
7. Conidiophores monoblastic, branched or verticillate, et al. (2015), but do not provide a key to genera and most genera
produce hyaline amero conidia……..….…. Mycogone are probably wrongly placed. Pseudomassaria is placed in
8. Conidiophores densely verticillate, conidia arranged as Pseudomassariaceae.
singles (one conidium on each conidiogenous cell)
….…….…….…...........……………………… Sibirina Hyponectria Sacc., Michelia 1(no. 2): 250 (1878)
8. Conidiophores branched, penicillate or verticillate, co- Facesoffungi number: FoF 02112; Fig. 72
nidia arrange as singles or in basipetal chains…… Pathogenic or saprobic on leaves of Buxus sp. Sexual
morph: Ascomata immersed, depressed globose, visible as
Cladobotryum orange to brown dots on the host surface, coriaceous,
Hyponectriaceae Petr., Annls mycol. 21(3/4): 305 (1923) ostiolate, solitary or mostly gregarious. Ostiole aperiphysate.
Facesoffungi number: FoF 01775 Peridium one stratum of 2 layers of cells of textura angularis
Saprobic or pathogenic on dead plant matter, in terrestrial with thickened, brown walls cells around the ostiole.
and aquatic habitats. Sexual morph: Appearing as black, Paraphyses not observed. Asci 8-spored, unitunicate,
shiny spots on host surface or small black lines arising from cylindric-clavate to clavate, short pedicellate, with an indis-
cracks in bark. Pseudostromata present or lacking, if present, tinct, J-, apical ring. Ascospores overlapping biseriate or
superficial, forming clypeus over the ascomata. Ascomata sol- obliquely uniseriate, hyaline, ellipsoidal or oblong, straight
itary or aggregated, immersed, erumpent or rarely somewhat or inequilateral, 1-celled, contents minutely guttulate, lacking
superficial, globose to ovoid, upright or horizontal, brown to a sheath. Asexual morph: Undetermined.
black. Papilla short, ostiolate, with or without periphyses. Type species: Hyponectria buxi (DC.) Sacc., Michelia
Peridium comprising two layers, outer layer of brown cells 1(no. 2): 250 (1878)
of textura angularis, inner layer of hyaline, cells of textura Notes: Hyponectria is typified by Hyponectria buxi, which
angularis, in some genera cells are of textura globosa. was introduced by Saccardo (1878). Hyponectria buxi needs to
Hamathecium of sparse, septate, paraphyses, tapering towards be recollected and sequenced as the sequence data in GenBank
the apex, often deliquescing at maturity. Asci 8-spored, may be inaccurate. The other genera in this family should also
unitunicate, oblong, cylindrical or ellipsoidal, short pedicel- be recollected and sequenced to establish affinities.
late, with a J+, or J-, apical ring. Ascospores overlapping
biseriate, or fasciculate, hyaline, yellow to light brown, fusoid,
Fig. 72 Hyponectria buxi (Material examined: UK, Surrey, Mickleham,„
isthmoid, elongate filiform, obovoid, or oblong, asymmetric on leaves of Buxus sp., 1927, E.W. Mason no. 365 (IMI 16895, IMI52229
or symmetric, unicellular or septate, smooth-walled or and IMI49466). a-c Package of herbarium material d Herbarium
verruculose, with or without a mucilaginous sheath. Asexual specimen e Ascomata on the upper leaf surface f Ascomata on the
morph: Undetermined. lower leaf surface g Close up of ascoma h Close up of ascomata under
microscope i Section of ascoma j Cells of peridium k–o Asci p–u
Type: Hyponectria Sacc. Ascospores v J- reaction of apical ring in Melzer’s reagent. Scale bars:
Notes: Hyponectriaceae was introduced by Petrak (1923) to e–f = 500 μm g = 200 μm, h-i = 50 μm, j = 20 μm, k–o = 10 μm, p–
accommodate both Hyponectria and Anisostomula. Seventeen u = 5 μm and v = 10 μm
Fungal Diversity
Fungal Diversity

Other genera included Xenothecium Höhn., Sber. Akad. Wiss. Wien, Math.-naturw.
Kl., Abt. 1 128: 589 (1919)
Apiothyrium Petr., Sydowia 1(1–3): 1 (1947) Type species: Xenothecium jodophilum Höhn., Sber.
Type species: Apiothyrium arcticum Petr., Sydowia 1(1–3): Akad. Wiss. Wien, Math.-naturw. Kl., Abt. 1 128: 589 (1919)
1 (1947)
Arecomyces K.D. Hyde, Sydowia 48(2): 227 (1996)
Type species: Arecomyces frondicola K.D. Hyde, Sydowia Key to genera of Hyponectriaceae
48(2): 232 (1996)
Arwidssonia B. Erikss., Svensk bot. Tidskr. 68: 199 (1974) 1. Ascospore unicellular…………………………...……… 2
Type species: Arwidssonia empetri (Rehm) B. Erikss., 1. Ascospore multicellular……………………………… 7
Svensk bot. Tidskr. 68: 200 (1974) 2. A mucilaginous sheath………………………………. 3
Cesatiella Sacc., Michelia 1(no. 2): 250 (1878) 2. Lacking a mucilaginous sheath……………………… 4
Type species: Cesatiella australis Sacc. & Speg., Michelia 3. Ellipsoidal, Cylindrical, J-, immersed under a clypeus or
1(no. 2): 250 (1878) pseudostroma ..….…………………..…… Arecomyces
Chamaeascus L. Holm et al., Blyttia 51(3–4): 121 (1993) 3. Cymbiform, Clavate, lacking an apical apparatus, im-
Type species: Chamaeascus arcticus L. Holm et al., Blyttia mersed, pseudostroma, globose to subglobose…………
51(3–4): 121 (1993) Phragmitensis
Charonectria Sacc., Michelia 2(no. 6): 72 (1880) 4. Oblong….….….….….….….….….….….….....……. 5
Type species: Charonectria consolationis Sacc., Michelia 4. Baciliform to lunate, clavate to ellipsoidal, lacking apical
2(no. 6): 72 (1880) apparatus immersed, non papillate, darker around the
Discosphaerina Höhn., Sber. Akad. Wiss. Wien, Math.- pore....…………….……….……..…...... Chamaeascus
naturw. Kl., Abt. 1 126(4–5): 353 (1917) 5. Oblong or ellipsoidal…………………….…...……… 6
Type species: Discosphaerina discophora Höhn., Sber. 5. Oblong to fusiform, clavate, lacking ostiole and paraph-
Akad. Wiss. Wien, Math.-naturw. Kl., Abt. 1 126(4–5): 353 yses………………………………...… Discosphaerina
(1917) 6. Cylindric-clavated to clavate, J-, immersed, globose,
Exarmidium P. Karst., Bidr. Känn. Finl. Nat. Folk 23: 29, 222 Ostiole aperiphyses, paraphyses……....… Hyponectria
(1873) 6. Cylindric-clavated to clavate, J+ or J-, immersed, glo-
Type species: Exarmidium hysteriiforme P. Karst., Bidr. bose or conical, clypeus……………...… Physalospora
Känn. Finl. Nat. Folk 23: 222 (1873) 7. 1-septate…………………………………...………… 8
Frondicola K. D. Hyde., J. Linn. Soc., Bot. 110(2): 100 7. More than 1 septate………………………...…….… 10
(1992) 8. Apiosporous, lacking a J+…………………....……… 9
Type species: Frondicola tunitricuspis K.D Hyde., Bot. J. 8. Oblong or ellipsoidal, J-, immersed, solitary, pallid clyp-
Linn. Soc. 110: 95–110, 1992 eus……………………………………… Charonectria
Micronectria Speg., Anal. Soc. cient. argent. 19(1): 45 (1885) 9. Ascospore with spine-like appendages, cylindric-cla-
Type species: Micronectria guaranitica Speg., Anal. Soc. vate, with inconspicuous apical ring, immersed under a
cient. argent. 19(1): 45 (1885) pseudostroma……...……………..….….. Frondispora
Papilionovela Aptroot, Mycol. Res. 101(3): 266 (1997) 9. Ascomata lie horizontally between the cuticle and epi-
Type species: Papilionovela albothallina Aptroot, Mycol. dermis of the host, with a laterally placed osti-
Res. 101(3): 266 (1997) ole……………...……………………..… Apiothyrium
Pellucida Dulym. et al., Mycol. Res. 105(2): 250 (2001) 10. 2-septate, hyaline, lacking a mucilaginous sheath,
Type species: Pellucida pendulina Dulym. et al., in cylindric-clavated to clavate, J-, immersed under the
Dulymamode et al., Mycol. Res. 105(2): 250 (2001) clypeus….....………………………........ Rachidicola
Physalospora Niessl, Verh. nat. Ver. Brünn 14: 170 (1876) 10. More than 2-septate……………………………..… 11
Type species: Physalospora alpestris Niessl, Verh. nat. Ver. 11. 1–3- septate or multiseptate………………………. 12
Brünn 14: 170 (1876) 11. (l-)3(−5)-septate, hyaline, cylindrical to slightly sac-
Phragmitensis M.K.M. Wong et al., Bot. Mar. 41(4): 379 cate, J+, immersed, opening by 3–5 lobes……
(1998) Arwidossonia
Type species: Phragmitensis marina M.K.M. Wong et al., 12. 1–3- septate, long ellipsoid, ornamented with two col-
Bot. Mar. 41(4): 379 (1998) our, gelatinous wings, cylindrical, lacking an apical
Rachidicola K.D. Hyde & J. Fröhl., in Hyde, Sydowia 47(2): apparatus, apothecia erumpent, discoid with periphyses
217 (1995) and paraphyses……………...............… Papilionovela
Type species: Rachidicola palmae K.D. Hyde & J. Fröhl., 12. 3-septate or more than………………………..…… 13
in Hyde, Sydowia 47(2): 218 (1995) 13. 3-septae……………………………………....…… 14
 Fungal Diversity

13. Multiseptate………………………………………. 15 basally immersed, globose to subglobose, lageniform to

14. 3-septae, fusoid, hyaline, cylindrical, immersed, global, obpyriform, brown to black or externally with yellowish pig-
filiform paraphyses. . . . . . . . . . . . . . . . . . . Cesatiella ments, solitary to gregarious, papillate with long upright neck.
14. 3-septage, ellipsoid to fusiform-oblong, hyaline-yel- Peridium 3-layered, comprising cells of textura angularis or
low, immersed in clusters under a clypeus, filiform textura prismatica or textura intricata, some with an orange,
paraphyses…………………………...... Exarmidium middle wall layer. Hamathecium with numerous, filamentous,
15. Filiform, hyaline ascospore……………………….. 16 septate, hyaline paraphyses. Asci 8-spored, unitunicate, cylin-
15. Oblong, hyaline ascospore, uniseriate, lacking a pa- drical to cylindric-clavate, short pedicellate, with a large, re-
raphyses………….....………………… Xenothecium fractive, J-, apical ring, rarely with delicate ring like structures.
16. Cylindrical asci, erumpent, globose, Ostiole with papil- Ascospores uniseriate to overlapping uniseriate, reddish
la, paraphyses……………………….…. Micronectria brown to brown, to greenish brown, ellipsoidal to oblong-
16. Filiform, hyaline ascospore……..…………. Pellucida elliptical, to fusiform, 1-septate, with germ pores at one or
both ends. Asexual morph: Undetermined.
Iodosphaeriaceae O. Hilber, in Hilber & Hilber, The Genus Type: Jobellisia M.E. Barr; Fig. 74
Lasiosphaeria and Allied Taxa (Kelheim): 7 (2002) Type species: Jobellisia luteola (Ellis & Everh.) M.E. Barr,
Facesoffungi number: FoF 01905 Mycotaxon 46: 61 (1993)
Saprobic on plant hosts. Sexual morph: Ascomata super- Notes: The monotypic family Jobellisiaceae was intro-
ficial, solitary, black, and easily removed from the substrate, duced by Réblová (2008) based on LSU rDNA sequence data.
ascomata covered with dark brown, setae-like, hairs, compris- This family is characterized by superficial or basally immersed,
ing agglutinated mycelial strands, with a stellate arrangement, astromatic, brown or yellowish, pigmented ascomata, and ellip-
arising from cells at the perithecial surface. Ostiole pore-like, soidal, 1-septate, reddish brown to brown ascospores, with
periphysate. Peridium outer region comprising angular, darker pigment at the median septum, with germ pores at one
pigmented, brown cells, inner region comprising flattened, hy- or both ends. The genus is found in tropical and temperate zones
aline cells. Paraphyses hypha-like, septate, flexuose, numerous, of the northern hemisphere (Réblová 2008). According to
slightly tapered towards the apex. Asci 8-spored, unitunicate, the phylogenetic analysis of Réblová (2008), the closest
narrowly clavate, short pedicellate or apedicellate, apex round- relatives of Jobellisia and the family Jobellisiaceae were
ed, with a J+, subapical ring. Ascospores biseriate, allantoid, members of the orders Diaporthales, Calosphaeriales and
unicellular, hyaline, smooth-walled, lacking sheaths or append- Togniniaceae. In the phylogenetic analyses carried out by
ages. Asexual morph: Ceratosporium-like conidia have been Maharachchikumbura et al. (2015), this family is raised to
observed on the surface of perithecia, but may not be related. a new order Jobellisiales and is phylogenetically closer to
Type: Iodosphaeria Samuels et al., Mycotaxon 28(2): 486 Togniniales and Calosphaeriales.
(1987); Fig. 73 The genus currently comprises nine species. Jobellisia was
Type species: Iodosphaeria phyllophila (Mouton) Samuels placed in Clypeosphaeriaceae by Barr (1994), and Huhndorf
et al., Mycotaxon 28(2): 486 (1987) et al. (1999) suggested that this genus should be placed in
Notes: The family Iodosphaeriaceae was introduced by Diaporthales. A new family Jobellisiaceae was introduced
Hilber and Hilber (2002) to accommodate the genus by Réblová (2008) to accommodate Jobellisia.
Iodosphaeria. The genus was placed in the order
Amphisphaeriales; as it has superficial ascomata covered with Juncigenaceae E.B.G. Jones et al., in Jones et al., Cryptog.
dark brown, setae-like hairs, comprising agglutinated mycelial Mycol. 35(2): 133 (2014)
strands, with a stellate arrangement (Hilber and Hilber 2002). Li Facesoffungi number: FoF 01665
et al. (2015b) introduced a new species, Iodosphaeria Saprobic growing on substrates, such as intertidal wood,
tongrenensis Li et al., bringing the total of species in the genus mangrove and herbaceous wood and roots, bark, leaves in
to nine. Li et al. (2015b) provided molecular data that showed marine habitats. Sexual morph: Ascomata perithecial,
this taxon clustered in Xylariales and in this study we confirm subglobose to pyriform, immersed to superficial, subcoria-
its distinctness as a family and placement in Xylariomycetidae. ceous to coriaceous, brown to dark-brown, ostiolate,
When Samuels et al. (1987) introduced the genus Iodosphaeria periphysate, papillate. Peridium comprising several cell layers
they deposited a specimen collected in New Zealand in PDD of ellipsoidal to subglobose cells forming textura angularis.
and this was used in this study to illustrate the family (Fig. 73). Hamathecium comprising numerous, narrow, unbranched,
persistent, paraphyses, connected to the apex and base of the
Jobellisiaceae Réblová, Mycologia 100 (6): 899 (2009) ascomatal wall. Asci 8-spored, unitunicate, thin-walled, per-
Facesoffungi number: FoF 01906 sistent, clavate, cylindrical to fusiform, with apical ring, short
Saprobic on wood in terrestrial and freshwater habitats. pedicellate. Ascospores uniseriate, hyaline, ellipsoidal, clavate
Sexual morph: Ascomata perithecial, superficial or erumpent, to fusiform, 1–3-septate with or without equatorial and polar
Fungal Diversity

Fig. 73 Iodosphaeria phyllophila (Material examined: NEW surface of the perithecia h–i Ascus apex with J+, subapical ring j–m
ZEALAND: Kaipara Harbour, Mt Auckland, Atuanui State Forest; 17 Asci with ascospores (m stained in Melzer’s reagent). n–q Ascospores.
November 1973, G.J. Samuels, GJS73-260, PDD 32622) a, b Scale bars: c = 1000 μm, d = 500 μm, e = 50 μm, f, j–m = 10 μm,
Herbarium package and material c, d Ascomata on the surface of host e g = 20 μm, h, i, n–q = 5 μm
Section of ascoma f Peridium g Ceratosporium-like conidium on the
 Fungal Diversity

Fig. 74 Jobellisia luteola (Material examined: USA, Ohio, J.P. Morgan, Section through ascomata g, h Section through peridium showing wall
1109, NY 914363, holotype) a Herbarium label b, c Ascomata on layer i-k Squash mount of asci l Ascus. m–t Ascospores. Scale bars:
substrate d, e Section through neck showing periphyses f b = 1 mm; c = 200 μm; f, g, i, j, k = 50 μm; d = 20 μm; e, m–t = 10 μm

appendages. Asexual morph: hyphomycetous. Hyphae sep- dark brown. Conidia brown, single, helicoid, septate, con-
tate, branched, hyaline to brown. Conidiogenous cells non- stricted at the septa.
specialized, short, lateral, solitary, helicoid, septate, light to Type: Juncigena Kohlm. et al.
Fungal Diversity

Notes: The family Juncigenaceae was introduced by Jones coding and ribosomal nuclear loci (Schoch et al. 2007) has
et al. (2014) with Juncigena as the type genus and included the shown that Juncigena adarca clusters with Swampomyces sp.
genera Fulvocentrum, Marinokulati and Moheitospora. They and is associated with Coronophorales with good support.
formed a monophyletic clade with high statistical support However, Jones et al. (2009b) referred Juncigena to
based on analysis of SSU and LSU sequence data. The family Hypocreales incertae sedis and Jones et al. (2014) transferred
forms sister clades to the Etheirophoraceae, Falcocladiacee and this genus in to a newly introduced family Juncigenaceae.
Torpedosporaceae in the subclass Hypocreomycetidae. Jones
et al. (2014) showed that the species Swampomyces aegyptiacus
Abdel-Wahab, El-Shar. & E.B.G. Jones and S. clavatispora Other genera included
Abdel-Wahab, El-Shar. & E.B.G. Jones did not group in the
genus Swampomyces sensu stricto and introduced the genus Fulvocentrum E.B.G. Jones & Abdel-Wahab, Cryptog.
Fulvocentrum to accommodate them. Likewise, the marine as- Mycol. 35(2): 131 (2014)
comycete Chaetosphaeria chaetosa Kohlm. did not group in Type species: Fulvocentrum aegyptiaca (Abdel-Wahab
Chaetosphaeria sensu stricto (Chaetosphaeriales) and was et al.) E.B.G. Jones & Abdel-Wahab, in Jones et al.,
transferred to a new genus Marinokulati (Jones et al. 2014). Cryptog. Mycol. 35(2): 131 (2014)
Moheitospora ruticosae Abdel-Wahab, Abdel-Aziz & Nagah., Marinokulati E.B.G. Jones & K.L. Pang, Cryptog. Mycol.
an asexual morph, groups in this family with high support. Jones 35(2): 132 (2014)
et al. (2014) referred the Juncigenaceae to the subclass Type species: Marinokulati chaetosa (Kohlm.) E.B.G.
Hypocreomycetidae, order incertae sedis with the Hypocreales Jones & K.L. Pang, Cryptog. Mycol. 35(2): 133 (2014)
as a sister order and this was followed in Maharachchikumbura Moheitospora Abdel-Wahab et al., Mycol. Progr. 9(4): 551
et al. (2015). Subsequently, Jones et al. (2015) introduced the (2010)
order Torpedosporales to accommodate the three marine fami- Type species: Moheitospora fruticosae Abdel-Wahab
lies: Etheirophoraceae, Juncigenaceae and Torpedosporaceae et al., Mycol. Progr. 9(4): 551 (2010)

Juncigena Kohlm. et al., Bot. Mar. 40(4): 291 (1997)

Facesoffungi number: FoF 02113; Figs 75, 76 Key to genera of Juncigenaceae
Saprobic, a salt marsh fungus on the sedge Juncus
roemerianus, which is obligate marine. Sexual morph: 1. Coelomycete with helicoid conidia . . . . . . . Moheitospora
Ascomata solitary, subglobose to pyriform, coriaceous, fus- 1. Ascomycete with fusiform to elongate ellipsoidal asco-
cous, immersed, ostiolate, papillate. Ostiolar canal cylindrical, spores …………………………………....……….......2
periphysate, hyaline. Peridium comprising several layers of 2. Asci with an attenuate at the base, ascospores with polar
ellipsoidal to subglobose cells, forming a textura angularis, and equatorial appendages . . . . . . . . . . . Marinokulati
darker to hyaline from outside to inside. Hamathecium com- 2. Asci with a short pedicel, ascospores lacking
posed of unbranched pseudoparaphyses attached to the top appendages. . . . . . . . . . . . . . . . . . . . . . . 3
and bottom of the ascomatal cavity. Asci 8-spored, fusiform 3. Asci with an apical ring . . . . . . . . . . . . . . . .Juncigena
to cylindrical, short pedicellate, unitunicate, with a J-, apical 3. Asci with thickened apex . . . . . . . . . . . .Fulvocentrum
ring. Ascospores uni or biseriate, hyaline, fusiform to ellipsoi-
dal, 3-septate, constricted at the septa. Asexual morph: Kathistaceae Malloch & M. Blackw., Can. J. Bot. 68(8): 1719
Saprobic on submerged bases of leaves in marine habitats. (1990)
Hyphae hyaline to light brown, septate, branched; in pure Facesoffungi number: FoF 01289
culture producing conidia and multi-celled intercalary chla- Saprobic on herbivore dung and insects. Sexual morph:
mydospores. Conidiogenous cells non-specialized, short, lat- Ascomata globose to subglobose, with a long neck composed
eral, similar to vegetative hyphae; solitary, irregularly heli- of parallel hyphae, straight curved ostiolar setae, with spherical
coid, light to dark brown, deeply constricted at the septa, in- spore-bearing structures (sporidiomata), cylindrical unicellular
creasing in diam. from base to apex. Conidia brown, single, extension present. Peridium thin walled, pseudoparenchyma-
helicoid, septate, constricted at the septa tous, forming cells of textura angularis in surface view and
Type species: Juncigena adarca Kohlm. et al., Bot. Mar. thick walled in cross section. Hamathecium lacking interascal
40(4): 291 (1997) filaments. Asci unitunicate, 8–spored, ellipsoidal to fusoid, thin-
Asexual morph. Cirrenalia adarca Kohlm. et al., Bot. Mar. walled, evanescent at maturity, arranged in a basal fascicle.
40(4): 292 (1997) Ascospores clavate to falcate, hyaline or pale brown, 0–1 or
Notes: This genus was introduced by Kohlmeyer et al. transversely multi septate. Asexual morph: Conidiomata su-
(1997). Eriksson (1999) included Juncigena in the family perficially similar to the ascomata, spherical, hyaline, without
Magnaporthaceae. Analysis of sequence data from protein hyphal attachments, single wall layer thick at base, with or
 Fungal Diversity

Fig. 75 Juncigena adarca

(USA, North Carolina, Broad
Creek (Atlantic Ocean), 34° 43′
00″N, 76° 55′ 07″W (34.717,
−76.919), on standing dead culms
of Juncus roemerianus Scheele
(Juncaceae), 9 October 1994J.
Kohlmeyer (NY01276214,
holotype) a Herbarium material b
Ascoma c, d Ascoma in cross
section e Asci and Ascospores f–l
Ascospores. Note: Figure e and i
are stained in Melzer’s reagent.
Scale bars: b = 200 μm, c–
d = 100 μm, e = 50 μm, f–l = 20

without ostiole, when present ostiole comprising with long and smooth. Conidia spherical or cylindrical, produced in the
neck. Conidiomatal wall composed of flattened cells of textura center of the conidiomata, hyaline, smooth, 0–1 or multi-sep-
angularis. Conidiogenous cells enteroblastic, phialidic, hyaline, tate, escaping in a continuous chain through the ostiolar neck.
Fungal Diversity

Maharachchikumbura et al. (2015) included three genera in

the family Kathistaceae, including both sexual and asexual
morphs. Kathistes is the only sexual genus in Kathistaceae,
while Mattirolella and Termitariopsis are asexual genera.

Kathistes Malloch & M. Blackw., Can. J. Bot. 68(8):

1712 (1990)
Facesoffungi number: FoF 01290; Fig. 77
Saprobic on herbivore dung. Sexual morph: Ascomata
subsphaerical to spherical, hyaline to pale brown, transparent,
glabrous, thin-walled, with a long neck, with hyphal attachments
extending into the substrate. Ostiolar neck long, dark reddish
brown, composed of parallel hyphal elements, terminated by
incurved setae to form a cuplike apex, often with a thickened
collar-like area joined to the perithecium. Peridium one cell
layer thick, comprising cells of textura angularis in surface
view. Hamathecium lacking. Asci 8-spored, unitunicate, ellipsoi-
dal to broadly clavate, short stipitate, tapered below, thin-walled,
evanescent at maturity, arranged in a basal fascicle. Ascospores
narrowly clavate to falcate, hyaline, smooth, initially unicellular,
one septum at initially and more septa at maturity, without pores
or slits, lacking gelatinous sheath, germinating by repetition of
budding yeast-like cells. Asexual morph: Conidiomata super-
ficially similar to the ascomata, spherical, hyaline, with a long
neck, without hyphal attachments. Peridium comprising flat-
tened cells of textura angularis, one cell layer thick. Ostiolar
neck unicellular, dark reddish brown, arising from a single pe-
ridium cell, continuous into the hollow center of the
conidiomata, open apically at maturity. Conidia spherical,
produced in the center of the conidiomata, hyaline, smooth,
escaping in a continuous chain through the ostiolar neck.
Type species: Kathistes calyculata Malloch & M. Blackw.,
Can. J. Bot. 68(8): 1712 (1990)
Notes: The genus Kathistes was introduced as the type
Fig. 76 Cirrenalia adarca (asexual morph of Juncigena adarca). a genus in Kathistaceae by Malloch and Blackwell (1990),
Conidia from decomposing insect larva in leaf of Juncus roemerianus. which was originally described with three species. The genus
b Conidia from culture on seawater agar, isolate made from ascospores of is characterized by long-necked perithecia with setose osti-
J. adarca (redrawn from Kohlmeyer et al. 1997). Scale bars: a, b = 10 μm
oles, basally arranged asci, with hyaline, transversely septate,
ascospores without pores, slits, or sheaths. A budding yeast-
Type: Kathistes Malloch & M. Blackw. like asexual morph is produced directly from the cells of the
Notes: The monotypic family Kathistaceae was established ascospores in culture.
invalidly by Malloch and Blackwell (1990), to accommodate In addition, preliminary data obtained from sequence data,
the type species Kathistes calyculata, and two more species, show that Kathistes species are different from both
K. fimbriata (Barrasa & G. Moreno) Malloch & Blackw., and Pyxidiophoraceae and Ophiostomataceae, hence a family
K. analemmoides (Malloch & Blackw.). Malloch and Kathistaceae was introduced to accommodate Kathistes
Blackwell (1990) placed Kathistaceae in order (Malloch and Blackwell 1990).
Ophiostomatales based on the characters such as, ellipsoidal
to spherical asci borne uniformly in the centrum. However,
Kathistes, which was characterized with globose to Other genera included
subglobose ascomata, long necked perithecia with setose osti-
oles and ascospore germinating by repetition to produce budding Mattirolella S. Colla, Boll. Lab. Zool. Portici 22: 44 (1929)
yeast-like cells, morphologically does not appear to be closely Type species: Mattirolella silvestrii S. Colla [as
related to other genera (Malloch and Blackwell 1990). ‘silverstrii’], Boll. Lab. Zool. Portici 22: 44 (1929)
 Fungal Diversity

Fig. 77 Kathistes calyculata

(Material examined: CANADA,
New Brunswick, Charlotte Co.,
St. James Parish, Upper Canoose
Flowage, on moose dung in moist
chamber, 8 August 1987; D.
Malloch and M. Blackwell,
TRCT51145, holotype) a, b
Material c Ascomata on substrate
d Ascoma terminated by incurved
setae to form a cup-like apex e
Ascoma neck show substructure f
Apex of ascoma neck g Peridium
h Asci i, j, k, l Ascospores m, n,
o, p Ascospores germinating q
Yeast cells r Sporidiomata
showing single-celled wall. Scale
bars: c = 200 μm, d = 30 μm,
e = 20 μm, f and g, = 10 μm, i, j,
k and l = 5 μm (redrawn from
Malloch and Blackwell 1990)
Fungal Diversity

Termitariopsis M. Blackw et al., Mycotaxon 12(1): 98 more or less incrusted with melanin. Paraphyses simple, rare-
(1980) ly branched, septate, filling immature ascomata, later sur-
Type species: Termitariopsis cavernosa M. Blackw et al., rounding the asci. Asci 8-spored, clavate to ellipsoidal, lacking
Mycotaxon 12(1): 98 (1980) an apical ring, with a short pedicel, dissolving at maturity,
unitunicate, arising from an ascogenous tissue at the base of
the locule. Ascospores overlapping, ellipsoidal to oval, with
Key to genera of Kathistaceae several cross septa, mostly septate near the apices, hyaline,
thick-walled, lacking a sheath or appendages, germinating
1. Conidiomata hyaline, with a long neck . . . . . . . .Kathistes apically. Asexual morph: Spermatia enteroblastic, with ba-
1. Conidiomata black, without neck . . . . . . . . . . . . . . . . 2 sipetal succession, subglobose, wall cup shaped, hyaline,
2. Phialoconidia . . . . . . . . . . . . . . . . . . . . . . .Mattirolella sticky.
2. Macroconidia . . . . . . . . . . . . . . . . . . . . .Termitariopsis Type species: Koralionastes ovalis Kohlm. & Volkm.-
Kohlm., Mycologia 79(5): 765 (1987)
Koralionastetaceae Kohlm. & Volkm.-Kohlm., Mycologia Notes: Koralionastes was introduced by Kohlmeyer and
79(5): 764 (1987) Volkmann-Kohlmeyer (1987) with three species,
Facesoffungi number: FoF 01391 Koralionastes angustus Kohlm. & Volkm.-Kohlm.,
Saprobic on coralline covered rocks, sponges or on algae, K. ellipticus Kohlm. & Volkm.-Kohlm. and K. ovalis
found in marine habitats. Sexual morph: Ascomata black, Kohlm. & Volkm.-Kohlm. (the type species) encountered from
subglobose, ovoid or ellipsoidal, superficial, ostiolate, papil- corals and placed under a novel family Koralionastetaceae.
late or epapillate, periphysate, subiculate, or without Subsequently, two new species, K. giganteus Kohlm. &
subiculum. Hamathecium composed of simple, septate pa- Volkm.-Kohlm. and K. violaceus Kohlm. & Volkm.-Kohlm.
raphyses. Asci 8-spored, unitunicate, clavate to ellipsoidal or were introduced by Kohlmeyer and Volkmann-Kohlmeyer
fusiform, pedicellate, deliquescent. Ascospores overlapping, (1990). Koralionastes species are found on dead coralline
hyaline, ellipsoidal to fusiform, multi-septate near the apices, covered rocks among crustose sponges and their ascospores
or filiform and evenly multi-septate, thick or thin walled, ger- have double walls and are ellipsoidal or fusiform (Campbell
minating apically, lacking appendages. Asexual morph: et al. 2009). The genera Koralionastes and Pontogenia differ
Spermatia enteroblastic, subglobose. from members of the Lulworthiales by the possession of
Type: Koralionastes Kohlm. & Volkm.-Kohlm. paraphyses and periphyses, and lack apical mucous-
Notes: The family Koralionastetaceae was introduced by containing chambers or gelatinous sheaths.
Kohlmeyer and Volkmann-Kohlmeyer (1987) to include a
single genus Koralionastes, based on the thick-walled asco-
spores that germinate into hyphae bearing phialidic antheridia Other genus included
with enteroblastic spermatia. However, the ordinal and higher
classification of the family Koralionastetaceae was un- Pontogeneia Kohlm., Bot. Jb. 96(1–4): 201 (1975)
clear and Koralionastes was referred to Ascomycota Facesoffungi number: FoF 02115; Fig. 79
genera incertae sedis (Eriksson 2006). A molecular study with Parasitic on marine algae (Phaeophyta). Sexual morph:
SSU and LSU sequence data, with supporting morphological Ascomata solitary to gregarious, subglobose or ovoid, super-
characteristics, enabled the assignment of the genera ficial or semi immersed ostiolate, papillate or epapillate, cori-
Koralionastes and Pontogeneia to the new order aceous, black. Paraphyses distinct, thick, septate, or indistinct.
Koralionastetales, which is a sister group to the order Asci 8-spored, clavate or fusiform, unitunicate, thin walled,
Lulworthiales (Campbell et al. 2009). The orders early deliquescing, attached in the lower part of the locule.
Lulworthiales and Koralionastetales clustered together in a Ascospores filiform, rearly ellipsoidal, hyaline, often curved,
well-supported clade based on combined sequence data and phragmosporous. Asexual morph: Undetermined.
a new subclass Lulworthiomycetidae was introduced by Type species: Pontogeneia padinae Kohlm., Bot. Jb. 96(1–
Maharachchikumbura et al. (2015). 4): 201 (1975)
Koralionastes Kohlm. & Volkm.-Kohlm., Mycologia 79(5): Notes: The genus Pontogeneia was introduced by
765 (1987) Kohlmeyer (1975) for five perithecial ascomycetes parasitiz-
Facesoffungi number: FoF 02114; Fig. 78 ing marine Phaeophyta. Two more species have since been
Saprobic on coralline covered rocks, sponges or on algae, added (Kohlmeyer and Kohlmeyer 1979; Kohlmeyer and
found in marine habitats. Sexual morph: Ascomata ellipsoi- Demoulin 1981). Later, Pontogeneia microdictyi Kohlm. &
dal, superficial, ostiolate, periphysate, papillate, black, Volkm.-Kohlm was introduced to this genus by Campbell
subiculate, single. Peridium composed of several layers of et al. (2009). All Pontogeneia species are associated with ma-
pseudoparenchymatous cells, forming a textura angularis, rine algae, belonging to the Chlorophyta or Phaeophyta
 Fungal Diversity

Fig. 78 Koralionastes ovalis (Material examined: BELIZE, back reef of Section through ascoma (NY 1317811) c Section through peridium d–g
South water Cay, sub tidal coral slab (0.2–0.5 m deep), November 1986, Asci (NY 1317828) h Paraphyses (NY 1317828) i–l Ascospores (NY
J. Kohlmeyer; NY 1317811, NY 1317823, NY 1317828, NY 1317838, 1317838). Scale bars: b, c, h = 100 μm, d-g, i–l = 50 μm
slides from holotype) a Microslides of Koralionastes ovalis b

(Kohlmeyer and Kohlmeyer 1979; Kohlmeyer and Demoulin order Koralionastetales (C am pbell et al . 2009).
1981). Based on the combined dataset of SSU and LSU rDNA Maharachchikumbura et al. (2015) and Jones et al. (2015)
sequences and morphological characters, the genera confirmed the placement of both Koralionastes and
Koralionastes and Pontogeneia were assigned to the new Pontogeneia under Koralionastetaceae, Koralionastetales.
Fungal Diversity
 Fungal Diversity

ƒFig. 79 Pontogeneia padinae (Material examined: Intertidal zone at the 1957; Carroll and Munk 1964; Lundqvist 1972; Kruys et al.
Marine Laboratory of the University of Arizona, on living plants near the 2014). The family Lasiosphaeriaceae forms a sister clade to
base among rhizoids of the holdfast, 22 June 1974, J. J. Kohlmeyer, NY
985934, holotype) a, b Herbarium material of Pontogeneia padinae c
Chaetomiaceae with high support (Maharachchikumbura
Ascomata on the base of algae d section through ascomata e Peridium f–i et al. 2015) and has 35 accepted genera. The family needs to
Ascospores. Scale bars: d = 100 μm, e = 20 μm, f–i = 10 μm be revised following a thorough molecular study.

Key to the genera of Koralionastetaceae Lasiosphaeria Ces. & De Not., Comm. Soc. crittog. Ital. 1(4):
229 (1863)
1. On coral rock, associated with crustose Facesoffungi number: FoF 01146, Fig. 80
sponges . . . . . . . . . . . . . . . . . . . . . . . . . . Koralionastes Saprobic on wood, rotting vegetation, dung of herbivores,
1. On algae . . . . . . . . . . . . . . . . . . . . . . . . . Pontogeneia in fresh water or terrestrial habitats, many coprophilous.
Sexual morph: Ascomata perithecial, gregarious or scattered,
Lasiosphaeriaceae Nannf., Nova Acta R. Soc. Scient. upsal., brown to black, solitary, superficial or sub-immersed, globose,
Ser. 4 8(no. 2): 50 (1932) subglobose to ovoid, carbonaceous, coriaceous or
Facesoffungi number: FoF 01145 membranaceous, ornamented or smooth, with variously
Saprobic on wood, rotting vegetation, dung of herbivores, coloured tomentum below neck, papillate or indistinct, sitting
in freshwater, marine or terrestrial habitats, many coprophi- on brown, septate subiculum, ostiolate and periphysate.
lous. Sexual morph: Ascomata perithecial or cleistothecial, Peridium thick (30–80 μm), composed of two layers, outer
gregarious or scattered, yellow, brown to black, solitary, su- layer comprising brown cells of textura angularis, carbona-
perficial, erumpent or immersed, globose, subglobose to ceous or coriaceous; inner layer comprising hyaline cells of
ovoid, carbonaceous, coriaceous or membranaceous, textura prismatica, thin, membranaceous. Hamathecium, with
ornamented, tuberculate or smooth, with setae or hairs, papil- numerous, hyaline, septate, filiform paraphyses. Asci 8-
late or papilla indistinct or absent, ostiolate, periphysate, the spored, unitunicate, thin walled, cylindrical to clavate, pedi-
apex collapsing when dry. Peridium thick (10–100 μm), com- cellate, J-. Ascospores uni to biseriate, hyaline, yellowish to
posed of two layers, outer layer comprising brown cells of brown, allantoid, cylindrical to ellipsoid, curved or not,
textura angularis or globulosa, carbonaceous, coriaceous or smooth, gelatinous appendages present or absent, with or
membranaceous; inner layer comprising hyaline cells of without guttules, one end swollen or not. Asexual morph:
textura prismatica or porrecta, thin, membranaceous. Undetermined.
Hamathecium with numerous, hyaline, septate, filiform or cy- Type species: Lasiosphaeria ovina (Pers.) Ces. & De Not.,
lindrical paraphyses. Asci 4- or 8-spored, unitunicate, thin or Comm. Soc. crittog. Ital. 1(4): 229 (1863)
thick-walled, cylindrical to clavate, pedicellate, J-. Ascospores Notes: The genus Lasiosphaeria was introduced by Cesati
uni-seriate to irregularly biseriate, allantoid, clavate, cylindri- and De Notaris (1863), to accommodate the type species
cal, ellipsoid to dumbbell-shaped, curved or not, 0–multi-sep- L. ovina. This genus was included in Lasiosphaeriaceae, un-
tate, concolourous or versicolorous, hyaline, brown or black, der the Sordariales based on its brown ascospores with pores
divided, ornamented or smooth, with or without germ pore, (Barr 1990). It is characterized by tuberculate ascomata, cov-
appendages present or absent, with or without guttules. ered with variously coloured tomenta, carbonaceous
Asexual morph: Hyphomycetous. Conidiophores ascomata, numerous paraphyses, clavate asci and diverse-
macronematous or mononematous, scattered or gregarious, shaped ascospores.
brown, straight, septate, branched, smooth-walled.
Conidiogenous cells enteroblastic or holoblastic, phialidic,
hyaline to dark brown, subglobose to ampulliform, proliferat- Other genera included
ing percurrently, with collarette. Conidia solitary, globose,
subglobose to cylindrical, hyaline to brown, smooth, aseptate. Angulimaya Subram. & Lodha, Antonie van Leeuwenhoek
Type: Lasiosphaeria Ces. & De Not 30: 329 (1964)
Notes: The family Lasiosphaeriaceae was introduced by Type species: Angulimaya sundara Subram. & Lodha,
Nannfeldt (1932) based on black ascomata and cylindrical Antonie van Leeuwenhoek 30: 329 (1964)
asci, with brown to hyaline ascospores, and is typified by Anopodium N. Lundq., Bot. Notiser 117: 356 (1964)
Lasiosphaeria (Cesati and de Notaris 1863). This family is Type species: Anopodium ampullaceum N. Lundq., Bot.
polyphyletic (Barr 1990) and this was confirmed by Notiser 117: 356 (1964)
Huhndorf (2004a) based on analysis of LSU sequence data. Apiosordaria Arx & W. Gams, Nova Hedwigia 13: 201
Lasiosphaeriaceae is similar to Nitschkiaceae and (1967)
Tripterosporaceae (Barr 1990), the former differs by having Type species: Apiosordaria verruculosa (C.N. Jensen) Arx
turbinate, ostiolate ascomata and septate ascospores (Munk & W. Gams, Nova Hedwigia 13: 201 (1967)
Fungal Diversity
 Fungal Diversity

ƒFig. 80 Lasiosphaeria ovina (Material examined: SWEDEN, Småland, Type species: Emblemospora monotrema Jeng & J.C.
Högsby par., Massemåla, on decayed branch of apple tree in old pasture, Krug, Can. J. Bot. 54(16): 1972 (1976)
N. Lundqvist, 3 October 1992, S F-123316). a Material label b
Specimens c Ascomata d Ascoma cross section e Peridium. f
Eosphaeria Höhn., Annls mycol. 15(5): 362 (1917)
Paraphyses g-j Asci k-o Ascospores Notes: Figs f-o soaked in 3 % Type species: Eosphaeria uliginosa (Kunze) Höhn., Annls
KOH. Scale bars: c = 200 μm, d = 100 μm, e, g-j = 20 μm, f, k–o = 5 μm mycol. 15(5): 362 (1917)
Fimetariella N. Lundq., Bot. Notiser 117: 239 (1964)
Type species: Fimetariella rabenhorstii (Niessl) N.
Apodospora Cain & J.H. Mirza, Can. J. Bot. 48(5): 891 Lundq., Bot. Notiser 117: 239 (1964)
(1970) Immersiella A.N. Mill. & Huhndorf, Mycol. Res. 108(1):
Type species: Apodospora simulans Cain & J.H. Mirza, 31 (2004)
Can. J. Bot. 48(5): 891 (1970) Type species: Immersiella immersa (P. Karst.) A.N. Mill.
Apodus Malloch & Cain, Can. J. Bot. 49(6): 872 (1971) & Huhndorf, Mycol. Res. 108(1): 31 (2004)
Type species: Apodus deciduus Malloch & Cain, Can. J. Jugulospora N. Lundq., Symb. bot. upsal. 20(no. 1): 256
Bot. 49(6): 872 (1971) (1972)
Arniella Jeng & J.C. Krug, Mycologia 69(1): 73 (1977) Type species: Jugulospora rotula (Cooke) N. Lundq.,
Type species: Arniella echoura Jeng & J.C. Krug, Symb. bot. upsal. 20(no. 1): 260 (1972)
Mycologia 69(1): 74 (1977) Lacunospora Cailleux, Cahiers de La Maboké 6(2): 93
Arnium Nitschke ex G. Winter, Bot. Ztg. 31: 450 (1873) (1969) [1968]
Type species: Arnium lanuginosum Nitschke [as Type species: Lacunospora stercoraria Cailleux, Cahiers
‘lanuginosa’], Bot. Ztg. 31: 450 (1873) de La Maboké 6(2): 96 (1969) [1968]
Bellojisia Réblová, Mycologia 100(6): 897 (2008) Mammaria Ces. ex Rabenh., Bot. Ztg. 12: 190 (1854)
Type species: Bellojisia rhynchostoma (Höhn.) Réblová, Type species: Mammaria echinobotryoides Ces., in
Mycologia 100(6): 897 (2008) Rabenhorst, Klotzschii Herb. Viv. Mycol., Editio novo.
Biconiosporella Schaumann, Veröff. Inst. Meeresf. Century 19: no. 1859 (1854)
Bremerhaven 14(1): 24 (1972) Melanocarpus Arx, Stud. Mycol. 8: 17 (1975)
Type species: Biconiosporella corniculata Schaumann, Type species: Melanocarpus albomyces (Cooney & R.
Veröff. Inst. Meeresf. Bremerhaven 14(1): 24 (1972) Emers.) Arx, Stud. Mycol. 8: 175 (1975)
Bombardia (Fr.) P. Karst., Bidr. Känn. Finl. Nat. Folk 23: Periamphispora J.C. Krug, Mycologia 81(3): 476 (1989)
20 (1873) Type species: Periamphispora phacelodes J.C. Krug,
Type species: Bombardia fasciculata Fr., Bidr. Känn. Finl. Mycologia 81(3): 476 (1989)
Nat. Folk 23: 20 (1873) Podospora Ces., in Rabenhorst, Klotzschii Herb. Viv.
Bombardioidea C. Moreau ex N. Lundq., Symb. bot. Mycol.: no. 259 (vel 258) (1856)
upsal. 20(no. 1): 274 (1972) Type species: Podospora fimiseda (Ces. & De Not.) Niessl,
Type species: Bombardioidea bombardioides (Auersw.) Hedwigia 22: 156 (1883)
Moreau, Encyclop. Mycol. 25(1): 317 (1954) Pseudocercophora Subram. & Sekar, Journal of the
Camptosphaeria Fuckel, Jb. nassau. Ver. Naturk. 23–24: Singapore National Academy of Science 15: 58 (1986)
140 (1870) [1869–70] Type species: Pseudocercophora ingoldii Subram. &
Type species: Camptosphaeria sulphurea Fuckel, Jb. nas- Sekar, Journal of the Singapore National Academy of
sau. Ver. Naturk. 23–24: 140 (1870) [1869–70] Science 15: 58 (1986)
Cercophora Fuckel, Jb. nassau. Ver. Naturk. 23–24: 244 Rinaldiella Deanna A et al., in Crous et al., Persoonia, Mol.
(1870) [1869–70] Phyl. Evol. Fungi 32: 301 (2014)
Type species: Cercophora mirabilis Fuckel, Jb. nassau. Type species: Rinaldiella pentagonospora Deanna A.
Ver. Naturk. 23–24: 245 (1870) [1869–70] Sutton et al., in Crous et al., Persoonia, Mol. Phyl. Evol.
Cladorrhinum Sacc. & Marchal, in Marchal, Bull. Soc. R. Fungi 32: 301 (2014)
Bot. Belg. 24(1): 64 (1885) Schizothecium Corda, Icon. fung. (Prague) 2: 29 (1838)
Type species: Cladorrhinum foecundissimum Sacc. & Type species: Schizothecium fimicola Corda [as
Marchal, in Marchal, Bull. Soc. R. Bot. Belg. 24(1): 64 (1885) ‘fimicolum’], Icon. fung. (Prague) 2: 29 (1838)
Diffractella Guarro et al., Syst. Ascom. 10: 107 (1991) Strattonia Cif., Sydowia 8(1–6): 245 (1954)
Type species: Diffractella curvata (Fuckel) Guarro et al., Type species: Strattonia tetraspora (R. Stratton) Cif.,
Syst. Ascom., Reprint of Volumes 1–4 (1982–1985) 10(2): Sydowia 8(1–6): 245 (1954)
108 (1991) Thaxteria Sacc., Syll. fung. (Abellini) 9: 687 (1891)
Emblemospora Jeng & J.C. Krug, Can. J. Bot. 54(16): Type species: Thaxteria kunkelii Giard, C. r. Seanc. Soc.
1971 (1976) Biol. 44(4): 156 (1892)
Fungal Diversity

Triangularia Boedijn, Annls mycol. 32(3/4): 302 (1934) 15. Ascospores versicolored. . . . . . . . . . . . . . . . . . . . . 16
Type species: Triangularia bambusae (J.F.H. Beyma) 16. Ascomata ovoid or pyriform . . . . . . . . . Triangularia
Boedijn, Annls mycol. 32(3/4): 302 (1934) 16. Ascomata globose or subglobose . . . . . . . . Strattonia
Tripterosporella Subram. & Lodha, Curr. Sci. 37: 246 17. Ascospores with 1 or 2 germ pores . . . . . . . . . . . . 18
(1968) 17. Ascospores without or with inconspicuous germ
Type species: Tripterosporella coprophila Subram. & pore . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
Lodha, Curr. Sci. 37: 246 (1968) 18. Ascospores with 2 germ pores . . . . . . . . . . . . . . . . 19
Zygopleurage Boedijn, Persoonia 2(3): 316 (1962) 18. Ascospores with a single germ pore . . . . . . . . . . . . 24
Type species: Zygopleurage zygospora (Speg.) Boedijn, 19. Ascospores ellipsoid to fusiform, surround by a gelat-
Persoonia 2(3): 316 (1962) inous sheath . . . . . . . . . . . . . . . . . . . .Bombardioidea
Zygospermella Cain, Mycologia 27(2): 227 (1935) 19. Ascospores comprising two ellipsoid, not surround
Type species: Zygospermella setosa (Cain) Cain, by a gelatinous sheath, with short
Mycologia 27(2): 227 (1935) appendages . . . . . . . . . . . . . . . Zygopleurage
19. Ascospores cylindrical, with long appendages at the
ends. . . . . . . . . . . . . . . . . . . . . . . . . . Zygospermella
Key to genera of Lasiosphaeriaceae 20. Ascomata with hairs or setae . . . . . . . . . . . . . . . . . 21
20. Ascomata without hairs . . . . . . . . . . . . . Immersiella
1. Hyphomycete . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 21. Ascomata concolourous . . . . . . . . . . . . . . . . . . . . . 22
1. Ascomycete . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 21. Ascomata versicolored . . . . . . . . . . . . . . .Anopodium
2. Conidia brown to dark brown. . . . . . . . . . . Mammaria 22. Ascomata ovoid to obpyriform. . . . . . . . . . . . . . . . 23
2. Conidia hyaline 3 22. Ascomata globose . . . . . . . . . . . . . . . . . . Diffractella
3. Conidia globose to subglobose. . . . . . . . . Angulimaya 22. Ascomata pyriform to subglobose . . . . . . Rinaldiella
3. Conidia ellipsoid to obovoid . . . . . . . . Cladorrhinum 23. Ascospores without appendages . . . . . . . Jugulospora
4. Asci 4-spored. . . . . . . . . . . . . . . . . . . . . Lacunospora 23. Ascospores with appendages at ends . . . . Podospora
4. Asci 8-spored. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 24. Ascomata smooth. . . . . . . . . . . . . . . . . . Apodospora
5. Ascomata carbonaceous . . . . . . . . . . . . . . . . . . . . . . . 6 24. Ascomata with hairs or tomentose . . . . . . . . . . . . . 25
5. Ascomata coriaceous or membranaceous . . . . . . . . . . 9 25. Ascomata tomentose, with cylindrical
6. Ascomata pyriform . . . . . . . . . . . . . . Camptosphaeria neck . . . . . . . . . . . . . . . . . . . . Bellojisia
6. Ascomata subglobose to ovoid. . . . . . . . . . . . . . . . . . 7 25. Ascomata with hairs. . . . . . . . . . . . . . . . . . . . . . . . 26
7. Ascomata covered with variously coloured tomentum 26. Ascomata with swollen agglutinated
below neck . . . . . . . . . . . . . . . . . . . . . .Lasiosphaeria hairs . . . . . . . . . . . . . . . . . . . . . . . . Schizothecium
7. Ascomata without tomentum below neck, with setae or 26. Ascomata with cylindrical hairs . . . . . . . . . . . . . . . 27
tuberculate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 27. Ascospores concolourous . . . . . . . . . . . . . . . . . . . . 28
8. Ascomata with hairy asexual morphs in 27. Ascospores versicolored. . . . . . . . . . . . .Apiosordaria
Mammaria . . . . . . . . . . . . . . . . . . Pseudocercophora 28. Ascospores surround by a gelatinous sheath . . . . . . 29
8. Ascomata without hairy asexual morphs, ascospores L- 28. Ascospores with appendage at ends . . . . . . . . . . . . 30
shaped, multi-septate, with short appendages at the ends, 28. Ascospores without appendages and sheath . . . . . . 31
sexual morphs Cladorrhinum-like . . . . . . . Eosphaeria 29. Asci with apical ring . . . . . . . . . . . . . . . . . . . Apodus
9. Ascomata coriaceous . . . . . . . . . . . . . . . . . . . . . . . . 10 29. Asci lacking apical ring . . . . . . . . . . . . . . . . Arniella
9. Ascomata membranaceous . . . . . . . . . . . . . . . . . . . . 17 30. Ascospores oval to ellipsoid . . . . . . . . . . . . . .Arnium
10. Marine fungi . . . . . . . . . . . . . . . . . . Biconiosporella 30. Ascospores cylindrical, straight or the lower cell curved
10. Not marine fungi . . . . . . . . . . . . . . . . . . . . . . . . . . 11 when mature, upper cell ventricose. . . . . Cercophora
11. Ascomata cleistothecial . . . . . . . . . . Tripterosporella 31. Ascospores oval to ellipsoid, divided in all
11. Ascomata perithecial . . . . . . . . . . . . . . . . . . . . . . . 12 directions . . . . . . . . . . . . . . . . . . . . . Emblemospora
12. Ascomata sitting on or in the subiculum. . . . . . . . . 13 31. Ascospores globose to subglobose, smooth-
12. Ascomata superficial to immersed on the bark, without walled . . . . . . . . . . . . . . . . . . . . . . . Melanocarpus
a subiculum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
13. Ascospores cylindrical to ellipsoid, Lautosporaceae Kohlm. et al., Bot. Mar. 38(2): 169 (1995)
guttulate . . . . . . . . . . . . . . . . . . . . . . . . Bombardia Facesoffungi number: FoF 01764
13. Ascospores allantoid . . . . . . . . . . . . . . . . . Thaxteria Saprobic on submerged mangrove wood in marine habi-
14. Ascomata with hairs. . . . . . . . . . . . . . . . . . . . . . . . 15 tats. Sexual morph: Ascomata fusiform to ellipsoidal,
14. Ascomata without hairs . . . . . . . . . . . . . Fimetariella ostiolate, brown, coriaceous, solitary. Hamathecium of sim-
15. Ascospores concolourous . . . . . . . . . Periamphispora ple, septate persistent paraphyses. Asci 4-spored, cylindrical,
 Fungal Diversity

short pedicellate, thick-walled, unitunicate, with an ocular ocular chamber, thick-walled, pedicellate. Ascospores
chamber. Ascospores uni- or biseriate, fusiform, muriform, muriform, fusiform, hyaline, septate, thick-walled. Asexual
distoseptate, hyaline, outer wall very thick. Asexual morph: morph: Undetermined.
Undetermined. Type species: Lautospora gigantea K.D. Hyde & E.B.G.
Type: Lautospora K.D. Hyde & E.B.G. Jones Jones, Bot. Mar. 32(3): 479 (1989)
Notes: Kohlmeyer et al. (1995) introduced this family for Notes: Lautospora was introduced to include taxa with
two marine Lautospora species, Lautospora gigantea K.D. immersed ascocarps, asci which are unitunicate, 4-spored,
Hyde & E.B.G. Jones and Lautospora simillima Kohlm. et al., and with an ocular chamber and a ring, cellular
thought to possess bitunicate asci and were not assigned to any pseudoparaphyses and large ascospores, which are muriform,
order in the Dothideomycetes (Kohlmeyer et al. 1995). Based on hyaline and thick walled (Hyde and Jones 1989). Kohlmeyer
LSU sequence data (S. Suetrong, unpublished data), L. simillima et al. (1995) introduced the family Lautosporaceae to include
was referred to an unnamed clade in Sordariomycetes and in a this genus and assigned it to Ascomycotina incertae sedis.
basal clade to the orders Diaporthales (Jones et al. 2015). Jones
et al. (2015) showed that L. simillima groups with the neotropical Lopadostomataceae Daranagama & K.D. Hyde, [as
ascomycete Mirannulata samuelsii Huhndorf et al. with weak ‘Lopadostomaceae’] in Senanayake et al., Fungal Divers 73:
support, but they share few similar morphological features 1 (2015)
(Huhndorf et al. 2003). The Lautospora/Mirannulata clade Facesoffungi number: FoF 00071
formed a sister group to Vertexicola caudatus K.D. Hyde et al. Saprobic on dead wood. Sexual morph: Pseudostromata
and Rhamphoria delicatula Niessl (Annulatascaceae) in an un- (for Lopadostoma) immersed to erumpent, appearing as dark-
supported clade hence, Jones et al. (2015) emended the diagnosis ened areas, densely gregarious, scattered, compact. Stromata
of the family Lautosporaceae. (for Creosphaeria) erumpent to nearly superficial, with abrupt
margins, multi-perithecial, coalescing in linear rows, dull
Lautospora K.D. Hyde & E.B.G. Jones, Bot. Mar. 32(3): 479 black, waxy, roughened, flattened at the top, with inconspic-
(1989) uous perithecial mounds. Ascomata multi-perithecial, single
Facesoffungi number: FoF 02116; Fig. 81 to multi-layered, clustered, having long ostiolar necks (for
Saprobic on submerged wood of Rhizophora spp. Sexual Lopadostoma) lined with whitish material (for
morph: Ascomata immersed, subglobose in frontal view, fu- Creosphaeria). Ostioles umbilicate or at the same level as
siform in side view, long axis parallel to the host surface, stromatal surface. Peridium amorphous, comprised of dark
coriaceous, ostiolate, brown above, light brown to hyaline brown to black cells. Hamathecium comprising numerous,
below. Peridium 17–65 μm thick composed of several layers long, hyaline, rarely branched paraphyses. Asci (4–)8-spored,
of thin-walled cells. Paraphyses hyaline, septate, persistent. cylindrical, pedicellate, rounded apex, with a J+, apical ring.
Asci uniseriate, 4-spored, cylindrical, unitunicate with an Ascospores uniseriate or partly biseriate, oblong, initially

Fig. 81 Lautospora spp.: a

L. simillima Ascoma immersed in
substratum b Two ascospores
with thick-walled apical region of
ascus (Material examined:
THAILAND, Chumporn
Province, Wetland Development
Office 12, mangrove forest, on
mangrove wood, 18 January 2012
S. Suetrong, BBH 31841) c
L. gigantea (Material examined:
THAILAND, Chachoengsao,
Bang Pakong
Bowornwitthayayon School,
mangrove forest, on mangrove
wood, 20 June 2011 S. Suetrong)
Thick-walled hyaline ascospore.
Scale bars: a = 100 μm,
b = 10 μm, c = 25 μm
Fungal Diversity

hyaline, turning light brown to nearly black, narrowly ellip- Key to the genera of Lopadostomataceae
soid, smooth-walled, with full length, straight germ slit.
Asexual morph: Libertella-like. Conidiomata convoluted or 1. Stromata nearly superficial, effuse, surface dull black,
multi-loculate, with black pycnidial wall. Conidiophores di- waxy, roughened, flattened at the top, depressed and paler
or trichotomously branched, stipes yellow, hyaline above, around the ostioles, the tissue between the perithecia waxy,
smooth-walled, arranged in dense palisade. Conidiogenous orange to black mixed with orange granules, asci with a dis-
cells terminal, cylindrical, smooth-walled, bearing conidial coid, amyloid apical ring . . . . . . . . . . . . . . . . Creosphaeria
secession scars. Conidia hyaline, falcate, strongly curved to 1. Stromata typically immersed in and erumpent from bark,
semicircular, unicellular. subglobose to bluntly conical, surrounded by a narrow, black,
Type: Lopadostoma (Nitschke) Traverso; Fig. 82 carbonized encasement, the tissue between the perithecia
Type species: Lopadostoma turgidum (Pers.) Traverso, Fl. black, lack orange granules, asci with a globose to ellipsoid
ital. crypt., Pars 1: Fungi. Pyrenomycetae. Xylariaceae, apical ring . . . . . . . . . . . . . . . . . . . . . . . . . . . Lopadostoma
Valsaceae, Ceratostomataceae 1 (2): 170 (1906)
Notes: Lopadostomataceae was introduced in Senanayake Lulworthiaceae Kohlm. et al., Mycologia 92(3): 456 (2000)
et al. (2015) in the order Xylariales to accommodate the gen- Facesoffungi number: FoF 01295
era Creosphaeria and Lopadostoma. Creosphaeria was intro- Saprobic on wood or growing on sea grasses and marsh
duced by Theissen (1910) with soft, light coloured stromata. plants. Sexual morph: Ascomata dark brown to black,
Petrini and Müller (1986) reported a Libertella asexual morph subglobose to cylindrical, swan-like, immersed or superficial,
from culture of Creosphaeria (Fig. 83), while Ju et al. (1993) coriaceous, ostiolate, papillate, sometimes with a long neck.
observed the same type of conidia associated with the sexual Hamathecium absent; centrum initially filled with a hyaline
morph. Jaklitsch et al. (2014) revised the genus Lopadostoma pseudoparenchyma, dissolving at maturity. Peridium mostly
and also reported a Libertella asexual morph. The species 2-layered, composed of an outer layer of cells of textura
L. americanum Jaklitsch et al., L. fagi Jaklitsch et al., angularis and an inner layer of elongate cells. Asci 8-spored,
L. linospermum (Durieu & Mont.) Jaklitsch et al., unitunicate, cylindrical to fusiform, short pedicellate, deli-
L. meridionale Jaklitsch et al. and L. turgidum (Pers.) quescent. Ascospores fasciculate, hyaline, filiform, septate or
Traverso, are known to have libertella-like asexual morphs. aseptate, with or without a mucus-containing, conical or semi-
Ju et al. (1993) believed the two genera Lopadostoma and globose chamber at each end. Asexual morph: hyphomy-
Creosphaeria should be represented separately as they are cetous. Hyphae hyaline, septate, branched. Conidiophores
not similar in morphology with other xylariaceous members. micronematous or semi-micronematous. Conidia ellipsoidal
In their phylogenetic analyses, Maharachchikumbura et al. or helicoid, septate or aseptate, spermatia, enteroblastic,
(2015) and Senanayake et al. (2015) placed Creosphaeria subglobose.
sassafras (Schwein.) and Lopadostoma species as a distinct Type: Lulworthia G.K. Sutherl.
monophyletic group outside Xylariaceae. Although they form Notes: Spatafora et al. (1998) demonstrated that the order
separate monophyletic groups, the close relationship between Halosphaeriales was polyphyletic and comprised two distinct
Lopadostoma and Creosphaeria noted in Senanayake et al. lineages. The first clade, Halosphaeriales, included eleven
(2015), is in agreement with Jaklitsch et al. (2014). representative genera, that were closely related to, and are
Lopadostoma turgidum was lectotypified by Lu and Hyde now included in the order Microascales. The second clade,
(2000), which turned out to be invalid because the material comprised the genera Lulworthia and Lindra, and was
selected was not from the original collection by Persoon (it assigned to Lulworthiaceae, Lulworthiales (Kohlmeyer et al.
was not possible to loan this material). Jaklitsch et al. (2014) 2000). Jones et al. (2008) showed that some asexual morph
was able to observe Persoon’s original material by going to species of Cirrenalia, Cumulospora, Zalerion and Orbimyces
herbarium (L) and epitypified L. turgidum with a fresh collec- grouped in Lulworthiaceae. Subsequently, Abdel-Wahab et al.
tion. We do not described the genus here as it was detailed in (2010) in an analysis of a larger data set of SSU and LSU
Senanayake et al. (2015). rDNA sequence data, showed that the genera Cirrenalia and
Cumulospora were polyphyletic and introduced a number of
new genera to accommodate taxa grouping in
Lulworthiaceae: Halazoon (= Cirrenalia fusca I. Schmidt),
Other genus included Hydea (= Cirrenalia pygmea Kohlm.), Matsusporium (=
Cirrenalia tropicalis Kohlm.), and Moromyces (=
Creosphaeria Theiss., Beih. bot. Zbl., Abt. 2 27: 396 (1910); Cumulospora varia Chatmala & Somrithipol). A further new
Fig. 83 genus Moleospora was introduced for the new species
Type species: Creosphaeria riograndensis Theiss., Beih. M. maritima Abdel-Wahab et al. (Abdel-Wahab et al. 2010).
bot. Zbl., Abt. 2 27: 396 (1910) Studies by Inderbitzin et al. (2004), Campbell et al.
 Fungal Diversity

Fig. 82 Lopadostoma turgidum (Material examined: AUSTRIA, Peridium h Paraphyses i Mature and immature asci j Apical ring bluing at
Niederösterreich, Gaaden, 3 December 2011, H. Voglmayr, WU 32085, Melzer’s reagent k–m Ascospores n-r Asci. Scale bars: b = 500 μm, c–
epitype). a Herbarium specimen b Stromata in face view c Ectostromatic d = 200 μm, e–f = 100 μm, g = 20 μm, j, h = 10 μm, I = 20 μm, i, n–
disc d Valsoid configuration of ascomata e-f Cross sections of stroma g r = 50 μm, k–m = 5 μm

(2009) showed that the sexual morph Haloguignardia in Lulworthiaceae with high statistical support.
with five species parasitic on brown seaweeds, grouped Currently, Lulworthiaceae comprises seven asexual
Fungal Diversity

Fig. 83 Creosphaeria sassafras

(redrawn from Ju et al. 1993) a-c
Creosphaeria sassafras a, c
Conidiophores and
conidiogenesis cells b Conidia d-
e Lopadostoma turgidum d
Conidia e Conidiophores and
conidiogenesis cells. Scale bar: a–
e = 10 μm

morphs genera. Maharachchikumbura et al. (2015) Type species: Lulworthia fucicola G.K. Sutherl., Trans. Br.
showed that the orders Lulworthiales and mycol. Soc.5(2): 259 (1916) [1915]
Koralionastetales clustered together in a well-supported Notes: The type genus Lulworthia was described by
clade, based on analysis of combined sequence data and Sutherland (1915) to accommodate the species L. fucicola
a new subclass was introduced as Lulworthiomycetidae. G.K. Sutherl., a taxon found on the brown seaweed, common-
ly known as the bladder wrack, at Lulworth Cove on the
Lulworthia G.K. Sutherl., Trans. Br. mycol. Soc. 5(2): 259 Dorset coast, UK. Later, the genera Lulworthia and Lindra
(1916) [1915] were included in the family Halosphaeriaceae, order
Facesoffungi number: FoF 02117; Fig. 84 Halosphaeriales by Müller and von Arx (1962). The type
Saprobic on wood or parasitic on algae. Sexual morph: species of Lulworthia (L. fucicola) has never been isolated
Ascomata sphaerical to ellipsoidal, often swan-shaped, im- from the marine alga Fucus vesiculosus, so a collection from
mersed to superficial, carbonaceous to coriaceous, ostiolate, Chile was designated as neotype by Campbell et al. (2005).
dark brown with or without a long neck up to 154 mm. Subsequently, based on molecular analysis, a new family
Peridium 1–2 layered, composed of an outer layer of Lulworthiaceae in Lulworthiales was designated to include
cells of textura angularis and an inner layer of elongate halosphaerialean species with filamentous ascospores
cells. Asci 8-spored, clavate to fusiform, short pedicel- (Kohlmeyer et al. 2000). Several new Lulworthia species were
late, unitunicate, deliquescing early. Ascospores filiform, described from marine habitats and the genus became one of
mostly hyaline, septate, with apical chambers. Asexual the largest in marine habitats (Kohlmeyer et al. 2000).
morph: Undetermined. Lulworthia has been subjected to revisions over many years
 Fungal Diversity

Fig. 84 Lulworthia sp. (Material

examined: TAIWAN, New Taipei
City, Tamsui, mangrove, on dead
mangrove wood, 20 September
2009, K.L. Pang) a Neck
protruding from the wood surface
b Immersed ascoma on bark c
Peridium composed of thick-
walled angular cells d
Filamentous ascospore e Polar
end chamber of ascospore. Scale
bars: a = 100 μm, b, d = 50 μm, c,
e = 10 μm

(Kohlmeyer 1972; Kohlmeyer and Kohlmeyer 1979; Koch Type species: Haloguignardia decidua Cribb & J.W.
and Jones 1984;). Johnson and Sparrow (1961) introduced Cribb, Pap. Dept. Bot. Univ. Qd.3: 97 (1956)
12 new Lulworthia species, but Cavaliere and Johnson Hydea K.L. Pang & E.B.G. Jones, in Abdel-Wahab et al.,
(1965) synonymized all species with L. medusa (Ellis & Mycol. Progr. 9(4): 549 (2010)
Everh.) Cribb & J.W. Cribb. Kohlmeyer (1972) recognized Type species: Hydea pygmea (Kohlm.) K.L. Pang &
at least three species; and Koch and Jones (1984) rec- E.B.G. Jones, in Abdel-Wahab et al., Mycol. Progr. 9(4):
ognized six species. Currently eleven species are includ- 549 (2010)
ed with a number of other taxa (Kohlmeyer et al. 2000). Kohlmeyeriella Jones et al., Soc., Bot. 87: 208 (1983)
As a result of sequence analysis of LSU, a number of Type species: Kohlmeyeriella tubulata (Kohlm.) E.B.G.
transfers have been proposed, such as Lulworthia crassa Jones et al., J. Linn. Soc., Bot. 87(2): 210 (1983)
Nakagiri to Kohlmeyeriella, Lulworthia lignoarenaria Lindra I.M. Wilson, Trans. Br. mycol. Soc. 39(4): 411
Jørg. Koch & E.B.G. Jones to Lulwoidea and (1956)
Lulworthia uniseptata Nakagiri to Lulwoana (Campbell Type species: Lindra inflata I.M. Wilson, Trans. Br. mycol.
et al. 2005). Currently only one species (the type spe- Soc. 39(4): 411 (1956)
cies L. fucicola) is accepted in Lulworthia sensu stricto, Lulwoana Kohlm. et al., Mycol. Res. 109(5): 562 (2005)
while eleven species are referred to Lulworthia sensu Type species: Lulwoana uniseptata (Nakagiri) Kohlm.
lato until they are recollected and sequenced (Jones et al., Mycol. Res.109(5): 562 (2005)
et al. 2015). Lulwoidea Kohlm. et al., Mycol. Res. 109(5): 564 (2005)
Type species: Lulwoidea lignoarenaria (Jørg. Koch &
E.B.G. Jones) Kohlm. et al., Mycol. Res.109(5): 564
(2005)
Other genera included Matsusporium E.B.G. Jones & K.L. Pang, in Abdel-
Wahab et al., Mycol. Progr. 9(4): 550 (2010)
Cumulospora I. Schmidt, Mycotaxon 24: 420 (1985) Type species: Matsusporium tropicale (Kohlm.) E.B.G.
Type species: Cumulospora marina I. Schmidt, Jones & K.L. Pang, in Abdel-Wahab et al., Mycol. Progr.
Mycotaxon 24: 421 (1985) 9(4): 550 (2010)
Halazoon Abdel-Aziz et al., Mycol. Progr. 9(4): 545 Moleospora Abdel-Wahab et al., Mycol. Progr. 9(4): 547
(2010) (2010)
Type species: Halazoon melhae Abdel-Aziz et al. in Type species: Moleospora maritima Abdel-Wahab et al., in
Abdel-Wahab et al., Mycol. Progr. 9(4): 546 (2010) Abdel-Wahab et al., Mycol. Progr. 9(4): 548 (2010)
Haloguignardia Cribb & J.W. Cribb, Pap. Dept. Bot.) Moromyces Abdel-Wahab et al., in Abdel-Wahab et al.,
Univ. Qd.3: 97 (1956) Mycol. Progr. 9(4): 555 (2010)
Fungal Diversity

Type species: Moromyces varius (Chatmala & Somrith.) 13. Ascomata superficial, with sterile hairs, parasitic on red
Abdel-Wahab et al., in Abdel-Wahab et al., Mycol. Progr. algae . . . . . . . . . . . . . . . . . . . . . . . . . Spathulospora
9(4): 555 (2010) 13. Ascomata immersed in a gall, lacking sterile hairs, on
Orbimyces Linder, Farlowia 1: 404 (1944) brown algae . . . . . . . . . . . . . . . . . . . Haloguignardia
Type species: Orbimyces spectabilis Linder, Farlowia1: 14. Ascomata with bell-like structure within the
404 (1944) centrum . . . . . . . . . . . . . . . . . . . . . . . . Rostrupiella
Rostrupiella Jørg. Koch et al., Bot. Mar. 50(5–6): 295 14. Ascomata lacking a bell-like structure . . . Lulworthia
(2007) 15. Ascospores 1-septate . . . . . . . . . . . . . . . . . Lulwoana
Type species: Rostrupiella danica Jørg. Koch et al., Bot. 15. Ascospores multi-septate . . . . . . . . . . . . . .Lulwoidea
Mar. 50(5/6): 295 (2007)
Spathulospora A.R. Caval. & T.W. Johnson, Mycologia Macrohilaceae Crous, in Crous et al., IMA Fungus 6(1): 180
57(6): 927 (1965) (2015)
Type species: Spathulospora phycophila A.R. Caval. & Facesoffungi number: FoF 01394
T.W. Johnson, Mycologia 57(6): 927 (1965) Pathogenic forming leaf spots. Sexual morph:
Undetermined. Asexual morph: Conidiomata pycnidial, im-
mersed, becoming erumpent, medium brown, globose.
C o n i d i o p h o re r e d u c e d t o c o n i d i o g e n o u s c e l l s .
Key to genera of Lulworthiaceae Conidiogenous cells lining the inner cavity, pale brown, cy-
lindrical, proliferating percurrently near the apex. Conidia sol-
1. Ascomycete with filamentous ascospores . . . . . . . . . . . . 9 itary, medium to dark brown, ovoid, smooth, guttulate, medi-
1. Hyphomycete. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 ally septate, apex obtuse, base truncate with a visible scar.
2. Conidia tetraradiate . . . . . . . . . . . . . . . . . . Orbimyces Type: Macrohilum H.J. Swart
2. Conidia helicoid. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 Notes: The family Macrohilaceae was introduced by Crous
3. Conidia initially spiral, becoming a tangled knot of cells et al. (2015a) based on a phylogenetic analysis of LSU se-
after cell division in several planes . . . . . . . . . . . . . . . 4 quence data and it was accommodated in Diaporthales. This
3. Conidia initially spiral, not forming a tangled knot of family is typified by Macrohilum. Macrohilaceae differs from
cells after cell division . . . . . . . . . . . . . . . . . . . . . . . . 5 other families of Diaporthales in having single, dark brown,
4. Conidia irregularly helicoid, muriform . . . . Moromyces guttulate, thick-walled, medially septate, oval conidia with
4. Conidia rosette-like, globose, forming pseudo- obtuse apex and truncate base. Species in this family are com-
chains . . . . . . . . . . . . . . . . . . . . . . . . . . Cumulospora monly associated with leaf spots of Eucalyptus species.
5. Cells of the conidia are more or less similar in size and
colour . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 Macrohilum H.J. Swart, Trans. Br. mycol. Soc. 90(2): 288
5. Cells increasing in diam. and pigmentation from the base (1988)
to the apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 Facesoffungi number: FoF 02118; Fig. 85
6. Conidia irregularly helicoid, cells of the conidia tightly Pathogenic forming leaf spots. Sexual morph:
joined and appearing muriform . . . . . . . Moheitospora Undetermined. Asexual morph: Coelomycetous. Conidiomata
6. Conidia helicoid when young, soon become a mass of immersed, becoming erumpent, medium brown, globose, coria-
cells . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Moleospora ceous. Conidiophore reduced to conidiogenous cells.
7. Conidia terminal or lateral . . . . . . . . . . . . . . Halazoon Conidiogenous cells lining the inner cavity, pale brown, cylindri-
7. Conidia acrogenous . . . . . . . . . . . . . . . . . . . . . . . . . . 8 cal, proliferating percurrently near the apex. Conidia solitary,
8. Conidia 3–4-septate, black or fuscous . . . . . . . . Hydea medium to dark brown, ovoid, smooth, guttulate, developing a
8. Conidia up to 12 septate, immature conidia surrounded single supramedian septum, thick-walled, frequently constricted
by a sheath. . . . . . . . . . . . . . . . . . . . . . Matsusporium at the septum, apex obtuse, base truncate with a visible scar.
9. Ascospores with apical chambers or appendages . . . 10 Type species: Macrohilum eucalypti H.J. Swart, Trans. Br.
9. Ascospores lacking appendages . . . . . . . . . . . . Lindra mycol. Soc. 90(2): 288 (1988)
10. Ascospores aseptate . . . . . . . . . . . . . . . . . . . . . . . . 11 Notes: The coelomycetous genus Macrohilum is typified
10. Ascospores septate . . . . . . . . . . . . . . . . . . . . . . . . . 15 by Macrohilum eucalypti. Swart (1988) provided a detailed de-
11. Ascospores filiform . . . . . . . . . . . . . . . . . . . . . . . . 14 scription for this genus. The New Zealand isolate (CPC 10945)
11. Ascospores fusiform, ellipsoidal . . . . . . . . . . . . . . . 12 of Macrohilum eucalypti differs from the Australian epitype
12. Ascospore appendages tubular, longer than isolate (CPC 19421) by four base pairs in the ITS region.
35 μm . . . . . . . . . . . . . . . . . . . . . . . Kohlmeyeriella
12. Ascospore appendages apiculate or conical, less than Magnaporthaceae P.F. Cannon, Syst. Ascom. 13(1): 26
35 μm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13 (1994)
 Fungal Diversity

Fig. 85 Macrohilum eucalypti (CPC 19421, epitype) (illustration based on Crous et al. 2015a). a, b Conidiogenous cells c Conidia. Scale bars: a–
c = 10 μm

Facesoffungi number: FoF 01101 Kirk et al. (2001) accepted nine genera, while Kirk et al.
Pathogenic on a wide range of monocotyledonous plants or (2008) accepted 13 genera. Luo et al. (2014) recently
saprobic on plant material, terrestrial some aquatic. Sexual established the genus Pseudophialophoda in the family.
morph: Ascomata perithecial, solitary or scattered, black, su- Maharachchikumbura et al. (2015) provided an updated out-
perficial or immersed in plant tissue, globose to subglobose, line of the family Magnaporthaceae with 20 genera, which
with long, unilateral, cylindrical, black, periphysate neck. included both sexual and asexual morphs. In the same report,
Peridium comprising a few to several layers of cells of textura they also confirmed that Magnaporthaceae belongs to the
epidermoidea. Hamathecium comprising hyaline, septate, pa- order Magnaporthales, which was introduced based on
raphyses, intermingled among asci. Asci 8-spored, molecular data and morphology (Maharachchikumbura
unitunicate, subcylindrical, short-pedicellate, with apical ring. et al. 2015).
Ascospores biseriate, hyaline to olivaceous, filiform or fusoid, Nakataea Hara, Diseases Rice Plant, Edn 2: 185 (1939)
curved to sigmoid, with or without transverse septate ends, Facesoffungi number: FoF 01102; Figs 86, 87
bluntly rounded, lacking sheaths. Asexual morph: Pathogenic or saprobic on plants. Sexual morph:
Hyphomycetous, at times formed from sclerotia. Ascomata perithecial, globose, dark brown, immersed, with
Conidiophores unbranched or branched. Conidiogenous cells ostiolar neck frequently protruding from the leaf tissue.
integrated, pigmented, phialidic with collarettes, or denticu- Peridium thick-walled, comprising 5–12 layers, with dark
late. Conidia variable in shape, hyaline to pale brown, straight cells. Asci 8-spored, unitunicate, thin-walled, subcylindrical,
or curved, with or without septa. short pedicellate, deliquescent at maturity. Ascospores parallel
Type: Nakataea Hara to spirally twisted, hyaline at median cells, turning yellowish
Notes: The family Magnaporthaceae was introduced by brown, fusiform, curved, 3-septate, slightly constricted at sep-
Cannon (1994). The placement of taxa of Magnaporthaceae ta, granular. Asexual morph: Conidiophores brown, solitary,
has long been problematic due to the lack of convincing mor- erect, smooth, septate, branched, with integrated terminal.
phological characters and inconclusive molecular data Conidiogenous cells forming a rachis with several denticles.
(Thongkantha et al. 2009). Magnaporthaceae was placed as Conidia solitary, 3-septate, falcate to sigmoid, smooth-walled,
a family within the class Sordariomycetes (Kirk et al. 2001, wider in the middle, end cells hyaline, median cells medium
Lumbsch and Huhndorf 2007a). Thongkantha et al. (2009) brown.
established a new order, Magnaporthales to accommodate Type species: Nakataea oryzae (Catt.) J. Luo & N. Zhang,
Magnaporthaceae based on morphological characters of both Mycologia 105: 1025 (2013)
sexual and asexual morphs together with phylogenetic Notes: The genus Nakataea was established by Hara
analyses of combined SSU and LSU sequence data. (1939) based on Nakataea oryzae (Catt.) Luo & Zhang
According to the phylogenetic analysis by Klaubauf et al. (2013) (synonym Nakataea sigmoidea (Cavara) Hara) isolat-
(2014) of combined LSU and RPB1 sequence data, ed from Oryza sativa in Italy. Krause and Webster (1972)
Magnaporthaceae is sister to Pyriculariaceae and showed that Nakataea and Magnaporthe are congeneric and
Ophioceraceae. their type species, Nakataea oryzae (Catt.) J. Luo & N. Zhang
The exact number of genera in this family is still unclear and Magnaporthe salvinii (Catt.) Krause & Webster (1972)
(Thongkantha et al. 2009, Bussaban et al. 2005, Thongkantha are the same species. Therefore Luo and Zhang (2013), based
et al. 2009, Zhang et al. 2011, 2014). The family was origi- on Article 59.1 of the The International Code of Nomenclature
nally described with six genera and 20 species (Cannon 1994). for algae, fungi, and plants (Melbourne Code; ICN; McNeill
Fungal Diversity

Fig. 86 Nakataea oryzae (Material examined: INDIA, New Orleans, on Sclerotium g, h Cross section of sclerotium. Scale Bars: d = 2 mm,
Oryzae sativa L. (Poaceae) (Rice Straw), 4 August 1967, B.E. Devis e = 1 mm, f = 300 μm, g and h = 100 μm
(621662)) a, b, c Herbarium material d, e Sclerotium on substrate f

et al. 2012), synonymized Magnaporthe oryzae under Type species: Bussabanomyces longisporus (Bussaban)
N. oryzae because the name Nakataea (1939) is older than Klaubauf et al., Stud. Mycol. 79: 99 (2014)
Magnaporthe (1972). This resulted in the introduction of Ceratosphaerella Huhndorf et al., Mycologia 100(6): 941
Nakataea as the correct name for the type of the family (2008)
(Murata et al. 2014, Klaubauf et al. 2014, Luo et al. 2015, Type species: Ceratosphaerella castillensis (C.L. Sm.)
Maharachchikumbura et al. 2015). Huhndorf et al., Mycologia 100(6): 944 (2008)
Ceratosphaeria Niessl, Verh. nat. Ver. Brünn 14: 203
(1876)
Type species: Ceratosphaeria lampadophora (Berk. &
Other genera included Broome) Niessl, Verh. nat. Ver. Brünn 14: 203 (1876)
Clasterosphaeria Sivan., Trans. Br. mycol. Soc. 83(4): 710
Budhanggurabania P. Wong et al., Persoonia, Mol. Phyl. (1984)
Evol. Fungi 34: 241 (2015) Type species: Clasterosphaeria cyperi Sivan., Trans. Br.
Type species: Budhanggurabania cynodonticola P. Wong mycol. Soc. 83(4): 710 (1984)
et al., Persoonia, Mol. Phyl. Evol. Fungi 34: 241 (2015) Clasterosporium Schwein., Trans. Am. phil. Soc., New
Buergenerula Syd., Annls mycol. 34(4/5): 392 (1936) Series 4(2): 300 (1832) [1834]
Type species: Buergenerula biseptata (Rostr.) Syd., Annls Type species: Clasterosporium caricinum Schwein. [as
mycol. 34(4/5): 392 (1936) ‘Clasterisporium’], Trans. Am. phil. Soc., New Series 4(2):
Bussabanomyces Klaubauf et al., Stud. Mycol. 79: 99 300 (1832) [1834]
(2014) Clavatisporella K.D. Hyde, Mycotaxon 55: 276 (1995)
 Fungal Diversity

Fig. 87 Nakataea oryzae a Ascomata b Cross section of ascomata c Peridium d Ascus e Apex f Conidiophore with conidia of asexual morph (redrawn
from Krause and Webster 1972)
Fungal Diversity

Type species: Clavatisporella musicola K.D. Hyde, Key to the sexual genera of Magnaporthaceae
Mycotaxon 55: 276 (1995)
Gaeumannomyces Arx & D.L. Olivier, Trans. Br. mycol. 1. Ascomata superficial . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
Soc. 35(1): 32 (1952) 1. Ascomata immersed or semi-immersed . . . . . . . . . . . 3
Type species: Gaeumannomyces graminis (Sacc.) Arx & 2. Peridium with several layers. . . . . . . . . . . . . . . . . . . . 4
D.L. Olivier, Trans. Br. mycol. Soc. 35(1): 32 (1952) 2. Peridium with two layers . . . . . . . . . Clasterosphaeria
Harpophora W. Gams, Stud. Mycol. 45: 192 (2000) 3. Peridium more than 4 layers. . . . . . . . . . . . . . . . . . . . 5
Type species: Harpophora radicicola (Cain) W. Gams, 3. Peridium 2–4 layers . . . . . . . . . . . . . . . . . . . . . . . . . . 6
Stud. Mycol. 45: 192 (2000) 4. Asci with an apical ring . . . . . . . . . . . . . . . . . . . . . . . 7
Herbampulla Scheuer & Nograsek, Mycotaxon 47: 419 4. Asci without an apical ring. . . . . . . . . . . Slopeiomyces
(1993) 5. Asci clavate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
Type species: Herbampulla crassirostris Scheuer & 5. Asci cylindrical . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
Nograsek, Mycotaxon 47: 419 (1993) 6. Asci clavate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
Kohlmeyeriopsis Klaubauf et al. Stud. Mycol. 79: 101 6. Asci cylindrical . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
(2014) 7. Ascospores 1–3-septate . . . . . . . . . . . . . . . . . . . . . . 12
Type species: Kohlmeyeriopsis medullaris (Kohlm. et al.) 7. Ascospores more than 3-septate . . . . Ceratosphaerella
Klaubauf et al., Stud. Mycol. 79: 101 (2014) 8. Ascospores oblong-fusoid, with 2-
Magnaporthiopsis J. Luo & N. Zhang, Mycologia 105(4): septa. . . . . . . . . . . . . . . . . . .Buergenerula
1021 (2013) 8. Ascospores narrowly fusiform, with 3-
Type species: Magnaporthiopsis poae (Landsch. & N. septa . . . . . . . . . . . . . . . . . . . . Muraeriata
Jacks.) J. Luo & N. Zhang, Mycologia 105(4): 1024 (2013) 9. Ascospores without septa . . . . . . . . . . . Phomatospora
Muraeriata Huhndorf et al., Mycologia 100(6): 948 9. Ascospores with septa . . . . . . . . . . . . . . . . . . . . . . . 13
(2008) 10. Ascospores with mucilaginous pad-like appendage at
Type species: Muraeriata collapsa Huhndorf et al., the end. . . . . . . . . . . . . . . . . . . . . . . . Clavatisporella
Mycologia 100(6): 949 (2008) 10. Ascospores without appendages . . Magnaporthiopsis
Mycoleptodiscus Ostaz., Mycologia 59(6): 970 (1968) 11. Ascospores filamentous, tapering towards the
[1967] base . . . . . . . . . . . . . . . . . . . . . . . . Kohlmeyeriopsis
Type species: Mycoleptodiscus terrestris (Gerd.) Ostaz., 11. Ascospores fusiform, curved, partially gelatinized at
Mycologia 59(6): 970 (1968) [1967] maturity . . . . . . . . . . . . . . . . . . . . . . . . Magnaporthe
Neogaeumannomyces D.Q. Dai & K.D. Hyde, in Liu 12. Asci obovoid to saccate; ascospores multi-
et al., Fungal Diversity: 10.1007/s13225-015-0324-y, [34] seriate. . . . . . . . . . . . . . . . . . . . Budhanggurabania
(2015) 12. Asci cylindric-clavate; ascospores bi-
Type species: Neogaeumannomyces bambusicola D.Q. seriate . . . . . . . . . . . . . . . . . . . . . . . . Omnidemptus
Dai & K.D. Hyde, in Liu et al., Fungal Diversity: 10.1007/ 13. Ascospores hyaline . . . . . . . . . . . . . . . . . . . . . . . . 14
s13225-015-0324-y, [35] (2015) 13. Ascospores median cells turning yellowish
Omnidemptus P.F. Cannon & Alcorn, Mycotaxon 51: 483 brown . . . . . . . . . . . . . . . . . . . . . . . . 15
(1994) 14. Ascus apical ring tall, with narrow or short furcate
Type species: Omnidemptus affinis P.F. Cannon & Alcorn, pedicel. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
Mycotaxon 51: 483 (1994) 14. Ascus apical ring small distinct (2 small refringent
Phomatospora Sacc., Nuovo G. bot. ital. 7: 306 (1875) dots) . . . . . . . . . . . . . . . . . . . . . . Gaeumannomyces
Type species: Phomatospora berkeleyi Sacc., Nuovo G. 15. Ascospores 3-septate, median cells turning yellowish
bot. ital. 7: 306 (1875) brow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nakataea
Pseudophialophora J. Luo & N. Zhang, Mycologia 15. Ascospores 5-septate with pallid at end
106(3): 581 (2014) cells. . . . . . . . . . . . . . . . . . . . . . . . . . Herbampulla
Type species: Pseudophialophora eragrostis J. Luo & N. 16. Ascospores 5–7-septate . . . . . . . . . . . Ceratosphaeria
Zhang, Mycologia 106(3): 581 (2014) 16. Ascospores 2–3-septate . . . . . Neogaeumannomyces
Pyriculariopsis M.B. Ellis, Dematiaceous Hyphomycetes
(Kew): 206 (1971)
Type species: Pyriculariopsis parasitica (Sacc. & Berl.) Key to the asexual genera of Magnaporthaceae
M.B. Ellis, Dematiaceous Hyphomycetes (Kew): 207 (1971)
Slopeiomyces Klaubauf et al., Stud. Mycol. 79: 102 (2014) 1. Conidiogenous cell forming phialides. . . . . . . . . . . . . . . 2
Type species: Slopeiomyces cylindrosporus (D. Hornby 1. Conidiogenous cell cylindrical, forming denticles. . . . 3
et al.) Klaubauf et al., Stud. Mycol. 79: 103 (2014) 2. Conidia aseptate. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
 Fungal Diversity

2. Conidia septate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 comprises a single genus Melanconis and its asexual morph
3. Conidia with 1-septum . . . . . . . . . . . . Kohlmeyeriopsis Melanconium based on their morphology and phylogeny.
3. Conidia multi-septate . . . . . . . . . . . . . . . . . . . . . . . . . 6 Voglmayr and Jaklitsch (2014) opined that a clear delineation
4. Conidiogenous cell straight . . . . . . . . . . . . . . . . . . . . 7 of Melanconium was not possible as similar asexual morphs
4. Conidiogenous cell curved . . . . . . . . . . . . . . . . . . . . . 8 occur in both Melanconis and Melanconiella.
5. Conidia solitary brown to dark brown, without Maharachchikumbura et al. (2015) listed 26 genera under this
setulae. . . . . . . . . . . . . . . . . . . . . . . . Clasterosporium family.
5. Conidia hyaline, with a simple setula at the apex and
often at the base. . . . . . . . . . . . . . . . . Mycoleptodiscus Melanconis Tul. & C. Tul., Selecta fungcarpol (Paris) 2:115
6. Conidia lacking mucoid cap . . . . . . . . . . . . . . . . . . . . 9 (1863).
6. Conidia with persistent mucoid cap . . . Pyriculariopsis Facesoffungi number: FoF 02119; Figs 88, 89
7. Conidia cylindrical, curved . . . . . . . . . . . . . . . . . . . 10 Saprobic or pathogenic on plants. Sexual morph:
7. Conidia subglobose to ovoid . . . . . Magnaporthiopsis Pseudostromata well-developed, obvious, erumpent.
8. Conidiogenous cells without Ectostromatic disc surrounded by bark or not, yellowish-
pigments . . . . . . . . . . . . . . . . . Budhanggurabania white, causing a coarse bark surface, inverted conical, ostioles
8. Conidiogenous cells pigmented . . Pseudophialophora open margin and middle of the disc. Ascomata perithecial,
9. Conidia obclavate, 4(−5)-septate . . . Bussabanomyces oblique or horizontal, globose to subglobose, coriaceous,
9. Conidia falcate to sigmoid, 3-septate . . . . . . Nakataea black, with long periphysate, lateral ostiolar canals.
10. Conidia phialidic, apex rounded, pointed towards Hamathecium comprising wide, hyphae-like, hyaline, septate
base. . . . . . . . . . . . . . . . . . . . . . . . . . . Slopeiomyces paraphyses, deliquescent at maturity. Peridium comprising
10. Conidia borne on slimy heads, outer, thick-walled, brown cells of textura globosa to textura
cylindrical . . . . . . . . . . . . . . Harpophora angularis and inner, thick-walled, flat, hyaline cells of textura
angularis. Asci 8-spored, unitunicate, oblong to fusiform,
Melanconidaceae G. Winter [as ‘Melanconideae’], Rabenh. short pedicellate, with distinct, J- apical ring. Ascospores over-
Krypt.-Fl., Edn 2 (Leipzig) 1.2: 764 (1886) lapping uniseriate to biseriate, hyaline, ellipsoid, 1-septate, not
Facesoffungi number: FoF 01395 or slightly constricted at the septum, smooth-walled, with or
Saprobic or pathogenic on plants and wood, in terrestrial without blunt or pointed appendages. Asexual morph:
aquatic habitats. Sexual morph: Pseudostromata well-devel- Coelomycetous. Melanconium-like. Conidiomata acervular,
oped, obvious, erumpent. Ectostromatic disc surrounded by scattered, solitary, superficial, black, coriaceous.
bark or not, yellowish-white, ostiolar canal opening around Conidiophores hyaline, branched at the base, septate, few-
the disc. Ascomata perithecial, arranged as circles around the celled, smooth walled. Conidiogenous cells hyaline, annellidic,
ectostromatic disc, oblique or horizontal, globose to cylindrical. Conidia hyaline to brown, ellipsoid or subglobose
subglobose, coriaceous, black, with long, periphysate, lateral smooth-walled, thick-walled, outer layer produced continuous-
ostiolar canals. Peridium comprising outer, thick-walled, ly, with the inner layer of the conidiogenous cell.
brown cells of textura globosa to textura angularis and inner, Type species: Melanconis stilbostoma (Fr.) Tul. & C. Tul.,
thick-walled, flat, hyaline cells of textura angularis. Select. fung. carpol. (Paris) 2: 115 (1863)
Hamathecium comprising wide, hyphae-like paraphyses, del- Notes: The genus Melanconis is typified by M. stilbostoma
iquescent at maturity. Asci 8-spored, unitunicate, oblong to and currently it comprises three species, including M. alni Tul.
fusiform, short pedicellate, with distinct, J- apical ring. & C. Tul. and M. marginalis (Peck) Wehm. (Castlebury et al.
Ascospores overlapping uniseriate to biseriate, hyaline, ellip- 2002). The asexual morph of Melanconis is placed in the
soid, 1-septate. Asexual morph: Coelomycetous. genus Melanconium and both of these genera have been
Melanconium-like. Conidiomata acervular, scattered, solitary, linked. Species in both Melanconis and Melanconium form
superficial, black, coriaceous. Conidiophores branched at the diseases on plants such as cankers and shoot blight.
base, septate, Conidiogenous cells annellidic, cylindrical.
Conidia hyaline to brown, ellipsoid or subglobose, smooth-
walled, thick-walled. Other genera included
Type: Melanconis Tul. & C. Tul.
Notes: The family Melanconidaceae was introduced by Botanamphora Nograsek & Scheuer, in Nograsek, Biblthca
Winter (1886) to accommodate species having yellowish- Mycol. 133: 50 (1990)
white ectostromatic discs surrounding ascomata in a circle. Type species: Botanamphora pachycarpa (Sacc. &
Members of this family are plant pathogens causing disease Marchal) Nograsek & Scheuer, in Nograsek, Biblthca
of economic plant species, as well as saprobes. Castlebury Mycol. 133: 51 (1990)
et al. (2002) and Rossman et al. (2007) showed this family Ceratoporthe Petr., Annls mycol. 23(1/2): 14 (1925)
Fungal Diversity

Fig. 88 Melanconis stilbostoma (Material examined: Stromata on substrate d, e Cross section of stroma f Peridium g-k Asci
CZECHOSLOVAKIA, Moravia, Ratschitz, on branches of Betula sp., l-q Ascospores. Scale bars: c, d = 200 μm, e = 100 μm, f–k = 20 μm, l–
Niessl, BPI 615061) a Herbarium packet b Herbarium specimen c q = 10 μm

Type species: Ceratoporthe didymospora Petr., Annls Cytomelanconis Naumov, Bot. Mater. Otd. Sporov. Rast.
mycol. 23(1/2): 14 (1925) Bot. Inst. Komarova Akad. Nauk S.S.S.R. 7: 108 (1951)
 Fungal Diversity

Fig. 89 Asexual morph of Melanconis stilbostoma (Material examined: RUSSIA, Sakhalin Island, Valley of the river Evay, on stems of Betula sp., 27
August 2000, A. Bogachova, BPI 748234, living cultures AR 3548 = CBS 109493) a-d Conidia and to the conidiogenous cells. Scale bars: a–d = 10 μm

Type species: Cytomelanconis systema-solare Naumov, Melanamphora Lafl., Sydowia 28(1–6): 243 (1976)
Bot. Mater. Otd. Sporov. Rast. Bot. Inst. Komarova Akad. [1975–1976]
Nauk S.S.S.R. 7: 109 (1951) Type species: Melanamphora spinifera (Wallr.) Lafl.,
Dicarpella Syd. & P. Syd., Annls mycol. 18(4/6): 181 Sydowia 28(1–6): 245 (1976) [1975–1976]
(1921) [1920] Melanconiella Sacc., Syll. fung. (Abellini) 1: 740 (1882)
Type species: Dicarpella bina (Harkn.) Syd. & P. Syd., Type species: Melanconiella spodiaea (Tul. & C. Tul.)
Annls mycol. 18(4/6): 181 (1921) [1920] Sacc., Syll. fung. (Abellini) 1: 740 (1882)
Dictyoporthe Petr., Sydowia 9(1–6): 556 (1955) Melanconiopsis Ellis & Everh., Bull. Torrey bot. Club 27:
Type species: Dictyoporthe ahmadii Petr., Sydowia 9(1–6): 575 (1900)
556 (1955) Type species: Melanconiopsis inquinans Ellis & Everh.,
Fremineavia Nieuwl., Am. Midl. Nat. 4: 501 (1916) Bull. Torrey bot. Club 27: 575 (1900)
Type species: Fremineavia berkeleyi (Berl.) Nieuwl., Am. Phragmodiaporthe Wehm., Mycologia 33(1): 54 (1941)
Midl. Nat. 4: 501 (1916) Type species: Phragmodiaporthe caryae (Peck) Wehm.,
Gibellia Sacc., Atti Inst. Veneto Sci. lett., ed Arti, Sér. 6 3: Mycologia 33(1): 55 (1941)
714 (1885) Plagiophiale Petr., Sydowia 9(1–6): 585 (1955)
Type species: Gibellia dothideoides Sacc. & Berl., Revue Type species: Plagiophiale eucarpa (P. Karst.) Petr.,
mycol., Toulouse 7: 94 (1885) Sydowia 9(1–6): 586 (1955)
Hypophloeda K.D. Hyde & E.B.G. Jones, Trans. mycol. Plagiostigme Syd., Annls mycol. 23(3/6): 341 (1925)
Soc. Japan 30(1): 61 (1989) Type species: Plagiostigme couraliae Syd., Annls mycol.
Type species: Hypophloeda rhizospora K.D. Hyde & 23(3/6): 342 (1925)
E.B.G. Jones, Trans. mycol. Soc. Japan 30(1): 62 Prostratus Sivan.et al., Mycol. Res. 97(10): 1179 (1993)
(1989) Type species: Prostratus cyclobalanopsidis Sivan.et al.,
Kensinjia J. Reid & C. Booth, Can. J. Bot. 67(3): 896 Mycol. Res. 97(10): 1179 (1993)
(1989) Pseudovalsella Höhn., Annls mycol. 16(1/2): 123 (1918)
Type species: Kensinjia umbrina (Jenkins) J. Reid & C. Type species: Pseudovalsella thelebola (Fr.) Höhn., Annls
Booth, Can. J. Bot. 67(3): 896 (1989) mycol. 16(1/2): 123 (1918)
Macrodiaporthe Petr., Annls mycol. 17(2/6): 94 (1920) Wehmeyera J. Reid & C. Booth, Can. J. Bot. 67(3): 895
[1919] (1989)
Type species: Macrodiaporthe occulta (Fuckel) Petr., Type species: Wehmeyera acerina (Wehm.) J. Reid & C.
Annls mycol. 17(2/6): 94 (1920) [1919] Booth, Can. J. Bot. 67(3): 895 (1989)
Massariovalsa Sacc., Michelia 2(no. 8): 569 (1882) Wuestneia Auersw. Ex Fuckel, Hedwigia 3: 159 (1864)
Type species: Massariovalsa sudans (Berk. & M.A. Type species: Wuestneia aurea (Fuckel) Auersw., in
Curtis) Sacc., Syll. fung. (Abellini) 2: LV (1883) Fuckel, Hedwigia 3: 159 (1864)
Mebarria J. Reid & C. Booth, Can. J. Bot. 67(3): 898 Wuestneiopsis J. Reid & Dowsett, Can. J. Bot. 68(11):
(1989) 2406 (1990)
Type species: Mebarria thujina (Nag Raj & DiCosmo) J. Type species: Wuestneiopsis georgiana (J.H. Mill. & G.E.
Reid & C. Booth, Can. J. Bot. 67(3): 898 (1989) Thomps.) J. Reid & Dowsett, Can. J. Bot. 68(11): 2406 (1990)
Fungal Diversity

Key to genera of Melanconidaceae 20. Ascospores hyaline at maturity. . . . . . . . . .Wuestneia

20. Ascospores yellowish brown to dark brown at
1. Stromata well-developed . . . . . . . . . . . . . . . . . . . . . . . . 5 maturity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
1. Stromata absent or reduced . . . . . . . . . . . . . . . . . . . . 2 21. Ascospores with hyaline basal or apical end cell/cells22
2. Ascospores aseptate . . . . . . . . . . . . . . . . . . .Dicarpella 21. Ascospores without hyaline basal or apical end
2. Ascospores 3–5-septate . . . . . . . . . . . . . . . . . . . . . . . 3 cell/cells . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23
3. Ascospores fusiform; brown with pale 22. Beak lateral; ascospores oval to globose. . .Prostratus
ends…………………………….…. Botanamphora 22. Beak erect; ascospores allantoid to
3. Ascospores broadly oblong to clavate or cylindrical; lunate . . . . . . . . . . . . . . . . . . . . . . .Melanamphora
hyaline . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 23. Appendages present . . . . . . . . . . . . . . Pseudovalsella
4. Associated with submerged marine wood; perithecia 23. Appendages absent. . . . . . . . . . . . . . . . . . . . . . . . . 24
with long necks and cylindrical 24. Ascospores hyaline, yellowish or brown, fusoid or
ascospores . . . . . . . . . . . . . . . . . . . . . . . Hypophloeda ellipsoid. . . . . . . . . . . . . . . . . . . . . . . .Melanconiella
4. Associated with plant leaves, perithecia without necks 24. Ascospores dark brown, globose to broadly
and with broadly oblong to clavate ellipsoid . . . . . . . . . . . . . . . . . . . . . . . . . . Mebarria
ascospores . . . . . . . . . . . . . . . . . . . . . . . .Plagiostigme
5. Asexual morph common in nature . . . . . . . . . . . . . . . 6 Meliolaceae G.W. Martin ex Hansf., Mycol. Pap. 15: 23
5. Asexual morph not common in nature; sexual morph (1946)
common . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 Facesoffungi number: FoF 00741
6. Stromatic tissues prosenchymatous. . . . . . . . . . . . . . . 7 Epiphytes, or pathogens on leaves, occasionally on stems,
6. Stromatic tissues pseudoparenchymatous Melanconium branches, bark, or fruits. Superficial hyphae branched, septate,
7. Conidia hyaline subfusoid . . . . . . . . . . . . . . Kensinjia brown to dark brown, hyphal setae present or absent.
7. Conidia dark brown, obovoid to Hyphopodia capitate on hyphae, variously shaped, alternate
subglobose . . . . . . . . . . . . . . . . . . . Melanconiopsis or opposite, 2-celled, brown. Hyphal setae when present, de-
8. Stromatic tissues pseudoparenchymatous . . . . . . . . . . 9 veloping from hyphae, septate, branched or unbranched at
8. Stromatic tissues prosenchymatous. . . . . . . . . . . . . . 12 apex, or with bulbous apices or apical part, brown to dark
9. Appendages present; elongate strap- brown. Sexual morph: Ascomata superficial on surface of
like. . . . . . . . . . . . . . . . . . Macrodiaporthe web-like colonies on host, globose to subglobose, or flattened.
9. Appendages absent . . . . . . . . . . . . . . . . . . . . . . . . . 10 Peridium comprising brown cells of textura angularis when
10. Asci 1-celled; ascospores dark brown, cylindrical to viewed in squash mounts, with two strata, outer stratum of
fusoid, 5–11-septate . . . . . . . . . . . . . . . . Fremineavia brown to dark brown cells, raised conoid cells, appendages
10. Asci 8-spored; ascospores hyaline, spindle-shaped to or setae, inner stratum of hyaline to pale brown cells.
ellipsoid, 0–1-septate . . . . . . . . . . . . . . . . . . . . . . . 11 Hamathecium comprising evanescent paraphyses. Asci 2–4-
11. Ascospores spindle-shaped, 1-septate . . Ceratoporthe spored, unitunicate, subglobose to broadly clavate.
11. Ascospores broadly ellipsoid, aseptate . . . . . .Gibellia Ascospores 2–3-seriate, hyaline to brown, ellipsoid or cylin-
12. Asci 2-spored. . . . . . . . . . . . . . . . . . Cytomelanconis drical to ovoid, 3–4-septate. Asexual morph: Phialides
12. Asci 4–8-spored . . . . . . . . . . . . . . . . . . . . . . . . . . . 13 ampulliform or flask-shaped, alternate or opposite on hyphae.
13. Sheath present . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 Conidiogenous cells formed directly from vegetative hyphae.
13. Sheath absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15 Conidia unicellular, small and hyaline (Cannon and Kirk
14. Ascospores brown . . . . . . . . . . . . . . . Massariovalsa 2007; Hongsanan et al. 2015).
14. Ascospores hyaline to yellow . . . . . . . . . Plagiophiale Type: Meliola Fr., Syst. orb. veg. (Lundae) 1: 111 (1825);
15. Ectostromatic disc present . . . . . . . . . . . . . . . . . . . 16 Fig. 90
15. Ectostromatic disc absent . . . . . . . . . . . . . . . . . . . . 20 Type species: Meliola nidulans (Schwein.) Cooke,
16. Appendages present . . . . . . . . . . . . . . . . . . . . . . . . 17 Grevillea 11(no. 57): 37 (1882)
16. Appendages lacking . . . . . . . . . . . . . . . . . . . . . . . . 18 Notes: The family Meliolaceae was revised by Hongsanan
17. Ascospores multi-septate . . . . . . . . . . . .Dictyoporthe et al. (2015) and is not discussed further here. The genus
17. Ascospores 1-septate . . . . . . . . . . . . . . . .Melanconis Meliola was introduced by Fries (1825), and is the largest
18. Stromatic tissues prosenchymatous genus in the family Meliolaceae, which comprises over
hyphae . . . . . . . . . . . . . . . . . . . Phragmodiaporthe 1200 species (Kirk et al. 2008). Most species have been intro-
18. Stromatic tissues pseudoparenchymatous hyphae . . 19 duced primarily on the basis of host association, and followed
19. Ectostromatic disc yellowish orange to pale by morphology of branching or habit of hyphae, shape and
brown. . . . . . . . . . . . . . . . . . . . . . . . . Wuestneiopsis size of hyphopodia, size and shape of phialides as well as
19. Ectostromatic disc black. . . . . . . . . . . . . .Wehmeyera distribution, or size and shape of ascospores (Mibey and
 Fungal Diversity

Fig. 90 Meliola thailandicum (Material examined: Thailand, Chiang forked apices f Peridium g Hyphae with capitate hyphopodia h Terminal
Rai, Mueang, Rai Chun Tawan Meditation Centre, on living leaves of hyphae i Upper wall of ascoma j Phialides k Immature ascospore when in
Dimocarpus longan Lour. (Sapindaceae), 3 January 2015, S. Hongsanan, Melzer’s reagent l, m Mature ascospores n, o, q Immature asci p
MFLU 15-0044, holotype) a Leaf specimens b Ascomata on host Immature ascus in Melzer’s reagent. Scale bars: c, d = 100 μm, e,
substrate c Section through ascoma d Hyphal setae e Hyphal setae with f = 20 μm, g–j = 10 μm, k–q = 20 μm
Fungal Diversity

Hawksworth 1997; Justavino et al. 2015). Molecular data are Melogrammataceae G. Winter [as ‘Melogrameae’],
needed to determine if species really are host-specific Rabenh. Krypt.-Fl., Edn 2 (Leipzig) 1.2: 797 (1886)
(Hongsanan et al. 2015). Facesoffungi number: FoF 00840
Saprobic on bark of woody plants. Sexual morph:
Stromata superficial, erumpent, pulvinate or discoid, red-
dish-brown, subglobose, soft-textured, with pseudoparenchy-
Other genera included
matous cells, with black ostiole. Ascomata perithecial, dark
brown to black, immersed, globose, ostiole periphysate.
Amazonia Theiss., Annls mycol. 11(6): 499 (1913)
Peridium comprising rows of cells, externally brown, inter-
Type species: Amazonia psychotriae (Henn.) Theiss.,
nally hyaline. Hamathecium of septate, paraphyses. Asci 8-
Annls mycol. 11(6): 499 (1913)
spored, unitunicate, clavate, or fusoid, short pedicellate, with
Appendiculella Höhn., Sber. Akad. Wiss. Wien, Math.-
a shallow, J-, apical ring. Ascospores 0-3-seriate, hyaline or
naturw. Kl., Abt. 1 128: 556 (1919)
brown, filiform, aseptate or 1–3 septate, cylindrical or falcate.
Type species: Appendiculella calostroma (Desm.) Höhn.,
Asexual morph: Coelomycetous. Mycelium hyphae to yel-
Sber. Akad. Wiss. Wien, Math.-naturw. Kl., Abt. 1 128: 556
low-brown, septate, branched. Conidiophores long, cylindri-
(1919)
cal, stiffly upright, septate, verticillate and whorled or not.
Asteridiella McAlpine, Proc. Linn. Soc. N.S.W. 22(1): 38
Conidiogenous cells holoblastic, proliferating sympodially,
(1897)
hyaline, thin-walled. Conidia elongate falcate or filiform.
Type species: Asteridiella solani McAlpine, Proc. Linn.
Type: Melogramma Fr.
Soc. N.S.W. 22(1): 38 (1897)
Notes: The family Melogrammataceae is characterized by
Cryptomeliola S. Hughes & Piroz., Mycol. Pap. 174: 14
parasitic species possessing oblong to cylindrical asci, a re-
(1997)
duced apical annulus at the ascal apex and narrow paraphyses
Type species: Cryptomeliola orbicularis (Berk. & M.A.
(Schatz 1983). Melogramma was introduced by Fries (1849),
Curtis) S. Hughes & Piroz., in Mibey & Hawksworth,
and represented in Nitschke ex Fuckel (1870) as
Mycol. Pap. 174: 15 (1997)
“Melogrammaeae” (Fuckel 1870; Laflamme 1976). Winter
Endomeliola S. Hughes & Piroz., N.Z. Jl Bot. 32(1): 53
(1886) placed “Melogrammaeae” in Sphaeriales and included
(1994)
the genera Botryosphaeria, Endiothia, Melogramma and
Type species: Endomeliola dingleyae S. Hughes & Piroz.,
Valsaria. Melogramma was placed in Diaporthaceae, also in
N.Z. Jl Bot. 32(1): 54 (1994)
the order Sphaeriales from 1957 to 1976 (Laflamme 1976).
Irenopsis F. Stevens, Annls mycol. 25(5/6): 411 (1927)
Munk (1957), Laflamme (1976) revised the family
Type species: Irenopsis tortuosa (G. Winter) F. Stevens,
Melogrammataceae. Barr (1990) transferred
Annls mycol. 25(5/6): 439 (1927)
Melogrammataceae to Xylariales, and accepted only
Melogramma, with Melogramma campylosporum as its type
species. She redescribed the family and its asexual morph
Key to genera of Meliolaceae (Barr 1990). Kirk et al. (2008) listed Melogrammataceae in
Diaporthales. The family Melogrammataceae was not listed
1. Hyphae with intercellular hyphopodia. . . . . . Endomeliola by Lumbsch and Huhndorf (2010), and the type of
1. Hyphae with superficial hyphopodia . . . . . . . . . . . . . 2 Melogramma was classified in the family Melanconidaceae
2. Ascomata flattened. . . . . . . . . . . . . . . . . . . . Amazonia under Diaporthales. Jaklitsch and Voglmayr (2012) used se-
2. Ascomata globose to subglobose . . . . . . . . . . . . . . . . 3 quence data to show that Melogrammataceae nested in
3. Hyphae with setae . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 Xylariales in a distinct genus and was used to accommodate
3. Hyphae without setae . . . . . . . . . . . . . . . . . . . . . . . . . 5 Melogramma. Maharachchikumbura et al. (2015) confirmed
4. Hyphal setae with bulbous tips, apical part curved, cov- the placement of Melogramma in Melogrammataceae in
ering ascomata . . . . . . . . . . . . . . . . . . . . Cryptomeliola Xylariales, while Senanayake et al. (2015) showed its place-
4. Hyphal setae straight or sometimes slightly ment in Xylariaceae.
curved . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Meliola
5. Ascomata with appendages or setae . . . . . . . . . . . . . . 6 Melogramma Fr., Summa veg. Scand., Section Post.
5. Ascomata without appendages and setae, raised conical (Stockholm): 386 (1849)
cells on the ascoma wall. . . . . . . . . . . . . . . Asteridiella Facesoffungi number: FoF 01320 Figs 91, 92
6. Ascomata with larviform to cylindrical Saprobic on woody plants. Sexual morph: Stromata red-
appendages . . . . . . . . . . . . . . . . . . . . Appendiculella dish-brown, solitary, scattered or aggregated in lines, im-
6. Ascomata with long setae, mostly curved at the mersed when young, erumpent from bark and becoming su-
apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Irenopsis perficial when mature, pulvinate or discoid, soft textured, with
 Fungal Diversity

Fig. 91 Melogramma
campylosporum (Material
examined: AUSTRIA, Styria,
Graz, distr. Mariatrost, on the
north-facing hillside below the
church, 47°06′29″N, 15°29′32″E,
MTB 8858/4, c. 440 m alt.;
clearing, on dead shoots of
Carpinus betulus L. (Betulaceae),
5 November 2006, leg. and det. C.
Scheuer (#5321), S F123341) a
Herbarium material b Substrate c
Stromata d Ostioles as black spots
on stroma e Periphyses f
Ascomata formed in a stroma g
Peridium h-l Asci m Paraphyses
n-q Ascospores. Scale bars: c, d,
= 500 μm, f = 200 μm, e, h–
l = 30 μm, g, m–q = 15 μm

pseudoparenchymatous cells, apex papillate with black osti- straight, mature ascospores 3-septate, falcate, often strongly
ole. Ascomata perithecial, dark brown to black, sometimes curved, ends subacute, guttulate, smooth-walled.
confluent, immersed, globose, ostiolate. Ostiole periphysate. Asexual morph: Coelomycetous. Conidiomata formed
Peridium comprising of two layers, outer layer dark brown to as pycnidia. Mycelium composed of hyphae hyaline to
black, composed of textura angularis to prismatica, inner yellow-brown, septate, branched. Conidiophores long,
layer hyaline, with thin-walled cells. Hamathecium of narrow, cylindrical, stiff, upright, septate, verticillate and
septate, hyphae-like paraphyses. Asci 8-spored, unitunicate, whorled at the tip or not. Conidiogenous cells holoblas-
straight, clavate, fusoid, sigmoid, short pedicellate, with a tic, polyblastic, hyaline, elongate, thin-walled, and pro-
shallow, J-, apical ring. Ascospores overlapping 2-seriate to liferating sympodially. Conidia acrogenous, elongate,
fasciculate, brown, young ascospores 0–1-septate, hyaline, falcate or filiform, tapering slightly to rounded ends,
Fungal Diversity

Fig. 92 Melogramma
campylosporum (redrawn from
Laflamme 1976) a. Germinated
ascospore b Pycnidia c Young
hyphae at the edge of culture d
Conidiophores with conidia e
Conidia. Scale bars: a = 15 μm,
b = 30 μm, c–e = 3 μm

hyaline, simple, smooth (Laflamme 1976; Barr 1990; Melogramma campylosporum and M. caucasicum and the
Sutton 1980; Fungi of Great Britain and Ireland). asexual morphs. Breitenbach and Kränzlin (1984) provided
Type species: Melogramma campylosporum Fr., Summa illustrations for Melogramma. Jaklitsch and Voglmayr
veg. Scand., Section Post. (Stockholm): 386 (1849) (2012) further illustrated and provided descriptions after
Notes: The genus Melogramma was established by Fries checking the herbarium of type species. Kirk et al. (2008)
(1849) with Melogramma campylosporum as its type species. estimated there were three Melogramma species, however,
Laflamme (1976) provided descriptions of two species other papers (Fries 1849; Lumbsch and Huhndorf 2010;
 Fungal Diversity

Maharachchikumbura et al. 2015) have rarely mentioned the darkly pigmented, usually hairy, rarely glabrous; asci
genus. arise singly or in chains, without croziers, evanescent,
and irregularly disposed throughout the centrum and as-
Microascaceae Luttr. ex Malloch, Mycologia 62(4): 734 cospores are reddish brown to copper coloured with
(1970) germ slits, dextrinoid when young, and smooth-walled,
Facesoffungi number: FoF 01798 extruded through the ostiole into a gelatinous drop or a
Saprobic on soil, decaying plant material and animal dung long cirrhus. The asexual morphs of this family are
or opportunistic pathogens of insects and animals, including annellidic, such as, Cephalotrichum and Scopulariopsis.
humans, in terrestrial and aquatic habitats. Sexual morph: Some produce aleurioconidia as in Petriella, and also
Ascomata perithecial or cleistothecial, immersed or superfi- arthroconidia as in Kernia (Malloch 1970, 1971). The
cial, scattered or aggregated, black, globose to ampulliform, Microascaceae forms a sister clade to the Halosphaeriaceae
glabrous or covered with scattered hairs, ostiolate. Ostiole with moderate support in phylogenies based on SSU
usually with a neck of variable length and shape, sometimes sequence data (Réblová et al. 2011). However, few taxa
with a tuft of ostiolar hairs. Peridium dark brown or black, in the family have been sequenced and further
composed of thick-walled, slightly flattened cells of textura collections, isolation and sequencing is warranted to
angularis or textura intricata. Hamathecium lacking resolve its relationship with the Halosphaeriaceae
interascal tissues. Asci 8-spored, very thin-walled, unitunicate, within the order Microascales. Maharachchikumbura
obovate, barrel-shaped or nearly globose, formed in basipetal et al. (2015) confirmed the placement of Microascus
rows, evanescent. Ascospores reddish brown to copper and Petriella within the family Microascaceae in the
coloured, 1-celled, asymmetrical, reniform, heart-shaped, tri- order Microascales, Hypocreomycetidae, with support
angular or quadrangular, dextrinoid when young, smooth- from molecular data.
walled, with germ slits. Asexual morph: Conidiophores sim-
ple, macronematous, mononematous or sporodochial to Microascus Zukal, Verh. zool.-bot. Ges. Wien 35: 342 (1886)
synnematous, with or without rhizoids. Conidiogenous cells Facesoffungi number: FoF 02120; Fig. 93
annellidic, borne singly and laterally on the vegetative hyphae, Saprobic on soil, decaying plant material and animal dung
or in groups of 2–5 on short simple or little branched conidi- or opportunistic pathogens of insects and animals, including
ophores, ampulliform or lageniform, subhyaline or darkening humans. Sexual morph: Ascomata superficial or immersed,
with age, smooth- or rough-walled with a distinct cylindrical subglobose to globose, solitary to gregarious, dark brown to
annellated zone. Conidia 1-celled, pale yellowish to dark black, carbonaceous, hairy or glabrous, ostiolate. Ostioles pa-
brown, globose to subglobose, obovate or clavate, with a trun- pillate to long necked. Peridium composed of several cell
cate base and rounded or pointed at the apex, smooth- and layers of hyaline, pseudoparenchymatous cells. Asci 8-spored,
thin-walled or finely roughened and thick-walled, produced ovoid to subglobose sometimes clavate, sessile or short pedi-
singly or in basipetal dry chains. Solitary conidia present in cellate, disposed at all levels within the centrum, evanescent.
some species, borne sessile or on short stalks from the vege- Ascospores seriate, pale red-brown to dark red-brown in mass,
tative hyphae. ellipsoidal, or mostly asymmetrical, sometimes triangular or
Type: Microascus Zukal tetra-angular, with two germ pores, dextrinoid when imma-
Notes: The family Microascaceae and order Microascales ture, smooth-walled. Asexual morph: Conidiophores often
were proposed by Luttrell (1951) and validly published with elongate, sometimes synnematous, dark and usually smooth-
Latin descriptions by Malloch (1970) and Benny and walled. Conidiogenous cells annellidic, borne singly and lat-
Kimbrough (1980). Luttrell (1951) described the family erally on the vegetative hyphae, or in groups of 2–5 on short
Microascaceae for taxa with ascomata with necks and simple or rarely branched conidiophores, ampulliform or
evanescent, apedicellate asci, disposed irregularly lageniform, subhyaline or darkening with age, smooth- or
throughout the filamentous centrum. The description by rough-walled with a distinct cylindrical, annellate zone.
Luttrell (1951) was confirmed by Corlett (1963, 1966) who Conidia 1-celled, pale yellowish to dark brown, globose to
showed that asci of Microascus and Petriella develop directly subglobose, obovate or clavate, with a truncate base and
from the cells of the ascogenous hyphae, and not from rounded or pointed at the apex, smooth, thin-walled or finely
croziers. Benny and Kimbrough (1980) suggested that mem- roughened, thick-walled, produced singly or in basipetal dry
bers of the family Microascaceae had evolved away from a chains. Solitary conidia present in some species, borne sessile
hymenial configuration; in the microascaceous centrum, or on short stalks from the vegetative hyphae.
a peripheral layer of paraphysoid elements that grows Type species: Microascus longirostris Zukal, Verh. zool.-
inward towards the ascogenous hyphae. Malloch bot. Ges. Wien 35: 339 (1886)
(1970) reviewed the family Microascaceae and included Notes: The genus Microascus was established by Zukal
both ostiolate and non-ostiolate taxa; ascomata are (1886) with Microascus longirostris as the type species.
Fungal Diversity

Fig. 93 Microascus alveolaris

CBS 139501 a. Ascomata b
Peridium c Asci d Ascospores e
Conidiophores, annellides and
conidia (redrawn from Sandoval-
Denis et al. 2016). Scale bars:
a = 30 μm, b–e = 5 μm

Microascus (1886) was initially characterized by carbona- (Emmons and Dodge 1931; Moreau 1953; Doguet
ceous, ostiolate, more or less spherical perithecia and ev- 1957). Barron et al. (1961) and Morton and Smith
anescent asci distributed at all levels throughout the ven- (1963) evaluated the taxonomy of the genus, and its ge-
ter of the ascomata. Microascus was initially included in neric concept remains unchanged. Microascus groups
the Eurotiales based on the close similarities in the devel- with Pseudallescheria with low bootstrap support
opment and organization of the perithecial centrum (Réblová et al. 2011).
 Fungal Diversity

Other genera included Type species: Pseudoscopulariopsis schumacheri (E.C.

Hansen) Sandoval-Denis et al., in Sandoval-Denis et al.,
Anekabeeja Udaiyan & V.S. Hosag., J. Econ. Taxon. Bot. Persoonia, Mol. Phyl. Evol. Fungi 36: 24 (2015)
15(3): 649 (1992) [1991] Scedosporium Sacc. ex Castell. & Chalm., Manual of trop-
Type species: Anekabeeja lignicola Udaiyan & V.S. ical medicine (London): 1122 (1919)
Hosag., J. Econ. Taxon. Bot. 15(3): 649 (1992) [1991] Type species: Scedosporium apiospermum Sacc. ex
Brachyconidiellopsis Decock et al., Cryptog. Mycol. Castell. & Chalm., Manual of tropical medicine (London):
25(2): 140 (2004) 1122 (1919)
Type species: Brachyconidiellopsis fimicola Decock et al., Scopulariopsis Bainier, Bull. Soc. Mycol. Fr. 23: 98 (1907)
Cryptog. Mycol. 25(2): 142 (2004) Type species: Scopulariopsis brevicaulis (Sacc.) Bainier,
Canariomyces Arx, Persoonia 12(2): 185 (1984) Bull. Soc. mycol. Fr. 23: 99 (1907)
Type species: Canariomyces notabilis von Arx, Persoonia Tinhaudeus K.L. Pang et al., in Ariyawansa et al., Fungal
12(2): 185 (1984) Diversity: 10.1007/s13225-015-0346-5, [134] (2015)
Cephalotrichum Link, Mag. Gesell. naturf. Freunde, Type species: Tinhaudeus formosanus K.L. Pang et al., in
Berlin 3(1–2): 20 (1809) Ariyawansa et al., Fungal Diversity: 10.1007/s13225-015-
Type species: Cephalotrichum stemonitis (Pers.) Nees, 0346-5, [138] (2015)
Mag. Gesell. naturf. Freunde, Berlin 3(1–2): 20 (1809) Wardomyces F.T. Brooks & Hansf., Trans. Br. mycol. Soc.
Doratomyces Corda, in Sturm, Deutschl. Fl., 3 Abt. (Pilze 8(3): 137 (1923)
Deutschl.) 2: 65 (1829) Type species: Wardomyces anomalus F.T. Brooks &
Type species: Doratomyces neesii Corda, in Sturm, Hansf. [as ‘anomala’], Trans. Br. mycol. Soc. 8(3): 137 (1923)
Deutschl. Fl., 3 Abt. (Pilze Deutschl.) 2(7): 65 (1829) Wardomycopsis Udagawa & Furuya, Mycotaxon 7(1): 92
Echinobotryum Corda, in Sturm, Deutschl. Fl., 3 Abt. (1978)
(Pilze Deutschl.) 3(12): 51 (1831) Type species: Wardomycopsis inopinata Udagawa &
Type species: Echinobotryum atrum Corda, in Sturm, Furuya, Mycotaxon 7(1): 92 (1978)
Deutschl. Fl., 3 Abt. (Pilze Deutschl.) 3(12): 51 (1831)
Enterocarpus Locq.-Lin., Revue Mycol., Paris 41(4): 510
(1977) Key to sexual genera of Microascaceae
Type species: Enterocarpus uniporus Locq.-Lin., Revue
Mycol., Paris 41(4): 513 (1977) 1. Ascomata with a peridium of textura epidermoidea . . . . 2
Kernia Nieuwl., Am. Midl. Nat. 4: 379 (1916) 1. Ascomata with a peridium of pseudoparenchymatous
Type species: Kernia nitida (Sacc.) Nieuwl., Am. Midl. cells. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
Nat. 4: 379 (1916) 2. Ascomata perithecial, ascospores asymmetrical . . . . . 2
Knoxdaviesia M.J. Wingf. et al., Mycologia 80(1): 26 (1988) 2. Ascomata cleistothecial, ascospores
Type species: Knoxdaviesia proteae M.J. Wingf. et al., symmetrical. . . . . . . . . . . . . . . . . . .Pseudallescheria
Mycologia 80(1): 26 (1988) 3. Ascospores reddish brown . . . . . . . . . . . . . . . Petriella
Lophotrichus R.K. Benj., Mycologia 41(3): 347 (1949) 3. Ascospores subhyaline . . . . . . . . Pseudoscopulariopsis
Type species: Lophotrichus ampullus R.K. Benj., 4. Ascospores embedded in a hyphal capillitium at maturi-
Mycologia 41(3): 347 (1949) ty, with a prominent germ pore. . . . . . . . Enterocarpus
Parascedosporium Gilgado et al., Int. J. Syst. Evol. 4. Ascospores not embedded in a capillitium, without or
Microbiol. 57(9): 2176 (2007) with 1–2 germ pores . . . . . . . . . . . . . . . . . . . . . . . . . 5
Type species: Parascedosporium tectonae (C. Booth) 5. Ascospores pale red-brown to dark red-brown when ar-
Gilgado et al., Int. J. Syst. Evol. Microbiol. 57(9): 2176 (2007) ranged in a mass extruded from the mature perithecium
Petriella Curzi, Boll. R. Staz. Patalog. Veget. Roma 10: as a gelatinous ball at the ostiole . . . . . . . . Microascus
384 (1930) 5. Ascospores hyaline, reddish, yellowish or brownish not
Type species: Petriella asymmetrica Curzi, Boll. R. Staz. in a mass extruded from the mature perithecium. . . . . 6
Patalog. Veget. Roma 10: 392 (1930) 6. Ascospores without or with one germ pore. . . . . . . . . 7
Pseudallescheria Negr. & I. Fisch., Revista Inst. Bacteriol. 6. Ascospores with two germ pores . . . . . . . . . . . . . . . . 8
Dr. Carlos G. Malbrán’ 12 (201): 5–9 (1944) 7. Ascospores ellipsoidal or broadly
Type species: Pseudallescheria shearii Negroni & I. fusiform . . . . . . . . . . . . . . . . . . . . . . Scopulariopsis
Fisch., Rev. Inst. bact., B. Aires 12: 201 (1944) 7. Ascospores broadly reniform or lunate . .Canariomyces
Pseudoscopulariopsis Sandoval-Denis et al., in 8. Ascomata non-ostiolate, with or without tufts of hairs,
Sandoval-Denis et al., Persoonia, Mol. Phyl. Evol. ascospores hyaline, reddish, yellowish or
Fungi 36: 24 (2015) brownish . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Kernia
Fungal Diversity

8. Ascospores usually ostiolate, with an apical tuft of thick- morph: Conidiomata if present, sporodochial. Conidiophores
walled hairs, ascospores yellowish or solitary or aggregated, mono- or biverticillate. Conidiogenous
brownish . . . . . . . . . . . . . . . . . . . . . . . . . Lophotricus cells solitary or in whorls, polyblastic, sympodial, denticulate,
cylindrical often ampulliform, lageniform with elongated
necks and minute annellides from percurrent proliferations,
Key to asexual genera of Microascaceae hyaline to pale brown. Conidia lunate, oblong, fusiform or
cylindrical, straight or curved, hyaline, flattened at base.
1. Conidia with germ slits. . . . . . . . . . . . . . . . . . . . . . . . . . 2 Chlamydospores if present, brown (Hernández-Restrepo
1. Conidia without germ slits . . . . . . . . . . . . . . . . . . . . . 3 et al. 2015b).
2. Conidia produce as short chains from annellidic Type: Microdochium Syd.
conidiogenous cells . . . . . . . . . . . . . . .Wardomycopsis Notes: Based on the results of LSU sequence data
2. Conidia single, produce from polyblastic conidiogenous Microdochium, Idriella and Selenodriella form a distinct
cells . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Wardomyces clade in Xylariales (Hernández-Restrepo et al. 2015b).
3. Conidiophores macronematous, mononematous, arising Therefore, the novel family Microdochiaceae was introduced
from well-developed rhizoids . . . . . . . . . Knoxdaviesia for the taxa producing monographella-like sexual morphs and
3. Conidiophores unbranched or sporodochial to an asexual morph characterized by polyblastic, sympodial or
synnematous, without rhizoids . . . . . . . . . . . . . . . . . 4 annellidic conidiogenous cells, with hyaline conidia without
4. Conidia greyish to greyish black, cheirosporous, with appendages (Hernández-Restrepo et al. 2015b). The genera
divergent branches, composed of chains of narrowly placed in Microdochiaceae are phytopathogenic and saprobic
ellipsoid to slightly barrel-shaped on leaves, seeds and soil.
cells. . . . . . . . . . . . . . . . . . . . . . Brachyconidiellopsis
4. Conidia subhyaline, pale brown to brown, without Microdochium Syd. & P. Syd., Annls mycol. 22(3/6): 267
branches, solitary, catenate or clustered . . . . . . . . . . . 5 (1924)
5. Conidia ovate, ellipsoidal to cylindrical, catenate, trun- Facesoffungi number: 02121; Figs 94, 95
cate at the base and pointed at the apex . . . . . . . . . . . 6 Saprobic or pathogenic on plants. Sexual morph:
5. Conidia not as above . . . . . . . . . . . . . . . . . . . . . . . . . 7 Monographella-like, on natural substrate. Ascomata peri-
6. Conidiophores synnematous, upper half of the synnema thecial, immersed, subepidermal, solitary or in groups,
covered with flask-shaped annellides . . . .Doratomyces pale brown to black, globose, subglobose to oval, with
6. Conidiophores synnematous with dark brown to black- central, papillate and often acute ostiole, ostioles usually
ish brown stipes, with cylindrical or ellipsoidal more distinctly pigmented than the perithecial body,
heads . . . . . . . . . . . . . . . . . . . . . . . . . Cephalotrichum filled with slightly clavate periphyses. Peridium brown,
7. Conidiophores flask-shaped with swollen base part and thin-walled, thickened and darker around the ostiole, in sur-
elongated neck . . . . . . . . . . . . . . . . . . . .Scedosporium face view of textura angularis-epidermoidea. Paraphyses
7. Conidiophores erect, simple, branched or irregularly filamentous, apically free, thin-walled. Asci unitunicate,
branched . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 oblong to clavate with 8 bi- to multi-seriate ascospores,
8. Conidia brown, pear-shaped, beaked, apex with an amyloid, refractive, flat, funnel-shaped ring.
roughened . . . . . . . . . . . . . . . . . . . . .Echinobotryum Ascospores clavate, fusoid or oblong, hyaline to brownish,
8. Conidia subhyaline to brown, sub cylindrical, smooth- straight or curved, smooth-walled and septate. Asexual
walled . . . . . . . . . . . . . . . . . . . . . . Parascedosporium morph: Coelomycetous. Mycelium branched and septate.
Sporodochia, if present, epidermal, subepidermal to erumpent
Microdochiaceae Hern.-Restr. et al., in Hernández-Restrepo through stromata; hyaline, pseudoparenchymatic, spreading
et al., Persoonia 36: 64 (2015) after egress. Conidiophores more or less verticillate, reduced
Facesoffungi number: FoF 01907 to conidiogenous cells, hyaline, smooth. Conidiogenous cells
Saprobic, endophytic or pathogenic on leaves, seeds and holoblastic, discrete, hyaline, smooth, solitary or aggregated
soil. Sexual morph: Stroma present or absent. Ascomata peri- in small sporodochia. Two kinds: with sympodial prolifera-
thecial, immersed, scattered, solitary, papilla of textura tion, cylindrical or slightly tapering, or clavate, denticulate
intricata, Peridium 3–4 layers, in surface view textura with one or more apical denticles. Or with percurrent prolif-
epidermoidea. Asci cylindrical, oblong, clavate, unitunicate, eration (annellidic), subcylindrical, obpyriform, ampulliform
with an amyloid funnel-shaped apical ring and 8 biseriate or to lageniform. Conidia dry or slimy in mass, unicellular or
uniseriate ascospores. Paraphyses persisting between asci, multi-septate, hyaline, smooth, lunate, falcate, fusiform, fili-
apically free, filamentous, unbranched or infrequently form, obovoid or subpyriform, straight or curved, apex round-
branched, thin-walled, septate. Ascospores ellipsoid or ob- ed, base flattened. Sometimes the conidia originate directly
long, fusoid, hyaline to pale brown, smooth-walled. Asexual from hyphae. Chlamydospores terminal or intercalary,
 Fungal Diversity

Other genera included

Idriella P.E. Nelson & S. Wilh., Mycologia 48: 550. 1956

Type species: Idriella lunata P.E. Nelson & S. Wilh.,
Mycologia 48: 550.
Selenodriella R.F. Castañeda & W.B. Kendr., Univ.
Waterloo Biol. Ser. 33: 34 (1990)
Type species: Selenodriella fertilis (Piroz. & Hodges) R.F.
Castañeda & W.B. Kendr., Univ. Waterloo Biol. Ser. 33: 34
(1990)

Key to genera of Microdochiaceae

1. Chlamydospores absent . . . . . . . . . . . . . . Microdochium

1. Chlamydospores present . . . . . . . . . . . . . . . . . . . . . . 2
2. Conidiophores setiform, conidiogenous cells and
branches are arranged in whorls along the main axis of
setiform conidiophores . . . . . . . . . . . . . Selenodriella
2. C o n i d i o p h o r e s r e d u c e d t o c o n i d i o g e n o u s
cells . . . . . . . . . . . . . . . . . . . . . . Idriella

Myelospermataceae K.D. Hyde & S.W. Wong [as

‘Myelospermaceae’], Mycol. Res. 44(1): 349 (1999)
Facesoffungi number: FoF 02098
Saprobic or endophytic on palms. Sexual morph:
Ascomata occurring in host tissues, in pseudostromata,
appearing as weakly raised, blister-like regions on the host
surface, immersed, solitary, or aggregated in a valsoid
configuration, if aggregated clustered around a common
central pore, subglobose to ellipsoidal, dark brown,
Fig. 94 Microdochium seminicola - sexual morph (CBS 139951, ex- necks mostly eccentric. Peridium comprising a thick
type) a, b Asci c Ascus ring in Melzer’s reagent; d Ascospores (redrawn
from Hernández-Restrepo et al. 2015b)
layer of pale brown-walled compressed cells of textura
angularis. Paraphyses filamentous, basal cell globose,
cylindrical, septate, unbranched, tapering distally. Asci
8-spored, unitunicate, thin-walled, cylindrical, short ped-
solitary, in chains or grouped in clusters, brown (Hernández- icellate, apically rounded, with a J-, or J+, refractive,
Restrepo et al. 2015b). subapical ring. Ascospores overlapping uniseriate, hya-
Type species: Microdochium phragmitis Syd. & P. Syd., line, olivaceous brown when mature, reniform to ellip-
Annls mycol. 22(3/6): 267 (1924) soidal, unicellular, guttulate, smooth walled and
Notes: Microdochium was introduced by Sydow (1924) surrounded by a mucilaginous sheath. Asexual morph:
with the type M. phragmitis Syd. for a fungus on living on Undetermined.
discoloured leaves of Phragmites australis. To date 31 Type: Myelosperma Syd. & P. Syd.; Fig. 96
Microdochium species have been described, which includes Type species: Myelosperma tumidum Syd. & P. Syd.,
saprotrophs, some endophytes that produce secondary metab- Annls mycol. 13(1): 38 (1915)
olites and pathogens of various plants (Zhang et al. 2015). The Notes: Myelospermataceae was introduced by Hyde and
sexual morphs of Microdochium were identified as Wong (1999) and is typified by Myelosperma with
Monographella and many of them cause plant diseases pre- Myelosperma tumidum as the type species. Characters include
dominantly on grasses and cereals (Jaklitsch & Voglmayr cylindrical asci, with a refractive subapical ring, and reniform,
2012. Since, Microdochium has more species and the name olivaceous brown ascospores (Hyde 1993; Hyde and Wong
is more frequently used in the literature, Hernández-Restrepo 1999). Hyde and Wong (1999) first placed this family under
et al. (2015b) gave priority to Microdochium over Diaporthales. Subsequently, Kirk et al. (2001) placement the
Monographella. genus in Xylariales, which seemed controversial due to lack of
Fungal Diversity

Fig. 95 Microdochium
phragmitis – asexual morph
(CBS 423.78) a, b Sporodochia c,
d Conidiophores with
conidiogenous cells e Conidia
(redrawn from Hernández-
Restrepo et al. 2015b). Scale bars:
a = 50 μm, b–e = 10 μm

suitable reference taxa and sequence analysis (Eriksson 1999; = Tuberculariaceae Fr. [as ‘Tubercularini’], Syst. orb. veg.
Kang et al. 2002). Later, Eriksson et al. (2003) placed (Lundae) 1: 169 (1825)
Myelosperma in Lasiosphaeriaceae based on molecular phy- Facesoffungi number: FoF 01396
logenetic analysis of LSU and SSU data. The taxonomic Folicolous, entomogenous, endophytic or saprobic mostly
placement of this genus is confused, thus further taxon sam- on woody plant hosts, a few species being human pathogens,
pling and multi-gene analysis is required (Smith et al. 2003). in terrestrial and aquatic habitats. Sexual morph: Ascomata
Currently, five species are recorded in Index Fungorum stromatic or non-stromatic, white, red, dark red, reddish-
(2016). brown, orange, orange-red, orange-brown, umber, yellow,
We collected a new species of Myelosperma from south- pale yellow, brown, greyish yellow-green, dark bluish, bluish
ern Thailand, also on a palm rachis. LSU sequence data purple, bluish black or black, solitary or aggregated in groups,
indicate the species belongs in the order Xylariales, which perithecial, globose to subglobose, ovoid, elongate-ovoid,
is in agreement with the asci having a J+, subapical ring. obpyriform, obovoid or pyriform, changing or not changing
However, multi-gene analysis is needed to confirm colour in KOH, surface smooth to papillate, striate, warted,
Myelospermataceae is a distinct family. Our taxon com- verrucose or scaly, with or without setae, periphysate, ostiolar
prised individual ascomata clustered in a raised region sometimes papillate. Paraphyses present or absent.
pseudostroma, and may be a new species, which will be Asci 4–8-spored, unitunicate, clavate to narrowly clavate, cy-
the subject of a future publication. lindrical or ellipsoidal, with or without apical ring, with point-
ed or pedicellate base. Ascospores uniseriate to biseriate or
Nectriaceae Tul. & C. Tul. [as ‘Nectriei’], Select. fung. overlapping, hyaline to yellow, yellow-brown, golden-brown,
carpol. (Paris) 3: 3 (1865) pale-brown or green, fusiform, long-fusiform, ellipsoidal,
 Fungal Diversity

Fig. 96 Myelosperma sp. (Material examined: THAILAND, Krabi bluing in Melzer’s reagent h-j Asci k-n Ascospores o Germinated
Province, 3 December 2014, S. Konta, MFLU 15-0269) a, b Ascomata ascospore p Colony on culture. Scale bars: a = 1 mm, b = 200 μm,
c Section of ascoma d Neck e Peridium f Paraphyses g +, apical ring, c = 100 μm, d, h–j = 20 μm, e = 10 μm, f-g, k–o = 5 μm
Fungal Diversity

oblong, biconic, pyriform, reniform or allantoid, aseptate to Curvicladium, Cyanochyta, Cyanophomella, Dacryoma,
multi-septate or muriform, constricted at the septum or not, Persiciospora, Stachybotryna and Xenonectriella) accepted
smooth-walled, spinulose, verruculose or striate. Asexual by Maharachchikumbura et al. (2015), but they did include
morph: Hyphomycetous or coelomycetous. Conidiomata a n a dd i t i o n a l fi ve ge n er a ( Cy li nd ro c ar p os ty l us ,
synnematous, sporodochial or pycnidial. Conidiophores un- Dactylonectria, Sarcopodium, Xenocylindrocladium and
branched or branched, penicillate, verticillate. Conidiogenous Xenogliocladiopsis). Lombard et al. (2015) accepted 47 gen-
cells monophialidic to polyphialidic, ampulliform to era in the family based on rDNA sequence data. The present
lageniform, cylindrical, elongate-ampulliform or study follows Lombard et al. (2015)
subcylindrical, hyaline, smooth-walled. Conidia globose, Nectria (Fr.) Fr., Summa veg. Scand., Section Post.
ovate, ellipsoidal, cylindrical to subcylindrical, fusiform, (Stockholm): 387 (1849)
long-fusiform, filiform, allantoid or falcate, straight to slightly Facesoffungi number: FoF 02122; Figs 97, 98
or strongly curved, hyaline, aseptate to multi-septate, con- Saprobic on decaying wood and occurring as pathogens of
stricted at septum or not, with or without visible abscission trees. Sexual morph: Stromata, erumpent. Ascomata perithe-
scars, sometimes guttulate, smooth-walled. Chlamydospores cial, on or nearly or completely immersed in a stroma, aggre-
present or absent. gated in groups, red to bay to sienna, turning bright red, blood
Type: Nectria (Fr.) Fr. red to purple in KOH, globose to subglobose, with a smooth to
Notes: Seaver (1909a, b, 1910a, b, 1911) divided the order warted surface. Asci 8-spored, unitunicate, clavate to narrowly
Hypocreales into two families (Nectriaceae and clavate or cylindrical, apical ring inconspicuous. Ascospores
Hypocreaceae) by considering stromatic and perithecial 0–4-septate, ellipsoidal, oblong, fusiform, pyriform or
characters. Petch (1938) also accepted Nectriaceae as a sepa- allantoid, with rounded ends, straight to slightly curved, hya-
rate family in Hypocreales, while Munk (1957) and Dennis line, smooth-walled or spinulose. Asexual morph:
(1960) placed it in Sphaeriales. Miller (1949); Bessy (1950); Conidiophores pycnidial, sporodochial, lateral phialidic pegs
Luttrell (1951); Dingley (1951a, b, 1952a, b, 1953, 1954, or acropleurogenous. Microconidia ellipsoid to fusoid or
1956); von Arx and Müller (1954); Müller and von Arx sometimes curved, aseptate, hyaline. Macroconidia 0–1-
(1962); Gäumann (1964); Rogerson (1970); Barr (1990) septate, ellipsoidal, oblong, cylindrical to allantoid or
placed Nectriaceae under Hypocreaceae as one family. subglobose to ellipsoidal, straight to slightly curved,
However, Kreisel (1969); Rossman et al. (1999) accepted that rounded at ends, hyaline, smooth-walled.
Nectriaceae and Hypocreaceae are two separate families Chlamydospores rare (description based on Hirooka
within Hypocreales. Nectriaceae was provisionally accepted et al. 2012; Lombard et al. 2015).
as a separate family based on cladistic analyses of rRNA data Type species: Nectria cinnabarina (Tode) Fr., Summa veg.
(Spatafora and Blackwell 1993). Nectriaceae is characterized Scand., Section Post. (Stockholm): 388 (1849)
by uniloculate, pigmented ascomata and phialidic Notes: Fries (1849) introduced the genus Nectria based on
amerosporous to phragmosporous conidia. Ascomata are yel- Hypocrea sect. Nectria Fr. Since then many species have been
low, orange-red or purple and KOH+ (Rossman et al. 1999; added to the genus. Based on morphological characteristics
Rossman 2000; Lombard et al. 2015). Some members of this such as structure of perithecial and asexual morphs (Hirooka
family are weak or even virulent pathogens, while some are et al. 2010; Rossman et al. 1999) and LSU sequence data
saprobes, fungicolous or insecticolous (Rossman et al. 1999; (Rehner and Samuels 1995) Nectria sensu lato was separated
Rossman 2000; Chaverri et al. 2011; Schroers et al. 2011; into a number of genera. Rossman et al. (1999) restricted
Hyde et al. 2014). Rossman et al. (1999) restricted Nectria Nectria sensu stricto to species considered congeneric with
sensu stricto to species considered congeneric with the type species Nectria cinnabarina and accepted 27 species
N. cinnabarina and accepted 20 genera in Nectriaceae. within the genus.
Lumbsch and Huhndorf (2010) listed 26 genera under ≡ Tubercularia vulgaris Tode, Fung. mecklenb. sel.
Nectriaceae, while Maharachchikumbura et al. (2015) accept- (Lüneburg) 1: 18 (1790)
ed 67 genera. Of these, Lombard et al. (2015) synonymized
ten (Chaetopsinectria, Dialonectria, Glionectria, Heliscus,
Antipodium, Viridispora, Actinostilbe, Pleonectria, Other genera included
Volutellonectria and Xenocalonectria) under other genera in
the Nectriaceae, while four (Pseudocosmospora, Stalagmites, Albonectria Rossman & Samuels, in Rossman et al., Stud.
Pleurocolla and Pleogibberella) were excluded or not treated Mycol. 42: 105 (1999)
due to lack of molecular data, and Rodentomyces was Type species: Albonectria rigidiuscula (Berk. & Broome)
transferred to the Hypocreales as genera incertae sedis. Rossman & Samuels, in Rossman et al., Stud. Mycol. 42: 105
However, Lombard et al. (2015) did not consider ten of the (1999)
genera (Allonectella, Baipadisphaeria, Chaetonectrioides, Allantonectria Earle, in Greene, Plant. Bak. 2(1): 11 (1901)
 Fungal Diversity

Fig. 97 Nectria cinnabarina

(Material examined: FRANCE:
Villiers en Bois, on dead twigs of
Aesculus sp., Lechat, C.
(CLL7152), 2008 February 13,
Detr: Lechat, C. BPI 879981,
epitype) a Herbarium material b
Ascomata and long stipitate
sporodochia on host substrate c
Ascomata on host substrate d, e
Transverse section through
ascoma f Transverse section
through peridium g Ascus h
Ascus in Melzer’s reagent i-k
Ascospores. Scale bars:
b = 1 mm, c = 2 mm, d = 500 μm,
e = 200 μm, f = 50 μm, g,
h = 20 μm, i–k = 10 μm

Type species: Allantonectria yuccae Earle, in Greene, Atractium Link, Mag. Gesell. naturf. Freunde, Berlin 3(1–
Plant. Bak. 2(1): 11 (1901) 2): 10 (1809)
Allonectella Petr., Sydowia 4(1–6): 345 (1950) Type species: Atractium stilbaster Link, Mag. Gesell.
Type species: Allonectella rubescens Petr., Sydowia 4(1– naturf. Freunde, Berlin 3(1–2): 10 (1809)
6): 345 (1950) Bisifusarium L. Lombard, Crous & W. Gams, in Lombard
Aquanectria L. Lombard & Crous, in Lombard et al., Stud. et al., Stud. Mycol. 80: 223 (2015)
Mycol. 80: 207 (2015) Type species: Bisifusarium dimerum (Penz.) L. Lombard
Type species: Aquanectria penicillioides (Ingold) L. & Crous, in Lombard et al., Stud. Mycol. 80: 224 (2015)
Lombard & Crous, Lombard et al., Stud. Mycol. 80: 207 Calonectria De Not., Comm. Soc. crittog. Ital. 2(3): 477
(2015) (1867)
Fungal Diversity

Fig. 98 Nectria cinnabarina (Material examined: FRANCE: Villiers en sporodochia on host substrate b Transverse section through
Bois, on dead twigs of Aesculus sp., Lechat, C. (CLL7152), 2008 sporodochium c Conidiophore d Conidia. Scale bars: a = 2 mm,
February 13, Detr: Lechat, C. BPI 879981, epitype) a Long stipitate b = 200 μm, c = 20 μm, d = 10 μm

=Cylindrocladium Morgan, Bot. Gaz. 17: 191 (1892) Type species: Corallomycetella heinsenii Henn. [as
Type species: Calonectria daldiniana De Not., Comm. ‘heinesii’], Hedwigia 43: 245 (1904)
Soc. crittog. Ital. 2(3): 477 (1867) Corallonectria C. Herrera & P. Chaverri, Mycosystema
Calostilbe Sacc. & P. Syd., Syll. fung. (Abellini) 16: 591 32(3): 539 (2013)
(1902) Type species: Corallonectria jatrophae (Möller) C.
= Calostilbella Höhn., Ber. dt. bot. Ges. 37: 160 (1919) Herrera & P. Chaverri, Mycosystema 32(3): 539 (2013)
Type species: Calostilbe longiasca (Möller) Sacc. & P. Cosmospora Rabenh., Hedwigia 2: 59 (1862)
Syd., Syll. fung. (Abellini) 16: 591 (1902) Type species: Cosmospora coccinea Rabenh., Fungi europ.
Campylocarpon Halleen et al., Stud. Mycol. 50(2): 448 exsicc.: no. 459 (1862)
(2004) Curvicladiella Decock & Crous, Stud. Mycol. 55: 225
Type species: Campylocarpon fasciculare Schroers et al., (2006)
in Halleen et al., Stud. Mycol. 50(2): 448 (2004) Type species: Curvicladiella cignea (Decock & Crous)
Chaetonectrioides Matsush., Matsush. Mycol. Mem. 9: 5 Decock & Crous, in Crous et al., Stud. Mycol. 55: 225
(1996) (2006)
Type species: Chaetonectrioides malaysiana Matsush., Curvicladium Decock & Crous, Mycologia 90(2): 276
Matsush. Mycol. Mem. 9: 5 (1996) (1998)
Chaetopsina Rambelli, Atti Accad. Sci. Ist. Bologna, Cl. Type species: Curvicladium cigneum Decock & Crous,
Sci. Fis. Rendiconti 3: 5 (1956) Mycologia 90(2): 277 (1998)
= Chaetopsinectria J. Luo & W.Y. Zhuang, Mycologia Cyanochyta Höhn., Sber. Akad. Wiss. Wien, Math.-
102(4): 979 (2010) naturw. Kl., Abt. 1 124: 92 (1915)
Type species: Chaetopsina fulva Rambelli, Diagn. IV 3: 5 Type species: Cyanochyta cyanogena (Speg.) Höhn., Öst.
(1956) bot. Z. 68: 120 (1915)
Coccinonectria L.Lombard & Crous, in Lombard et al., Cyanonectria Samuels & P. Chaverri, in Samuels et al.,
Stud. Mycol. 80: 218 (2015) Mycol. Progr. 8(1): 56 (2009)
Type species: Coccinonectria pachysandricola (B.O. Type species: Cyanonectria cyanostoma (Sacc. &
Dodge) L. Lombard & Crous. in Lombard et al., Stud. Flageolet) Samuels & P. Chaverri, Mycol. Progr. 8(1): 56
Mycol. 80: 218 (2015) (2009)
Corallomycetella Henn., Hedwigia 43: 245 (1904) Cyanophomella Höhn., Hedwigia 60: 156 (1918)
 Fungal Diversity

Type species: Cyanophomella acervalis (Sacc.) Höhn., Macroconia (Wollenw.) Gräfenhan, Seifert & Schroers, in
Hedwigia 60: 157 (1918) Gräfenhan et al., Stud. Mycol. 68(1): 101 (2011)
Cylindrocarpostylus R. Kirschner & Oberw., Mycol. Res. Type species: Macroconia leptosphaeriae (Niessl)
103: 1155. 1999. Gräfenhan & Schroers, in Gräfenhan et al., Stud. Mycol.
Type species: Cylindrocarpostylus gregarius (Bres.) R. 68(1): 102 (2011)
Kirschner & Oberw., Mycol. Res. 103: 1155. 1999. Mariannaea G. Arnaud, Bull. trimest. Soc. mycol. Fr. 68:
Cylindrocladiella Boesew., Can. J. Bot. 60(11): 2289 196 (1952)
(1982) Type species: Mariannaea elegans G. Arnaud, Bull.
Type species: Cylindrocladiella parva (P.J. Anderson) trimest. Soc. mycol. Fr. 68: 196 (1952)
Boesew., Can. J. Bot. 60(11): 2289 (1982) Microcera Desm., Annls Sci. Nat., Bot., sér. 3 10: 359
Cylindrodendrum Bonord., Handb. Allgem. mykol. (1848)
(Stuttgart): 98 (1851) Type species: Microcera coccophila Desm., Annls Sci.
Type species: Cylindrodendrum album Bonord., Handb. Nat., Bot., sér. 3 10: 359 (1848)
Allgem. mykol. (Stuttgart): 48 (1851) Nalanthamala Subram., J. Indian bot. Soc. 35: 478 (1956)
Dacryoma Samuels, Brittonia 40(3): 328 (1988) Type species: Nalanthamala madreeya Subram., J. Indian
Type species: Dacryoma album Samuels [as ‘alba’], bot. Soc. 35: 478 (1956)
Brittonia 40(3): 328 (1988) Nectricladiella Crous & C.L. Schoch, Stud. Mycol. 45: 54
Dactylonectria L. Lombard & Crous, Phytopathol. Medit. (2000)
53: 348. 2014. Type species: Nectricladiella camelliae (Shipton) Crous &
Type species: Dactylonectria macrodidyma (Halleen et al.) C.L. Schoch, in Schoch et al., Stud. Mycol. 45: 54 (2000)
L. Lombard & Crous, Phytopathol. Medit. 53: 352. 2014. Neonectria Wollenw., Annls mycol. 15(1/2): 52 (1917)
Dematiocladium Allegr.et al., Mycol. Res. 109(7): 836 Type species: Neonectria ramulariae Wollenw., Annls
(2005) mycol. 15(1/2): 52 (1917)
Type species: Dematiocladium celtidis Allegr. et al., Neocosmospora E.F. Sm., U.S.D.A. Div. Veg. Pathol. Bull.
Mycol. Res. 109(7): 836 (2005) 17: 45 (1899)
Fusarium Link, Mag. Gesell. naturf. Freunde, Berlin 3(1– Type species: Neocosmospora vasinfecta E.F. Sm., Bull.
2): 10 (1809) U.S. Department of Agriculture 17: 45 (1899)
Type species: Fusarium roseum Link, Mag. Gesell. naturf. Ophionectria Sacc., Michelia 1(no. 3): 323 (1878)
Freunde, Berlin 3(1–2): 10 (1809) Type species: Ophionectria trichospora (Berk. & Broome)
Fusicolla Bonord., Handb. Allgem. mykol. (Stuttgart): 150 Sacc., Michelia 1(no. 3): 323 (1878)
(1851) Paracremonium L. Lombard & Crous, in Lombard et al.,
Type species: Fusicolla betae Bonord., Handb. Allgem. Stud. Mycol. 80: 233 (2015)
mykol. (Stuttgart): 150 (1851) Type species: Paracremonium inflatum L. Lombard &
Geejayessia Schroers et al., in Schroers et al., Stud. Mycol. Crous, in Lombard et al., Stud. Mycol. 80: 233 (2015)
68(1): 124 (2011) Payosphaeria W.F. Leong, in Leong, Tan, Hyde & Jones,
Type species: Geejayessia cicatricum (Berk.) Schroers, in Bot. Mar. 33: 511 (1990)
Schroers et al., Stud. Mycol. 68(1): 124 (2011) Type species: Payosphaeria minuta H.Y.M. Leung, in
Gliocephalotrichum J.J. Ellis & Hesselt., Bull. Torrey bot. Leong, Tan, Hyde & Jones, Bot. Mar. 33: 511 (1990)
Club 89: 21 (1962) Penicillifer Emden, Acta bot. neerl. 17: 54 (1968)
= Leuconectria Rossman, Samuels & Lowen, Mycologia Type species: Penicillifer pulcher Emden, Acta bot. neerl.
85(4): 686 (1993) 17: 54 (1968)
Type species: Gliocephalotrichum bulbilium J.J. Ellis & Persiciospora P.F. Cannon & D. Hawksw., J. Linn. Soc.,
Hesselt., Bull. Torrey bot. Club 89: 22 (1962) Bot. 84: 133 (1982)
Gliocladiopsis S.B. Saksena, Mycologia 46: 662 (1954) Type species: Persiciospora moreaui P.F. Cannon & D.
= Glionectria Crous & C.L. Schoch, Stud. Mycol. 45: 58 Hawksw., J. Linn. Soc., Bot. 84(2): 134 (1982)
(2000) Pleogibberella Sacc. ex Berl. & Voglino, in Saccardo, Syll.
Type species: Gliocladiopsis sagariensis S.B. Saksena, fung., Addit. I-IV (Abellini): 217 (1886)
Mycologia 46: 662 (1954) Type species: Pleogibberella calamia (Cooke) Berl. &
Ilyonectria P. Chaverri & C. Salgado, in Chaverri et al., Voglino, in Saccardo, Syll. fung., Addit. I-IV (Abellini): 217
Stud. Mycol. 68(1): 69 (2011) (1886)
Type species: Ilyonectria radicicola (Gerlach & L. Pleonectria Sacc., Nuovo G. bot. ital. 8(2): 178 (1876)
Nilsson) P. Chaverri & C. Salgado, in Chaverri et al., Stud. Type species: Pleonectria lamyi (Desm.) Sacc. [as
Mycol. 68(1): 71 (2011) ‘lamyii’], Michelia 1(no. 3): 325 (1878)
Fungal Diversity

Pleurocolla Petr., Annls mycol. 22(1/2): 15 (1924) Type species: Nectria serpens Decock, Hennebert &
Type species: Pleurocolla tiliae Petr., Annls mycol. Crous, Mycol. Res. 101(7): 788 (1997)
22(1/2): 15 (1924) Xenogliocladiopsis Crous & W.B. Kendr., Canad. J. Bot.
Pseudocosmospora C. Herrera & P. Chaverri, Mycologia 72: 63. 1994.
105: 1291 (2013) Type species: Xenogliocladiopsis eucalyptorum Crous &
Type species: Pseudocosmospora eutypellae C. Herrera & W.B. Kendr., Can. J. Bot. 72: 63. 1994.
P. Chaverri, Mycologia 105: 1293 (2013) Xenonectriella Weese, Sber. Akad. Wiss. Wien, Math.-
Pseudonectria Seaver, Mycologia 1(2): 48 (1909) naturw. Kl., Abt. 1 128: 749 (1919)
Type species: Pseudonectria rousseliana (Mont.) Type species: Xenonectriella lutescens (Arnold) Weese,
Wollenw., Z. ParasitKde 3(3): 489 (1931) Sber. Akad. Wiss. Wien, Math.-naturw. Kl., Abt. 1 128: 749
Rectifusarium L. Lombard et al., in Lombard et al., Stud. (1919)
Mycol. 80: 229 (2015)
Type species: Rectifusarium ventricosum (Appel &
Wollenw.) L. Lombard & Crous., in Lombard et al., Stud. Key to the genera of Nectriaceae with sexual morph
Mycol. 80: 229 (2015)
Rugonectria P. Chaverri & Samuels, in Chaverri et al., 1. Ascospores always aseptate . . . . . . . . . . . . . . . . . . . . . . 2
Stud. Mycol. 68(1): 73 (2011) 1. Ascospores 1 to multi-septate or muriform, rarely
Type species: Rugonectria rugulosa (Pat. & Gaillard) aseptate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
Samuels et al., in Chaverri et al., Stud. Mycol. 68(1): 73. (2011) 2. Ascomata on well-developed erumpent stromata, . . . . 3
Sarcopodium Ehrenb. ex Schlecht., Synop. Pl. Crypt. 2: 2. Ascomata with basal stromata, superficial . . . . . . . . . 5
101. (1824). 3. Ascospores fusoid with rounded ends. . . . .Calonectria
Type species: Sarcopodium circinatum Ehrenb. ex 3. Ascospores allantoid to cylindrical with rounded
Schlecht., Synop. Pl. Crypt. 2: 101. 1824. corners . . . . . . . . . . . . . . . . . . . . . . . . .Allantonectria
Stachybotryna Tubaki & T. Yokoy., Trans. Mycol. Soc. 5. Ascomata, pale yellow to greyish yellow-green, not
Japan 12(1): 18 (1971) changing color in KOH . . . . . . . . . . . . .Pseudonectria
Type species: Stachybotryna columaris Tubaki & T. Yokoy. 5. Ascomata brilliant red, turning purple in
[as ‘columare’], Trans. Mycol. Soc. Japan 12(1): 18 (1971) KOH . . . . . . . . . . . . . . . . . . . . . Gliocephalotrichum
Stalagmites Theiss. & Syd., Annls mycol. 12(2): 189 6. Ascospores green . . . . . . . . . . . . . . . . . . . . Penicillifer
(1914) 6. Ascospores hyaline or pigmented, not green. . . . . . . . 7
Type species: Stalagmites tumefaciens (Syd. & P. Syd.) 7. Ascomata white to pale yellow, pale brownish-yellow or
Theiss. & Syd., Annls mycol. 12(2): 189 (1914) greyish yellow-green, never red, black, blue or
Stylonectria Höhn., Sber. Akad. Wiss. Wien, Math.- purple. . . . . . . . . . . . . . . . . . . . . . . . . . . . .Albonectria
naturw. Kl., Abt. 1 124: 52 (1915) 7. Ascomata yellow, red, redish orange, orange, or-
Type species: Stylonectria applanata Höhn., Sber. Akad. ange-brown, black, dark blue or purple, rarely
Wiss. Wien, Math.-naturw. Kl., Abt. 1 124: 52 (1915) white . . . . . . . . . . . . . . . . . . . . . . . . . . 8
Thelonectria P. Chaverri & C. Salgado, in Chaverri et al., 8. Ascomata stromatic, erumpent, fully or partially im-
Stud. Mycol. 68(1): 76 (2011) mersed in a slimy, pale orange sheet of hyphae over
Type species: Thelonectria discophora (Mont.) P. the substrate . . . . . . . . . . . . . . . . . . . . . . . . Fusicolla
Chaverri & C. Salgado, in Chaverri et al., Stud. Mycol. 8. Ascomata stromatic or not, if stromatic not immersed in
68(1): 76 (2011) a slimy sheet of hyphae . . . . . . . . . . . . . . . . . . . . . . . 9
Thyronectria Sacc., Grevillea 4(no. 29): 21 (1875) 9. Ascospores more than 1-septate or muriform, rarely 1
Type species: Thyronectria patavina Sacc., Grevillea 4(no. septate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
29): 21 (1875) 9 Ascospores always 1-septate, not muriform or multi-
Volutella Fr., Syst. mycol. (Lundae) 3(2): 458, 466 (1832) septate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
Type species: Volutella ciliata (Alb. & Schwein.) Fr., Syst. 10. Ascospores sometimes budding in the ascus to produce
mycol. (Lundae) 3(2): 467 (1832) oblong to allantoid, aseptate, hyaline,
Xenoacremonium L. Lombard & Crous, in Lombard et al., ascoconidia . . . . . . . . . . . . . . . . . . . . . .Thyronectria
Stud. Mycol. 80: 234 (2015) 10. Ascospores do not produce ascoconidia . . . . . . . . . 33
Type species: Xenoacremonium recifei (Leão & Lõbo) L. 11. Ascomata seated on short red stalks . . Corallonectria
Lombard & Crous., in Lombard et al., Stud. Mycol. 80: 235 11. Ascomata not seated on stalks . . . . . . . . . . . . . . . . 12
(2015) 12. Ascomata broadly ampulliform with a short necks, or
Xenocalonectria Crous & C.L. Schoch, Stud. Mycol. 45: broadly ellipsoidal, pale orange, brownish to reddish
50 (2000) orange, bright red or black . . . . . . . . . . . .Geejayessia
 Fungal Diversity

12. Ascomata globose, ovoid, pyriform to obpyriform, yel- smooth or finely warted, sometimes becoming tubercu-
low to orange-brown to red, but never black. . . . . . 13 late when mature . . . . . . . . . . . . . . . . . . . . . . . . . . 23
13. Found only in aquatic habitats, ascomata ovate to 23. Ascomata globose with a blunt papilla, orange to
subglobose . . . . . . . . . . . . . . . . . . . . . . Aquanectria dark red, ascomatal surface finely
13. Mainly in terrestrial habitats, ascomata globose to roughened . . . . . . . . . . . . . . . . . . Microcera
subglobose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 23. Ascomata ampulliform to obpyriform or pyriform,
14. Ascomata with a rounded or broad, circular, flat disc on dark bluish or red to red-brown, ascomatal surface
a venter-like neck, ascospores cylindrical to allantoid to smooth . . . . . . . . . . . . . . . . . . Cyanonectria
ellipsoidal . . . . . . . . . . . . . . . . . . . . . . . . Stylonectria 24. Ascomata red-brown with a dark red stromatic
14. Ascomata papillate or non-papillate, without flat disc base . . . . . . . . . . . . . . . . . . . . . . . . . . Gliocladiopsis
on a venter-like neck, ascospores ellipsoidal to fusi- 24. Ascoamat pale yellow, orange, red to dark red or
form, oblong or globose. . . . . . . . . . . . . . . . . . . . . 15 brown . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25
15. Ascomatal wall with hyphal hairs (sometimes forming 25. Ascomata not reacting in KOH . . . . . . . . . . . . . . . 26
a tomentum on the ascomatal surface) or with minute 25. Ascomata changing color in KOH . . . . . . . . . . . . . 28
brown setae, very rarely smooth, not warted, scaly or 26. Asci clavate, ascospores light brown,
scruffy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16 verrucose. . . . . . . . . . . . . . . . . . . . . .Rectifusarium
15. Ascomatal wall smooth, warted, scaly or scruffy . . 17 26. Asci cylindrical to narrowly clavate, ascospores hyaline
16. Ascomata developing on old sporodochia, previously or becoming brownish with age, smooth, spinulose or
conidial stroma, or at the base of a synnema . . . . . 35 rarely striate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27
16. Ascomata with a inconspicuous stroma, basal stro- 27. Ascomata globose with a flat apex, pale yellow, orange
ma present or absent, not associated with old or brown . . . . . . . . . . . . . . . . . . . . . . . . Mariannaea
sporodochia, previously conidial stroma or 27. Ascomata globose, subglobose, or pyriform to elongat-
synnema . . . . . . . . . . . . . . . . . . . . . . . 36 ed, red . . . . . . . . . . . . . . . . . . . . . . . . . . Thelonectria
17. Ascomata with orange (rarely green) or red 28. Ascomata dark red, becoming purple-red in
scales . . . . . . . . . . . . . . . . . . . . . . . . 18 KOH . . . . . . . . . . . . . . . . . . . . . . . . Dactylonectria
17. Ascomatal surface not scaly . . . . . . . . . . . . . . . . . . 19 28. Ascomata orange red or bright red, red, yellow to or-
18. Ascomata with orange, rarely green scales, ange or orange-brown, becoming dark red, dark yellow
turning dark red in KOH, ascospores pale to dark orange-brown. . . . . . . . . . . . . . . . . . . . . . . 29
brown to golden-brown, coarsely 29. Ascomata orange-red, surface conspicuously warted,
striate . . . . . . . . . . . . . . . . Nalanthamala sometimes smooth . . . . . . . . . . . . . . . . . . . . . . . . . 30
18. Ascomata with red scales, turning purple-red in KOH, 29. Ascomata yellow to orange-brown to red, surface gen-
ascospores hyaline, smooth . . . . . . . . . . . Ilyonectria erally smooth, very rarely scruffy . . . . . . . . . . . . . . 31
19. Ascospores aggregated in the upper third of the ascus, 30. Ascospores initially hyaline, becoming yellow brown
fusoid. . . . . . . . . . . . . . . . . . . . Xenocylindrocladium to reddish brown, striate. . . Cosmospora/Dialonectria
19. Ascospores not aggregated in the upper third of the 30. Ascospores hyaline or sometimes yellow, becoming
ascus, ellipsoidal, oblong, fusiform, pyriform or tuberculate when mature . . . . . . . . . . . . Rugonectria
allantoid . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20 31. Ascomata globose to pyriform, yellow to orange-
20. Ascomata associated with reddish rhizomorphs or brown . . . . . . . . . . . . . . . . . . . . . . . Neocosmospora
synnemata, changing to purple in 31. Ascomata subglobose to obpyriform or broadly
KOH . . . . . . . . . . . . . . . . . . . . . . . .Corallomycetella obpyriform, generally red . . . . . . . . . . . . . . . . . . . 32
20. Ascomata not associated with reddish rhizomorphs or 32. Ascomata non-stromatic, obpyriform, with an acute
synnemata, changing or not changing color in KOH, if apex . . . . . . . . . . . . . . . . . . . . . . . . . . . Chaetopsina
change becoming dark red, purple-red, violet or 32. Ascomata seated on an erumpent stroma,
sienna . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21 subglobose to broadly obpyriform, rarely
21. Stromata well-developed and/or byssus covering host, papillate . . . . . . . . . . . . . . . . . Neonectria
rarely reduced . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22 33. Ascomata astromatic, or on a thin stroma or
21. Stromata inconspicuous, absent, or with a basal seated on a subiculum of thick-walled, minutely
stroma, . . . . . . . . . . . . . . . . . . . . . . . 24 warted septate hyphae . . . . . . . . . . . . . . 34
22. Ascomata turning sienna in KOH, ascospores yellow- 33. Ascomata on or nearly or completely immersed in an
brown, coarsely striate, appearing as longitudinal erumpent stroma . . . . . . . . . . . . . . . . . . . . . . Nectria
furrows . . . . . . . . . . . . . . . . . . . . . . . . . . . Calostilbe 34. Ascomata ovoid to elongate-ovoid to cylindrical, asco-
22. Ascomata turning dark red, dark red-brown or violet in spores long-fusiform, often somewhat bent,
KOH, ascospores hyaline to pale yellow-brown, vermiform . . . . . . . . . . . . . . . . . . . . . . Ophionectria
Fungal Diversity

34. Ascomata globose to subglobose to pyriform asco- 8. Phialides cylindrical or narrowly flask-shaped,
spores mostly ellipsoidal . . . . . . . . . . . . . . Fusarium macroconidia cylindrical, curved, (1–)3–4(−5)-septate,
35. Ascomata orange to orange-red to carmine red, becom- with minutely tapered, obtuse ends . . Campylocarpon
ing pink to purple in KOH, ascospores finely 8. Phialides cylindrical to allantoid, conidia cylindrical,
verrucose . . . . . . . . . . . . . . . . . . . . . . Coccinonectria straight or slightly curved, 0–3-septate, rounded at both
35. Ascomata red, becoming dark red in KOH, ascospores ends. . . . . . . . . . . . . . . . . . . . . . . Cylindrocarpostylus
striate . . . . . . . . . . . . . . . . . . . . . . . . . . Sarcopodium 9. Conidiophores elongate doliiform to reniform to
36. Stroma inconspicuous or absent, ascospores obpyriform, conidia do not form slimy heads on the
yellowish . . . . . . . . . . . . . . . . . . . . . . . Macroconia conidiophore . . . . . . . . . . . . . . . . . . Cylindrodendrum
36. With or without basal stroma, ascospores 9. Conidiophores elongate-ampulliform or subcylindrical,
hyaline . . . . . . . . . . . . . . . . . . . . . . . 37 conidia forming slimy heads on the conidiophore . . 10
37. Basal stroma absent, ascospores ellipsoid to fusoid with 10. Forming sterile coils with conidiophores radiating out-
obtuse ends . . . . . . . . . . . . . . . . . . Cylindrocladiella wards, hyphal septa inconspicuously
37. Ascomata on a thin basal stromata, Ascospores fusi- swollen . . . . . . . . . . . . . . . . . . . . . . Paracremonium
form to biconic . . . . . . . . . . . . . . . . . . . . . . Volutella 10. Not forming sterile coils, hyphae inconspicuously
swollen septa. . . . . . . . . . . . . . . . .Xenoacremonium

Niessliaceae Kirschst., Annls mycol. 37(1/2): 89 (1939

Key to the genera of Nectriaceae with asexual morph Facesoffungi number: FoF 01126
Saprobic or parasitic on wood, leaves and stems in fresh-
1. Conidiophores separate or aggregated into sporodochia or water and terrestrial habitats, some possibly fungicolous and
synnemata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 lichenicolous. Sexual morph: Ascomata perithecial or
1. Conidiophores simple, not produce sporodochia. . . . . 6 cleistothecial, yellow, brown, dark brown to black, solitary
2. Conidiophores sporodochial or synnematous, con- or gregarious, superficial, erumpent to immersed in a
sist of fertile branches and sterile vesiculate stipe subiculum or crustose stroma, cupulate, globose to
extensions, generally not pathogenic to other fungi subglobose, membranaceous, tuberculate, with or without cir-
or insects . . . . . . . . . . . . . . . . . . . . . . . . 3 cinately coiled, lightly pigmented, brown, septate, apical se-
2. Conidiophores simple sporodochial, consist of tae, collabent, or collapsing laterally, or not collapsing, papilla
conidiogenous ring only, sterile stipe extensions absent, present or lacking, when present with periphysate ostiole.
generally pathogenic to other fungi or insects . . . . . . 5 Peridium thick, membranaceous, outer layer composed of yel-
3. Conidia (0–)1–5-septate, clavate, obovoid or gently low, light brown or brown cells of textura angularis; inner
curved, rarely ellipsoidal . . . . . . . . . . . . . . . Atractium layer composed of hyaline cells of textura prismatica.
3. Conidia 0–3-septate, cylindrical to fusiform . . . . . . . . 4 Hamathecium of filiform, hyaline, septate paraphyses, or pa-
4. Conidia 0–3-septate, cylindrical, stipe with a raphyses absent. Asci 8-spored, unitunicate, oblong to clavate,
verruculose, pale brown apical cell . . . . Curvicladiella long or short pedicellate, with narrowly discoid J- apical ring
4. Conidia aseptate, cylindrical to fusiform, stipe smooth, or absent, evanescent at maturity. Ascospores 1–3-seriate, hya-
hyaline. . . . . . . . . . . . . . . . . . . . . . Xenogliocladiopsis line to brown, ellipsoid to fusiform, slightly curved, aseptate to
5. Microconidia absent or present, 0–1(−2)-septate; 1-septate, striate or verrucose, smooth-walled. Asexual
macroconidia uniseptate to multi-septate, falcate, with morph: Hyphomycetous. Mycelium white to light brown, su-
or without a hooked apical cell . . . . . . . . . . Fusarium perficial, effused, thick subiculum. Conidiophores thin-walled,
5. Microconidia 0(−1)-septate, ellipsoidal and straight or hyaline, unbranched, aseptate, swollen, with a short and distinct
allantoid, broadly lunate to reniform or curved and ta- collarette, teminating in a single conidium. Phialides straight,
pering at both ends; macroconidia (0–)1–2(−3)-septate, smooth. Conidia oval to ellipsoid, aseptate to 1-septate, hya-
curved to lunate . . . . . . . . . . . . . . . . . . . .Bisifusarium line, smooth. Coelomycetous, Conidiomata sub-immersed,
6. Conidiophores/Conidiogenous cells bearing 1 to several globose to subglobose, dark brown. Peridium thick-walled,
phialides. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 wall composed of brown to hyaline cells of textura angularis,
6. Conidiogenous cells monophialidic . . . . . . . . . . . . . . 9 becoming hyaline towards inner conidiogenous region.
7. Conidiophores arising from pseudoparenchymatous Conidiophores hyaline, subcylindrical, branched apically,
cells in a basal stroma, phialides elongate doliiform to 1–2-septate. Conidiogenous cells phialidic, hyaline, fusiform
reniform or subcylindrical . . . . . . . . . Dematiocladium to ellipsoid, straight to curved, tapering towards a subtruncate
7. Conidiophores as lateral phialidic pegs or arising later- apex. Conidia oval to ellipsoid, aseptate to 2-septate, hyaline,
ally from somatic hyphae, phialides cylindrical to with guttulate, smooth-walled (Descriptions based on Samuels
allantoid or narrowly flask-shaped . . . . . . . . . . . . . . . 8 and Barr 1997, Crous et al. 2007b).
 Fungal Diversity

Type: Niesslia Auersw. Other genera included

Notes: The family Niessliaceae was introduced by
Kirschstein (1939) based on superficial, dark, setose peri- Atronectria Etayo, in Etayo & Rosato, Biblthca Lichenol. 98:
thecial ascomata, and is typified by Niesslia. Barr (1990) 52 (2008)
proposed that the family be accepted in a narrow sense Type species: Atronectria magellanica Etayo, in Etayo &
and removed it from the Sphaeriaceae (Müller and von Sancho, Biblthca Lichenol. 98: 52 (2008)
Arx 1962, 1973) and Trichosphaeriaceae (Hawksworth Circinoniesslia Samuels & M.E. Barr, Can. J. Bot. 75(12):
et al. 1983; Barr 1983). Samuels and Barr (1997) referred 2166 (1998) [1997]; Fig. 100
the family to the order Hypocreales, based on the short Type species: Circinoniesslia nectriae Samuels & M.E.
paraphyses, periphysate ostiole, dark pigmented peridium Barr, Can. J. Bot. 75(12): 2166 (1998) [1997]
and phialidic conidiogenesis cells. Eriksson and Cryptoniesslia Scheuer, Mycol. Res. 97(5): 543 (1993)
Hawksworth (1993) and Hawksworth et al. (1995) accept- Type species: Cryptoniesslia setulosa Scheuer, Mycol.
ed its disposition in the Hypocreales based on morpholog- Res. 97(5): 543 (1993)
ical features. Species are generally saprobic or parasitic Eucasphaeria Crous, in Crous et al., Fungal Divers 25: 21
on wood, leaves or lichens. Lumbsch and Huhndorf (2007b)
(2010) accepted 17 genera in the family. The placement Type species: Eucasphaeria capensis Crous, in Crous
of Niessliaceae in the Hypocreales is supported by mo- et al., Fungal Divers 25: 21 (2007b)
lecular data with Jaklitsch and Voglmayr (2012) showing Hyaloseta A.W. Ramaley, Mycotaxon 79: 269 (2001)
Niesslia exilis (Alb. & Schwein.) G. Winter, grouping Type species: Hyaloseta nolinae A.W. Ramaley,
with Eucasphaeria capensis Crous with high support Mycotaxon 79: 269 (2001)
and the new genus Rosasphaeria in a sister group. Malmeomyces Starbäck, Bih. K. svenska Vetensk Akad.
Valetoniellopsis was transfered to Hypocreales genera Handl., Afd. 3 25(no. 1): 32 (1899)
incertae sedis (Maharachchikumbura et al. 2015). Type species: Malmeomyces pulchellus Starbäck [as
Eighteen genera were recognized in Niessliaceae by ‘pulchella’], Bih. K. svenska Vetensk Akad. Handl., Afd. 3
Maharachchikumbura et al. (2015). 25(no. 1): 32 (1899)
Niesslia Auersw., in Gonnermann & Rabenhorst, Myc. Melanopsamma Niessl, Verh. nat. Ver. Brünn 14: 200 (1876)
Europ. Pyren. 5–6: 30 (1869) Type species: Melanopsamma pomiformis (Pers.) Sacc.,
Facesoffungi number: FoF 01127; Figs 99, 100, 101 Michelia 1(no. 3): 347 (1878)
Saprobic on leaves in terrestrial habitats. Sexual Melchioria Penz. & Sacc., Malpighia 11(9–10): 399
morph: Ascomata perithecial, solitary, superficial, (1897)
cupuliform, dark brown to black, membranaceous, tuber- Type species: Melchioria leucomelaena Penz. & Sacc.,
culate, collabent when dry, surround by brown, septate Malpighia 11(9–10): 399 (1897)
setae, with spiny termination, with periphysate ostiole. Miyakeomyces Hara, Bot. Mag., Tokyo 27(no. 317): (248)
Peridium thick, outer layer hard, composed of dark tissues, (1913)
basal part mixed with host cells; inner layer composed of Type species: Miyakeomyces bambusae Hara [as
brown to dark brown cells of textura angularis. ‘Miyakeamyces’], Bot. Mag., Tokyo 27(no. 317): (248) (1913)
Hamathecium of filiform, hyaline, septate paraphyses or Myrmaeciella Lindau, in Engler & Prantl, Nat.
paraphyses absent. Asci 8-spored, unitunicate, oblong to Pflanzenfam., Teil. I (Leipzig) 1: 478 (1897)
clavate, stipitate, apical ring present or absent, evanescent Type species: Myrmaeciella endoleuca (Sacc.) Lindau, in
at maturity. Ascospores ellipsoid to fusiform, aseptate to 1- Engler et al., Teil. I (Leipzig) 1(1): 478 (1897)
septate, hyaline, slightly curved, smooth, mostly with Paraniesslia K.M. Tsui et al., Mycologia 93(5): 1002 (2001)
guttules. Asexual morph: Undetermined. Type species: Paraniesslia tuberculata K.M. Tsui et al.,
Type species: Niesslia chaetomium (Corda) Auersw., in Mycologia 93(5): 1002 (2001)
Gonnermann & Rabenhorst, Myc. Europ. Pyren. 5–6: 30 Pseudonectriella Petr., Sydowia 13(1–6): 127 (1959)
(1869) Type species: Pseudonectriella ahmadii Petr., Sydowia
Current name: 13(1–6): 128 (1959)
Niesslia exosporioides (Desm.) G. Winter, Rabenh. Krypt.- Pseudorhynchia Höhn., Sber. Akad. Wiss. Wien, Math.-
Fl., Edn 2 (Leipzig) 1.2: 197 (1885) naturw. Kl., Abt. 1 118: 1206 [50 repr.] (1909)
Notes: The genus Niesslia was introduced by Auerswald Type species: Pseudorhynchia polyrhyncha (Penz. &
and includes 39 epithets according to Index Fungorum (2016). Sacc.) Höhn. [as ‘Pseudorhynchia polyrrhyncha’],
The type species, is characterized by tuberculate perithecia, (1909)
surrounded by brown, septate setae, clavate asci and filiform Rosasphaeria Jaklitsch & Voglmayr, Fungal Diversity
ascospores. 52(1): 93 (2012)
Fungal Diversity

Fig. 99 Niesslia exosporioides

(Material examined: GERMANY,
Hainburg, Hundsheimer Kogel,
Lower Danube, on dead leaves of
Carex pilosa Scop. (Cyperaceae),
Petrak F, May 1940, PRM
717659) a Material label b
Material c, d Ascoma of the side
view d Ascoma e Peridium f
Septate seta g-j Ascospores.
Notes: Figs e-j soaked in 3 %
KOH. Scale bars: c–d = 200 μm,
e-f = 10 μm, g–j = 5 μm

Type species: Rosasphaeria moravica (Petr.) Jaklitsch & Valetoniella Höhn., Sber. Akad. Wiss. Wien, Math.-
Voglmayr, Fungal Divers 52(1): 93 (2012) naturw. Kl., Abt. 1 118: 1499 [39 repr.] (1909)
Taiwanascus Sivan. & H.S. Chang, Mycol. Res. 101(2): Type species: Valetoniella crucipila Höhn., Sber. Akad. Wiss.
176 (1997) Wien, Math.-naturw. Kl., Abt. 1 118: 1500 [40 repr.] (1909)
Type species: Taiwanascus tetrasporus Sivan. & H.S.
Chang, Mycol. Res. 101(2): 176 (1997)
Trichosphaerella E. Bommer et al., Syll. fung. (Abellini) Key to genera of Niessliaceae
9: 604 (1891)
Type species: Trichosphaerella decipiens E. Bommer 1. Lichenicolous . . . . . . . . . . . . . . . . . . . . . . . . . Atronectria
et al., in Saccardo, Syll. fung. (Abellini) 9: 604 (1891) 1. Not lichenicolous . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
 Fungal Diversity

Fig. 100 Niesslia exosporioides

(redrwn from a description by
Saccardo 1882) a Ascoma b
Ascoma in cross section c Asci.
Scale bars: a–b = 100 μm,
c = 10 μm

2. Fungicolous . . . . . . . . . . . . . . . . . . . . . Circinoniesslia 9. Ascomata carbonaceous . . . . . . . . . . . . . . . Melchioria

2. Saprobic or parasitic on wood, leaves or stems. . . . . . 3 9. Ascomata not carbonaceous . . . . . . . . . . . . . . . . . . . 10
3. Ascomata sitting on a subiculum . . . . . . . . . . . . . . . . 4 10. Parasitic on leaves . . . . . . . . . . . . . . . . Eucasphaeria
3. Ascomata sitting on the substrate . . . . . . . . . . . . . . . . 7 10. Saprobic on wood . . . . . . . . . . . . . . . . . . . . . . . . . 12
4. Ascomata sitting on a thick subiculum, . . . . . . . . . . . 5 10. Saprobic on leaves . . . . . . . . . . . . . . . . . . . . . . . . . 11
4. Ascomata sitting on a thin subiculum. . . . . . . . . . . . . 6 11. Ascomata immersed . . . . . . . . . . . . . . .Cryptoniesslia
5. Bambusicolous, ascomata sitting on a black 11. Ascomata superficial, setae elongate. . . . . . . .Niesslia
subiculum. . . . . . . . . . . . . . . . . . . . . . Miyakeomyces 12. Ascomata erumpent, yellow, papillate orange to
5. S a p r o b i c o n s t e m s , p a l m m i d r i b s , w o o d , black . . . . . . . . . . . . . . . . . . . . . . . . . . Myrmaeciella
ascomata sitting on white to brown hyphal 12. Ascomata superficial . . . . . . . . . . . . . . . . . . . . . . . 13
subiculum, with hyphomycetous asexual 12. Ascomata immersed, with hyphomycetous asexual
morph . . . . . . . . . . . . . . . Trichosphaerella morphs . . . . . . . . . . . . . . . . . . . . . . . . Rosasphaeria
6. Ascomata sitting on the hyaline hyphal subiculum, asex- 13. Ascomata without setae . . . . . . . . . . . . . . . . . . . . . 14
ual morph Monocillium . . . . . . . . . . . . . . . . Hyaloseta 13. Ascomata with setae. . . . . . . . . . . . . . . . . . . . . . . . 15
6. Ascomata sitting on light brown hyphal subiculum, 14. Ascomata globose, smooth or bearing
asexul morph undetermined. . . . . . . . Pseudorhynchia conidiophores . . . . . . . . . . . . . . . . Melanopsamma
7. Bambusicolous. . . . . . . . . . . . . . . . . . . .Malmeomyces 14. Ascostroma cupulate . . . . . . . . . . . . Pseudonectriella
7. Saprobic or parasitic on wood, leaves and stems . . . . 8 15. Ascomata cleistothecial . . . . . . . . . . . . .Taiwanascus
8. Ascomata on wood submerged in freshwater 15. Ascomata perithecial . . . . . . . . . . . . . . . .Valetoniella
habitats. . . . . . . . . . . . . . . . . . . . . . . . . . Paraniesslia
8. Ascomata on wood, leaves and stems in terrestrial Nitschkiaceae Nannf., Nova Acta R. Soc. Scient. upsal., Ser.
habitats . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 4 8(no. 2): 56 (1932
Fungal Diversity
 Fungal Diversity

ƒFig. 101 Circinoniesslia nectriae (Material examined: PUERTO RICO, Facesoffungi number: FoF 02123; Fig. 102
Caribbean National Forest, Luquillo Mts., El Verde Research Area, alt. Saprobic on wood in freshwater and terrestrial habitats.
350 m; Bionectria sp., Samuels, G. J. Schroers, H.-J. (8028), on decaying,
February, 1996, BPI 748359, holotype) a Material label b Material c
Sexual morph: Ascomata perithecial, black, gregarious, su-
Ascomata d Ascoma e Vertical section of ascoma f Peridium g–i Asci perficial or erumpent, sitting on or in a subiculum or absent,
j–m Ascospores. Note: Figures h, i and k are stained in Melzer’s reagent. globose to subglobose, carbonaceous or membranaceous,
Scale bars: c = 500 μm, d–e = 200 μm, f = 50 μm, g–i = 20 μm, j– rough, with a short ostiole, the apex collapsing when dry,
m = 10 μm
shallowly cupulate when moist or not collabent. Ostiole lined
with hyaline periphyses. Subiculum thin or thick, numerous or
Facesoffungi number: FoF 01399 scanty, brown to dark brown, septate, branched hyphae, with
Saprobic or parasitic on wood, leaves or lichens, in freshwa- spiny termination. Peridium (20–70 μm), munk pores present,
ter, marine and terrestrial habitats. Sexual morph: Ascomata composed of three layers, outer layer comprising dark tissues,
perithecial, dark brown to black, gregarious, superficial to thin, carbonaceous; middle layer comprising dark brown to
erumpent, sitting on or in a subiculum or absent, cupuliform, brown cells of textura angularis, thick, membranaceous; the
globose to subglobose, carbonaceous, coriaceous or inner layer has hyaline cells of textura prismatica, thin,
membranaceous, rough, tuberculate, smooth or with short membranaceous. Hamathecium comprising septate, simple
spines, with or without an ostiole, the apex collapsing when paraphyses,. Asci 8-spored, unitunicate, clavate to cylindrical,
dry or shallowly cupulate when moist, periphyses inside of the long pedicellate, apex blunt. Ascospores 2–3-seriate or irreg-
ostiole or absent. Subiculum thin or thick, numerous or scanty, ularly arranged, hyaline, allantoid or subcylindrical, slightly
brown to dark brown, septate, branched hyphae, with spiny curved, aseptate, smooth-walled, with or without guttules.
termination. Peridium thick (20–150 μm), munk pores present Asexual morph: Undetermined.
or absent, composed of three layers, outer layer comprising dark Type species: Nitschkia cupularis (Pers.) P. Karst., Bidr.
tissues, thin, carbonaceous; middle layer comprising dark brown Känn. Finl. Nat. Folk 23: 81 (1873)
to brown cells of textura angularis, thick, membranaceous, Notes: The taxonomy of the genus Nitschkia introduced by
while the inner layer has hyaline cells of textura prismatica, thin, G.H. Otth ex Karsten (1873) is confused, and currently in-
and membranaceous. Hamathecium with or without hyaline cludes 65 epithets (Index Fungorum 2016). The name
paraphyses, septate, simple or branched. Asci 4- to multi-spored, Nitschkia fuckelii was first introduced by Fuckel (1870), but
unitunicate, thin-walled, clavate to cylindrical, long or short Saccardo (1873) suggested Coelosphaeria to replace
pedicellate or sessile, blunt. Ascospores 2–3-seriate or irregular- Nitschkia, because the spelling of “Nitschkia” was similar to
ly arranged, hyaline or brown, fusiform, allantoid or the generic name for a diatom. Fitzpatrick (1923) reviewed the
subcylindrical, slightly curved, 0–4-aseptate, concolourous, nomenclature of Coelosphaeria and the name was rejected.
smooth-walled, sheath and appendages present or absent, with Confusion as to the type species arises because Fuckel (1870)
or without guttules. Asexual morph: Undetermined. introduced the type species N. fuckelii based on Sphaeria
Type: Nitschkia G.H. Otth ex P. Karst. cupularis Fr. These two taxa, N. fuckelii and N. cupularis,
Notes: The family Nitschkiaceae (Coronophorales) was differ morphologically with respect to the production of a
introduced by Nannfeldt (1932) based on erumpent, black, definite pseudoparenchymatous stroma/ subiculum, respec-
rough ascomata and clavate asci, with hyaline allantoid asco- tively. Neither taxon has been sequenced, and molecular anal-
spores, and is typified by Nitschkia fuckelii G.H. Otth ex P. ysis is based on Nitschhia calyculus (Mont.) Kuntze,
Karst. (Karsten 1873, based on Coelosphaeria fuckelii). N. grevillei (Rehm) Nannf., N. meniscoidea Huhndorf et al.
Nannfeldt (1975) concluded that Nitschkiaceae comprised and N. tetraspora Nannf. (Mugambi and Huhndorf 2010).
three separate groups, but the differences between them were
not sufficient to place them in separate families. He accepted
five genera in this family and provided a detailed morphology Other genera included
description. Mugambi and Huhndorf (2010) accepted twelve
genera based on a multi-gene dataset and a morphological Acanthonitschkea Speg., Anal. Mus. nac. B. Aires, Ser. 3
study (Lumbsch and Huhndorf 2010). The monotypic genera 17(10): 116 (1908)
Neochaetosphaerella and Tortulomyces were introduced Type species: Acanthonitschkea argentinensis Speg.,
based on morphological observations (Vasilyeva et al. 2012, Anal. Mus. nac. B. Aires, Ser. 3 17(10): 116 (1908)
2013). In our study, 14 genera are included in this family. Biciliosporina Subram. & Sekar, Kavaka 18(1–2): 69
Most taxa assigned to the family need to be evaluated at the (1993) [1990]
molecular level. Type species: Biciliosporina karwarensis Subram. &
Sekar, Kavaka 18(1–2): 69 (1993) [1990]
Nitschkia G.H. Otth ex P. Karst., Bidr. Känn. Finl. Nat. Folk Botryola Bat. & J.L. Bezerra, in Batista et al., Publicações
23: 13 (1873) Inst. Micol. Recife 431: 11 (1964)
Fungal Diversity

Fig. 102 Nitschkia cupularis (Material examined: on the branches of Ascospores. Notes: Figs e-g. i-o are stained in KOH. Scale bars:
Prunus mahaleb L. (Rosaceae) and Carpinus betulus L. (Betulaceae), BR c = 500 μm, e = 100 μm, f = 40 μm, g = 30 μm, d. h–k = 10 μm, l–
5020096960571) a Material label b Material c Ascomata d Munk pores e o = 5 μm
Ascomata in cross section f Peridium g Paraphyses h–k Asci l–o
 Fungal Diversity

Type species: Botryola tetrasperma Bat. & J.L. Bezerra, in 4. Ascomata superficial, ascospores
Batista et al., Publicações Inst. Micol. Recife 431: 13 (1964) cylindrical. . . . . . . . . . . . . . . . . . . Rhagadostomella
Fracchiaea Sacc., Atti Soc. Veneto-Trent. Sci. Nat. 2: 163 4. Ascomata sub-immersed, ascospores allantoid, slightly
(1873) curved . . . . . . . . . . . . . . . . . . . . . . . . . Rhagadostoma
Type species: Fracchiaea heterogenea Sacc., Atti Soc. 5. Ascospores brown . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
Veneto-Trent. Sci. Nat. 2: 163 (1873) 5. Ascospores hyaline . . . . . . . . . . . . . . . . . . . . . . . . . . 7
Groenhiella Jørg. Koch et al., Bot. Mar. 26(6): 265 (1983) 6. Lichenicolous, ascomata superficial . . .Lasiosphaeriopsis
Type species: Groenhiella bivestia Jørg. Koch et al., Bot. 6. Corticolous, ascomata sub-immersed . . . . . . . . . . . . . 8
Mar. 26(6): 265 (1983) 7. Ascomata with setae . . . . . . . . . . . . . . . . . . . . . . . . . 9
Janannfeldtia Subram. & Sekar, Kavaka 18(1–2): 69 7. Ascomata without setae . . . . . . . . . . . . . . . . . . . . . . 11
(1993) [1990] 8. Ascomata ostiolate, with spiny septate
Type species: Janannfeldtia karwarensis Subram. & setae . . . . . . . . . . . . . . . . . . . . Neochaetosphaerella
Sekar, Kavaka 18(1–2): 70 (1993) [1990] 8. Ascomata lacking ostiole, with
Lasiosphaeriopsis D. Hawksw. & Sivan., in Hawksworth, tubercules . . . . . . . . . . . . . . . Tortulomyces
Trans. Br. mycol. Soc. 74(2): 371 (1980) 9. Asci multi-spored. . . . . . . . . . . . . . . . . . . . . . . . . . . 10
Type species: Lasiosphaeriopsis salisburyi D. Hawksw. & 9. Asci more than 8-spored . . . . . . . . .Acanthonitschkea
Sivan., in Hawksworth, Trans. Br. mycol. Soc. 74(2): 373 10. Quellkörper absent . . . . . . . . . . . . . . . . . . Fracchiaea
(1980) 10. Quellkörper present . . . . . . . . . . . . . . . . Neotrotteria
Loranitschkia Lar. N. Vassiljeva, Mikol. Fitopatol. 24(3): 11. Ascomata immersed and erumpent through the
207 (1990) bark . . . . . . . . . . . . . . . . . . . . . . . . . 12
Type species: Loranitschkia viticola Lar. N. Vassiljeva [as 11. Ascomata superficial . . . . . . . . . . . . . . Loranitschkia
‘Loranitschki’], Mikol. Fitopatol. 24(3): 207 (1990) 12. Quellkörper absent . . . . . . . . . . . . . . . . . . . Nitschkia
Neochaetosphaerella Lar. N. Vassiljeva et al., in Vasilyeva 12. Quellkörper present . . . . . . . . . . . . . . . . . . . . . . . . 13
et al., Fungal Diversity 52(1): 192 (2012) 13. Asci 8-spored, irregularly uniseriate or overlapping
Type species: Neochaetosphaerella thaxteriospora Lar. N. biseriate. . . . . . . . . . . . . . . . . . . . . . . . Biciliosporina
Vassiljeva et al., in Vasilyeva et al., Fungal Diversity 52(1): 13. Asci more than 8-spored, not regularly
192 (2012) arranged . . . . . . . . . . . . . . . . . . . . . . Janannfeldtia
Neotrotteria Sacc., Bulletino dell’orto Botanico della R.
Universitá di Napoli 6: 45 (1921) Ophioceraceae Klaubauf et al., Studies in Mycology 79: 85–
Type species: Neotrotteria pulchella Sacc., Bulletino 120 (2014).
dell’orto Botanico della R. Universitá di Napoli 6: 45 (1918) Facesoffungi number: FoF 01254
Rhagadostoma Körb., Parerga lichenol. (Breslau) 5: 472 Saprobic on wood and other plant material, commonly iso-
(1865) lated in aquatic habitats. Sexual morph: Ascomata perithe-
Type species: Rhagadostoma corrugatum Körb., Parerga cial, solitary or in small groups, immersed or semi-immersed,
lichenol. (Breslau) 5: 473 (1865) dark brown to black, globose to subglobose, coriaceous,
Rhagadostomella Etayo, Biblthca Lichenol. 84: 109 ostiolate. Ostiole a long neck, lined with periphyses.
(2002) Hamathecium comprising numerous, septate, tapering, hy-
Type species: Rhagadostoma corrugatum Körb., Parerga phae-like, thin-walled, filamentous, branched paraphyses,
lichenol. (Breslau) 5: 473 (1865) intermingled among asci. Peridium outwardly comprising
Tortulomyces Lar. N. Vassiljeva et al., Mycoscience 54(1): small angular dark brown cells, inwardly comprising several
110–115 (2013) layers of dark brown to pale brown cells of textura angularis.
Type species: Tortulomyces thailandicus Lar. N. Vassiljeva Asci 8-spored, unitunicate, subcylindrical to narrowly fusoid,
et al., Mycoscience 54(1): 110–115 (2013) apedicellate, with thimble-shaped, J-, apical ring. Ascospores
fasciculate, hyaline, yellowish in mass, filiform, bluntly
rounded at the apices, 3–or multi-septate. Asexual morph:
Key to genera of Nitschkiaceae Undetermined.
Type: Ophioceras Sacc.
1. In marine environments . . . . . . . . . . . . . . . . . . . . . . . . . 2 Notes: The family Ophioceraceae was introduced as a
1. In freshwater and terrestrial environments . . . . . . . . . 3 monotypic family by Klaubauf et al. (2014) in the order
2. Ascospores hyaline . . . . . . . . . . . . . . . . . . . . Botryola Magnaporthales to accommodate Ophioceras. Previously,
2. Ascospores light brown with sheath . . . . . Groenhiella Ophioceras was placed in Magnaporthaceae following the
3. Lichenicolous. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 phylogenetic analysis of Chen et al. (1999) and Inderbitzin
3. Saprobic on wood . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 and Berbee (2001). However, based on combined LSU and
Fungal Diversity

RPB1 analysis, it was found that Ophioceras clusters sepa- Purpureocillium. Most species in this family produce dark
rately from Magnaporthaceae within the order pigmented, tough to pliant stromata, that often possess
Magnaporthales (Thongkantha et al. 2009) and hence a new aperithecial apices (Sung et al. 2007). The main distinguishing
monotypic family Ophioceraceae was introduced to accom- characters of this genus are that the ascospores do not break
modate Ophioceras (Klaubauf et al. 2014). into part-spores at maturity and asci have thin apical caps
(Petch 1931a, b). The type species is Ophiocordyceps blattae
Ophioceras Sacc., Syll. fung. (Abellini) 2: 358 (1883); (Petch) Petch. This taxon was collected from Sri Lanka with
Fig. 103 only one collection and without cultural or molecular data.
Type species: Ophioceras dolichostomum (Berk. & M.A. Asexual generic names associated with Ophiocordyceps in-
Curtis) Sacc., Syll. fung. (Abellini) 2: 358 (1883) clude Sorosporella (Sorokin 1888), Hirsutella (Doassans
Notes: Saccardo (1883) introduced Ophioceras based on and Patouillard 1892; Gams and Zare 2003), Hymenostilbe
O. dolichostomum (Berk. & M.A. Curtis) Sacc., characterized (Petch 1931b), Stilbella (Seifert 1985), Syngliocladium
by black ascomata with long necks, cylindrical asci with (Petch 1932b), and Paraisaria (Samson and Brady 1983).
small, refractive, apical rings and filiform ascospores Based on the morphology and sequence data,
(Teng 1934; Conway and Barr 1977; Shearer et al. Ophiocordyceps was protected and other names suppressed
1999; Tsui et al. 2001; Thongkantha et al. 2009; in accordance with “one fungus one name” (Quandt et al.
Klaubauf et al. 2014). Ophioceras species are common- 2014).
ly encountered on decaying woody substrates in fresh- Asexual morphs of the family are Tolypocladium sensu
water habitats all over the world (Hyde 1992; Hyde and Gams (1971), Chaunopycnis sensu Gams (1980), or
Goh 1998; Shearer et al. 1999; Tsui et al. 2001; verticillium-like (Quandt et al. 2014). Many asexual morphs
Thongkantha et al. 2009). Currently 39 records of are associated with this family, some of which are restricted in
Ophioceras are listed in Index Fungorum (2016). their phylogenetic distribution, while others are often found in
Morphologically Ophioceras is reminiscent of disparate lineages (Quandt et al. 2014). For example,
Gaeumannomyces, however, the two genera can be distin- Verticillium is a common asexual morph of many species in
guished easily by the differences in their habitats. several hypocrealean families, including
Ophioceras is usually associated with wood and herbaceous Ophiocordycipitaceae, Cordycipitaceae and Clavicipitaceae
material in aquatic habits, while Gaeumannomyces species are (see Zare et al. 2000; Sung et al. 2001, 2007; Gams and
plant pathogens (Chen et al. 1999; Klaubauf et al. 2014). Zare 2001). Ophiocordycipitaceae are also associated
with Hymenostilbe and Hirsutella asexual morphs
Ophiocordycipitaceae G.H. Sung et al., in Sung et al., Stud. (Kepler et al. 2013).
Mycol. 57: 35 (2007) Tolypocladium was proposed by Quandt et al. (2014) for
Facesoffungi number: FoF 01315 protection over Elaphocordyceps (Sung et al. 2007) and
Parasitic on arthropods, protozoans, rotifers, nematodes, Chaunopycnis (Gams 1980). This genus forms a single mono-
humans, animals and fungi worldwide. Sexual morph: phyletic clade that includes truffle parasites, several insect
Stromata or subiculum darkly pigmented or rarely brightly pathogens, rotifer pathogens, mycoparasitic fungi, and general
coloured, tough, fibrous, pliant to wiry, rarely fleshy, often soil-inhabitants (Quandt et al. 2014). This genus is character-
with aperithecial apices or lateral pads. Ascomata superficial ized by robust stipitate stroma with clavate to capitate to high-
to completely immersed, ordinal or oblique in arrangement. ly reduced stroma, comprising rhizomorphs and aggregated
Asci unitunicate, cylindrical, rarely fusoid to ellipsoid, usually ascomata, which may be immersed and produced on highly
with thickened and almost bulbous ascus apex. Ascospores reduced stromatic pads (Quandt et al. 2014). Quandt et al.
fasciculate, usually filiform, hyaline, multi-septate, (2014) list 26 new combinations within this genus.
disarticulating into part-spores or non-disarticulating. Purpureocillium was described as a genus by Luangsa-ard
Asexual morph: see notes. et al. (2011) for the medically important Paecilomyces
Type: Ophiocordyceps Petch lilacinus (Thom) Samson. There are only two species in this
Notes: The family Ophiocordycipitaceae was introduced genus: P. lilacinum (Thom) Samson (Luangsa-ard et al. 2011)
by Sung et al. (2007) based on phylogenetic analyses and later and P. lavendulum (Thom) Samson (Perdomo et al. 2013a).
emended by Quandt et al. (2014). Quandt et al. (2014) includ-
ed Ophiocordyceps plus six genera within Fig. 103 Ophioceras sp. (Material examined: THAILAND, Chiang Rai,„
Ophiocordycipitaceae based on morphological and Tham Luang Nang Non Cave, on submerged wood, 25 November 2014,
phylogenetic analyses. Spatafora et al. (2015) confirmed this Jing Yang, YJ-11 (MFLU 15–0378) a Substrate b Colonies c Fruiting
body d Section of ascoma e Peridium f Paraphyses g–l Asci m–o
classification system and introduced necessary species com- Ascospores p Apical ring of ascus q Germinating ascospore r–s
binations into four genera of this family, including Colonies on PDA (S from below). Scale bars: b = 500 μm, c = 200 μm,
Drechmeria, Harposporium, Ophiocordyceps and d = 100 μm, e = 15 μm, f, j–l, q = 30 μm, g–I, m–p = 25 μm
Fungal Diversity
Fungal Diversity

According to phylogenetic analyses, the nematophagous name Hirsutella (Evans and Samson 1982). According
clade separated in two clades; one containing Harposporium to morphological analyses and economic value of the
(Lohde 1874) and Podocrella (Seaver 1928a), and the other genus, Ophiocordyceps should be protected over the
comprising Drechmeria (Gams and Jansson 1985), other names.
Haptocillium (Gams and Zare 2001), and Cordyceps gunnii
(Berk.) Berk (Quandt et al. 2014). Harposporium was record-
ed as the asexual morph of Podocrella based on morpholog-
ical and phylogenetic analyses (Chaverri et al. 2005). Other genera included
Harposporium is an older name, and the morphology of at
least somewhat crescent-shaped conidia is a shared character Drechmeria W. Gams & H.-B. Jansson, Mycotaxon 22(1): 36
for this clade (Quandt et al. 2014). Harposporium was (1985)
protected over Podocrella as the earlier genus name. For the Type species: Drechmeria coniospora (Drechsler) W.
other clade, Drechmeria is an older name than Haptocillium. Gams & H.-B. Jansson, Mycotaxon 22(1): 37 (1985)
Drechmeria was protected as the genus name, which includes Harposporium Lohde, Tagbl. Versamml. Ges. Deutsch.
Cordyceps gunni (Berk.) Berk (Quandt et al. 2014). Naturf. 47: 206 (1874)
Polycephalomyces was introduced by Kobayasi (1941), Type species: Harposporiuman guillulae Lohde, Tagebl.
accepted as circumscribed by Seifert (1985) and emended by Versamm dt. naturf. Ärzte (Breslau) 47: 203–206 (1874)
Kepler et al. (2013). Polycephalomyces is an incertae sedis Polycephalomyces Kobayasi, Sci. Rep. Tokyo Bunrika
genus currently within Hypocreales, as its placement has Daig., Sect. B 5: 245 (1941)
lacked support in previous phylogenetic studies (Kepler Type species: Polycephalomyces formosus Kobayasi, Sci.
et al. 2013). Many morphological characters are shared be- Rep. Tokyo Bunrika Daig., Sect. B 5: 245 (1941)
tween Ophiocordycipitaceae and Polycephalomyces as nu- Purpureocillium Luangsa-ard et al., in Luangsa-ard et al.,
merous species produce hirsutella-like anamorphs with co- FEMS Microbiol. Lett. 321(2): 144 (2011)
nidia often borne in a slimy mass (Seifert 1985), sexual spores Type species: Purpureocillium lilacinum (Thom)
of Polycephalomyces often possess a wiry, tough, carbona- Luangsa-ard et al., in Luangsa-ard et al., FEMS Microbiol.
ceous stipe, which is a common morphology of Lett. 321(2): 144 (2011)
Ophiocordyceps (Kepler et al. 2013). The asexual morph in- Tolypocladium W. Gams, Persoonia 6(2): 185 (1971)
cludes acremonium-like, hirsutella-like and Type species: Tolypocladiumin flatum W. Gams,
Polycephalomyces species (Kepler et al. 2013). Persoonia 6(2): 185 (1971)
Ophiocordyceps Petch, Trans. Br. mycol. Soc.16(1): 73
(1931)
Facesoffungi number: FoF 01318; Fig. 104
Key to genera of Ophiocordycipitaceae
Parasitic on arthropods, worldwide. Sexual morph:
Stromata or subiculum darkly pigmented or rarely brightly
1. Sexual morph . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
coloured, tough, fibrous, pliant to wiry, rarely fleshy,
1. Asexual morph. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
often with aperithecial apices or lateral pads. Ascomata
2. Ascospores breaking into part spores . . . . . . . . . . . . . 3
superficial to completely immersed, ordinal or oblique
2. Ascospores not breaking into part
in arrangement. Asci cylindrical, 8-spored, rarely fusoid
spores . . . . . . . . . . . . . . . . . . . . . . . Ophiocordyceps
to ellipsoid, usually with thickened ascus apex.
3. Ascospores breaking into many small part spores. . . . 4
Ascospores fasciculate, hyaline usually cylindrical,
3. Ascospores breaking into two lanceolate multi-septate
multi-septate, disarticulating into part-spores or non-
part spores . . . . . . . . . . . . . . . . . . . . . . Harposporium
disarticulating.
4. Stromata multi-furcate, stromata apex
Type species: Ophiocordyceps blattae (Petch) Petch, Trans.
infertile . . . . . . . . . . . . . . . . . . . . Polycephalomyces
Br. mycol. Soc.16(1): 74 (1931)
4. Stromata with single stalk, apex fertile . . . . . . . . . . . . 5
Notes: This genus was introduced by Petch (1931a) and
used by Sung et al. (2007) as the type genus of
Fig. 104 Ophiocordyceps nutans (Material examined: CHINA.„
Ophiocordycipitaceae. Quandt et al. (2014) emended Province of Jiang-Xi, on dead stinkbug. 18 June 2014, Yuan-Pin Xiao
this genus based on morphological and phylogenetic JGS 2014061810 (MFU 15-3201). a Stromata growing out from leaf
analyses. According to phylogenetic analyses, the oldest litter b Overview of stromata and the host. c The infected host d Cross
sections showing the complete stromata and perithecia e, f Sections of
name in Ophiocordycipitaceae clade is Sorosporella, a
ascomata g Part of perithecia h-i Asci n Part of ascospore. j-n Asci stained
synonym of Syngliocladium. Sorosporella was sup- in methylene blue o Asci with apical cap p Asci stained in methylene blue
pressed (Quandt et al. 2014) while Syngliocladium was with apical cap q Part spores r Part spores in methylene blue. Scale Bars:
previously treated with respect to the second oldest e = 2 mm, f, h–l, n = 1 mm, g = 20 μm, m = 500 μm, o–r = 10 μm
Fungal Diversity
Fungal Diversity

5 . S t r o m at a w i t h p e r i t h e c i a l ap i c e s or la t e r a l Jacobs and Wingfield 2001; Zipfel et al. 2006). Many species

pads . . . . . . . . . . . . . . . . . . . . . . . . . . Ophiocordyceps of ophiostomatoid fungi cause sap stain or blue stain fungi on
5. Stromata with a highly reduced stromatic freshly cut logs and affect timber quality (Seifert 1993), and
pad. . . . . . . . . . . . . . . . . . . . . . . . . . . Tolypocladium several species are important pathogens of Protea spp., Pinus
6. Conidia straight to slightly curved or ellipsoidal to fusi- spp., Larix spp., Tsuga spp. and Pistacia spp. (Wingfield et al.
form, host arthropod or vertebrate . . . Purpureocillium 1988; Jacobs and Wingfield 2001; Roets et al. 2013).
6. Conidia crescent-shaped, helical, host This family was originally described with three genera in-
protozoan. . . . . . . . . . . . . . . . . . . . . . . . Drechmeria cluding Ophiostoma, Ceratostomella and Endoconidiophora
(Nannfeldt 1932). Recently, De Beer et al. (2013) accepted
Ophiostomataceae Nannf., Nova Acta R. Soc. Scient. upsal., Ceratocystiopsis, Fragosphaeria, Graphium,
Ser. 4 8(no. 2): 30 (1932) Leptographium, Ophiostoma and Raffaelea based on phylo-
Facesoffungi number: FoF 01807 genetic inference. Maharachchikumbura et al. (2015) provid-
Saprobic or parasitic on woody plants, occasionally on ed an updated outline of the family Ophiostomataceae with 12
herbaceous plants. Sexual morph: Ascomata superficial, peri- genera, which included sexual (Subbaromyces, Klasterskya,
thecial or cleistothecial, globose to subglobose, with external Fragosphaeria, Ophiostoma, Ceratocystiopsis and
hairs, pale-brown, dark-brown or black, with long ostiolar Spumatoria) and asexual (Leptographium, Pesotum,
neck or sometimes lacking ostioles. Ostiolar neck long and Phialographium, Raffaelea, Hyalobelemnospora and
slender, brown to dark brown at the base, pale-brown to hya- Hyalorhinocladiella) morph genera. In phylogenetic analysis
line at the apex, comprising parallel hyphae with pigmented of combined LSU, SSU, TEF and RPB2 sequence data,
cell walls, and numerous, filamentous, septate, pale-brown to Ophiostomataceae formed a well-supported sister clade to
hyaline ostiolar hyphae. Peridium pseudoparenchymatous, Ophioceraceae (Magnaporthales) and Annulatascaceae
the outer wall layers composed of angular cells with thick- (Annulatascales). It was also confirmed that
ened, pigmented walls, and lined by thin-walled, hyaline, cells Ceratocystiopsis, Fragosphaeria, Ophiostoma and Raffaelea
of textura angularis. Asci 8–spored, unitunicate, thin-walled, belong in the family Ophiostomataceae
fusiform, globose to subglobose, oblong to clavate, deliques- (Maharachchikumbura et al. 2015).
cent. Ascospores bi- to multi-seriate, variable in shape, fre-
quently asymmetrical, oval, cylindrical, ellipsoidal, reniform, Ophiostoma Syd. & P. Syd., Annls mycol. 17(1): 43 (1919)
or falcate, aseptate or 1-septate, hyaline or appearing yellow- Facesoffungi number: FoF 01808; Fig. 105
ish in mass, some with sheath-like structure, a hyaline outer Saprobic or weak parasites on decaying woody substrates.
layer of wall. Asexual morph: Hyphomycetous. Sexual morph: Ascomata superficial, perithecial, globose to
Conidiophores synnematous or mononematous, simple, subglobose, dark brown to black, with long ostiolar neck.
branched, bearing a brush-like apical, branching structure, Ostiole occasionally comprising two or three necks, long, be-
erect, hyaline or dark brown to black. Conidiogenous cells coming pale brown to hyaline at the apex, including ostiolar
holoblastic phialides, some proliferating at or somewhat be- hyphae, hyphae pale brown to hyaline, filamentous, septate,
low the apex and giving rise to another series of denticles or with an outer layer of thin-walled, dark cells. Peridium pseu-
conidiogenous cells. Conidia solitary or aggregated, fusiform, doparenchymatous, comprising hyaline to pale-brown, cells
obovate to oblong, cylindrical, hyaline to pale reddish-brown, of textura angularis. Asci 8-spored, unitunicate, pear-shaped,
aseptate, some forming in a viscid head. evanescent. Ascospores bi- to multi-seriate, allantoidal in side
Type: Ophiostoma Syd. & P. Syd. view, elliptical in face view and circular in end view,
Notes: The family Ophiostomataceae was introduced by surrounded by a thick, hyaline sheath, in end view apparently
Nannfeldt (1932). Prior to 1980, this family was treated in triangular, aseptate. Asexual morph: Hyphomycetous.
the order Plectascales (Nannfeldt 1932), Microascales Conidiophores synnematous or mononematous, erect, dark
(Luttrell 1951), Sphaeriales (Ainsworth and Bisby 1954; brown to black at the base, becoming light brown or
von Arx 1978), or as a synonym of the Endomycetaceae in subhyaline towards the apex. Conidiogenous cells annellidic,
the Endomycetales (Redhead and Malloch 1977). Later, the cylindrical, hyaline. Conidia unicellular, oblong or clavate to
order Ophiostomatales, was introduced by Benny and obovate, hyaline.
Kimbrough (1980) to accommodate the family Type species: Ophiostoma pilifera (Fr.) C. Moreau, Revue
Ophiostomataceae. Mycol., Paris 17 (Suppl. Colon. no. 1): 22 (1952)
The members of this family are mostly saprobes on sap- Notes: The genus Ophiostoma was introduced by Sydow
wood. These fungi are specialized with sticky spores to facil- and Sydow (1919), with Ophiostoma piliferum (Fr.) Sydow &
itate insect dispersal. Many bark beetles act as vectors of Sydow (1919) as the type species. De Beer et al. (2013) ac-
ophiostomatoid fungi, especially species of Ophiostoma, cepted 134 species in the genus. Some species are important
Leptographium and Ceratocytiopsis (Upadhyay 1981; tree pathogens. The best known examples are Ophiostoma
 Fungal Diversity

Fig. 105 Ophiostoma pilifera a

Ascomata b Ostiolar region with
ostiolar hyphae c Ascospores d
Conidiophores and conidia of the
Raffaelea state e Conidiophores
and conidia of the
Hyalorhinocladiella state
(redrawn from Osorio 1985)

ulmi (Buisman) Melin & Nannf and O. novo-ulmi Brasier, the Type species: Equicapillimyces hongkongensis S.S.Y.
causal agents of Dutch elm disease, which are effectively Wong et al., Veterinary Microbiology 115(2–4): 406 (2012)
transmitted by elm bark beetles (Webber and Gibbs 1989; Fragosphaeria Shear, Mycologia 15(3): 124 (1923)
Brasier 1991). Type species: Fragosphaeria purpurea Shear, Mycologia
15(3): 124 (1923)
Hyalobelemnospora Matsush., Matsush. Mycol. Mem. 7:
54 (1993)
Other genera included Type species: Hyalobelemnospora amazonica Matsush.,
Matsush. Mycol. Mem. 7: 54 (1993)
Ceratocystiopsis H.P. Upadhyay & W.B. Kendr., Mycologia Hyalorhinocladiella H.P. Upadhyay & W.B. Kendr.,
67(4): 799 (1975) Mycologia 67(4): 800 (1975)
Type species: Ceratocystiopsis minuta (Siemaszko) H.P. Type species: Hyalorhinocladiella minuta–bicolor (R.W.
Upadhyay & W.B. Kendr., Mycologia 67(4): 800 (1975) Davidson) H.P. Upadhyay & W.B. Kendr., Mycologia 67(4):
Equicapillimyces S.S.Y. Wong et al., Veterinary 800 (1975)
Microbiology 115(2–4): 406 (2012) Klasterskya Petr., Annls mycol. 38(2/4): 225 (1940)
Fungal Diversity

Type species: Klasterskya acuum (Mouton) Petr., Annls 3. Conidia formed in a viscid head. . . . . . . . . . . Pesotum
mycol. 38(2/4): 227 (1940) 3. Conidia not formed in a viscid . . . . . . Phialographium
Leptographium Lagerb. & Melin, in Lagerberg et al., 4. Conidiophores aggregated . . . . . . . . . . . . . . . . . . . . . 5
Svensk Skogsvårdsförening Tidskr. 25: 257 (1927) 4. Conidiophores single . . . . . . . . . . . . . . . . . . Raffaelea
Type species: Leptographium lundbergii Lagerb. & Melin, 5. Conidia aggregated . . . . . . . . . . . Hyalorhinocladiella
in Lagerberg et al., Svensk Skogsvårdsförening Tidskr. 25: 5. Conidia solitary . . . . . . . . . . . . . . Hyalobelemnospora
248 (1927)
Pesotum J.L. Crane & Schokn., Am. J. Bot. 60: 347 (1973) Papulosaceae Winka & O.E. Erikss., Mycoscience 41 (2):
Type species: Pesotum ulmi (M.B. Schwarz) J.L. Crane & 102 (2000)
Schokn., Am. J. Bot., Suppl. 60(4): 348 (1973) Facesoffungi number: FoF 01284
Phialographium H.P. Upadhyay & W.B. Kendr., Saprobic in freshwater or marine habitats. Sexual morph:
Mycologia 66(1): 183 (1974) Ascomata occurring singly, immersed or semi-immersed,
Type species: Phialographium sagmatosporae H.P. black to dark brown, globose to ellipsoidal, coriaceous,
Upadhyay & W.B. Kendr., Mycologia 66(1): 183 (1974) ostiolate. Necks long, periphysate. Peridium thick, composed
Raffaelea Arx & Hennebert, Mycopath. Mycol. appl. 25: of cells of textura angularis or brown-walled compressed
310 (1965) cells. Hamathecium comprising tapering, simple or branched,
Type species: Raffaelea ambrosiae Arx & Hennebert, septate, hyaline paraphyses. Asci 8-spored, unitunicate, cylin-
Mycopath. Mycol. appl. 25: 310 (1965) drical, short pedicellate, with a J-, refractive, bipartite or dis-
Spumatoria Massee & E.S. Salmon, Ann. Bot., Lond. 15: coid, apical ring. Ascospores uniseriate to biseriate, hyaline or
350 (1901) brown, ellipsoidal or fusiform, unicellular to 3-septate,
Type species: Spumatoria longicollis Massee & E.S. smooth or verruculose, with or without cup-like, bipolar ap-
Salmon, Ann. Bot., Lond. 15: 350 (1901) pendages. Asexual morph: Undetermined.
Subbaromyces Hesselt., Bull. Torrey bot. Club 80: 511 Type: Papulosa Kohlm. & Volkm.-Kohlm.
(1953) Notes: Papulosaceae is a monotypic family introduced by
Type species: Subbaromyces splendens Hesselt., Bull. Winka and Eriksson (2000) based on morphology and SSU
Torrey bot. Club 80: 511 (1953) rDNA sequence data. Winka and Eriksson (2000) concluded
that the exact phylogenetic position of the genus Papulosa
could not be established, due to the small number of SSU
Key to the sexual genera of Ophiostomataceae sequences available for unitunicate pyrenomycetes at that
time. Papulosa seemed to be closely related to Diaporthales,
1. Ascospores with septate . . . . . . . . . . . . . . . . . . . . . . . . . 2 Ophiostomatales, Phyllachorales and Sordariales based on
1. Ascospores aseptate . . . . . . . . . . . . . . . . . . . . . . . . . . 3 molecular data, thus the genus was accommodated in the sub-
2. Ascospores 1-septate . . . . . . . . . . . . . . . . . . . . . . . . . 4 class Sordariomycetidae. Phylogenetic analyses of
2. Ascospores more than 1-septate . . . . Equicapillimyces Maharachchikumbura et al. (2015) placed Papulosaceae as a
3. Ascomata perithecial . . . . . . . . . . . . . . . . . . . . . . . . . 5 sister clade to Coniochaetaceae and the order Annulatascales.
3. Ascomata cleistothecial . . . . . . . . . . . . Fragosphaeria The genera Brunneosporella and Fluminicola are predomi-
4. Perithecium forming a disc or collar-like nantly found in freshwater habitats and resemble other genera
structure . . . . . . . . . . . . . . . . . . . . . . . Subbaromyces in Annulatascaceae with cylindrical asci and a large bipartite
4. Perithecium not forming a disc or collar-like apical ring and were placed in family Annulatascaceae by
structure . . . . . . . . . . . . . . . . . . . . . . . . . . Klasterskya Ranghoo et al. (2001); Wong et al. (1999); Zelski et al.
5. Asci fusiform to clavate . . . . . . . . . . . . . . . . . . . . . . . 6 2011. Brunneosporella differs from Fluminicola in having
5. Asci pear-shaped . . . . . . . . . . . . . . . . . . . Ophiostoma brown, 1-septate, smooth ascospores, whereas Fluminicola
6. Ascospores falcate, with sheath . . . . . Ceratocystiopsis has hyaline, 3-septate ascospores with cup-like, bipolar ap-
6. Ascospores obovoid, without sheath. . . . . .Spumatoria pendages. Abdel-Wahab et al. (2011) analyzed only LSU se-
quence data and found that Brunneosporella and Fluminicola
clustered with Papulosa amerospora and Cataractispora
Key to the asexual genera of Ophiostomataceae receptaculorum W.H. Ho, K.D. Hyde & Hodgkiss. A similar
clustering was seen in phylogenetic analyses carried out by
1. Conidiophores mononematous . . . . . . . . . . . . . . . . . . . . 2 Réblová et al. (2013) using LSU, SSU, rpb-2 sequence data,
1. Conidiophores synnematous. . . . . . . . . . . . . . . . . . . . 3 wherein both Brunneosporella and Fluminicola appear in the
2. Conidiophores hyaline, with simple branching . . . . . . 4 same clade as that of Papulosa and Pleurophragmium
2. Conidiophores dark, bearing a brush-like apical parvisporum (Preuss) Hol.-Jech. and were marked as
branching structure . . . . . . . . . . . . . . .Leptographium Papulosaceae. In LSU and SSU sequence analyses carried
 Fungal Diversity

out by Boonyuen et al. (2012), Brunneosporella (as 2. Ascospores 3-septate, with cup-like, bipolar
Ascobrunneispora aquatica Ranghoo, K.D. Hyde & E.C.Y. appendages. . . . . . . . . . . . . . . . . . . . . . . Fluminicola
Liew) clustered with Papulosa. Maharachchikumbura et al.
(2015) placed Brunneosporella and Fluminicola in Pestalotiopsidaceae Maharachch. & K.D. Hyde, in
Papulosaceae. The phylogenetic tree prepared by Boonyuen Senanayake et al., Fungal Divers 73: 107 (2015)
et al. (2012) shows that many annulatascaceous genera, which Facesoffungi number: FoF 0666
were previously placed in Annulatascaceae, are scattered and Saprobic or pathogenic on plants. Sexual morph:
do not cluster with the familial type, Annulatascus velatispora Ascomata scattered, solitary or in small groups, immersed,
K.D. Hyde. These genera were placed in Sordariomycetidae apex short erumpent. Ostiole subglobose, papillate ostiolar
incertae sedis. Thus a molecular phylogenetic approach is canal periphysate. Peridium comprising several layers of dark
needed to resolve the position of these genera in brown to blackish, compressed cells. Hamathecium compris-
Annulatascaceae and confirm the composition of ing tapering paraphyses. Asci 8-spored, unitunicate, cylindri-
Annulatascaceae sensu stricto. cal, short to long pedicellate, with a J+, or J-, apical ring.
Ascospores uniseriate, brown, ovoid to elliptic, straight or
Papulosa Kohlm. & Volkm.-Kohlm., Syst. Ascom. 11(2): 96 inequilateral, 2–3-septate. Asexual morph: Coelomycetous.
(1993) Conidiomata acervular or pycnidial, subglobose, globose, cla-
Facesoffungi number: FoF 01285; Figs 106, 107 vate, solitary or aggregated, dark brown to black, immersed to
Saprobic on culms of saltmarsh plant Juncus roemerianus erumpent, unilocular or irregularly pluri-loculate. Peridium
Scheele in marine habitats. Sexual morph: Ascomata occur- comprising 2–3 strata of textura angularis, outer layer
ring singly, dark brown, immersed, subglobose, ostiolate, with of pale brown, thick-walled cells, becoming hyaline in
a long periphysate neck. Peridium composed of textura the inner layer. Conidiophores indistinct, often reduced
angularis. Hamathecium comprising tapering, simple, sep- to conidiogenous cells, when present peripheral, hyaline,
tate, hyaline paraphyses. Asci 8-spored, cylindrical, pedicel- branched or unbranched, cylindrical or lageniform.
late, with a bipartite, IKI positive, apical ring. Ascospores Conidiogenous cells discrete, holoblastic, annellidic, in-
uniseriate, unicellular, hyaline, ellipsoidal, verruculose, with- determinate, integrated, cylindrical, hyaline, smooth.
out appendages. Asexual morph: Undetermined. Conidia ellipsoid to clavate, or fusiform, straight or
Type species: Papulosa amerospora Kohlm. & Volkm.- curved, 3–4-euseptate, hyaline, pale olivaceous or
Kohlm., Syst. Ascom. 11(2): 96 (1993) brown, bearing cellular, filiform or attenuated
Notes: The monotypic genus Papulosa was established by appendages.
Kohlmeyer and Volkmann-Kohlmeyer (1993) and occurs on Type: Pestalotiopsis Steyaert
Juncus roemerianus in salt marshes in the USA. Notes: Pestalotiopsidaceae is a recently introduced family
that possesses pestalotiopsis-like asexual morphs
(Senanayake et al. 2015). The other genera belonging to
Pestalotiopsidaceae are Ciliochorella, Lepteutypa,
Other genera included
Monochaetia, Neopestalotiopsis, Pseudopestalotiopsis and
Seiridium. Previously, all these genera were assigned to
Brunneosporella Ranghoo & K.D. Hyde, Mycol. Res.
Amphisphaeriaceae. Members of the Pestalotiopsidaceae
105(5): 625 (2001)
are common phytopathogens that cause a variety of diseases
Type species: Brunneosporella aquatica Ranghoo & K.D.
in plants or are often isolated as saprobes or endophytes and
Hyde, in Ranghoo et al., Mycol. Res. 105(5): 625 (2001)
are widely distributed throughout tropical and temperate re-
Fluminicola S.W. Wong et al., Fungal Diversity Res. Ser.
gions (Guba 1961; Barr 1975; Nag Raj 1993;
2: 190 (1999)
Maharachchikumbura et al. 2014). The sexual morph of
Type species: Fluminicola bipolaris S.W. Wong et al.,
Pestalotiopsis is Pestalosphaeria (Barr 1975) and Seiridium
Fungal Diversity 2: 190 (1999)
has a Lepteutypa sexual morph (Nag Raj 1985).

Pestalotiopsis Steyaert, Bull. Jard. bot. État Brux. 19: 300

Key to the genera of Papulosaceae (1949)
Facesoffungi number: FoF 02124; Figs 108, 109
1. Ascospores smooth-walled, brown; asci with discoid apical Saprobic or pathogenic on plants. Sexual morph:
ring. . . . . . . . . . . . . . . . . . . . . . . . . . . . .Brunneosporella Ascomata scattered or gregarious, immersed, subglobose to
1. Ascospores verruculose, hyaline; asci with bipartite api- globose, with central black irregular ostioles. Peridium com-
cal ring. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 prising 3–5 layers of brown, relatively thick-walled cells of
2. Ascospores 1-celled, without appendages. . . Papulosa textura angularis, inner cells flattened and thin-walled.
Fungal Diversity

Fig. 106 Papulosa amerospora (Material examined: USA, North Squash mount of ascoma d Vertical section through ostiole e Immature
Carolina, Carteret County, Broad Creek, on dead standing culms of ascus f–h Developing asci with paraphyses. Scale bars: c = 100 μm; d,
Juncus roemerianus Scheele (Juncaceae), 20 December 1992, J. f = 50 μm; e, g, h = 20 μm
Kohlmeyer 5491 (NY 01348450, holotype) a–b Herbarium packet c

Paraphyses with few septa, base relatively wide and tapering Ascospores uniseriate, or 1-seriate in the upper part and 2-
to free ends. Asci 8-spored, unitunicate, cylindrical, pedicel seriate at the base, oblong to ellipsoidal or fusiform, smooth
short, with a distinct J+, amyloid apical ring in ascus apex. or verrucose, pale yellowish brown. Asexual morph:
 Fungal Diversity

Fig. 107 Papulosa amerospora

(redrawn from Kohlmeyer and
Volkmann-Kohlmeyer 1993) a
Asci at different stages of
development b, c Ascospores in
surface view and optical section d
Apex of ascus, showing apical
ring e Paraphyses (left) and hairs
on ascomata (right). Scale bars: a,
e = 20 μm; b, c, d = 5 μm

Conidiomata acervular or pycnidial, subglobose, globose, cla- Maharachchikumbura et al. (2011) pointed out that the com-
vate, solitary or aggregated, dark brown to black, immersed to mon Pestalotiopsis name should be applied to both morphs.
erumpent, unilocular to pluri-loculate. Conidiophores branded Pestalotiopsis species have often been isolated as endophytes
and septate, occasionally reduced to conidiogenous cells, hy- and due to their ability to switch life-modes, many pathogens
aline, smooth. Conidiogenous cells discrete, cylindrical, or endophytes may persist as saprobes (Zhang et al. 2012a,b;
ampulliform to lageniform, hyaline, smooth- and thin- Maharachchikumbura et al. 2014).
walled; conidiogenesis initially holoblastic, percurrent prolif-
erations to produce additional conidia at slightly higher levels.
Conidia fusoid, ellipsoid, subcylindrical, straight to slightly Other genera included
curved, 4-septate, slightly constricted at septa; basal cell con-
ical to cylindric with a truncate base; three median cells Ciliochorella Syd., in Sydow & Mitter, Annls mycol. 33(1/2):
doliiform, concolourous, light brown to olivaceous, wall ru- 62 (1935)
gose to verruculose; apical cell conic to cylindrical, thin- and Type species: Ciliochorella mangiferae Syd., Annls
smooth-walled; with tubular apical appendages, one to many, mycol. 33(1/2): 63 (1935)
filiform or attenuated, flexuous, branched or unbranched, with Lepteutypa Petr., Annls mycol. 21(3/4): 276 (1923)
or without spatulate tips; basal appendage single, tubular, Type species: Lepteutypa fuckelii (Nitschke) Petr., Annls
centric. mycol. 21(3/4): 276 (1923)
Type species: Pestalotiopsis maculans (Corda) Nag Raj, Monochaetia (Sacc.) Allesch., Rabenh. Krypt.-Fl., Edn 2
Mycotaxon 22(1): 47 (1985) (Leipzig) 1(7): 665 (1902) [1903]
Notes: Based on the conidial forms, Steyaert (1949) split Type species: Monochaetia monochaeta (Desm.) Allesch.,
Pestalotia into three genera, namely Pestalotia, Pestalotiopsis Rabenh. Krypt.-Fl., Edn 2 (Leipzig) 1(7): 667 (1902) [1903]
and Truncatella. Pestalotiopsis was introduced for species Neopestalotiopsis Maharachch. et al., in
with 5-celled conidia and P. maculans is regarded as the type Maharachchikumbura et al., Stud. Mycol. 79: 135 (2014)
species. The sexual morph of Pestalotiopsis is Type species: Neopestalotiopsis protearum (Crous & L.
Pestalosphaeria and only 13 species are known as compared Swart) Maharachch. et al., in Maharachchikumbura et al.,
to the asexual morph (295 species names). Stud. Mycol. 79: 147 (2014)
Fungal Diversity

Fig. 108 Pestalotiopsis trachicarpicola – sexual morph (Material layers c Section of ascomata d Asci and paraphyses e-f Mature and
examined: CHINA, Yunnan Province, Kunming, Kunming Botanical immature unitunicate asci g Asci in Melzer’s reagent, note the distinct
Gardens, on leaf spots on living leaves of Trachycarpus fortunei J+ apical ring h Ascospores. Scale Bars: a = 200 μm, d, e–g = 20 μm,
Beccari (Arecaceae), March 2011, K.D. Hyde OP068 (IFRD 9026, c = 50 μm, h = 10 μm
holotype) a Conidiomata on living leaves b Peridium with five cell

Fig. 109 Pestalotiopsis clavata – asexual morph (Material examined: 0268, ex-type) a Conidiomata sporulating on PDA b–c Conidiogenous
CHINA, Yunnan Province, Kunming, Kunming Botanical Garden, living cells d–f Conidia. Scale bars = 20 μm
leaf of Buxus sp., 19 March 2002, Wenping Wu KBG26-5 (MFLUCC 12-
 Fungal Diversity

Pseudopestalotiopsis Maharachch. et al., in genera (Serenomyces, Cocoicola) previously placed in the

Maharachchikumbura et. al., Stud. Mycol. 79: 180 (2014) Phyllachoraceae. Later, Phaeochoropsis was also included
Type species: Pseudopestalotiopsis theae (Sawada) in this family based on morphology (Hyde and Cannon
Maharachch. et al. in Maharachchikumbura et al., Stud. 1999). Although Hyde et al. (1997) tentatively placed this
Mycol. 79: 183 (2014) family in the order Phyllachorales, no molecular data are
Seiridium Nees, Syst. Pilze (Würzburg): 22 (1816) available to confirm its phylogenetic placement and assign it
[1816–17] to any order. This family is characterized by ascomata devel-
Type species: Seiridium marginatum Nees, Syst. Pilze oping in a pseudostromata aseptate brown ascospores, and
(Würzburg): 23 (1816) [1816–17] clavate to saccate evanescent asci Hyde et al. 1997; Hyde
and Cannon 1997). The description above is a combination
of characters from Hyde et al. (1997), Hyde and Cannon
Key to asexual genera of Pestalotiopsidaceae (1999) and Taylor and Hyde (2003).

1. Conidia 3-septate . . . . . . . . . . . . . . . . . . . . .Ciliochorella Phaeochora Höhn., Sber. Akad. Wiss. Wien, Math.-naturw.
1. Conidia more than 3-septate. . . . . . . . . . . . . . . . . . . . 2 Kl., Abt. 1 118: 1513 [53 repr; in clave] (1909)
2. Conidia 4-septate . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 Facesoffungi number: FoF 02125; Fig. 110
2. Conidia 5-septate . . . . . . . . . . . . . . . . . . . . . Seiridium Saprotrophic or biotrophic on leaves of palms. Sexual
3. Conidia with 1 apical appendage . . . . . . Monochaetia morph: Stromata scattered as blackened regions, shiny black,
3. Conidia with 1 to many apical appendage . . . . . . . . . 4 conspicuous on upper and lower leaf surfaces, rarely coalesc-
4. Conidia concolourous . . . . . . . . . . . . . . . . . . . . . . . . 5 ing, uniloculate to multi-loculate, composed of brown cells of
4. Conidia versicolourous. . . . . . . . . . . Neopestalotiopsis textura globosa and crystalline granules. Ostiole inconspicu-
5. Conidia light concolourous . . . . . . . . . . Pestalotiopsis ous. Ascomata oblate to subglobose, dark brown. Peridium
5. Conidia dark concolourous. . . . . .Pseudopestalotiopsis several layered, composed of strongly flattened, brown-
Phaeochoraceae K.D. Hyde et al., in Hyde et al., Syst. walled cells, thinner at the base. Periphyses and interascal
Ascom. 15(1–2): 118 (1997) tissue lacking. Asci 8-spored, unitunicate, thin-walled, sac-
Facesoffungi number: FoF 01910 cate, short pedicellate, without an apical ring, evanescent.
Saprotrophic or biotrophic on leaves of palms. Sexual Ascospores biseriate, triseriate to multi-seriate or irregular.
morph: Stromata scattered as blackened regions on the host, golden to mid-brown, aseptate, ellipsoidal to fusiform, with
rarely inconspicuous, usually raising the substrate surface, 1– one flattened face, thick-walled, smooth or minutely
to multi-loculate, ostiole conspicuous or not. Ascomata ellip- verruculose, with globose, hyaline appendages near the apices
soidal to subglobose, dark brown, clustered or solitary, if sol- of the flattened surface. Asexual morph: Coelomycetous.
itary single ostiolate, when clustered ostiolar necks together to Peridium single layered, composed of brown cells of textura
form a multi-ostiolate cavity. Ostioles conspicuous or incon- angularis. Conidiogenous cells narrowly conical, usually pro-
spicuous. Peridium several layered, composed of flattened, liferating percurrently, occasionally sympodially, periclinal
brown to dark brown-walled cells, thinner at the base. thickenings visible, without collarette. Conidia aseptate, nar-
Interascal tissue present only in some taxa, composed of rowly fusiform to bacillar, hyaline, thin-walled.
paraphyses-like hyphae, wide, septate, tapered, thin-walled, Type species: Phaeochora chamaeropis (Cooke) Höhn.
guttulate, evanescent at maturity. Asci 6–8-spored, [as ‘chamaeropsidis’], Sber. Akad. Wiss. Wien, Math.-naturw.
unitunicate, fusiform or saccate, very thin-walled, long or Kl., Abt. 1 118: 1513 [53 repr; in clave] (1909)
short pedicellate, usually without apical structures, deliquesc- Notes: von Höhnel (1909) introduced the genus
ing early. Ascospores uniseriate to biseriate, yellow to oliva- Phaeochora to accommodate Auerswaldia chamaeropis
ceous or various shades of brown, aseptate, ellipsoidal, fusi- (Cooke) Sacc. (Syn. Dothidea chamaeropis Cooke). It was
form or cylindrical, thick-walled, sometimes with one flat- recognized by ‘phyllachoroid’ stromata embedded in the epi-
tened face, sometimes delicately striate, with or without ap- dermis and parenchyma, and not disrupting the covering epi-
pendages. Asexual morph: Coelomycetous. Conidiogenous dermal layer. Hohnel consider A. chamaeropis as the type and
cells narrowly conical, usually proliferating percurrently, but introduced another species, A. densa E. Bormmer & M.
occasionally sympodially, periclinal thickening visible, with- Rousseau to the genus. However A. densa was transferred to
out a collarette. Conidia aseptate, narrowly fusiform to bacil- Sphaerodothis (Sacc. & P. Syd.) Shear by Shear (1909), but its
lar, hyaline, thin-walled (Hyde et al. 1997; Hyde and Cannon identity remains unclear. Study by Joly (1961) only provide a
1999; Taylor and Hyde 2003). short description to the Sphaerodothis and the host species
Type: Phaeochora Höhn. was not recorded by the original authors, the fungaus has
Notes: The family Phaeochoraceae was introduced by described simply as occurring on Coriaceous leaves in Costa
Hyde et al. (1997) to accommodate Phaeochora and two other Rica. Müller (1965) clarified the earliar name of the type
Fungal Diversity

Fig. 110 Phaeochora calamigena (Material examined: SRI LANKA, c Transverse section through stroma showing ascoma d Immature asci e
Peradeniya, on leaves of Calamus rudentum Lour. (Arecaceae), G.H.K. Ascus in Melzer’s reagent f, g Immature and mature ascospores. Scale
Thwaites 442 (ex herb. M.J. Berkeley), January 1868 (K(M) 35077, bars: b, c = 200 μm, d, e = 20 μm, f, g = 10 μm
holotype) a Herbarium material with label b Stromata on host substrate

species of Phaeochora as Sphaeria steinhelii Mont. and Type species: Phaeochoropsis neowashingtoniae (Shear)
transferred it as Phaeochora steinhelii Mont. E. Mull. K.D. Hyde & P.F. Cannon, Mycol. Pap. 175: 33 (1999)
Phaeochora is superficially similar to Sphaerodothis, and Serenomyces Petr., Sydowia 6(1–4): 296 (1952)
the two genera were treated as synonyms by von Arx & Type species: Serenomyces shearii Petr., Sydowia 6(1–4):
Müller (1954) and Joly (1961). However, the Genus 296 (1952)
Phaeochora is characterized by oblate to subglobose
ascomata and golden to mid-brown, aseptate, ellipsoidal to
fusiform, ascospores with appendages near the apices of the
flattened surface (Hyde et al. 1997). Key to the genera of Phaeochoraceae

1. Ascospores asymmetrical . . . . . . . . . . . . . . . . . . . . . . . . 2
Other genera included 1. Ascospores symmetrical. . . . . . . . . . . . . . . . . . . . . . . 3
2. Ascospores, striate, surrounded by mucilage when
Cocoicola K.D. Hyde, Nova Hedwigia 60(3–4): 600 (1995) young, without polar appendages . . . Phaeochoropsis
Type species: Cocoicola cylindrospora (C. Booth & D.E. 2. Ascospores smooth or verrucose, not surrounded by mu-
Shaw) K.D. Hyde, Nova Hedwigia 60(3–4): 600 (1995) cilage, with polar appendages. . . . . . . . . . Phaeochora
Phaeochoropsis K.D. Hyde & P.F. Cannon, Mycol. Pap. 3. Ascospores light brown . . . . . . . . . . . . . .Serenomyces
175: 30 (1999) 3. Ascospores yellow or olivaceous . . . . . . . . . Cocoicola
 Fungal Diversity

P. eucalyptorum Crous and P. uniforme Crous & Summerell

Phlogicylindriaceae Senan. & K.D. Hyde, in Senanayake all occuring on species of Eucalyptus, from Australia (Farr
et al., Fungal Diversity 73: 35 (2015) and Rossman 2016; Index Fungorum 2016).
Facesoffunginumber: FoF 00681
Saprobic on leaves, twigs and branches, presently known
only from dicotyledons. Sexual morph: Pseudostroma Other genera included
scattered, solitary, immersed, black, thick around papilla, thin-
ner towards the base. Ascomata forming under pseudostroma, Ciferriascosea Senan.et al., in Senanayake et al., Fungal
scattered, solitary, globose, coriaceous, brown to black, Divers 73: 35 (2015); Fig. 112
ostiolate, papillate. Papilla short, narrow. Peridium compris- Type species: Ciferriascosea rectamurum Senan. et al., in
ing outer, brown to light brown, thick-walled cells of textura Senanayake et al., Fungal Diversity 73: 107 (2015)
angularis and inner, hyaline, thin-walled cells of textura
angularis. Hamathecium comprising septate, hyaline paraph- Phyllachoraceae Theiss. & P. Syd., Annls mycol. 13(3/4):
yses. Asci 8-spored, unitunicate, cylindrical, short pedicellate, 168 (1915)
rounded at apex, with J+ discoid subapical ring. Ascospores Facesoffungi number: FoF 01329
overlapping uniseriate, hyaline, fusiform, rounded at ends, Parasitic on plants in terrestrial and rarely aquatic habitats.
equally 1-septate, slightly constricted at septa. Asexual Sexual morph: Leaf spots on host black, abundant, scattered,
morph: Coelomycetous. Conidiomata slimy, erect tufts of raised, mostly rounded to oblong or elongated, sometimes
hyaline conidial masses, synnematous, indeterminate, turning parallel with leaf venation, surrounded by light brown necrotic
brown with age. Conidiophores consisting of an intricate net- region. Ascomata flattened, globose, with thin walls, lying in
work of brown, smooth, branched cells. Conidiogenous cells leaf tissues or in a stroma and maturing in living leaves,
hyaline, annellidic, smooth, becoming pale brown with age, ostiolate, ostiolar canal conical, wide, lacking periphyses.
ampulliform with elongated necks, with percurrent prolifera- Peridium clypeate, thickest adjacent to ostiolar canal, com-
tions. Conidia formed apically, hyaline, cylindrical with ob- posed of a deeply melanized, brown-black, amorphous layer
tusely rounded apex, aseptate to 1-septate. of host cuticle and epidermal cells, often merging with a ligh-
Type: Phlogicylindrium Crous et al. ter pigmented region of brownish, distorted parenchyma cells
Notes: The family Phlogicylindriaceae was introduced by infiltrated with fungal hyphae, beneath the clypeus. Lower
Senanayake et al. (2015) to accommodate the genus and lateral peridium composed of two layers; an outer region
Phlogicylindrium Crous et al. (Summerell et al. 2006) and comprising several layers of dark brown, flattened, thin-
Ciferriascosea Senan. et al. The type species, walled fungal cells, which merge inwardly with several layers
Phlogicylindrium eucalypti Crous et al. has no known sexual of hyaline, flattened, thin-walled fungal cells. Lateral peridi-
morph and was reported occurring in association with a um fuses outwardly with an irregular, narrow region of
Mycosphaerella species causing lesions on living leaves of distorted host parenchyma infiltrated by fungal cells. The bas-
Eucalyptus. Phlogicylindrium has unique characteritics that al peridium merges outwardly with either a narrow zone of
include slimy conidiomata, with erect tufts of hyaline conidial infiltrated and distorted host parenchyma and occasionally
masses, and hyaline, unicellular to 1-septate, cylindrical co- lower epidermal cells, or integrates directly with a lower clyp-
nidia, with obtusely rounded apices (Summerell et al. 2006). eus, similar in construction to that basal peridium. Paraphyses
The genus Ciferriascosea, which had sexual characteristics, numerous, persistent, filiform, branched or unbranched, sep-
showed a close affinity to the species of Phlogicylindrium tate, slightly longer than asci. Asci 8-spored, persistent, cylin-
based on their phylogeny, and was placed in this family by drical to fusiform, short pedicellate, with walls uniform in
Senanayake et al. (2015). The genus comprises two species, thickness, not especially thickened at apex, apical ring often
the type C. rectamurum Senan. et al. (illustrated) and present rarely absent, J-, or J+ Ascospores 1–3 seriate, fusi-
C. fluctamurum Senan. et al., both of which occured on form to narrowly oval, usually hyaline, aseptate or rarely sep-
Spartium junceum, in Italy. The description and illustration tate, often with a mucilaginous sheath, with or without an
provided of the asexual morph, is based on Summerell et al. attenuated base, sometimes with pad like appendages.
(2006). Asexual morph: Coelomycetous, with conidia in locules
within a stroma, spermatial or disseminative. Conidiomata
Phlogicylindrium Crous et al., in Summerell et al., Fungal similar to ascomata, immersed, subcuticular, strongly raising
Diversity 23: 340 (2006); Fig. 111 the host surface, occasionally pycnidial, occupying the region
Type species: Phlogicylindrium eucalypti Crous et al., in between the cuticle and epidermis, ostiolate. Conidiogenous
Summerell et al., Fungal Diversity 23: 340 (2006) cells arising from the basal and lateral walls, cylindrical,
Notes: The genus comprises three species, the type phialidic, aseptate, hyaline, forming conidia singly at the apex.
P. e u c a l y p t i C r o u s , S u m m e r b . & S u m m e r e l l , Conidia hyaline, filiform, aseptate.
Fungal Diversity

Fig. 111 Phlogicylindrium

eucalypti (CBS H-19749,
redrawn from Summerell et al.
2006). a Colony sporulating on
MEA b–d Conidiophores and
conidiogenous cells e–g
Conidiogenous cells h–k
Conidia. Scale bars: d, g = 20 μm

Type: Phyllachora Nitschke Phyllachoraceae, whereas Barr (1990), who provided a

Notes: Phyllachoraceae was introduced by Theissen and key to genera of Phyllachoraceae, included only 12
Sydow (1915) and has been included in several orders, includ- genera. Eriksson and Hawksworth (1993) recognized
ing Dothideales (Horst 1990), Sphaeriales (Nannfeldt 1932; 39 genera and Hawksworth et al. (1995) accepted 42
Miller 1949; Müller and von Arx 1962; Wehmeyer 1975), genera. In a recent study based on available molecular
Xylariales (Luttrell 1951; Barr 1990), Glomerellales data, Maharachchikumbura et al. (2015) listed 58 genera
(Chadefaud 1960; Locquin 1984), Phyllachorales (Barr in the family Phyllachoraceae.
1976a, b, 1983), Polystigmatales (Eriksson 1982; Phyllachora Nitschke ex Fuckel, Jb. nassau. Ver. Naturk.
Hawksworth et al. 1983), and Diaporthales (Cannon 1988). 23–24: 216 (Fuckel 1870) [1869–70]
Members of Phyllachoraceae are characterised by Facesoffungi number: FoF 02126; Fig. 113
ascohymenial development with paraphyses, thin-walled asci, Parasitic on plants. Sexual morph: Leaf spots on host
which may have an apical ring, that does not stain blue black, abundant, scattered, raised, mostly rounded to oblong
in iodine (J-) and ascospores that are often hyaline and or elongated, sometimes parallel with leaf venation,
1-celled (Cannon 1991). Asexual morphs are surrounded by light brown necrotic region. Ascomata flat-
coelomycetes, spermatial or disseminative (Hawksworth tened, globose, with thin walls, lying in leaf tissues or in a
et al. 1995). Munk (1957) and Barr (1990) had a dif- stroma and maturing in living leaves, ostiolate and sometimes
ferent concept of the family, including genera with J+, with a clypeus formed by the lateral proliferation of ostiolar
apical rings, in the ascus. The number of genera recog- hyphae. Peridium clypeate, thickest adjacent to ostiolar canal,
nized within the family varies according to the authori- composed of a deeply melanized, brown-black, amorphous
ty. Hawksworth (1985) recognized 23 genera of layer of host cuticle and epidermal cells, often merging with
 Fungal Diversity

Fig. 112 Ciferriascosea rectamurum (Material examined: ITALY, ascoma d Peridium e Paraphyses f–h Asci with ascospores j–l
Province of Forlì-Cesena, Santa Sofia, Castellaccio di Corniolino, on Ascospores. Scale bars: a, b = 1 mm, c, = 100 μm, d = 20 μm, e–
branches of Spartium junceum L. (Fabaceae), 29 December 2012, Erio h = 50 μm, j–l = 25 μm, i = 20 μm
Camporesi, IT 986, MFLU 15-0726). a, b Ascomata on host c Section of

a lighter pigmented region of brownish, distorted parenchyma conidia in locules within a stroma. Conidiomata similar to
cells infiltrated with fungal hyphae, beneath the clypeus. ascomata, immersed, subcuticular, strongly raising the host
Lower and lateral peridium composed of two layers; an outer surface, occasionally pycnidial, occupying the region between
region comprising several layers of dark brown, flattened, the cuticle and epidermis, ostiolate. Conidiophores cylindri-
thin-walled fungal cells, which merge inwardly with several cal, hyaline, thin-walled, formed directly from the strongly
layers of hyaline, flattened, thin-walled fungal cells. Lateral contorted cells of the lower wall Conidiogenous cells arising
peridium fuses outwardly with an irregular, narrow region of from the basal and lateral walls, cylindrical, phialidic,
distorted host parenchyma infiltrated by fungal cells. The bas- aseptate, hyaline, forming conidia singly at the apex.
al peridium merges outwardly with either a narrow zone of Conidia hyaline, clavate, aseptate.
infiltrated and distorted host parenchyma and occasionally Type species: Phyllachora graminis (Pers.) Fuckel, Jb.
lower epidermal cells, or integrates directly with a lower clyp- nassau. Ver. Naturk. 23–24: 216 (1870) [1869–70]
eus, similar in construction to that of basal peridium. Notes: Phyllachora is a large genus with about 1500 spe-
Paraphyses numerous, persistent, filiform, branched, slightly cies currently recognized (Index Fungorum 2016). This large
longer than asci. Asci 8-spored, persistent, cylindrical to fusi- number of species is a result of naming species on the basis of
form, short pedicellate, with walls fairly uniform in thickness, host association (Cannon 1988). There have been several mor-
not specially thickened at apex, apical ring often present rarely phological studies of Phyllachora on various groups of host
absent, J-, or J+. Ascospores 1–3 seriate, fusiform to narrowly plants. For example, Phyllachora species causing leaf tar spot
oval, hyaline, with or without attenuated base, sometimes with diseases on Leguminosae (Cannon 1991), on Duranta spp. in
pad like appendages. Asexual morph: Coelomycetous, with the tropics (Hanlin and Tortolero 1991), small scabby
Fungal Diversity
 Fungal Diversity

ƒFig. 113 Phyllachora nigeriensis (Material examined: AFRICA, West Type species: Deshpandiella jambolana (T.S. Ramakr.
Tropical Africa, Nigeria, on leaf surface of Calamus sp., 1977 September, et al.) Kamat & Ullasa, in Ullasa & Rao, Bull. Torrey bot.
E.U. Okpala, IMI 216890, holotype) a Herbarium specimen b Ascomata
on host c Section through ascoma d Peridium e Paraphyses f-i Asci j-m
Club 100: 42 (1973)
Ascospores. Scale bars: c, d = 100 μm, e–m = 50 μm Diachora Müll. Arg., Jb. wiss. Bot. 25: 623 (1893)
= Diachorella Höhn., Hedwigia 60: 192 (1918).
Type species: Diachora onobrychidis (DC.) Jul. Müll. [as
leafspots or “lixa-pequena” on coconut palms in Brazil ‘onobrychis’], Jb. wiss. Bot. 25: 693 (1893)
(Subileau et al. 1993), and leaf-spots on grasses and sedges Diatractium Syd. & P. Syd., Annls mycol. 18(4/6): 183
in northern regions (Seaver 1928b), on plants of family (1921) [1920]
Asclepiadaceae, tar spots on grasses in Australia (Sivanesan Type species: Diatractium cordiae (F. Stevens) Syd. & P.
and Shivas 2002). Phyllachora retains its uniqueness from Syd., Annls mycol. 18(4/6): 183 (1921) [1920]
most other members of the family by the development of Erikssonia Penz. & Sacc., Malpighia 11(11–12): 526
ascomata embedded between a clypeus and epidermis (1898) [1897]
(Cannon 1991). However, differences in depth of ascomata Type species: Erikssonia pulchella Penz. & Sacc.,
can be influenced by the consistency of the host tissue Malpighia 11(11–12): 526 (1898) [1897]
(Cannon 1991), therefore, this is not a reliable character on Fremitomyces P.F. Cannon & H.C. Evans, Mycol. Res.
which to distinguish genera. 103(5): 585 (1999)
Type species: Fremitomyces punctatus P.F. Cannon &
H.C. Evans, Mycol. Res. 103(5): 587 (1999)
Geminispora Pat., in Patouillard & Lagerheim, Bull. Soc.
Other genera included mycol. Fr. 9: 151 (1893)
Type species: Geminispora mimosae Pat., in Patouillard &
Acerviclypeatus Hanlin, Mycotaxon 37: 380 (1990) Lagerheim, Bull. Soc. mycol. Fr. 9: 151 (1893)
Type species: Acerviclypeatus poriformans Hanlin, Gibellina Pass., in Roumeguère, Revue mycol., Toulouse
Mycotaxon 37: 381 (1990) 8: 177 (1886)
Apiosphaeria Höhn., Sber. Akad. Wiss. Wien, Math.- Type species: Gibellina cerealis (Pass.) Pass., Revue
naturw. Kl., Abt. 1 118: 1218 [62 repr.] (1909) mycol., Toulouse 8: 177 (1886)
Type species: Apiosphaeria guaranitica (Speg.) Höhn. [as Imazekia Tak. Kobay. & Y. Kawabe, Japanese Journal of
‘guarantica’], Sber. Akad. Wiss. Wien, Math.-naturw. Kl., Tropical Agriculture 36(3): 201 (1992)
Abt. 1 118: 1218 [62 repr.] (1909) Type species: Imazekia ryukyuensis Tak. Kobay. & Y.
Ascovaginospora Fallah et al., Mycologia 89(5): 813 Kawabe, Japanese Journal of Tropical Agriculture 36(3):
(1997) 201 (1992)
Type species: Ascovaginospora stellipala Fallah et al., Isothea Fr., Summa veg. Scand., Section Post.
Mycologia 89(5): 813 (1997) (Stockholm): 421 (1849)
Brobdingnagia K.D. Hyde & P.F. Cannon, Mycol. Pap. Type species: Isothea rhytismoides (Bab.) Fr., Summa veg.
175: 47 (1999) Scand., Section Post. (Stockholm): 421 (1849)
Type species: Brobdingnagia nigeriensis (Sivan. & Lichenochora Hafellner, Nova Hedwigia 48(3–4): 358
Okpala) K.D. Hyde & P.F. Cannon, Mycol. Pap. 175: 47 (1989)
(1999) Type species: Lichenochora thallina (Cooke) Hafellner,
Camarotella Theiss. & Syd., Annls mycol. 13(3/4): 370 Nova Hedwigia 48(3–4): 363 (1989)
(1915) Lindauella Rehm, Hedwigia 39: 82 (1900)
Type species: Camarotella astrocaryae (Rehm) Theiss. & Type species: Lindauella pyrenocarpoidea Rehm,
Syd., Annls mycol. 13(3/4): 370 (1915) Hedwigia 39: 82 (1900)
Coccodiella Hara, Bot. Mag., Tokyo 25: 224 (1910) Linochora Höhn., Sber. Akad. Wiss. Wien, Math.-naturw.
Type species: Coccodiella arundinariae Hara, Bot. Mag., Kl., Abt. 1 119: 638 [22 repr.] (1910)
Tokyo 24: 224 (1910) Type species: Linochora leptospermi (Cooke) Höhn., Sber.
Cyclodomus Höhn., Sber. Akad. Wiss. Wien, Math.- Akad. Wiss. Wien, Math.-naturw. Kl., Abt. 1 119: 639 [23
naturw. Kl., Abt. 1 118: 1527 [67 repr.] (1909) repr.] (1910)
Type species: Cyclodomus umbellulariae Höhn., Sber. Lohwagia Petr., Bot. Arch. 43: 205 (1942) [1941]
Akad. Wiss. Wien, Math.-naturw. Kl., Abt. 1 118: 1528 [2 Type species: Lohwagia intermedia (Speg.) Petr., Bot.
repr.] (1909) Arch. 43: 205 (1942) [1941]
Deshpandiella Kamat & Ullasa, in Ullasa & Rao, Bull. Maculatifrondes K.D. Hyde [as ‘Maculatifrondis’],
Torrey bot. Club 100: 41 (1973) Mycol. Res. 100(12): 1509 (1996)
Fungal Diversity

Type species: Maculatifrondes aequatoriensis K.D. Hyde, Phylleutypa Petr., Annls mycol. 32(5/6): 429 (1934)
in Hyde et al., Mycol. Res. 100(12): 1509 (1996) Type species: Phylleutypa dioscoreae (Wakef.) Petr., Annls
Malthomyces K.D. Hyde & P.F. Cannon, Mycol. Pap. 175: mycol. 32(5/6): 429 (1934)
69 (1999) Phyllocrea Höhn., Annls mycol. 16(1/2): 38 (1918)
Type species: Malthomyces calamigena (Berk. & Broome) Type species: Phyllocrea quitensis (Pat.) Höhn., Annls
K.D. Hyde & P.F. Cannon, Mycol. Pap. 175: 70 (1999) mycol. 16(1/2): 38 (1918)
Muelleromyces Kamat & Anahosur, in Anahosur, Polystigma DC., in de Candolle & Lamarck, Fl. franç., Edn
Experientia 24: 849 (1968) 3 (Paris) 6: 164 (1815)
Type species: Muelleromyces indicus Kamat & Anahosur, = Polystigmina Sacc., Syll. fung. (Abellini) 3: 622 (1884)
Experientia 24: 849 (1968) Type species: Polystigma rubrum (Pers.) DC., in de
Mycohypallage B. Sutton, Mycol. Pap. 88: 4 (1963) Candolle & Lamarck, Fl. franç., Edn 3 (Paris) 6: 164
Type species: Mycohypallage congesta (Berk. & Broome) (1815)
B. Sutton, Mycol. Pap. 88: 5 (1963) Pseudothiella Petr., Hedwigia 68(5): 257 (1928)
Neoflageoletia J. Reid & C. Booth, Can. J. Bot.44: 450 Type species: Pseudothiella hirtellae (Henn.) Petr.,
(1966) Hedwigia 68(5): 257 (1928)
Type species: Neoflageoletia bambusina (Syd.) J. Reid & Pseudothiopsella Petr., Hedwigia 68(5): 259 (1928)
C. Booth 1966 Type species: Pseudothiopsella hirtellae Petr., Hedwigia
Ophiodothis (Berk. & M.A. Curtis) Sacc., Syll. fung. 68(5): 259 (1928)
(Abellini) 2: 652 (1883) Pterosporidium W.H. Ho & K.D. Hyde, Can. J.
Type species: Ophiodothis vorax (Berk. & M.A. Curtis) Bot.74(11): 1826 (1996)
Sacc., Syll. fung. (Abellini) 2: 652 (1883) Type species: Pterosporidium rhizophorae (Vizioli) W.H.
Ophiodothella Henn., Hedwigia 43: 258 (1904) Ho & K.D. Hyde, Can. J. Bot. 74(11): 1828 (1996)
Type species: Ophiodothella atromaculans (Henn.) Höhn., Rehmiodothis Theiss. & Syd., Annls mycol. 12(2): 192
Sber. Akad. Wiss. Wien, Math.-naturw. Kl., Abt. 1 119: 940 (1914)
[64 repr.] (1910) Type species: Rehmiodothis osbeckiae (Berk. & Broome)
Orphnodactylis Malloch & A. Mallik, Can. J. Bot.76(7): Theiss. & Syd., Annls mycol. 12(2): 192 (1914)
1267 (1998) Retroa P.F. Cannon, Mycol. Pap. 163: 201 (1991)
Type species: Orphnodactylis kalmiae Malloch & A. Type species: Retroa dimorphandrae (F. Stevens) P.F.
Mallik, Can. J. Bot. 76(7): 1267 (1998) Cannon, Mycol. Pap. 163: 202 (1991)
Oswaldina Rangel, Archivos da Escola Superior de Rhodosticta Woron., Izv. Imp. St.-Peterburgsk. Bot. Sada
Agricultura e Medicina Veterinaria, Nictheroy, Rio de 11: 13 (1911)
Janeiro 5: 37 (1921) Type species: Rhodosticta caraganae Woron., Consp.
Type species: Oswaldina icarahyensis Rangel, Archivos System. Corticiac. (Tartu) 11: 13 (1911
da Escola Superior de Agricultura e Medicina Veterinaria, Rikatlia P.F. Cannon, Syst. Ascom. 11(2): 83 (1993)
Nictheroy, Rio de Janeiro 5: 37 (1921) Type species: Rikatlia lungusaensis (Henn.) P.F. Cannon,
Oxodeora K.D. Hyde & P.F. Cannon, Mycol. Pap. 175: 75 Syst. Ascom. 11(2): 84 (1993)
(1999) Schizochora Syd. & P. Syd., Annls mycol. 11(3): 265
Type species: Oxodeora petrakii (Cif.) K.D. Hyde & P.F. (1913)
Cannon, Mycol. Pap. 175: 75 (1999) Type species: Schizochora elmeri Syd. & P. Syd., Annls
Parberya C.A. Pearce & K.D. Hyde, Fungal Diversity 6: mycol. 11(3): 265 (1913)
90 (2001) Sphaerodothella C.A. Pearce & K.D. Hyde, Fungal
Type species: Parberya kosciuskoa C.A. Pearce & K.D. Diversity 6: 85 (2001)
Hyde, Fungal Diversity 6: 91 (2001) Type species: Sphaerodothella danthoniae (McAlpine)
Petrakiella Syd., Annls mycol. 22(1/2): 230 (1924) C.A. Pearce & K.D. Hyde, Fungal Diversity 6: 86 (2001)
Type species: Petrakiella insignis Syd., Annls mycol. Sphaerodothis (Sacc. & P. Syd.) Shear, Mycologia 1(4):
22(1/2): 230 (1924) 162 (1909)
Phaeochorella Theiss. & Syd., Annls mycol. 13(3/4): 405 Type species: Sphaerodothis arengae (Racib.) Shear ex
(1915) Theiss. & Syd., Annls mycol. 13(5/6): 577 (1915)
Type species: Phaeochorella parinarii (Henn.) Theiss. & Stigmatula (Sacc.) Syd. & P. Syd., Bull. Herb. Boissier, 2
Syd., Annls mycol. 13(3/4): 405 (1915) sér. 1: 77 (1901)
Phycomelaina Kohlm., Phytopath. Z. 63: 350 (1968) Type species: Stigmatula sutherlandiae (Kalchbr. &
Type species: Phycomelaina laminariae (Rostr.) Kohlm., Cooke) Syd. & P. Syd., Bull. Herb. Boissier, 2 sér. 1: 78
Phytopath. Z. 63: 350 (1968) (1901)
 Fungal Diversity

Stigmochora Theiss. & Syd., Annls mycol. 12(3): 272 6. Ascospores flattened on one side, with a narrow oblong
(1914) longitudinal depression in the middle . . Sphaerodothis
Type species: Stigmochora controversa (Starbäck) Theiss. 6. Ascospores not as above . . . . . . . . . . . . . . . . . . . . . . 7
& Syd., Annls mycol. 12(3): 272 (1914) 7. Paraphyses persistent . . . . . . . . . . . . Ascovaginospora
Stromaster (Pat.) Höhn., in Weese, Mitt. bot. Inst. tech. 7. Paraphyses absent . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
Hochsch. Wien 7(3): 93 (1930) 8. Apical ring absent . . . . . . . . . . . . . . . Maculatifrondes
Type species: Stromaster tuberculatus (Pat.) Höhn., in 8. Apical ring conspicuous, not amyloid. . . . . . . Trabutia
Weese, Mitt. bot. Inst. tech. Hochsch. Wien 7(3): 93 (1930) 9. Ascospores mostly filiform . . . . . . . . . . . Ophiodothis
Telimena Racib., Parasit. Alg. Pilze Java’s (Jakarta) 1: 18 9. Ascospores cylindrical, obovoid or fusiform. . . . . . . 10
(1900) 10. Paraphyses present . . . . . . . . . . . . . . . . . . . . . . . . . 11
Type species: Telimena erythrinae Racib., Parasit. Alg. 10. Paraphyses absent . . . . . . . . . . . . . . . . . . . . . . . . . 14
Pilze Java’s (Jakarta) 1: 18 (1900) 11. Ascospores pigmented . . . . . . . . . . . . . . . . . . . . . . 12
Telimenella Petr., in Rechinger et al., Annln K. K. naturh. 11. Ascospores hyaline. . . . . . . . . . . . . . . . . . . . . . . . . 19
Hofmus. Wien 50: 468 (1940) [1939] 12. Immature ascospores hyaline, becoming brown when
Type species: Telimenella persica Petr., Annln K. K. mature . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
naturh. Hofmus. Wien 50: 469 (1940) [1939] 12. Ascospores brown at all stages. . . . . . Deshpandiella
Telimenochora Sivan., Trans. Br. mycol. Soc. 88(4): 474 13. Ascomata flask-shaped, with black, protruding
(1987) beaks . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Gibellina
Type species: Telimenochora abortiva (F. Stevens) Sivan., 13. Ascomata globose without a beak . . . Telimenochora
Trans. Br. mycol. Soc. 88(4): 474 (1987) 14. Ascomata with well-developed clypeus . . . . . . . . . 15
Trabutia Sacc. & Roum., Revue mycol., Toulouse 3(no. 14. Ascomata with poorly developed clypeus or without
9): 27 (1881) clypeus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
Type species: Trabutia quercina (F. Rudolphi ex Fr.) Sacc. 15. Asci readily gelatinizing at maturity, with thick apical
& Roum., Revue mycol., Toulouse 3(no. 9): 27 (1881) canal . . . . . . . . . . . . . . . . . . . . . . . . . Muelleromyces
Tribulatia Joanne E. Taylor et al., in Taylor & K.D.Hyde, 15. Asci cylindric-clavate at maturity . . . . . . . . . . . . . . 16
Fungal Divers. Res. Ser. 12: 189 (2003) 16. Asci bulbous at immature stage . . . . . .Phycomelaina
Type species: Tribulatia appendicospora Joanne E. Taylor 16. Asci not as above. . . . . . Stigmochora, Rehmiodothis
et al., in Taylor & Hyde, Fungal Diversity Res. Ser. 12: 191 17. Ascospores apiosporous . . . . . . . . . . . . . . . .Imazekia
(2003) 17. Ascospores non-apiosporous . . . . . . . . . . . . . . . . . 18
Uropolystigma Maubl., Bull. Soc. mycol. Fr. 36: 36 (1920) 18. Ascospores hyaline . . . . . . . . . . . . . . . . . Phyllocrea
Type species: Uropolystigma atrotestaceum Maubl., Bull. 18. Ascospores pigmented . . . . . . . . . . . . . Malthomyces
Soc. mycol. Fr. 36: 36 (1920) 19. Paraphyses branched . . . . . . . . . . . . . . . . . . . . . . . 20
Vitreostroma P.F. Cannon, Mycol. Pap. 163: 206 (1991) 19. Paraphyses unbranched . . . . . . . . . . . . . . . . . . . . . 23
Type species: Vitreostroma desmodii (Henn.) P.F. Cannon, 20. Persistent periphysoids present . . . . . . . . . . . . . . . . 22
Mycol. Pap. 163: 207 (1991) 20. Periphysoids absent . . . . . . . . . . . . . . . . . . . . . . . . 21
Zimmermanniella Henn., Hedwigia 41: 142 (1902) 21. Stroma pseudostromatic, . . . . . . . . . . . . Phyllachora
Type species: Zimmermanniella trispora Henn., Hedwigia 21. Stroma eustromatic. . . . . . . . . . . . . . . . . .Coccodiella
41: 142 (1902) 22. Ascospores asymmetrically 3-septate, often constricted
in the middle . . . . . . . . . . . . . . . . . . . . . . . .Telimena
22. Ascospores 2-septate, not constricted at the
Key to genera of Phyllachoraceae septa. . . . . . . . . . . . . . . . . . . . . . . . . . . . Telimenella
23. Ascospores with appendages . . . . . . . . . Schizochora
1. Ascomycete . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 23. Ascospores without appendages . . . . . . . . . . . . . . . 24
1. Coelomycete . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35 24. Ascomata with well-developed
2. Ascospores aseptate . . . . . . . . . . . . . . . . . . . . . . . . . . 3 clypeus . . . . . . . . . . . . . . . . . Diatractium
2. Ascospores 1- multi-septate . . . . . . . . . . . . . . . . . . . . 9 24. Ascomata with poorly developed clypeus or without
3. Ascospores hyaline . . . . . . . . . . . . . . . . . . . . . . . . . . 4 clypeus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25
3. Ascospores pigmented . . . . . . . . . . . . . . . . . . . . . . . 28 25. Ascospores 3-septate . . . . . . . . . . . . . . . . Petrakiella
4. Ascospores with appendages and/or sheaths. . . . . . . . 5 25. Ascospores uni septate . . . . . . . . . . . . . . . . . .Rikatlia
4. Ascospores without appendages or sheaths. . . . . . . . 26 26. Ascospores ovoid to globose or cylindrical-ellipsoid to
5. Ascospores with appendages and ellipsoid or fusiform . . . . . . . . . . . . . . . . . . . . . . . . 27
sheath . . . . . . . . . . . . . . . . . . Apiosphaeria 26. Ascospores filiform . . . . . Linochora, Ophiodothella
5. Ascospores without appendages but, with sheaths . . . 6 27. Paraphyses inconspicuous . . . . . . . . . . . . . . . . . . . 28
Fungal Diversity

27. Paraphyses well-developed. . . . . . . . . . . . . . . . . . . 29 solitary or occurring in small groups, subhyaline to pale

28. Ascomata usually clearly distinguishable from the sur- brown to black, immersed to superficial, papillate or with a
face even in old colonies, from the surface usually long, hyaline or black, upright or obliquely lying neck,
strongly domed, ostiole inconspicuous . . . . . . Isothea ostiolate, periphysate. Peridium leathery to fragile, com-
28. Ascomata immersed in host mesophyll between prising two layers, partly carbonaceous in the outer layers.
vascular bundles, often occupying the entire leaf Hamathecium comprising abundant, persistent, cylindri-
thickness, globose to irregularly shaped, occasion- cal, septate, hyaline paraphyses. Asci 8-spored,
ally almost stellate, with a central to slightly off- unitunicate, cylindrical-clavate, pedicellate, persistently
centre, ostiolate with conical ostiolar attached to the ascogenous hyphae at maturity, with a
canal. . . . . . . . . . . . . . . . . Sphaerodothella pronounced J+, or J-, apical ring. Ascospores overlapping
29. Ascomata with well-developed uniseriate to biseriate, hyaline, fusiform, cylindrical to
clypeus . . . . . . . . . . . . . . . Brobdingnagia cymbiform, transversely multi-septate, lacking a mucilag-
29. Ascomata with poorly develop/without clypeus . . . 30 inous sheath or appendages, with or without guttules.
30. Leaf spots cream buff to deep olive buff. . Erikssonia Asexual morph: Undetermined.
30. Leaf spots dark grey or brown to black . . . . . . . . . 31 Type genus: Pisorisporium Réblová & J. Fourn.
31. Ascomata flask shaped. . . . . . . . . . . . . . . . . . . . . . 32 Notes: The family Pisorisporiaceae was introduced by
31. Ascomata globose to sub-globose. . . . . . . . . . . . . . 33 Réblová et al. (2015) based on morphology and analysis of
32. Paraphyses filamentous, septate, disintegrating at ma- combined LSU, SSU and RPB2 sequence data and presently
turity to form a dense amorphous includes the genera Achroceratosphaeria and Pisorisporium.
matrix . . . . . . . . . . . . . . . . . . . . . . . . Orphnodactylis Pisorisporiaceae shows a sister relationship with the orders
32. Paraphyses longer than the asci, thin-walled, slightly Koralionastetales and Lulworthiales, but is weakly supported
tapered. . . . . . . . . . . . . . . . . . . . . . . . . Fremitomyces by Bayesian and maximum likelihood analyses. Although
33. Ascomata embedded in a stroma . . . . . . . . . . . . . . 34 currently placed in Hypocreomycetidae, Réblová et al.
33. Ascomata not embedded in a stroma . . . . Lindauella (2015) suggested it represents a new lineage of aquatic fungi
34. Stromata unilocular . . . . . . . . . . . . . . . . Camarotella and that the three orders might represent a novel sub-class.
34. Stromata bi- to multi-loculate . . . . . . . . . . . . . . . . . 35 The taxa in this family are found in Europe and Asia.
35. Stromata epiphyllous . . . . . . . Lohwagia, Polystigma, Réblová et al. (2010) introduced the genus
Phylleutypa, Pseudothiella, Retroa, Rhodosticta, Achroceratosphaeria based on analysis of combined LSU
Vitreostroma and SSU sequence data. Achroceratosphaeria formed a
35. Stromata hypophyllous. . . . . . . . . .Zimmermanniella sister clade to Lulworthiales and Koralionastetales, both
36. Present only one type of conidia. . . . . . . . . . . . . . . 37 orders being predominantly marine. In Réblová et al.
36. Present macro and micro conidia . . . . Phaeochorella (2015), Achroceratosphaeria lies as a sister clade to
37. Conidia with appendages . . . . . . . . . . . . . . . . . . . . 40 P i so r i s p o ri u m i n t h e fa m i l y P i s o r i s p o r i a ce a e .
37. Conidia without appendages. . . . . . . . . . . . . . . . . . 38 Achroceratosphaeria has hyaline to pale brown
38. Conidiophores present . . . . . . . . . . . . . . . . . . . . . . 39 ascomata, membranous ascomata, and asci with J- apical
38. Conidiophores absent . . . . . . . . . . . . . . . Cyclodomus ring as compared to black, carbonaceous ascomata with
39. Conidiogenous cells slender, tapering towards the J+ apical ring in Pisorisporium.
apex . . . . . . . . . . . . . . . . . . . . . . . . Acerviclypeatus
39. Conidiogenous cells lageniform to Pisorisporium Réblová & J. Fourn., Persoonia 34: 45 (2015)
cylindrical . . . . . . . . . . . . . . . . . . . . . . Oswaldina Facesoffungi number: FoF 02127; Fig. 114
40. Conidia with apical and basal Saprobic on submerged wood in freshwater habitats.
appendages . . . . . . . . . . . . . . . . . . Mycohypallage Ascomata immersed, gradually becoming superficial, solitary
40. Conidia with an apical appendage . .Pseudothiopsella or in groups, glabrous, subglobose to conical, with short
periphysate neck. Peridium carbonaceous, 2-layered.
Note: Some of genera are kept together under one character Hamathecium comprising of persistent, hyaline, septate pa-
and some genera are not included in the tree due to lack of data raphyses. Asci unitunicate, 8-spored, cylindrical-clavate,
to further separate them. short-stipitate, with a pronounced thimble-shaped J+ apical
ring. Ascospores fusiform, cylindrical to cymbiform, some-
Pisorisporiaceae Réblová & J. Fourn., Persoonia 34: 43 times falcate, hyaline, transversely multi-septate, smooth-
(2015). walled, with numerous guttules. Asexual morph:
Facesoffungi number: FoF 01286 Undetermined.
Saprobic on submerged, woody substrates, in freshwater Type species: Pisorisporium cymbiforme Réblová & J.
habitats. Sexual morph: Ascomata perithecial, astromatic, Fourn., Persoonia 34: 45 (2015)
 Fungal Diversity

Other genus included a single species Chordomyces antarcticum Bilanenko

et al. in the Plectosphaerellaceae. Currently, nine genera
Achroceratosphaeria Réblová, et al., Fungal Diversity are recognized in the Plectosphaerellaceae including two
43: 79 (2010) h o l o m o r p h i c g e n e r a v i z . P l e c t o s p h a e re l l a a n d
Type species: Achroceratosphaeria potamia Réblová Sodiomyces. Only the asexual morphs are known for the
et al., Fungal Diversity 43: 79 (2010) remaining genera viz. Acrostalagmus, Chordomyces,
Gibellulopsis, Lectera, Musicillium, Stachylidium and
Verticillium (Kirk et al. 2008; Zare et al. 2007; Réblová
Key to genera of Pisorisporiaceae et al. 2011; Cannon et al. 2012a; Wijayawardene et al.
2012; Grum-Grzhimaylo et al. 2013; Hirooka et al.
1. Ascomata hyaline, membranous; apical ring J- 20 14; Maha rach chiku m bura e t al. 20 15; G rum -
……………………………………Achroceratosphaeria Grzhimaylo et al. 2016). The family is referred to the
1. Ascomata black, carbonaceous; apical ring J+ order Glomerellales.
…………………………………….…Pisorisporium
Plectosphaerella Kleb., Phytopath. Z. 1: 43 (1929)
Plectosphaerellaceae W. Gams et al., Nova Hedwigia 85(3– Facesoffungi number: FoF 01335, Figs 114, 115, 116
4): 476 (2007) Saprobic on dead plant m aterial, in soil or
Facesoffungi number: FoF 01334 pathogenic on plants causing root and collar rots.
Saprobic on dead plant material, in soil or pathogenic Sexual morph: Ascomata perithecial, superficial, pale
on plants causing root and collar rots. Sexual morph: brown or dark brown, subglobose to pyriform, thin-
Ascomata perithecial or cleistothecial, superficial, dark walled, with elongated neck. Peridium multi-layered,
brown, subglobose to pyriform or globose, with elongate composed of dark brown cells of textura angularis.
neck. Peridium multi-layered, composed of dark brown Paraphyses conspicuous in young stages or absent.
cells of textura angularis. Paraphyses conspicuous in Asci 8-spored, unitunicate, clavate, without an apical
young ascomata or absent. Asci 8-spored, unitunicate, ring, J−. Ascospores irregularly arranged, hyaline, fusi-
clavate or saccate, without an apical ring. Ascospores form, both ends rounded, 1–2-celled, wall asperulate.
irregularly arranged, hyaline or pale brown, fusiform or Asexual morph: Hyphomycetous. Conidiophores vege-
ellipsoidal or ovoid, 1-celled or 2-celled, wall asperulate tative hyphae, hyaline or centrally pale brown or pale
or smooth. Asexual morph: Hyphomycetous. olivaceous, solitary, sometimes branched, apical or lateral
Conidiophores vegetative hyphae, hyaline or centrally with verticillate or single conidiogenous. Conidiophores
pale brown or pale olivaceous. Conidiophores cells cells phialidic, determinate, discrete, hyaline, thin-
phialidic, hyaline, some become orange-brown or oliva- walled, smooth. Conidia aggregated in slimy heads, hy-
ceous-brown, partly melanized or single. Conidia aggre- aline, bilaterally symmetric, oblong-ellipsoidal, slightly
gated in slimy heads, hyaline to pale olive, shapes vari- curved, 0–1-celled, smooth-walled.
able, ranging from subglobose to oval or cylindrical, Type species: Plectosphaerella cucumeris Kleb.,
slightly curved, or fusiform with pointed ends, 1–2- Phytopath. Z. 1: 43 (1929)
celled, smooth-walled (Description based on Uecker Notes: Plectosphaerella cucumerina (Lindfors) W.
1993 and Zare et al. 2007) Gams is a holomorphic fungus (Zare et al. 2007;
Type: Plectosphaerella Kleb. Carlucci et al. 2012; Arzanlou et al. 2013a, b). The sex-
Notes: The family Plectosphaerellaceae was intro- ual morph Plectosphaerella was described first by
duced by Zare et al. (2007), typified by Plectosphaerella Klebahn in 1929 (Palm et al. 1995). Elbakyan (1970)
(Kirk et al. 2008; Zare et al. 2007). In a phylogenetic considered P. cucumeris conspecific with Venturia
analysis based on combined LSU SSU and RPB2 cucumerina Lindf. (Carlucci et al. 2012). It was regarded
s e q u e n c e d a t a , R é b l o v á e t a l . ( 2 0 11 ) i n c l u d e d as a member of the Hypocreales, until Uecker (1993)
Stachylidium in the family Plectosphaerellaceae. Based suggested to place it in Sordariaceae (Uecker 1993;
on analysis of ITS and GAPDH sequence data, Cannon Palm et al. 1995; Liu et al. 2013; Arzanlou et al.
et al. (2012b) reported that Lectera belongs in the family 2013a, b; Carlucci et al. 2012). Palm et al. (1995) firstly
Plectosphaerellaceae rather than the order Hypocreales. linked Plectosphaerella to the hyphomycetous genus
Grum-Grzhimaylo et al. (2013) introduced Sodiomyces Plectosporium. Zare et al. (2007) introduced a new fam-
as the second holomorphic, alkaliphilic genus within the ily Plectosphaerellaceae based on Plectosphaerella.
Plectosphaerellaceae based on a multi-locus phylogeny Plectosphaerella has been suggested as the accepted ge-
(ITS, LSU, SSU, RPB2 and TEF). Grum-Grzhimaylo neric name because it was published earlier than
et al. (2016) established a new genus Chordomyces with Plectosporium (Carlucci et al. 2012). Plectosphaerella
Fungal Diversity

Fig. 114 Pisorisporium

cymbiforme (redrawn from
Réblová et al. 2015) a Ascomata
b Peridium c Paraphyses d Asci e
Ascospores. Scale bars:
a = 200 μm, b = 20 μm, c–
e = 10 μm

is characterized by off-white to pale yellow, or pale gray- Type species: Acrostalagmus cinnabarinus Corda, Icon.
brown colonies, 1–2-celled, ellipsoidal conidia and deter- fung. (Prague) 2: 15 (1838)
minate phialides (Carlucci et al. 2012). Chordomyces E.N. Bilanenko et al., Fungal Diversity 76:
31 (2015)
Type species: Chordomyces antarcticum E.N. Bilanenko
Other genera included et al., Fungal Diversity 76: 31 (2015)
Gibellulopsis Bat. & H. Maia, Anais Soc. Biol.
Acrostalagmus Corda, Icon. fung. (Prague) 2: 15 (1838) Pernambuco 16(1): 153 (1959)
 Fungal Diversity

Fig. 115 Plectosphaerella

cucumeris (redrawn from the
Uecker 1993) a Ascomata b
Section of ascomata c-g Asci h-l
Ascospores. Scale bars: a,
b = 50 μm, c–g = 10 μm, h–
l = 5 μm

Type species: Gibellulopsis piscis Bat. & H. Maia, Anais Stachylidium Link, Mag. Gesell. naturf. Freunde, Berlin
Soc. Biol. Pernambuco 16(1): 154 (1959) 3(1–2): 15 (1809)
Lectera P.F. Cannon, in Cannon et al., MycoKeys 3: 28 (2012) Type species: Stachylidium bicolor Link, Mag. Gesell.
Type species: Lectera colletotrichoides (J.E. Chilton) P.F. naturf. Freunde, Berlin 3(1–2): 15 (1809)
Cannon, in Cannon et al., MycoKeys 3: 28 (2012) Verticillium Nees, Syst. Pilze (Würzburg): 57 (1816)
Musicillium Zare & W. Gams, in Zare et al., Nova [1816–17]
Hedwigia 85(3–4): 482 (2007) Type species: Verticillium dahliae Kleb., Mykol. Zentbl. 3:
Type species: Musicillium theobromae (Turconi) Zare & 66 (1913)
W. Gams, Nova Hedwigia 85(3–4): 482 (2007)
Sodiomyces A.A. Grum-Grzhimaylo et al., Persoonia,
Mol. Phyl. Evol. Fungi 31: 154 (2013) Key to genera of Plectosphaerellaceae
Type species: Sodiomyces alkalinus (Bilanenko & M.
Ivanova) A.A. Grum-Grzhimaylo et al., Persoonia, Mol. 1. Ascomycete . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
Phyl. Evol. Fungi 31: 157 (2013) 1. Hyphomycete. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
Fungal Diversity

Fig. 116 Plectosphaerella

cucumeris (redrawn from
Carlucci et al. 2012, Pascoe et al.
1984 and Palm et al. 1995) a
Conidiophore b, d Phialides c
Hyphal coil with phialides e-j
Conidia. Scale bars: a, c = 10 μm,
b, d–j = 5 μm

2. Ascomata perithecial . . . . . . . . . . . . . Plectosphaerella 7. Phialides, tapering to the apex without

2. Ascomata cleistothecial . . . . . . . . . . . . . . .Sodiomyces collarette . . . . . . . . . . . . . . . . . . . . . . . . .Verticillium
3. Conidiophores basitonously verticillate . . . . . . . . . . . 4 8. Collarette with a sinuous apex, conidia bilaterally sym-
3. Conidiophores dichotomous or multi-branched metric, appear slightly curved in side
verticillate. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 view. . . . . . . . . . . . . . . . . . . . . . . . . . Plectosphaerella
4. Conidiogenous cell narrowly 8. Collarettes inconspicuous, conidia ellipsoidal to
ellipsoidal . . . . . . . . . . . . . . . . Stachylidium cylindrical . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
4.Conidiogenous cell tapering towards the 9. Phialides, in whorl of 3–6, slender aculeate and hardly
apex. . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Sodiomyces tapering towards the tip . . . . . . . . . . . . . . Musicillium
5. Conidiogenous cell cylindrical . . . . . . . . . . . . . Lectera 9. Phialides, in whorl of 1–3 . . . . . . . . . . . . . . . . . . . . 10
5. Conidiogenous cell phialidic . . . . . . . . . . . . . . . . . . . 6 10. Phialides 28–30 μm, often proliferating sympodially,
6. Conidiogenous 4–5 whorled branches at a node, separate tapering from 2.5–3 μm near the base to 0.8–1 μm at
with warty tips . . . . . . . . . . . . . . . . . . . Acrostalagmus the tip . . . . . . . . . . . . . . . . . . . . . . . . . Chordomyces
6. Conidiogenous tapering to the apex . . . . . . . . . . . . . . 7 10. Phialides 23–50 μm, aculeate, tapering from 1–2 μm
7. Phialides, tapering to the apex with collarette . . . . . . . 8 near the base to 0.5–1 μm at the apex . .Gibellulopsis
 Fungal Diversity

Pleurostomataceae Réblová et al., Stud. Mycol. 50: = Pleurostomophora Vijaykr.et al.

540 (2004) Type species: Pleurostoma candollei Tul. & C. Tul. [as
Facesoffungi number: FoF 01136 ‘candollii’], Select. fung. carpol. (Paris) 2: 247 (1863)
Saprobic on wood in terrestrial habitats. Sexual morph: Notes: Tulasne and Tulasne (1863a) introduced the type
Ascomata perithecial, black, gregarious or scattered, im- species, Pleurostoma candollei. Maharachchikumbura et al.
mersed to superficial, globose to subglobose, coriaceous, (2015), based on a multi-gene dataset, showed that
smooth, without setae, papillate. Peridium thick (30– Pleurostoma and Pleurostomophora formed a highly support-
100 μm), composed of two to four layers, outer layer com- ed clade in the Pleurostomataceae.
prising of brown cells of textura intricata or epidermoidea,
thick, coriaceous; inner layer comprising of hyaline cells of Pseudomassariaceae Senanayake et al., in Senanayake et al.,
textura prismatica or angularis, thin, membranaceous; some- Fungal Diversity 73: 132 (2015)
times middle layers comprised of two types of cells, forming Facesoffungi number: FoF 00843
thin-walled and brown cells of textura epidermoidea to in the Saprobic on recently dead twigs attached to the trees,
outer part, merging with thin-walled and dark brown cells of appearing as black dots, below small raised areas of bark,
textura angularis in the inner part, thick, coriaceous. opening through long fissures. Sexual morph: Ascomata
Hamathecium composed of numerous, hyaline, filamentous perithecial, scattered, solitary or aggregated, immersed, de-
paraphyses or paraphyses absent, with hyaline spicate pressed globose to ellipsoid, coriaceous, black, ostiolate, pa-
ascogenous hyphae in clusters. Asci multi-spored, pillate. Ostiolar papilla short, cylindrical, wide at the apex,
unitunicate, reniform to oblong, with short pedicel or periphysate. Peridium comprising strongly compressed, nar-
sessile, apical ring lacking. Ascospores crowded, hya- row, light to medium brown, thin-walled cells of textura
line, oblong to allantoid, curved, aseptate, smooth- angularis. Hamathecium comprising numerous, apically nar-
walled. Asexual morph: Hyphomycetous. Mycelium row, basally wide, hyaline paraphyses. Asci 8-spored,
composed of branched, septate, hyaline or brown hy- unitunicate, clavate to fusoid, short pedicellate, apically
phae. Conidiophores single, from aerial or submerged rounded, with J+, or J– apical ring. Ascospores biseriate or
hyphae, hyaline, straight or flexuous, aseptate to 2-sep- partially uniseriate, hyaline, broadly ellipsoid, oblong or nar-
tate, tuberculate or smooth. Conidiogenous cells rowly clavate, mostly apiosporous, with a rounded to
monophialidic or polyphialidic, cylindrical, hyaline, subconical, small, lower cell or equally uniseptate, straight
smooth-walled. Conidia aggregated in round, hyaline, or curved, thick-walled, smooth-walled. Asexual morph:
aseptate, oblong to allantoid, curved or straight, smooth, Hyphomycetous, Setae erect, dark brown, straight to
with or without guttules. flexuous, arising from superficial hyphae, branched at
Type: Pleurostoma Tul. & C. Tul. base, subcylindrical, tapering to obtuse apex, 3–7-sep-
Notes: The monotypic family Pleurostomataceae was intro- tate, basal cell slightly swollen. Conidiophores
duced by Réblová (2004). It was placed in Calosphaeriales subcylindrical to setiform, with radially lobed basal
based on allantoid ascospores, ascogenous hyphae and other cells, pale brown, smooth, arising from superficial my-
characteristics similar to Calosphaeriaceae, as well as SSU celium, straight to flexuous, 1–10-septate.
nrDNA and LSU nrDNA sequence data, based on Conidiogenous cells terminal or lateral, polyblastic,
Pleurostoma ootheca (Berk. & M.A. Curtis) M.E. Barr subcylindrical to somewhat clavate, pale brown, smooth,
(Réblová et al. 2004). Berlese (1900) introduced with 1–4 denticulate loci. Conidia lageniform, distal end
Neoarcangelia via N. ootheca (Berk. & M.A. Curtis) Berl., free, truncate, rostrate at proximal end, pale brown, with
based on upright papillae on the ascomata, but Barr (1985) a subhyaline transverse band at equatorial zone, smooth,
maintained there were insufficient features to separate the spe- guttulate, aseptate.
cies from Pleurostoma candollei Tul. & C. Tul. Neoarcangelia Type: Pseudomassaria Jacz.
was accepted as the synonym of Pleurostoma (von Höhnel Notes: Single gene sequence analyses of ITS and LSU
1918; Barr 1985). Shear (1937) reexamined Sphaeria ootheca, regions by Jaklitsch and Voglmayr (2012) indicate a separate
the synonym of P. ootheca from Virginia, and decided that the taxonomic clade grouping for Pseudomassaria and
species was similar with P. candollei. However, P. ootheca was Leiosphaerella. However, they did not formally establish this
included as the second species, and Pleurostomophora was family. Leiosphaerella also shows molecular and morphological
accepted as the asexual morph of Pleurostoma based on mor- similarities with Pseudomassaria (Jaklitsch and Voglmayr
phological and phylogenetic analyses (Réblová et al. 2004; 2012). Pseudomassariaceae was introduced by Senanayake
Vijaykrishna et al. 2004; Najwa et al. 2012). et al. (2015) to accommodate Pseudomassaria and
Leiosphaerella. This family is typified by Pseudomassaria Jacz.
Pleurostoma Tul. & C. Tul., Select. fung. carpol. (Paris) 2: Pseudomassaria Jacz., Bull. Herb. Boissier 2: 663 (1894)
247 (1863); Figs 117, 118 Facesoffungi number: FoF 00843; Fig. 119
Fungal Diversity
 Fungal Diversity

ƒFig. 117 Pleurostoma candollei (Material examined: FRANCE, assigned to the family Hyponectriaceae and introduced a
Chaville, on dead wood of Quercus sp., 8 March 1859, PC new species P. huwerae (Hyde et al. 1998). This was followed
PC0167640). a Material label b Herbarium material c Ascomata d-e
Ascoma cross section f Peridium g Outer layer cross section h Middle
by Lumbsch and Huhndorf (2010) and Maharachchikumbura
and inner layers cross section i-l Asci m-q Ascospores. Notes: Figs f-q et al. (2015). Combined gene analysis by Senanayake et al.
soaked in 3 % KOH. Scale bars: c = 200 μm, d–e = 100 μm, f = 20 μm, g– (2015), shows the taxonomic placement of Pseudomassaria
h = 10 μm, i–l = 5 μm, m–q = 1 μm as different from Hyponectriaceae. This result confirms the
taxonomic placement suggested by Jaklitsch and Voglmayr
(2012).
Saprobic on dead twigs attached to trees, appearing as
black dots, below small bumps on the bark. Sexual morph:
Ascomata perithecial, scattered, solitary or aggregated, im- Other genus included
mersed, depressed globose, coriaceous, black, ostiolate, papil-
late. Ostiolar papilla short, cylindrical, wide at the apex, even Leiosphaerella Höhn., Sber. Akad. Wiss. Wien, Math.-
with the bark surface, periphysate. Peridium comprising nar- naturw. Kl., Abt. 1 128: 579 (1919)
row, compressed, light brown, thin-walled, cells of textura Type species: Leiosphaerella praeclara (Rehm) Höhn.,
angularis at the base and thick-walled, light brown to hyaline, Sber. Akad. Wiss. Wien, Math.-naturw. Kl., Abt. 1 128: 579
isodiametric cells around the ostiole. Hamathecium compris- (1919)
ing numerous, apically narrow, basally wide, hyaline paraph-
yses. Asci 8-spored, unitunicate, clavate to fusoid, short ped-
icellate, apically rounded, with a J+ or J– apical ring. Key to genera of Pseudomassariaceae
Ascospores biseriate or partially uniseriate, hyaline, broadly
ellipsoid, oblong or narrowly clavate, apiosporous with a 1. Ascospores apiosporous . . . . . . . . . . . . . Pseudomassaria
rounded to subconical small lower cell, straight or curved, 1. Ascospores elongate, bicelled. . . . . . . . Leiosphaerella
thick-walled, smooth-walled. Asexual morph:
Hyphomycetous, Setae erect, dark brown, straight to flexuous, P s e u d o p l a g i o s t o m a t a c e a e C h e e w. e t a l . [ a s
arising from superficial hyphae, branched at base, ‘Pseudoplagiostomaceae’], in Cheewangkoon et al., Fungal
subcylindrical, tapering to obtuse apex, 3–7-septate, basal cell Diversity 44: 95 (2010)
slightly swollen. Conidiophores subcylindrical to setiform, Facesoffungi number: FoF 01403
with radially lobed basal cells, pale brown, smooth, arising Pathogen on leaves, forming spots. Sexual morph:
from superficial mycelium, straight to flexuous, 1–10-septate. Ascomata solitary, scattered, immersed, slanted to horizontal
Conidiogenous cells terminal or lateral, polyblastic, on host tissue, globose or elliptical, black, coriaceous, papil-
subcylindrical to somewhat clavate, pale brown, smooth, with late, ostiolate. Papilla short, internally covered with hyaline,
1–4 denticulate loci. Conidia pale brown, lageniform to rhom- filamentous, periphyses. Peridium comprising a few layers of
boid, distal end free, proximal end rostrate, with a subhyaline thick-walled, brown cells of textura angularis. Hamathecium
transverse band at equatorial zone, smooth-walled, guttulate, lacking paraphyses. Asci 8-spored, unitunicate, cylindrical,
aseptate. sessile, with J-, subapical ring. Ascospores overlapping
Type species: Pseudomassaria chondrospora (Ces.) Jacz., uniseriate to biseriate, hyaline, fusiform to ellipsoid, 1-septate,
Bull. Herb. Boissier 2: 663 (1894) with terminal, elongate, hyaline appendages. Asexual morph:
Notes: Pseudomassaria was introduced by Jaczewski Coelomycetous. Conidiomata acervular or pycnidial, brown.
(1894) to place Sphaeria chondrospora. This genus was Peridium comprising small, brown cells of textura angularis.
Conidiophores absent. Conidiogenous cells cylindrical to
ampulliform, enteroblastic, percurrently proliferating with
periclinal thickening and collarette. Conidia holoblastic, hya-
line to brown, ellipsoid, unicellular, subglobose to broadly-
allantoid, with obtuse apex and a flat protruding scar at
the base.
Type: Pseudoplagiostoma Cheew. et al.
Notes: The family Pseudoplagiostomataceae was intro-
duced by Cheewangkoon et al. (2010) to accommodate the
type species, P. eucalypti Cheew.et al. and two other new
Fig. 118 Pleurostomophora anamorph of Pleurostroma ootheca
species (P. oldii Cheew. et al. and P. variabile Cheew. et al.)
(From Vijaykrishna et al. 2004). Conidiogenous cells and conidia. Scale isolated from Eucalyptus leaf spots. This monotypic family
bars = 10 μm comprises foliar pathogens and has been recorded only from
Fungal Diversity

Fig. 119 Pseudomassaria chondrospora (Material examined: ITALY, MFLU 15-0729, reference specimen). a Appearance of ascomata on
Province of Forlì-Cesena, Premilcuore, Fiumicello, on branch of Tilia host substrate b Cross section of ascoma c Peridium. d Paraphyses e–h
cordata Mill. (Tiliaceae), 24 April 2013, Erio Camporesi, IT 1200 Asci i–o Ascospores. Scale bars: a = 500 μm, b = 100 μm, c, e–p = 10 μm

Eucalyptus. Ascospore morphology, in particular, is distinct horizontal, globose to elliptical, coriaceous, brown to black,
and morphologically this family differs from other families in papillate, ostiolate. Papilla erumpent, internal wall lined by
the order in having astromatic, slanted to horizontal, globose hyaline periphyses. Peridium comprising a few layers of
ascomata with an aparaphysate hamathecium and ascospores t h i c k - w a l l e d , b r o w n c e ll s o f t e x t u r a a n g u l a r i s .
with terminal, elongate, hyaline appendages. A maximum par- Hamathecium lacking paraphyses. Asci 8-spored, unitunicate,
simony analysis of LSU sequence data of the order subcylindrical to long obovoid, sessile, apex blunt, with
Diap ortha les, sho wed th e d istinct plac emen t of wedge-shaped, J-, subapical ring. Ascospores overlapping
Pseudoplagiostomaceae (Cheewangkoon et al. 2010). uniseriate to biseriate, hyaline, fusiform to ellipsoid, tapering
towards the rounded ends, 1-septate, with terminal, elongate,
Pseudoplagiostoma Cheew. et al., in Cheewangkoon et al., hyaline appendages. Asexual morph: Coelomycetous.
Fungal Diversity 44: 96 (2010) Conidiomata acervular to pycnidial, subcuticular to sub-
Facesoffungi number: FoF 02128; Fig. 120 epidermal. Peridium comprising small, brown cells of
Pathogenic on leaves, forming spots. Sexual morph: textura angularis. Conidiophores absent. Conidiogenous
Ascomata perithecial, immersed in host tissues, slanted to cells cylindrical t o am pulli form , enterobl astic,
 Fungal Diversity

Fig. 120 Pseudoplagiostoma eucalypti (Material examined: Ascospores o Conidiomata p Cross section though conidiomata q, r
VENEZUELA, on living leaves of Eucalyptus urophylla S. T. Blake Conidia attached to conidiogenous cells with percurrent proliferation s
(Myrtaceae), October 2006, M.J. Wingfield, CBS H-20303, holotype). Conidia t Colony on MEA u, v Conidia and conidiogenous cells w, y
a Leaf spot b, c Ascomata d Ascomatal wall e Cross section though Microcyclic conidiation. Scale bars: a = 5 mm, b = 1 mm, c, e = 50 μm,
ascomata f Ostiole g Asci h, i Young asci j Mature ascus k, l Asci d = 5 μm, f–j = 30 μm, k, s–u = 20 μm, o = 200 μm, p = 70 μm, q–s, w–
strained in Melzer’s reagent, showing non-amyloid subapical ring m, n y = 15 μm

proliferating with periclinal thickening. Conidia holo- Ty p e s p e c i e s : P s e u d o p l a g i o s t o m a e u c a l y p t i

blastic, hyaline to brown, ellipsoid, unicellular, with ob- Cheew.et al., in Cheewangkoon et al., Fungal Diversity
tuse apex and a flat protruding scar at the base. 44: 98 (2010)
Fungal Diversity

Notes: The genus Pseudoplagiostoma was introduced to C. lanciformis (Fr.) Voglmayr & Jaklitsch. Coryneum
accommodate a distinct foliar pathogen from Eucalyptus that species are reported to be the asexual morphs of
has astromatic, slanted to horizontal, globose perithecia with- Pseudovalsa (Rossman et al. 2015). Coryneum species
out paraphyses, ascospores with terminal, elongate, hyaline commonly occur on temperate hardwood trees.
appendages and a cryptosporiopsis-like asexual morph. The Maharachchikumbura et al. (2015) also listed
generic type of Cryptosporiopsis, C. nigra (Dermateaceae, Apoharknessia Crous & S.J. Lee under
Helotiales) is not closely related and Cryptosporiopsis Pseudovalsaceae.
eucalypti falls within Diaporthales in the phylogenetic analy- Coryneum Nees, Syst. Pilze (Würzburg): 34 (1816)
ses. Hence Pseudoplagiostoma was introduced to accommo- [1816–17]
date this taxon. Combined gene analysis of ITS and TUB gene = Murogenella Goos & E.F. Morris, Mycologia 57(5):
sequence data resolved the species within Pseudoplagiostoma 776 (1965)
(Cheewangkoon et al. 2010). = Pseudovalsa Ces. & De Not., Comm. Soc. crittog. Ital.
1(4): 206 (1863)
Pseudovalsaceae M.E. Barr, Mycol. Mem. 7: 151 (1978) Facesoffungi number: FoF 02129; Figs 121, 122
Facesoffungi number: FoF 01404 Saprobic or rarely pathogenic on dead wood. Sexual
Saprobic on dead wood. Sexual morph: Stromata solitary, morph: Stromata solitary, erumpent, comprising pseudopar-
erumpent, comprising pseudoparenchymatous cells. enchymatous cells. Ectostromatic disc well or poorly devel-
Ectostromatic disc well or poorly developed, brown to black, oped, brown to black, comprising small cells of textura
comprising small cells of textura prismatica. Ascomata peri- prismatica. Ascomata perithecial, immersed, aggregated, glo-
thecial, immersed, aggregated, globose to subglobose, coria- bose to subglobose, coriaceous, brown to black, papillate,
ceous, brown to black, papillate, ostiolate. Papilla upright, ostiolate. Papilla upright, central, broad, sometimes converg-
central, broad, sometimes converging. Peridium comprising ing. Peridium comprising outer, thick-walled, brown cells of
outer, thick-walled, brown cells of textura angularis and in- textura angularis and inner, thick-walled, hyaline, com-
ner, thick-walled, hyaline, compressed cells of textura pressed cells of textura angularis. Hamathecium comprising
angularis. Hamathecium comprising broad, cellular, septate broad, cellular, septate paraphyses. Asci 8–spored,
paraphyses. Asci 8-spored, unitunicate, ellipsoid to cylindri- unitunicate, ellipsoid to cylindrical, thin-walled, pedicellate,
cal, thin-walled, pedicellate, apex rounded with a J-, apical apex rounded, with J-, apical ring. Ascospores overlapping
ring. Ascospores overlapping uniseriate to biseriate, initially uniseriate to biseriate, initially hyaline, brown at maturity, el-
hyaline, brown at maturity, ellipsoid, fusoid or elongate, one lipsoid, fusoid or elongate, one to several septate, often
to several septate, often distoseptate, end cells light brown or distoseptate, end cells light brown or hyaline, straight or
hyaline, straight or curved. Asexual morph: Coelomycetous. curved. Asexual morph: Conidiomata acervular, solitary,
Conidiomata acervular, solitary, erumpent through the outer erumpent through the outer periderm layers of host, scattered,
periderm layers of host or immersed, scattered, surface tissues surface tissues above slightly domed. Conidiomatal wall
above slightly domed. Conidiomatal wall composed of thin- comprising thin-walled, vertically arranged, dark brown,
walled, vertically arranged dark brown cells of textura cells of textura angularis. Conidiophores branched at
angularis. Conidiophores branched at the base or not, cylin- the base, cylindrical, septate, hyaline at the apex, pale
drical to globose, septate or aseptate, hyaline or hyaline at the brown at the base. Conidiogenous cells formed from the
top, pale brown at the base. Conidiogenous cells formed from apical cell of the conidiophore, hyaline, annellidic, cy-
the apical cells of the conidiophores, hyaline, annellidic, cy- lindrical. Conidia dark brown, curved, broadly fusiform
lindrical, sometimes with setulose apical appendages. Conidia to cylindrical or clavate, smooth-walled, distoseptate
hyaline to dark brown, curved, globose, broadly fusiform to with 4–6 septa, the apical and basal cell darker than
cylindrical or clavate, smooth-walled, unicellular to other cells, apical cell hyaline.
distoseptate, with 4–6 septa, sometimes the apical and basal Type species: Coryneum umbonatum Nees, Syst. Pilze
cell darker than other cells with hyaline apex in apical cell or (Würzburg): 34 (1816) [1816–17]
hyaline, apical apiculus. Basionym: Sphaeria lanciformis Fr., Observ. Mycol.
Type: Coryneum Nees (Havniae) 2: 324 (1818)
Notes: The family Pseudovalsaceae was introduced by Notes: Coryneum was introduced and typified by
Barr (1978) to include diaporthalean taxa having upright, C. umbonatum Nees. Type species of Coryneum,
erumpent ascomata with central necks. However, many gen- C. umbonatum has been linked to the type species of
era previously included in Pseudovalsaceae have been placed Pseudovalsa, P. lanciformis (Rossman et al. 2015) and
in other families (Castlebury et al. 2002) and the only genus Coryneum (1816) has been conserved over Pseudovalsa
r em ai n i n g i n t h e f am i l y P s e u d ov a l s ac e a e w a s (1863) giving priority to the older name (Rossman et al.
Coryneum. This genus is based on the type species 2015). Both sexual and asexual morphs of Coryneum
 Fungal Diversity

Fig. 121 Coryneum lanciformis - sexual morph (Material examined: stroma e Longitudinal section of stroma f Longitudinal section of
USA, Maryland, Beltsville, Narragansett Parkway, on stem of Betula sp., ascoma g Peridium h–j Asci k Paraphyses l-o Ascospores. Scale bars:
7 June 2002, L. Vasilyeva BPI 871006A) a Herbarium packet b c = 1 mm, d = 200 μm, e, f = 100 μm, g = 20 μm, h–j = 20 μm, k–
Herbarium specimen c Stromata on substrate d Transverse section of o = 10 μm

cause plant disease, such as Coryneum-blight on leaves, Other genus included

twigs and buds of cherry, stone fruits, peach and
apricot. Index Fungorum (2016) lists 41 epithets under Apoharknessia Crous & S.J. Lee, Stud. Mycol. 50(1): 239
Pseudovalsa within the family Pseudovalsaceae and 159 (2004)
epithets are listed under Coryneum (Index Fungorum Type species: Apoharknessia insueta (B. Sutton) Crous &
2016). S.J. Lee, in Lee et al., Stud. Mycol. 50(1): 240 (2004)
Fungal Diversity

Fig. 122 Coryneum longipes - asexual morph (UK England, Devon, Conidiomata on substrate e Cross section of conidioma f Conidia
Slapton, near Kingsbridge, Slapton Ley National Nature Reserve, site J1, attached to conidiophores g-i Conidia. Scale bars: c = 1 mm,
on twigs of Quercus petraea (Mattuschka) Liebl. (Fagaceae), 11 d = 500 μm, e = 100 μm, f–j = 10 μm
May 1994, B.C. Sutton, IMI 362542) a, b Herbarium packet c, d

Key to genera of Pseudovalsaceae perithecial, immersed in plant tissue, solitary, scattered

or gregarious, black, surface smooth, coriaceous, brittle,
1. Conidiomata immersed and globose, unicellular conidia with long cylindrical necks, covered in setae. Peridium
with a hyaline, apical apiculus . . . . . . . . . Apoharknessia comprising of several layers of brown textura angularis.
1. Conidiomata erumpent and broadly fusiform to cylindri- Hamathecium comprising septate, hyaline, paraphyses
cal or clavate conidia, with 4–6 distosepta, apical cell intermingled among asci. Asci 8-spored, unitunicate,
hyaline . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Coryneum subcylindrical, short pedicellate, with a large, J+, apical
ring. Ascospores ellipsoid to obclavate, fusiform, often
Pyriculariaceae Klaubauf et al., Studies in Mycology 79: 85– with median cells pigmented, with 3–septa, pale-brown,
120 (2014) lacking a sheath. Asexual morph: Hyphomycetous.
Facesoffungi number: FoF 01801 Conidiophores solitary or in fascicles, subcylindrical,
Pathogenic or saprobic on various plant hosts, commonly erect, olivaceous, pale brown or brown, in some genera
o n m o n oc o t y l e do n s . Se x u al m o rp h : A s c o m a t a branched (Pseudopyricularia and Xenopyricularia) or
 Fungal Diversity

un branc hed (Ba mb usicula ria, Barretomyce s, conidia, which is a specific character of this genus
D e i g h t o n i e l l a , M a c g a r v i e o m y c e s , N e o c o rd a n a , (Bussaban et al. 2005). Klaubauf et al. (2014) synonymized
Neopyricularia, Proxipyricularia and Pyricularia). M. grisea under P. grisea, by clarify the taxonomic relation-
Conidiogenous cells integrated, pigmented, denticulate. ships among species that are magnaporthe- or pyricularia-like
Conidia hyaline to brown, 1–5-transversely septate, using molecular phylogeny (LSU, ITS, RPB1, Actin and
some genera (Neopyricularia, Proxipyricularia and Calmodulin).
Pyricularia) have apical mucoid appendages. This genus includes species that are pathogenic on a wide
Type: Pyricularia Sacc. range of monocotyledonous plants (Klaubauf et al. 2014), and
Notes: The family Pyriculariaceae was introduced by of these P. oryzae Cavara (1892) (synonym: Magnaporthe
Klaubauf et al. (2014), as a sister family to the oryzae) is responsible for the foliar disease of wheat and millet
Ophioceraceae, based on morphological and phylogenetic as well as the cause of the major rice blast disease, which is
analyses. All member of this family cause diseases on highly destructive and causes up to 30 % yield loss worldwide
monocotyledons plants. Klaubauf et al. (2014) accepted nine (Skamnioti and Gurr 2009). Pyricularia grisea is responsible
genera, of which seven were new genera (Bambusicularia, for a foliar disease of Digitaria (Klaubauf et al. 2014).
Barretomyces, Macgarvieomyces, Neopyricularia,
Proxipyricularia, Pseudopyricularia and Xenopyricularia) in-
to Pyriculariaceae based on morphology and molecular anal- Other genera included
ysis of the combined ACT, ITS and RPB1 genes (Klaubauf
et al. 2014). Bambusicularia Klaubauf et al., Stud. Mycol. 79: 104 (2014)
Klaubauf et al. (2014) re-evaluated generic and species Type species: Bambusicularia brunnea Klaubauf et al.
concepts within Pyriculariaceae, and novelties was proposed Stud. Mycol. 79: 104 (2014)
based on morphological and phylogenetic data. The taxonom- Barretomyces Klaubauf et al., Stud. Mycol. 79: 104 (2014)
ic relationships among magnaporthe-like or pyricularia-like Type species: Barretomyces calatheae (D.J. Soares et al.)
species were clarified using phylogenetic analyses of isolates Klaubauf et al., Stud. Mycol. 79: 104 (2014)
representing a wide range of host plants by using partial DNA Deightoniella S. Hughes, Mycol. Pap. 48: 27 (1952)
sequences of multiple genes (LSU, ITS, RPB1, Actin and Type species: Deightoniella africana S. Hughes, Mycol.
Calmodulin). Pap. 48: 27 (1952)
Pyricularia Sacc., Michelia 2(no. 6): 20 (1880) Macgarvieomyces Klaubauf et al., Stud. Mycol. 79: 106
Faces of fungi number: FoF 01802; Fig. 123 (2014)
Pathogenic on plants Sexual morph: Ascomata perithe- Type species: Macgarvieomyces borealis (de Hoog &
cial, immersed in host tissue, solitary and gregarious, Oorschot) Klaubauf et al., Stud. Mycol. 79: 107
subsphaerical, brown to black, with long neck protruding (2014)
above host surface. Peridium comprising several layers of Neocordana Hern.-Restr. & Crous, in Hernández-Restrepo
textura angularis. Paraphyses intermingled among asci, un- et al., Phytotaxa 205(4): 233 (2015)
branched, septate. Asci 8-spored, unitunicate, subcylindrical Type species: Neocordana musae (Zimm.) M. Hern.-Rest.
to clavate, short-stipitate, with prominent, with a large, J+, & Crous, in Hernández-Restrepo et al., Phytotaxa 205(4): 234
apical ring. Ascospores bi- to multi-seriate, hyaline, fusiform, (2015)
curved, with rounded ends, transversely 3-septate, slightly Neopyricularia Klaubauf et al., Stud. Mycol. 79: 108
constricted at septa, guttulate, smooth-walled. Asexual (2014)
morph: Hyphomycetous, Conidiophores solitary or in fasci- Type species: Neopyricularia commelinicola (M.J. Park &
cles, hyaline to pale brown, sub cylindrical, smooth, rarely H.D. Shin) Klaubauf et al., Stud. Mycol. 79: 108 (2014)
branched, with sympodial proliferation. Conidiogenous cells Proxipyricularia Klaubauf et al., Stud. Mycol. 79: 109
terminal and intercalary, pale brown, with denticulate (2014)
conidiogenous loci and rhexolytic secession. Conidia solitary, Type species: Proxipyricularia zingiberis (Y. Nisik.)
pyriform to obclavate, 2-septate, narrowed toward tip, round- Klaubauf et al., Stud. Mycol. 79: 109 (2014)
ed at the base, hyaline to pale brown, with a distinct basal Pseudopyricularia Klaubauf et al., Stud. Mycol. 79: 109
hilum, sometimes with marginal frill (2014)
Type species: Pyricularia grisea Sacc., Michelia 2(no. 6): Type species: Pseudopyricularia kyllingae Klaubauf et al.,
20 (1880) Stud. Mycol. 79: 109 (2014)
Notes: The genus Pyricularia was established by Saccardo Xenopyricularia Klaubauf et al., Stud. Mycol. 79: 116
(1880) with Pyricularia grisea Sacc. (synonym: (2014)
Magnaporthe grisea) as the type species. The name of the Type species: Xenopyricularia zizaniicola (Hashioka)
type genus Pyricularia, refers to the pyriform shape of the Klaubauf et al., Stud. Mycol. 79: 116 (2014)
Fungal Diversity

Fig. 123 Pyricularia grisea a, b,

c Conidiophores and conidia d
Conidia with apical marginal frill,
which is a remnant of the apical
mucoid cap. e Microconidia.
Scale bars: a–e = 10 μm (redrawn
from Klaubauf et al. 2014)

Key to genera of Pyriculariaceae 5. Conidia pale brown . . . . . . . . . . . . . . . . . . . . . . . . . . 7

5. Conidia hyaline (narrowly
1. Conidiophores branched . . . . . . . . . . . . . . . . . . . . . . . . . 2 obclavate) . . . . . . . . . . . . Macgarvieomyces
1. Conidiophores unbranched. . . . . . . . . . . . . . . . . . . . . 3 6. Conidia ellipsoid to obclavate, finely
2. C o n i d i a o b c l a v a t e , f i n e l y roughened . . . . . . . . . . . . . . . . . . . . Bambusicularia
roughened. . . . . . . . . . . . Pseudopyricularia 6. Conidia pyriform to obclavate, narrowed toward tip,
2. Conidia obovoid, narrowed toward rounded at the base . . . . . . . . . . . . . . . . . . . . . . . . . . 8
apex . . . . . . . . . . . . . . . . . . . . . . . . Xenopyricularia 7. Conidia ellipsoid, thin-walled. . . . . . . . . Deightoniella
3. Conidia with 2(−5)-septa . . . . . . . . . . . . . . . . . . . . . . 4 7. Conidia obclavate, thick-walled . . . . . . . . Neocordana
3. Conidia with 1-septa . . . . . . . . . . . . . . . . . . . . . . . . . 5 8. Conidiogenous loci slightly darkened. . Neopyricularia
4. Conidia 4(−5)-septate. . . . . . . . . . . . . . . Barretomyces 8. Conidiogenous loci not darkened . . . . . . . . . . . . . . . . 9
4. Conidia 2-septate . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 9. Conidiophores olivaceous to brown . . Proxipyricularia
 Fungal Diversity

9. Conidiophores pale brown to yellow . . . . . Pyricularia between Porosphaerellopsis and Sporoschismopsis has been
established by Réblová (2014) who synonymized the sexual
Reticulascaceae Réblová & W. Gams, in Réblová et al., Stud. genus Porosphaerellopsis under the asexual genus
Mycol. 68(1): 180 (2011) Sporoschismopsis. The link between Reticulascus and
Facesoffungi number: FoF 01311 Cylindrotrichum has also been established by Réblová and
Saprobic on terrestrial tree branches, wood and bark. Gams (1999) and Maharachchikumbura et al. (2015) recom-
Sexual morph: Stromata minute, sometimes present. mended the use of Cylindrotrichum over Reticulascus, which
Ascomata superficial, gregarious to solitary, brown to black, is followed in this paper.
subglobose to conical, base flattened, slightly verruculose, Kylindria has oblong, longer, wider, 1-several-sepatate, of-
glabrous with minute papilla, ostiole periphysate. Peridium ten asym metrical conidia that are in contrast to
26–39 μm wide, fragile, 2-layered, the whole wall heavily Cylindrotrichum that has cylindrical, narrower, 1-septate,
sclerotized in the upper part, poorly developed towards the symmetrical conidia. Therefore, we follow Réblová (2012)
rim, absent at the base. Hamathecium composed of copious, and keep these two genera separate, even though very Little
1–2 μm wide, filiform, sparsely septate, hyaline paraphyses. information is available on the genus Kylindria.
Asci 8-spored, unitunicate, cylindrical to clavate, short-pedi-
cellate, apex truncate to broadly rounded, with inamyloid, Cylindrotrichum Bonord., Handb. Allgem. mykol.
apical ring. Ascospores uni to bi-seriate, hyaline or dark (Stuttgart): 88 (1851) Fig. 124, 125
brown, septate, mostly 2-celled, with a delayed formation of Synonym: Reticulascus Réblová & W. Gams, in Réblová
the 2 additional septa, smooth-walled, ellipsoidal to fusiform, et al. Stud. Mycol. 68(1): 180 (2011)
sometimes with end pores. Asexual morph: Hyphomycetous. Type species: Cylindrotrichum oligospermum (Corda)
Cylindrotrichum: Setae straight, cylindrical, septate, dark Bonord., Handb. Allgem. mykol. (Stuttgart): 88 (1851)
b r o w n , p a l e r t o w a r d s t h e a p e x . C o n i d i o p h o re s Notes: The genus Reticulascus was introduced by Réblová
macronematous, mononematous, solitary, erect, unbranched, et al. (2011) for the two holomorphic species, namely
2-layered. Conidiogenous cells monophialidic, collarette hya- R. tulasneorum Réblová & W. Gams with the asexual morph
line to subhyaline, disappearing early. Conidia cylindrical, Cylindrotrichum oligospermum (Corda) Bonord and
rounded at the apex, 1-septate, not constricted at the septum, Reticulascus clavatus Réblová & J. Fourn. with the asexual
smooth-walled, hyaline. Kylindria: Conidiophores setiform, morph Cylindrotrichum clavatum W. Gams. The asexual
solitary, erect, sometimes curved, branched, gregarious, morph of Reticulascus generally resembles the dematiaceous,
scattered, septate, thick-walled, dark brown, becoming paler phialidic hyphomycete, linked with Chaetosphaeria, but dif-
towards the apex, smooth-walled. Conidia blastic-phialidic, 1- fers in having cylindrical, 1–septate conidia (Réblová et al.
several-septate, oblong-ellipsoid, apex rounded, tapering to- 2004; Réblová et al. 2011). Cylindrotrichum hennebertii W.
wards the truncate base, hyaline, smooth-walled. Gams & Hol.-Jech. was synonymized under C. oligospermum
Sporoschismopsis: Conidiophores erect, straight to slightly due to their identical ITS sequences (Réblová et al. 2011).
flexuous, un-branched, brown, septate, thick-walled, terminat- Reticulascus tulasneorum produces minute, black, astromatic
ing in a cylindrical to slightly flask-shaped monophialide, ascomata, growing on decaying wood. Reticulascus clavatus
collarette funnel-shaped, which encloses not more than one is common on submerged wood. It differs from Reticulascus
conidium. Conidia clavate, cuneiform, obovate, pyriform to tulasneorum and its asexual morph by having verruculose,
subcylindrical-clavate, distal end bluntly rounded, basal end mature ascospores and in the absence of setae among the
truncate, septate, uniformly brown or with paler distal and/or conidiophores. Physical separation of the conidiogenous
basal cells, sometimes the septa are obscured by a darker band, cells and setae makes Cylindrotrichum setosum Seifert
without or with conspicuous pores at the septa. unique in the genus. Microconidia have also been reported
Type: Cylindrotrichum Bonord., Handb. Allgem. mykol. only from C. setosum. Maharachchikumbura et al. (2015) rec-
(Stuttgart): 88 (1851) ommended that Cylindrotrichum and Reticulascus should be
Notes: The family Reticulascaceae was introduced to ac- treated as congeneric and as the Cylindrotrichum is the older
commodate two holomorphic genera Reticulascus and name, Reticulascus should be synonymized under this genus,
Porosphaerellopsis Samuels & E. Müll., supported by analy- which is followed in this paper.
sis of combined ITS, LSU, SSU and RPB2 sequence data.
Even though the characters and ontogeny of these genera dif-
fer, the centrum and interthecial tissues are quite similar
(Réblová et al. 2011). The asexual dematiaceous hyphomy- Other genus included
cete genera Cylindrotrichum, Kylindria and Sporoschismopsis
are linked with the family (Réblová et al. 2011; Kylindria DiCosmo, S.M. Berch & W.B. Kendr., Mycologia
Maharachchikumbura et al. 2015). The sexual-asexual link 75(6): 970 (1983)
Fungal Diversity

Fig. 124 Reticulascus tulasneorum (CZECH REPUBLIC, South- of the asexual morph on the host c Cells of peridium d Median,
western Bohemia, Javornická, Hornatina Mts, Strašín, near Sušice, on longitudinal section of ascomata with sclerotized lateral wall e
dead branch of Sambucus nigra L. (Adoxaceae), 21 October 1997, M. Interthecial filaments f Asci with ascospores g Immature hyaline ascus
Svrček, (PRM 842978, holotype), Median, longitudinal section of h–i Matured, hyaline asci j–n Hyaline 1–4 septate ascospores. Scale bars:
ascomata of Reticulascus tulasneorum redrawn from Réblová and a–b = 100 μm, c–d = 50 μm, g–I = 25 μm, e–f, j–n = 10 μm
Gams (1999) a Ascomata on the host b Ascomata with conidiophores
 Fungal Diversity

Fig. 125 Different

hyphomycetous asexual
morphs of Reticulascaceae
asexual morph of Reticulascus
tulasneorum redrawn from
Réblová and Gams (1999),
asexual morph Kylindria trisepta
redrawn from DiCosmo et al.
(1983) a Conidiophores and
hyaline conidia of Reticulascus
tulasneorum b Conidiophores
and conidia of Kylindria trisepta c
Conidiogenous cell and conidia
of K. trisepta Scale bars:
b = 50 μm, a, c = 20 μm

Type species: Kylindria triseptata (Matsush.) DiCosmo, conidiogenous cells lining the cavity of the locule.
S.M. Berch & W.B. Kendr., Mycologia 75(6): 971 (1983) Conidiogenous cells discrete, ampulliform to lageniform, hy-
Sporoschismopsis Hol.-Jech. & Hennebert, Bull. Jard. Bot. aline, smooth; proliferating sympodially. Conidia composed
natn. Belg. 42(4): 385 (1972) of a conidium body and a separate apical cell modified into a
Synonym: Porosphaerellopsis Samuels & E. Müll., in branched appendage; conidium body ellipsoid or fusiform, 1-
Müller & Samuels, Sydowia 35: 143 (1982) euseptate, wall smooth, hyaline to pale brown; apical cell di-
Type species: Sporoschismopsis moravica Hol.-Jech. & viding into appendages.
Hennebert, Bull. Jard. Bot. natn. Belg. 42(4): 387 (1972) Type: Robillarda Sacc.
Notes: This is a recently introduced monotypic family
based on ITS sequence data supplemented with HIS sequence
Key to genera of Reticulascaceae data. Robillarda showed a distinct lineage from other taxa in
Xylariales and therefore Crous et al. (2015a) introduced a
1. Conidiophores often seta-like, with a terminal monotypic family Robillardaceae with the Robillarda as the
monophialide at the apex . . . . . . . . . . . . . . . . . . . . . . . 2 type genus.
1. Setae straight, sterile, septate, dark
brown . . . . . . . . . . . . . . . . . . . . . . . Cylindrotrichum Robillarda Sacc., Michelia 2(no. 6): 8 (1880)
2. Conidia hyaline, 1-several septate, oblong- Facesoffungi number: FoF 02130; Fig. 126
ellipsoidal. . . . . . . . . . . . . . . . . . . . . . . . . . Kylindria Saprobic or pathogenic on plants. Sexual morph:
2. Conidia, uniformly brown or with paler distal and/or Undetermined. Asexual morph: Conidiomata pycnidial to
basal cells, 3-septate, clavate-sub-cylindrical pycnidioid, scattered to gregarious, occasionally confluent,
clavate . . . . . . . . . . . . . . . . . . . . . . . Sporoschismopsis immersed to partly erumpent, unilocular to variably loculate
with the locule often convoluted, glabrous; wall thick of
Robillardaceae Crous, in Crous et al., IMA Fungus 6: 184 textura angularis to textura prismatica. Conidiophores re-
(2015) duced to conidiogenous cells or with 1–2 supporting cells
Facesoffungi number: FoF 01912 lining the cavity of the locule. Conidiogenous cells discrete,
Saprobic or pathogenic on plants in terrestrial and aquatic ampulliform to lageniform, hyaline, smooth; proliferating
habitats. Sexual morph: Undetermined. Asexual morph: sympodially or percurrently near apex. Conidia com-
Conidiomata stromatic, pycnidial to pycnidioid or indetermi- posed of a conidium body and a separate apical cell
nate, immersed to partially erumpent, unilocular to convolut- modified into a branched appendage; conidium body
ed, dehiscing by an ostiole or by an irregular split in the apical ellipsoid or fusiform, 1-euseptate, wall smooth and oc-
wall and overlying host tissue; wall thick of textura angularis casionally constricted at the septum, hyaline to pale
to textura prismatica. Conidiophores reduced to brown, often guttulate; apical cell short-cylindrical at
Fungal Diversity

Fig. 126 Robillarda sessilis

(CBS 114312, epitype) (redrawn
from Crous et al. 2015a) a–d
Conidiogenous cells e Conidia.
Scale bars = 10 μm

base, then dividing into 2–5 branches, branches thin- with or without polar mucilaginous pads or appendages.
walled, tubular, ends pointed or swollen, flexuous, di- Asexual morph: hyphomycetous. Hyphae septate, branched,
vergent, smooth, hyaline, devoid of contents. white to pale brown. Conidiogenous cells absent, if present
Type species: Robillarda sessilis (Sacc.) Sacc., Michelia erect, smooth and thick-walled, cylindrical, flask-shaped.
2(no. 6): 8 (1880) Conidia solitary, dry, smooth, applanate or rounded, globose
Notes: There are 38 names referred to Robillarda Sacc. to subglobose or obovate to oval, holoblastic, with 3–6 longi-
(Index Fungorum 2016). However, many of them are still tudinal septa and 4–6 transverse septa or 1–4 transverse septa,
without critical reappraisal and need to be transferred to slightly constricted at the septa, with a pale brown small basal
Chaetoconis Hyalotiella, Neottiospora, Pestalotiopsis and cell, olive green to brown or blackish brown to black, with
Pseudorobillarda (Crous et al. 2015a). The type of prominent scar at the point of secession from the
Robillarda, R. sessilis was isolated from wilted leaves of conidiogenous cell.
Rubus caesius in Italy. Crous et al. (2015a) designated an Type: Savoryella E.B.G. Jones & R.A. Eaton
epitype collected from soil dust in Germany. Notes: Savoryella has been placed in the order Sphaeriales
family incertae sedis by Kohlmeyer and Kohlmeyer (1979),
Savoryellaceae Jaklitsch & Réblová, Index Fungorum 209: 1 ascomycetes incertae sedis by Kohlmeyer (1986), Eriksson
(2015) and Hawksworth (1986), Amphisphaeriaceae Eriksson and
Facesoffungi number: FoF 01283 Hawksworth (1987) and Sordariales (Jones and Hyde
Saprobic on submerged wood in freshwater, marine and 1992). Barr (1990) transferred this genus to the order
brackish habitats. Sexual morph: Ascomata perithecial, Halosphaeriales (now Microascales) based on morphology
brown to black, immersed or superficial, globose to pyriform, (the catenophyses-like paraphyses) and ultrastructural obser-
coriaceous, periphysate, ostiolate, papillate. Papilla central, or vations. Vijaykrishna et al. (2006) and Cai et al. (2006a) re-
eccentric when lying horizontally to the host. Peridium mem- ferred Savoryella elongata and S. longispora to the order
branous, comprising several layers of brown, thick-walled Hypocreales, subclass Hypocreomycetidae, based on a phylo-
cells of textura angularis, hyaline inwardly. Hamathecium genetic analysis of LSU rDNA data, although with weak sta-
comprising hypha-like, numerous or sparse, septate paraphy- tistical support. Jones et al. (2009b) referred the genus to the
ses. Asci 2–8-spored, unitunicate, clavate to cylindrical, pedi- Sordariales genera incertae sedis based on morphological ob-
cellate, persistent, with a small or relatively large, J-, apical servations. Boonyuen et al. (2011) introduced the new order
ring. Ascospores uniseriate or overlapping biseriate, Savoryellales to accommodate the genera Ascotaiwania,
versicolorous with brown middle cells and hyaline end cells, Ascothailandia (and its asexual morph, Canalisporium) and
ellipsoid, fusiform, 3 to many septate, smooth, thick-walled, Savoryella based on multi-gene dataset. The genus
 Fungal Diversity

Ascothailandia was introduced to accommodate a sexual refractive, J-apical ring, cylindrical, persistent.
morph that grouped with the asexual morph Canalisporium, Ascospores uniseriate or overlapping uniseriate, fusiform,
forming a sister clade to Savoryella and Ascotaiwania. straight or curved, 3-euseptate and versicolurus
Subsequently, Jaklitsch (2015) erected the family (Modified description of Sri-indrasutdhi et al. 2010).
Savoryellaceae and this was accepted by Asexual morph: Colonies sporodochial, punctiform,
Maharachchikumbura et al. (2015) and Jones et al. (2015). scattered, glistening, white to yellow when young, dark
Savoryella E.B.G. Jones & R.A. Eaton, Trans. Br. mycol. brown to black when mature. Mycelium immersed,
Soc. 52(1): 161 (1969) consisting of branched, septate, smooth, thin-walled, hy-
Facesoffungi number: FoF 02131; Fig. 127, 128, 129 aline to pale brown hyphae. Conidiophores semi-
Saprobic on submerged wood in cooling towers, riv- macronematous, unbranched, septate, smooth, thin-
ers, streams and marine environments. Sexual morph: walled, hyaline, becoming vesiculate and disintegrating
Ascomata perithecioid, globose, subglobose or ellipsoi- as conidia mature. Conidiogenous cells monoblastic, in-
dal, immersed, partly immersed or superficial, ostiolate, tegrated, terminal, determinate, oblong or often becoming
papillate, membranous, and pale to dark brown, long swollen, hyaline or pale brown. Conidia acrogenous, sol-
necked, brown coloured, periphysate. Peridium brown, itary, brown or pale olive brown, smooth, thick-walled,
composed of several layers of thick-walled angular cells broadly ellipsoidal to obovoid in surface view, cylindri-
forming textura angularis. Paraphyses present, but cal to clavate in lateral view, muriform, with ordered
sparse. Asci 2–8-spored, cylindrical or clavate, short- arrangement of a single, straight or slightly curved column
stalked, unitunicate, persistent, with an apical truncate vertical septum and 2–6 rows transverse septa, slightly con-
non-amyloid apical thickening containing a pore. stricted at the septa; septa becoming darker progressively with
Ascospores uni- or biseriate, ellipsoidal, 3-septate, not conidial maturity; basal cell pale brown, cuneiform, Conidial
markedly constricted at the septa, central cells brown, secession rhexolytic. Modified description of Nawawi and
apical cells smaller and hyaline. Asexual morph: Kuthubutheen (1989).
Undetermined. Type species: Canalisporium caribense (Hol.-Jech. &
Type species: Savoryella lignicola E.B.G. Jones & R.A. Mercado) Nawawi & Kuthubutheen, Mycotaxon 34 (2): 479
Eaton, Trans. Br. mycol. Soc. 52(1): 162 (1969) (1989)
Notes: Jones and Eaton (1969) established the genus Notes: Canalisporium species are saprobes on rotten wood
Savoryella with S. lignicola as the type species. Savoryella and have a pantropical distribution (Goh et al. 1998).
has been placed in the order Sphaeriales family incertae sedis The genus has been recorded in Cuba (Holubová-
by Kohlmeyer and Kohlmeyer (1979), ascomycetes incertae Jechová and Sierra 1984), India (Rao and de Hoog
sedis by Kohlmeyer (1986), 1986), Kenya (Kirk 1985), Malaysia (Nawawi and
Kuthubutheen 1989), Taiwan (Matsushima 1987), and
Uganda (Matsushima 1987). The genus Canalisporium
Other genera included Nawawi & Kuthubutheen (1989) was first introduced
to accommodate Berkleasmium caribense Hol.-Jech. &
Ascotaiwania Sivan. & H.S. Chang, Mycol. Res. 96(6): 481 Mercado, Berklea smiumpulchrum Hol.-Jech. &
(1992) Mercado (Holubová-Jechová and Sierra 1984), and a
Type species: Ascotaiwania lignicola Sivan. & H.S. new species, Canalisporium elegans Nawawi &
Chang, Mycol. Res. 96(6): 481 (1992) Kuthubutheen (1989). Canalisporium species are char-
Canalisporium Nawawi & Kuthubutheen, Mycotaxon acterized by having scattered, punctiform, pulvinate,
34(2): 477 (1989); Fig. 128 granular, black, glistening sporodochia that contain
= Ascothailandia Sri-indr. et al., Mycoscience 51(6): 414 acrogenous, holoblastic conidia developing in a hyaline
(2010) gelatinous sheath (Goh et al. 1998). At present, 12
Saprobic on submerged wood. Sexual morph: species are included under this genus including the
Ascomata immersed to semi-immersed or superficial, recently introduced Canalisporium microsporum G.Z.
perithecial, globose to sub-globose, dark brown to black, Zhao (Zhao et al. 2012). Canalisporium grenadoidia
solitary, scattered, long neck, ostiolate. Ostiole mostly has borne on ascospore isolates of Ascothailandia on
central but if ascomata are lying horizontal to the host the culture (Sri-indrasutdhi et al. 2010). Combined
surface, then at one end and curving upwards, long or analysis of SSU, LSU and ITS showed that seven
short, mostly brown or black, periphysate. Paraphyses Canalisporium species forms a highly supported mono-
hypha like, numerous, tapering distally, not embedded phyletic clade in the Hypocreomycetidae
in a gelatinous matrix. Asci 8-spored, long cylindrical, (Sordariomycetes) together with Ascothailandia. Thus
pedunculate, unitunicate, apically truncate, with a Ascothailandia, which is a recent name, becomes a
Fungal Diversity

Fig. 127 Savoryella lignicola

(Material examined: UK,
Flintshire, Connahis Quay, Test
block, 1966, IMI 129784, IMI
129785, holotype) a Herbarium
specimen b, c Ascomata on host
d, e Ascomata f Paraphyses g–i
Asci j–n Ascospores. Scale bars:
c = 200 μm, d–e = 100 μm, g–
i = 50 μm, j–n = 20 μm

synonym of Canalisporium (Sri-indrasutdhi et al. 2010, Taxonomic position in the family needs verification
Maharachchikumbura et al. 2015).
Monotosporella S. Hughes, Can. J. Bot. 36: 786 (1958) Helicoon Morgan, J. Cincinnati Soc. Nat. Hist. 15: 49 (1892)
Type species: Monotosporella setosa (Berk. & M.A. Type species: Helicoon sessile Morgan, J. Cincinnati Soc.
Curtis) S. Hughes, Can. J. Bot. 36: 787 (1958) Nat. Hist. 15: 50 (1892)
 Fungal Diversity

Fig. 128 Savoryella lignicola

(Material examined:
MALAYSIA, Tioman island,
Mangrove, on dead mangrove
wood, 12 July 2010, K. L. Pang) a
Immersed ascoma b Peridium
composed of a few layers of
angular cells forming textura
angularis c Cylindrical asci d–e
Ascospores with brown central
and hyaline end cells. Scale bars:
a, c = 30 μm, b, d–e = 10 μm

Key to genera of Savoryellaceae hyaline when young, becoming medium brown to dark brown
at maturity, broad or narrowly ellipsoidal, with tapering api-
1. Sexual morph perithecioid . . . . . . . . . . . . . . . . . . . . . . . 4 ces, with or without a longitudinal germ slit.
1. Asexual morph hyphomycetous . . . . . . . . . . . . . . . . . 2 Type: Pilidiella Petr. & Syd.
2. Conidia applanate or helicoid . . . . . . . . . . . . . . . . . . . 3 Notes: The family Schizoparmaceae was introduced to ac-
2. Conidia obovate to oval . . . . . . . . . . . Monotosporella commodate the genus Schizoparme and the asexual-morph
3. Conidia applanate . . . . . . . . . . . . . . . . Canalisporium genera Coniella and Pilidiella. (Castlebury et al. 2002; van
3. Conidia helicoid. . . . . . . . . . . . . . . . . . . . . . .Helicoon Niekerk et al. 2004). Some species in this family are plant
4. Asci with massive, refractive, apical ring; ascospores pathogens that cause leaf, stem and root diseases on a wide
fusiform, with pointed end cells. . . . . . . Ascotaiwania variety of hosts (van Niekerk et al. 2004). Schizoparme has
4. Asci with small, discoid, apical pore/ring; ascospores been synonymized under Pilidiella giving priority to the older
ellipsoid, with rounded end cells . . . . . . . . .Savoryella name Pilidiella (Maas et al. 1979; Rossman et al. 2015).
Sutton (1980) and Nag Raj (1993) treated Pilidiella as a syn-
Schizoparmaceae Rossman, Mycoscience 48 (3): 137 (2007) onym of Coniella but recent phylogenetic analyses showed
Facesoffungi number: FoF 01405 that these genera have distinct phylogenetic lineages in
Saprobic, parasitic or pathogenic on woody, herbaceous Schizoparmeaceae (van Niekerk et al. 2004; Wijayawardene
plants. Sexual morph: Ascomata perithecial, solitary, et al. 2016). Coniella has dark brown conidia while Pilidiella
scattered, subepidermal, erumpent to superficial, globose, co- having hyaline to pale brown conidia (Wijayawardene et al.
riaceous, brown to black, short-papillate, ostiolate, with hya- 2016). However, Pilidiella eucalyptorum Crous & M.J.
line periphyses. Peridium comprising thick-walled, brown Wingf.and P. tibouchinae B.E.C. Miranda et al. have dark
cells of textura angularis. Hamathecium lacking paraphyses. brown conidia which resemble Coniella (van Niekerk et al.
Asci 8-spored, unitunicate, ellipsoid to fusiform, sessile, with 2004; Miranda et al. 2012; Wijayawardene et al. 2016).
a J-, apical ring. Ascospores bi-seriate, hyaline, ellipsoidal, Nevertheless, Miranda et al. (2012) and Wijayawardene
aseptate. Asexual morph: Coelomycetous. Conidiomata pyc- et al. (2016) showed that these both species cluster in
nidial, subepidermal, erumpent, globose, slightly depressed Pilidiella sensu stricto in their phylogenetic analyses.
globose to subglobose. Conidiomata wall comprising thick-
walled, brown cells of textura angularis. Conidiophores Pilidiella Petr. & Syd., Beih. Reprium nov. Spec. Regni veg.
densely aggregated, slender, subulate, simple or branched, 42(1): 462 (1927)
0–3-septate. Conidiogenous cells annellidic, simple, smooth, Basionym: Macroplodia quercicola Oudem., Ned. kruidk.
with prominent periclinal thickening. Conidia hyaline or Archf, 3 sér. 2(3): 752 (1902)
Fungal Diversity
 Fungal Diversity

ƒFig. 129 Canalisporium caribense (Material examined: THAILAND, thick-walled, J-, apical ring. Ascospores bi-seriate, ellipsoidal,
Chiang Rai Province, stream flowing in Tham Luang Nang Non Cave, on hyaline, aseptate, thick-walled, granulate, with or without ter-
submerged wood, 25 November 2014, J. Yang, MFLU 15-3581,
holotype) a Substrate b–d Sporodochia on wood e–f Squash mount of
minal mucous caps. Asexual morph: Coelomycetous.
a sporodochium g Conidiophores h–i Vesiculate conidiogenous cell j Conidiomata pycnidia, subepidermal, erumpent, globose,
Germinated conidium on nature substrate k Germinated conidium on slightly depressed globose to subglobose, sometimes tapering
PDA medium l–o Conidia. Scale bars: b = 500 μm, c–d = 50 μm, e, i– slightly towards the ostiole, smooth, initially hyaline, becom-
k = 20 μm, f = 30 μm, g–h = 15 μm, l–o = 10 μm
ing medium to dark brown. Ostiole central, comprising 3–6
layers of dark brown cells of textura angularis and central
Facesoffungi number: FoF 01569; Fig. 130 cushion of hyaline cells at the base, giving rise to conidio-
Saprobic, parasitic or pathogenic on woody, herbaceous phores. Conidiophores densely aggregated, slender, simple
plants. Sexual morph: Ascomata perithecial, solitary, or branched, 0–3-septate, surrounded by a mucous coating.
scattered, subepidermal, erumpent to superficial, globose, co- Conidiogenous cells simple, slender, hyaline, smooth, with
riaceous, brown to black, warty structures forming around the prominent periclinal thickening. Conidia hyaline when imma-
erumpent parts, short papillate, ostiolate. Papilla wide, short, ture hyaline to medium brown at maturity, inequilateral,
internally lined by hyaline periphyses. Peridium comprising smooth-walled, frequently with a hyaline, lateral appendage,
thick-walled, brown, cells of textura angularis. Hamathecium narrowly ellipsoidal, apices tapering, subobtusely rounded,
lacking paraphyses. Asci unitunicate, 8-spored, unitunicate, acutely rounded bases subtruncate, guttulate, with a longitu-
ellipsoid to fusiform, sessile, apex wide and blunt, with dinal germ slit, straight to slightly curved.

Fig. 130 Pilidiella quercicola a Ascomata on host substrate b Cross Wijayawardene G0928-2 (HGUP502, MFLU 13-0006); living culture
section of ascoma (redrawn from Rossman et al. 2007) Pilidiella GDUP N77, ICMP 19810, MFLUCC 13-0299) e Disease symptoms on
destruens c Asci d Ascospores (redrawn from Van Niekerk et al. 2004) host tissues f Conidiomata on host substrate g Cross section of conidioma
Pilidiella castenicola (CHINA, Guizhou Province, Guiyang, Huaxi, on h Peridium i Conidia attached to the conidiogenous cells j Conidia. Scale
Acer buergerianum Miq. (Sapindaceae) leaves, 28 September 2013, N.N. Bars: a, d, e, h = 10 μm, b, g = 100 μm, c, i, j = 20 μm, f = 500 μm
Fungal Diversity

Type species: Pilidiella quercicola (Oudem.) Petr., Beih. light brown, allantoid, ellipsoid to oval, straight to
Reprium nov. Spec. Regni veg. 42(1): 462 (1927) [1926] slightly curved, aseptate, with or without appendages.
Notes: Pilidiella was introduced and typified by Asexual morph: Undetermined.
Pilidiella quercicola (Oudem.) Petr., and currently com- Type: Scortechinia Sacc.
prises twenty two epithets (Index Fungorum 2016). Notes: The family Scortechiniaceae was introduced
Pilidiella species are commonly associated with a wide by Huhndorf et al. (2004b) based on superficial, black,
variety of woody and herbaceous hosts (Van Niekerk turbinate ascomata with a quellkörper (this structure is
et al. 2004). Type species of Pilidiella, P. quercicola and made by a mucilaginous mass of thick-walled cells
Type species of Schizoparme, S. straminea are congeneric within the ascoma and is normally conical). Kirk et al.
and Pilidiella proposed to use over Schizoparme (2008) considers this characteristic as inducing rupture
(Rossman et al. 2015). Miranda et al. (2012) was intro- of the ascoma. Asci are clavate with hyaline ascospores.
duced a new species Pilidiella tibouchinae B.E.C. The genus is typified by Scortechinia (Saccardo 1891).
Miranda et al. which has brown conidia. Crous et al. Huhndorf et al. (2004b) introduced this family based on
(2015c) was introduced sexual morph of Pilidiella LSU sequence data and morphological observations. The
eucalyptigena Crous & M.J. Wingf from Eucalyptus. quellkörper was accepted as a main structure of this
family, and the family is based on this character.
Euacanthe, Neofracchiae and Scortechinia were includ-
Other genus included ed in the family (Huhndorf et al. 2004b). Mugambi and
Huhndorf (2010) included Biciliospora, Coronophorella,
Coniella Höhn., Ber. dt. bot. Ges. 36: 316 (1918) Cryptosphaerella, Scortechiniella, Scortechiniellopsis
Type species: Coniella pulchella Höhn., Ber. dt.bot. Ges. and Tympanopsis in the family based on multi-gene
36: 316 (1918) analyses.

Scortechinia Sacc., Syll. fung. (Abellini) 9: 604 (1891)

Key to genera of Schizoparmaceae Facesoffungi number: FoF 01124, Figs 131, 132
Saprobic on woody substrates in terrestrial habitats. Sexual
1. Conidia hyaline to pale brown, narrowly ellipsoidal, with morph: Ascomata perithecial, scattered or gregarious, super-
acutely rounded apices . . . . . . . . . . . . . . . . . . . . Pilidiella ficial on host, immersed or sub-immersed in the thick
1. Conidia dark brown, ellipsoidal, with narrowly, obtusely subiculum, globose to subglobose, black, coriaceous or
rounded apices . . . . . . . . . . . . . . . . . . . . . . . . Coniella membranous, turbinate or tuberculate, with or without
brown branched setae, collabent or not collapsing.
Scortechiniaceae Huhndorf et al., in Huhndorf et al., Mycol. Subiculum thin or thick, brown to dark brown, septate,
Res. 108(12): 1387 (2004) crisp when dry, branched or unbranched hyphae, with
Facesoffungi number: FoF 01123 spiny termination. Peridium thick (20–30 μm), munk
Saprobic on woody substrates in terrestrial habitats. pores present or absent, outer layer composed of black
Sexual morph: Ascomata perithecial, scattered or gregar- to brown cells of textura angularis, thick-walled; inner
ious, superficial, sub-immersed or immersed on host, sit- layer composed of hyaline cells of textura prismatica,
ting on or in a subiculum or subiculum absent, globose, thin-walled. Hamathecium quellkörper present, conical,
obpyriform or ovoid, dark brown to black, coriaceous to wide at the base, paraphyses absent. Asci 8-spored,
membranous, turbinate or tuberculate or smooth, with or unitunicate, cylindrical to clavate, long or short stipitate,
without brown branched setae, collabent or not collaps- apex rounded, apical ring indistinct or absent, evanes-
ing. Subiculum thin or thick, brown to dark brown, sep- cent. Ascospores 2-seriate to irregularly arranged, hya-
tate, crisp when dry, branched or unbranched hyphae, line, allantoid, ellipsoid to oval, straight to slightly
with spiny termination or smooth. Peridium thick (20– curved, aseptate, smooth-walled, without appendages,
80 μm), munk pores present or absent, outer layer com- sometimes with guttules. Asexual morph:
posed of black to brown cells of textura angularis, thick- Undetermined.
walled; inner layer composed of hyaline cells of textura Type species: Scortechinia culcitella (Berk. & Ravenel)
prismatica, thin-walled. Hamathecium quellkörper pres- Speg., Anal. Soc. cient. argent. 26(1): 29 (1888)
ent, conical, wide at the base, paraphyses absent or in- Basionym: Sphaeria culcitella Berk. & Ravenel, in
distinct, filiform, hyaline. Asci 8- to multi-spored, Berkeley & Curtis 1860
unitunicate, cylindrical to clavate, long or short pedicel- Notes: The genus Scortechinia was introduced by
late, apical ring indistinct or absent, evanescent. Saccardo (1891) and includes nine epithets according
Ascospores 2-seriate to irregularly arranged, hyaline or to Index Fungorum (2016). The type species
 Fungal Diversity

Fig. 131 Scortechinia culcitella (Material examined: USA, on the old e Ascomata in cross section f Section through perithecium g–h Spinulose
trunk of Ostrya sp., PC MNHN-PC-PC0121050) a Material label b subicular hyphae j–l Ascospores Notes: Figs k-l stained in Melzer’s
Specimen c-d Ascomata on substrate sub-immersed in thick subiculum reagent. Scale bars: c–e = 200 μm, f–g = 50 μm, h = 20 μm, i–l = 5 μm

Scortechinia culcitella, is characterized by tuberculate Other genera included

ascomata with a quellkörper, clavate asci and hyaline
ascospores (Huhndorf et al. 2004b). Biciliospora Petr., Sydowia 6(5–6): 429 (1952)
Fungal Diversity

Fig. 132 Scortechinia culcitella

(Based on the description by
Saccardo 1891) a Section through
perithecium with quellkörper
(arrowhead) b Asci with
ascospores c Munk pores between
cells of the ascoma wall
(arrowhead). Scale bars:
a = 100 μm, b = 20 μm

Type species: Biciliospora velutina Petr., Sydowia 6(5–6): Type species: Scortechiniella similis (Bres.) von Arx & E.
429 (1952) Müll., Beitr. Kryptfl. Schweiz 11(no. 1): 383 (1954)
Coronophorella Höhn., Sber. Akad. Wiss. Wien, Math.- Scortechiniellopsis Sivan., Trans. Br. mycol. Soc. 62(1):
naturw. Kl., Abt. 1 118: 1507 [47 repr.] (1909) 36 (1974)
Type species: Coronophorella chaetomioides (Penz. & Type species: Scortechiniellopsis leonensis Sivan., Trans.
Sacc.) Höhn. 1909 Br. mycol. Soc. 62(1): 37 (1974)
Cryptosphaerella Sacc., Syll. fung. (Abellini) 1: 186 Tympanopsis Starbäck, Bih. K. svenska Vetensk Akad.
(1882) Handl., Afd. 3 19(no. 3): 24 (1894)
Type species: Cryptosphaerella nitschkei (Auersw.) Sacc., Type species: Tympanopsis euomphala (Berk. & M.A.
Syll. fung. (Abellini) 1: 186 (1882) Curtis) Starbäck, Bih. K. svenska Vetensk Akad. Handl.,
Euacanthe Theiss., Annls mycol. 15(3/4): 272 (1917) Afd. 3 19(no. 3): 24 (1894)
Type species: Euacanthe usambarensis (Rehm) Theiss.,
Annls mycol. 15(3/4): 272 (1917)
Neofracchiaea Teng, Sinensia, Shanghai 9: 255 (1938) Key to genera of Scortechiniaceae
Type species: Neofracchiaea callista (Berk. & M.A.
Curtis) Teng, Sinensia, Shanghai 9: 255 (1938) 1. Ascomata immersed and becoming erumpent in the bark of
Scortechiniella von Arx & E. Müll., Beitr. Kryptfl. host . . . . . . . . . . . . . . . . . . . . . . . . . . . Cryptosphaerella
Schweiz 11(no. 1): 382 (1954) 1. Ascomata superficial on host . . . . . . . . . . . . . . . . . . . 2
 Fungal Diversity

2. Ascospores with appendage at the ends . . . . . . . . . . . 3 Sordaria Ces. & De Not., Comm. Soc. crittog. Ital. 1(4):
2. Ascospores without appendages . . . . . . . . . . . . . . . . . 4 225 (1863)
3. Asci more than 8-spores. . . . . . . . . . . . . . Biciliospora Facesoffungi number: FoF 01149, Fig. 133
3. Asci multi-spored. . . . . . . . . . . . . . . . . Scortechiniella Saprobic on wood, on dung and rotting vegetation in ter-
4. Ascomata with setae . . . . . . . . . . . . . . . . . Euacanthe restrial habitats. Sexual morph: Ascomata perithecial, brown
4. Ascomata without setae . . . . . . . . . . . . . . . . . . . . . . . 5 to black, gregarious or scattered, solitary, superficial or sub-
5. Ascomata with curved tomentum . . . . . Neofracchiaea immersed, subglobose to ovoid, membranaceous, or tubercu-
5. Ascomata smooth. . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 late, with setae or hairs, papillate or ostiole or indistinct, with
6. Asci 8-spored. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 hyaline periphyses or periphyses absent. Peridium thick (10–
6. Asci more than 8-spores. . . . . . . . . Scortechiniellspsis 30 μm), comprising brown to hyaline cells of textura
7. Ascospores allantoid . . . . . . . . . . . . . Coronophorella angularis, globulosa or prismatica, membranaceous.
7. Ascospores ellipsoid to globose . . . . . . . . . . . . . . . . . 8 Hamathecium comprising numerous, septate, filiform
8. Ascomata on thick subiculum, hyphae with spiny pseudoparaphyses or pseudoparaphyses absent. Asci 8-
termination . . . . . . . . . . . . . . . . . . . . . . . Scortechinia spored, unitunicate, thin-walled, cylindrical to clavate,
8. Ascomata on smooth subiculum. . . . . . . Tympanopsis pedicellate, with apical ring or apical ring instinct, not
amyloid. Ascospores hyaline, brown or dark brown, oval
Sordariaceae G. Winter [as ‘Sordarieae’], Rabenh. Krypt.-Fl., to ellipsoid, 0–1-septate, concolourous, smooth-walled,
Edn 2 (Leipzig) 1.2: 162 (1885) with reticulate pattern or with sheath, with or without
Facesoffungi number: FoF 01148 germ pores, appendages present or absent, with or with-
Saprobic on wood, rotting vegetation and dung in terrestri- out guttules. Asexual morph: Undetermined
al habitats, also associated with food. Sexual morph: Type species: Sordaria fimicola (Roberge ex Desm.) Ces.
Ascomata perithecial or cleistothecial, brown to black, gregar- & De Not., Comm. Soc. crittog. Ital. 1(4): 226 (1863)
ious or scattered, solitary, superficial, erumpent or immersed, Notes: The genus Sordaria was introduced by Cesati and
globose, subglobose to ovoid, carbonaceous, coriaceous or De Notaris (1863), to accommodate the type species
membranaceous, tuberculate or smooth, with setae or hairs, S. fimicola and 109 epithets are listed in Index Fungorum
papillate or indistinct or absent, ostiole, with hyaline (2016). This genus was included in Sordariaceae, in the
periphyses or absent. Peridium thick (5–60 μm), comprising Sordariales based on its smooth-walled ascospores with a
brown to hyaline cells of textura angularis, globulosa or pore and sheath (Barr 1990). Sordaria is characterized by dark
prismatica, carbonaceous, coriaceous or membranaceous. ascomata, cylindrical asci and brown ascospores, with gelati-
Hamathecium comprising numerous, septate, branched nous sheath or appendages.
or unbranched, filiform or cylindrical paraphyses or pa- Other genera included
raphyses absent. Asci 8-spored, unitunicate, thin-walled, Copromyces N. Lundq., Ark. Bot. 6: 327 (1967)
cylindrical to clavate, pedicellate, not amyloid. Type species: Copromyces bisporus N. Lundq., Ark. Bot.
Ascospores hyaline, yellowish, brown or black, stellate, 6: 328 (1967)
oval or ellipsoid, 0–1-septate, concolourous or Effetia Bartoli et al., Mycotaxon 19: 517 (1984)
versicolorous, with uneven thickness giving a striate to Type species: Effetia craspedoconidica Bartoli et al.,
reticulate pattern, or with a sheath, with or without Mycotaxon 19: 517 (1984)
germ pores, appendages present or absent, with or with- Gelasinospora Dowding, Can. J. of Res., Section C 9: 294
out guttules. Asexual morph: Undetermined. (1933)
Type: Sordaria Ces. & De Not Type species: Gelasinospora tetrasperma Dowding, Can.
Notes: The family Sordariaceae was introduced by Winter J. of Res., Section C 9: 294 (1933)
(1885) based on brown to black ascomata and cylindrical asci Guilliermondia Boud., Bull. Soc. mycol. Fr. 20: 19 (1904)
with brown to black ascospores, and is typified by Sordaria. Type species: Guilliermondia saccoboloides Boud., Bull.
This family is closely related to Lasiosphaeriaceae Soc. mycol. Fr. 20: 20 (1904)
(Lundqvist 1972; Huhndorf et al. 2004a). Species in this fam- Neurospora Shear & B.O. Dodge, J. Agric. Res.,
ily have been used as model organisms in various biological, Washington 34: 1025 (1927)
biochemical, ecological, genetic and evolutionary studies Type species: Neurospora sitophila Shear & B.O. Dodge,
(Randall and Metzenberg 1995; Nelson 1996; Coppin et al. J. Agric. Res., Washington 34: 1026 (1927)
1997; Dettman et al. 2003; Jacobson et al. 2004; Cai et al. Pseudoneurospora Dania García et al., Mycol. Res.
2006c). Most species in this family are coprophilous. They 108(10): 1139 (2004)
can be heterothallic, homothallic or pseudo-homothallic (Cai Type species: Pseudoneurospora amorphoporcata
et al. 2006c). Maharachchikumbura et al. (2015) accepted (Udagawa) Dania García et al., in García et al., Mycol. Res.
eight genera in this family. 108(10): 1139 (2004)
Fungal Diversity

Fig. 133 Sordaria fimicola (Material examined: ITALY, on decaying mature j–l Ascospores Note: Figures f, i and j are stained in Melzer’s
Vitis sp., 22 March 1930, BPI 580866) a Material label b Ascoma c reagent, Figure h is stained in cotton blue. Scale bars: b–c = 500 μm,
Ascomata d Vertical section of ascoma e Peridium f-i Asci immature to d = 100 μm, e = 25 μm, f–h = 50 μm, i–k = 10 μm

Stellatospora Tad. Ito & Nakagiri, Mycoscience 35(4): 413 Type species: Stellatospora terricola Tad. Ito & Nakagiri,
(1994) Mycoscience 35(4): 413 (1994)
 Fungal Diversity

Key to genera of Sordariaceae Facesoffungi number: FoF 02133; Fig. 134

Parasitic on red algae. Sexual morph: Ascomata superfi-
1. Ascomata cleistothecial . . . . . . . . . . . . . . . . Copromyces cial to immersed, globose to subsphaerical, carbonaceous or
1. Ascomata perithecial . . . . . . . . . . . . . . . . . . . . . . . . . 2 subcarbonaceous, brown or black, ostiolate. Ostiole con-
2. Ascospores oval to ellipsoid. . . . . . . . . . . . . . . . . . . . 3 ical, papillate. Peridium composed of several cell layers
2. Ascospores stellate, with germ pore . . . . .Stellatospora of sscleroplectenchymatous cells. Asci 8-spored, overlap-
3. Paraphyses absent . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 ping, unitunicate, deliquescent. Ascospores elongate,
3. Paraphyses present, filiform . . . . . . . . . . . . . . . . . . . . 6 straight or curved, aseptate, with an equatorial, cytoplas-
4. Ascospores with gelatinous sheath . . . . . . . . . Sordaria mic band, ends spatulate, provided with a gelatinous,
4. Ascospores lacking gelatinous sheath. . . . . . . . . . . . . 5 lateral, appressed appendage. Asexual m orph:
5. Ascomata without hairs, asci without ring at Undetermined.
apex . . . . . . . . . . . . . . . . . . . . . . . Effetia Type species: Spathulospora phycophila A.R. Caval. &
5. Ascomata with hairs, asci with ring at T.W. Johnson, Mycologia 57(6): 927 (1965)
apex. . . . . . . . . . . . . . . . . . . . . . . . . . Gelasinospora Notes: Spathulospora is a marine genus on seaweed with
6. Paraphyses branched . . . . . . . . . . . . . . Guilliermondia Spathulospora phycophila A.R. Caval. & T.W. Johnson as the
6. Paraphyses unbranched, evanescent . . . . . . . . . . . . . . 7 type species. It was introduced by Cavaliere and Johnson
7. Ascospore ornamentation unevenly thickened, giving a (1965) and referred variously to the Spathulosporomycetes,
striate to reticulate pattern . . . . . . . . . . . . Neurospora Spathulosporomycetidae (Locquin 1984), Spathulosporales
7. Ascospore ornamentation irregular, rounded, crest (Kohlmeyer 1973), Spathulosporaceae (Kohlmeyer 1973)
pattern. . . . . . . . . . . . . . . . . . . . . . Pseudoneurospora based on morphological observations. However, two
S p a t h u l o s p o r a s p e c i e s ( S . a n t a rc t i c a K o h l m . ,
Spathulosporaceae Kohlm., Mycologia 65(3): 615 (1973) S. adelpha Kohlm.), have shown a relationship to the
Facesoffungi number: FoF 01798 order Lulworthiales with weak support (Inderbitzin et al.
Parasitic on marine algae. Sexual morph: Ascomata 2004; Campbell et al. 2005; Jones et al. 2009b). Further
subglobose, ovoid, pyriform, subiculate, coriaceous or leath- collections, isolation and sequencing are required to de-
ery, dark brown, sterile hairs enclosing ascoma, ostiolate, termine the phylogenetic placement of this genus in the
ostiolar canal filled surrounded by tube-like projection extend- Ascomycota, especially as the type species is yet to be
ing into the ascomatal cavity, papillate or epapillate, lacking sequenced.
paraphyses. Asci clavate to subglobose, thin-walled,
unitunicate, without apical ring, deliquescing early.
Ascospores hyaline, fusiform, cylindrical, ellipsoidal, 0–3- Other genus included
septate, with an appendage at each end. Reproductive
structures: Antheridial. Spermatia ellipsoidal to fusiform, Retrostium Nakagiri & Tad. Ito, Mycologia 89(3): 485 (1997)
without appendages. Trichogynes simple or branched, septate, Type species: Retrostium amphiroae Nakagiri & Tad. Ito,
arising from the margin of young ascoma. Mycologia 89(3): 485 (1997)
Type: Spathulospora A.R. Caval. & T.W. Johnson
Notes: The family Spathulosporaceae was introduced by
Kohlmeyer (1973) and currently comprises the genera Key to genera of Spathulosporaceae
Spathulospora and Retrostium. Members of this family are
characterized by crustose, dark thalli, peg-like penetrating 1. Ascospores elongate, with spatulate ends, with a gelatinous,
cells, intracellular crusts and stromata, antheridia with lateral, appressed appendage . . . . . . . . . . . Spathulospora
spermatia and trichogynes, ostiolate ascomata without paraph- 1. Ascospores oblong to ellipsoidal, with one mucilaginous
yses, deliquescing asci, and 1-celled, appendaged ascospores. appendage at each end . . . . . . . . . . . . . . . . Retrostium
Kohlmeyer (1973) placed this family in the order
Spathulosporales. Later, Inderbitzin et al. (2004); Campbell Sporocadaceae Corda (as ‘Sporocadeae’), Icon. Fungorum
et al. (2005); Jones et al. (2009b) placed it in the order (Prague) 5: 34. 1842
Lulworthiales. However, Maharachchikumbura et al. (2015) = Discosiaceae Maharachch. & K.D. Hyde
confirmed the placement of Spathulosporaceae in the order Saprobic or pathogenic on leaves of flowering plants and
Spathulosporales by molecular analysis using available se- gymnosperms. Sexual morph: Ascomata scattered, immersed
quence data from GenBank. to erumpent. Ostiole circular, papillate. Peridium comprising
several layers of cells of textura prismatica, brown at the base,
Spathulospora A.R. Caval. & T.W. Johnson, Mycologia dark brown outwardly. Hamathecium comprising numerous,
57(6): 927 (1965) hypha-like, flexuose, paraphyses. Asci 8-spored, unitunicate,
Fungal Diversity

Fig. 134 Spathulospora phycophila (Material examined: Herbarium specimen b Ascomata on host surface c Hairs d, e
AUSTRALIA, Warrnambool, Victoria, on Ballia callitricha C. Agardh Section through ascoma f Peridium g Young asci h Young ascospores
(Balliaceae), April 1959, Norris L. R. E, NY 01350180, NY 01350182 showing equatorial cytoplasmic bands i Mature ascospores. Scale bars: c–
NY 01350186, NY 01350185, NY 01350187, NY 01350178, NY d = 100 μm. e–f = 50 μm. g–h = 25 μm
01350183, slides from the holotype); Ibid. BPI 580727, holotype). a

obclavate to cylindrical, with a discoid, J+ subapical ring. with or without vertical septa, hyaline. Asexual morph:
Ascospores overlapping uniseriate or biseriate, obovoid to el- Coelomycetous. Conidiomata stromatic, scattered to gregari-
lipsoid, transversally septate, longitudinal septum in mid cells, ous, subepidermal or subepidermal in origin, immersed to
 Fungal Diversity

erumpent. Peridium composed of 3–4 layers of cells of textura until further studies based on fresh collections and molecular
angularis, outer 1–2-layers brown and inner 1–2 layers hya- data are conducted (Maharachchikumbura et al. 2015).
line or pale brown. Conidiophores arising from the inner layer
cells of basal stroma, absent or reduced to conidiogenous
cells, when present long, septate and branched. Other genera included
Conidiogenous cells holoblastic, annellidic, cylindrical or
lageniform, hyaline. Conidia cylindrical to clavate, or fusi- Adisciso Kaz. Tanaka et al., in Tanaka et al., Persoonia, Mol.
form, straight or curved, 2–4-euseptate, hyaline, pale oliva- Phyl. Evol. Fungi 26: 90 (2011)
ceous, or brown, bearing cellular, filiform or attenuated Type species: Adisciso yakushimense Kaz. Tanaka et al., in
appendages. Tanaka et al., Persoonia, Mol. Phyl. Evol. Fungi 26: 92 (2011)
Type: Seimatosporium Corda Discosia Lib., Pl. crypt. Arduenna, fasc. (Liège) 4: no. 346
= Sporocadus Corda (1837)
Notes: Members of the family Sporocadaceae are Type species: Discosia artocreas (Tode) Fr., Summa veg.
saprobes, endophytes, or foliar pathogens in tropical Scand., Section Post. (Stockholm): 423 (1849)
and temperate regions (Nag Raj 1993; Tanaka et al. Discostroma Clem., Gen. fung. (Minneapolis): 50 (1909)
2011). The genus Sporocadus was synonymized under Type species: Discostroma rehmii (Schnabl) Clem., Gen.
Seimatosporium. The genera Adisciso, Discosia, fung. (Minneapolis): 1–227 (1909)
D i s c o s t ro m a , Sa rco s t ro m a an d t he ty pe g en us Sarcostroma Cooke, Journal of the Quekett microsc. Club
Seimatosporium are placed in Sporocadaceae following 2: 267 (1871)
the studies by Maharachchikumbura et al. (2015) and Type species: Sarcostroma berkeleyi Cooke, Journal of the
Senanayake et al. (2015). In addition, the genus Quekett microsc. Club 2: 267 (1871)
Strickeria was included into the family by Jaklitsch Strickeria Körb., Parerga lichenol. (Breslau) 5: 400 (1865)
et al. (2016) based on morphology and phylogeny. Type species: Strickeria kochii Körb., Parerga lichenol.
Seimatosporium Corda, in Sturm, Deutschl. Fl., 3 Abt. (Pilze (Breslau) 5: 400 (1865)
Deutschl.) 3(13): 79 (1833)
Facesoffungi number: FoF 02134; Figs 135, 136
Saprobic on leaves, stem and bark of various plants, path-
ogenic on leaves of Eucalyptus spp. Sexual morph: Key to the sexual genera of Sporocadaceae
Discostroma-like. Asexual morph: Conidiomata acervular,
solitary to gregarious, superficial to immersed, unilocu- 1. Ascospores arranged biseriately . . . . . . . . . . . . . Adisciso
lar, globose to subglobose, dark brown to black. 1. Ascospores arranged uniseriately . . . . . . . . . . . . . . . . 2
Peridium composed of brown thin or thick-walled 2. Ascospores hyaline . . . . . . . . . . . . . . . . . Discostroma
textura angularis. Conidiophores cylindrical, filiform, 2. Ascospores brown, end cells subhyaline. . . . Strickeria
septate, branched, hyaline. Conidiogenous cells holo-
blastic, annellidic, integrated or discrete, hyaline, deter-
minate. Conidia variable, cylindrical, fusiform or clavate Key to the asexual genera of Sporocadaceae
or obovoid, (2 −)3(−5)-septate, eguttulate, medium cells
brown, basal cell hyaline, with or without basal append- 1. Conidiomata not stromatic, immersed, conidia subhyaline
age, or with the apical cell provided with a single, cel- to pale brown . . . . . . . . . . . . . . . . . . . . . . . . . . . Discosia
lular, simple or branched appendage, basal cell truncate, 1. Conidiomata stromatic, semi-immersed to erumpent, co-
continuous or occasionally constricted at the septa. nidia brown . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
Type species: Seimatosporium rosae Corda, in Sturm, 3. Conidiophores unbranched or irregularly branched,
Deutschl. Fl., 3 Abt. (Pilze Deutschl.) 3(13): 79 (1833) conidiogenous cells with minute periclinal thickenings,
Notes: The genus Seimatosporium was introduced by conidia fusiform, 3–5-septate, occasionally 7-
Corda (1833) based on the type S. rosae. von Arx (1981) septate . . . . . . . . . . . . . . . . . . . . . . . . . . . Sarcostroma
treated Bartalinia, Bartaliniopsis, Bleptosporium, 3. Conidiophores branched, conidiogenous cells without
Doliomyces and Hyalotia as synonyms of Seimatosporium. minute periclinal thickenings, conidia ellipsoid to cla-
This was not accepted by Nag Raj (1993) and furthermore is vate, 2–4-septate, occasionally 5-, 6-
not supported by molecular data (Maharachchikumbura et al. septate . . . . . . . . . . . . . . . . . . . . . . . . Seimatosporium
2015). Seimatosporium species have been linked to
Discostroma sexual morphs through molecular data (Tanaka Stachybotriaceae L. Lombard & Crous, in Crous et al.,
et al. 2011), however the type of Discostroma lacks molecular Persoonia, Mol. Phyl. Evol. Fungi 32: 283 (2014)
data to confirm this. Therefore both names have been retained Facesoffungi number: FoF 01245
Fungal Diversity

Fig. 135 Discostroma fuscellum (Material examined: ITALY. Province Peridium e Paraphyses. f–h Immature asci i Mature ascus j Ring-like
of Forlì-Cesena [FC], Poggio Baldo - Predappio Alta, on dead stem of apical ring of ascus in Congo red k-o ascospore. Scale bars: b, f–
Rosa canina L. (Rosaceae), 3 March 2013, E. Camporesi, IT 1093 i = 200 μm, c = 100 μm, d–e, j = 50 μm, f–I = 20 μm, k–o = 30 mm
(MFLU 15-0750)) a-b Ascomata on host surface c Section of ascoma d

Saprobic or pathogenic on plants and commonly isolat- apex rounded to nearly truncate, deliquescent at maturi-
ed from soil. Sexual morph: Ascomata superficial or ty, lacking an apical ring. Ascospores 2–3-seriate, hya-
immersed, solitary or scattered, sometimes fused, glo- line or brown to dark-brown, ellipsoidal to fusiform or
bose to subglobose, bright to dark yellow or black, broadly reniform, 0–1-septate, with or lacking a muci-
sometimes covered with intertwined hyphae. Papilla laginous sheath. Asexual morph: Hyphomycetous.
central, short, black, without periphyses. Ostiole conical, Conidiophores single, sporodochial or synnematous, if
thin, black. Peridium composed of several layers of conidiophore simple, macronematous, mononematous,
dark brown, thin walled cells of textura angularis solitary or gregarious, clavate to broadly reniform
(Stachybotrys), or textura intricata (Scopinella); or out- phialides with conspicuous collarettes, simple or irregu-
wardly composed of thick-walled textura angularis and larly branched, hyaline or pale brown, smooth-walled; if
inwardly with textura prismatica (Peethambara). conidiophores sporodochial and synnematous scattered,
Hamathecium comprising moniliform or filiform, hya- amphigenous, pulvinate, gelatinous, hyaline to dark
line, septate or aseptate paraphyses, intermingled with green, with or without setae, with irregularly penicillate,
asci. Asci 4–8-spored, unitunicate, clavate to cylindrical, bi- or triverticillately branched conidiogenous ring.
 Fungal Diversity

Fig. 136 Seimatosporium rosae (Material examined: RUSSIA, Rostov Fruiting bodies on host substrate d Cross section of fruiting body e, f
Region, Krasnosulinsky District, Donskoye Forestry, edge of ravine Peridium g Conidiogenous cells h-j Conidia Scale bars: c = 200 μm;
forest, on dying and dead branches of Rosa kalmiussica, 21 May 2014, d = 300 μm; e, f, = 20 μm; h–j = 10 μm
T. Bulgakov T−056, MFLU 14-0771, epitype) a Habit in wood b, c

Conidiogenous cells enteroblastic, monophialidic, dis- Type: Stachybotrys Corda

crete, cylindrical, ellipsoidal or clavate, proliferating Notes: Stachybotriaceae was introduced by Crous et al.
percurrently, with terminal phialides, clavate to broadly (2014b) as a new family in the order Hypocreales to accom-
reniform, initially hyaline becoming pale brown at ma- modate the genera Myrothecium, Peethambara and
turity. Conidia arising from the apices of the phialides, Stachybotrys. Previously, Castlebury et al. (2004) based on
elongate, cylindrical or fusiform or ellipsoidal, 1–3-sep- combined SSU and LSU gene analysis, classified these three
tate or aseptate, smooth-walled, sometimes striate or genera in Hypocreales genera incertae sedis, which were dis-
roughened, hyaline or brown to dark brown, aggregated tinct from other families in the order Hypocreales.
in slimy, dark green to black masses. Summerbell et al. (2011) refrained from introducing a new
Fungal Diversity

family to accommodate these taxa, considering that more sex- 2004). The phylogenetic analysis by Wang et al.
ual morphs belonging to these genera are needed to be studied. (2015) based on ITS sequence data also supported the
Maharachchikumbura et al. (2015) provided an updated out- opinion that Melanopsamma is the sexual morph of
line of the family Stachybotriaceae with seven genera, which Stachybotrys. The name Stachybotrys was accepted over
included both sexual and asexual morphs. Melanopsamma based on phylogenetic analysis (ITS
gene) (Wang et al. 2015).
Stachybotrys Corda, Icon. fung. (Prague) 1: 21 (1837)
Faces of fungi number: FoF 01246; Figs 137, 138
Saprobic on plant material and pathogens on human and Other genera included
animals in indoor environments. Sexual morph: Ascomata
superficial, globose, sunken when dry, black, coriaceous, lack- Albosynnema E.F. Morris, Mycopath. Mycol. appl. 33: 179
ing or covered with numerous setae, papillate. Papilla short, (1967)
or a neck, black, shiny, with or without periphyses. Peridium Type species: Albosynnema elegans E.F. Morris,
thin-walled, comprising 4–8 layers of dark brown cells of Mycopath. Mycol. appl. 33: 179 (1967)
textura angularis. Hamathecium comprising filiform, septate Myrothecium Tode, Fung. mecklenb. sel. (Lüneburg) 1: 25
or aseptate paraphyses. Asci 8-spored, unitunicate, thin- (1790)
walled, clavate, pedicellate, lacking an apical ring, deliques- Type species: Myrothecium inundatum Tode, Fung.
cent at maturity. Ascospores 2–3-seriate, hyaline, ellipsoidal, mecklenb. sel. (Lüneburg) 1: 25 (1790)
1-septate, verrucose, with or lacking a mucilaginous sheath. Parasarcopodium Melnik et al., Mycol. Progr. 3(1): 22
Asexual morph: Hyphomycetous. Conidiophores (2004)
macronematous, simple or cymosely branched, with clusters Type species: Parasarcopodium ceratocaryi Melnik et al.,
of several ellipsoidal or subclavate phialides at the apex in Mel’nik et al., Mycol. Progr. 3(1): 24 (2004)
formed in succession. Phialides terminal, clavate to broadly Peethambara Subram. & Bhat, Revue Mycol., Paris 42(1):
reniform, hyaline becoming pale brown. Conidia 1-celled, 52 (1978)
ellipsoidal, cylindrical, reniform or fusiform, ornamented or Type species: Peethambara sundara Subram. & Bhat,
smooth-walled, pigmented or hyaline, held together in slimy Revue Mycol., Paris 42(1): 53 (1978)
drops or chains. Sarcopodium Ehrenb., Sylv. mycol. berol. (Berlin): 12, 23
Type species: Stachybotrys chartarum (Ehrenb.) S. (1818)
Hughes, Canad. J. Bot. 36: 812. 1958. Type species: Sarcopodium circinatum Ehrenb., Sylv.
Facesoffungi number: FoF 01247 mycol. berol. (Berlin): 23 (1818)
Notes: The genus Stachybotrys was introduced by Corda Scopinella Lév., in Orbigny, Dict. Univ. Hist. Nat. 8: 493
(1837) with S. chartarum as the type species, and was isolated (1846)
from the walls of a bedroom in Prague. Stachybotrys is char- Type species: Scopinella pleiospora (J. Schröt.) Sacc. [as
acterized by macronematous, mononematous, single or ‘plejospora’], Syll. fung. (Abellini) 9: 953 (1891)
branched conidiophores, with discrete, phialidic
conidiogenous cells, producing aseptate conidia, in a usually
dark colored slimy mass (Jong and Davis 1976; Mercado-
Sierra et al. 1997). Stachybotrys was earlier classified in Key to the sexual genera of Stachybotryaceae
Hypocreales genera incertae sedis based on sequence data
(SSU and LSU) (Castlebury et al. 2004). Crous et al. 1. Ascospores unicellular, barrel-shaped . . . . . . . .Scopinella
(2014b) transferred it to the newly established family 1. Ascospores didymosporous . . . . . . . . . . . . . . . . . . . . 2
Stachybotriaceae based on analysis of ITS and LSU 2. Ascomata superficial; ascospores surrounded by a mu-
sequence data. Wang et al. (2015) discussed and accepted 74 cilaginous sheath . . . . . . . . . . . . . . . . . . . Stachybotrys
species in Stachybotrys. Stachybotrys, is known as ‘black 2. Ascomata immersed or semi-immersed; ascospores lack-
mold’ or’toxic mold’, due to its ability to produce some rather ing sheath. . . . . . . . . . . . . . . . . . . . . . . . Peethambara
potent mycotoxins that cause stachybotrytoxicosis and its as-
sociation with long-term water-damaged surfaces on buildings
(Wang et al. 2015) Key to the asexual genera of Stachybotryaceae
Stachybotrys albipes (Berk. & Broome) S.C. Jong &
Davis, has been linked to the sexual morph reported 1. Conidiophores macronematous, mononematous . . . . . . . 2
as Melanopsamma pomiformis (Pers.) Sacc. (type), 1. Conidiophores micronematous or synnematous . . . . . 5
based on combined LSU, SSU, EF1, ATP6 and RPB1 2. Stroma with setae always present . . . . . . . . . . . . . . . . 3
gene analysis, with strong support (Castlebury et al. 2. Stroma without setae . . . . . . . . . . . . . . . . . . . . . . . . . 4
 Fungal Diversity

Fig. 137 Stachybotrys palmicola – sexual morph (Material examined: Papilla f The base of the ascoma illustrating the stromatic tissue which
ECUADOR, Oriente, Reserva de Produccion Faunistica Cuyabeno firmly anchors it to the substrate g, h Peridium i Intertwined hyphae j
(Cuyabeno Reserve), Rio Cayabeno, forest near the Laguna Grande, Paraphyses k-q Ascospores r Ascospores with verrucose wall and a large
Canangucho, Path A, on dead rachis of a palm, Aug. 1993, K. D. Hyde mucilaginous sheath. Scale Bars: d = 100 um, f, i = 50 um, e, h, j = 20 um,
(HKU(M) 2684, holotype). a Herbarium packet and specimen b g, k–r = 10 um
Ascomata on the surface of material c Ascomata d Section of ascoma e

3. Conidia aggregated in brown to dark brown in slimy 4. Conidia without appendages . . . . . . . . . . Stachybotrys
masses. . . . . . . . . . . . . . . . . . . . . . . . . . Sarcopodium 5. Conidia 3-septate . . . . . . . . . . . . . . . . . . Albosynnema
3. Conidia aggregated in dark green or black in slimy 5. Conidia 1-septate . . . . . . . . . . . . . . . . . . Peethambara
masses. . . . . . . . . . . . . . . . . . . . . . . . . . Myrothecium
4. Conidia with amorphous mucoid appendages at both Stilbosporaceae Link [as ‘Stilbosporei’], Abh. Königl. Akad.
ends. . . . . . . . . . . . . . . . . . . . . . . . . Parasarcopodium Wiss. Berlin 1824: 180. 1826, emend.
Fungal Diversity

Fig. 138 Stachybotrys chartarum - asexual morph (Material examined: formation redrawn from Jong and Davis (1976). G–j Conidia. Scale
JAPAN, from coal, 1952, Y. Sasaki, PDD 40007). a Herbarium packet bars: b = 1 mm, c = 200 μm, d and e = 40 μm, f = 10 μm, g = 20 μm,
and material b, c Conidiophores on the surface of material d, e h–j = 10 μm
Conidiophores and conidiogenous cells f Phialides with conidia

Facesoffungi number: FoF 01411 absent or present, if present inconspicuous, light brown,
Saprobic on bark of trees and shrubs. Sexual morph: rarely dark brown. Entostroma prosenchymatous, pale-
Pseudostromata inconspicuous, immersed. Ectostromatic disc coloured, slightly differentiated from the surrounding
 Fungal Diversity

bark tissue. Ascomata loosely arranged as valsoid Notes: The type species of Stilbospora, S. macrosperma
groups in a single layer, immersed, aggregated, globose Pers., has been linked to its asexual morph
to subglobose, coriaceous, black, ostiolate, papillate. Prosthecium ellipsosporum Fresen., the generic type of
Ostiole not obvious, convergent in groups. Prosthecium (Voglmayr and Jaklitsch 2008). This genus
Hamathecium comprising filiform, aseptate, hyaline pa- comprises opportunistic or moderately phytopathogenic
raphyses. Asci 8-spored, unitunicate, cylindrical, initially fungal species that cause branch dieback or twig blight
attached to the base, later floating in centrum, with J- of various plants.
refractive, apical ring. Ascospores overlapping uniseriate Type species: Stilbospora macrosperma Pers., Syn. meth.
to biseriate, brown, ellipsoid to oblong, distoseptate. fung. (Göttingen) 1: 96 (1801)
A s e x u a l m o r p h: C o e l o m y c e t o u s . C o n i d i o m a t a = Prosthecium ellipsosporum Fresen., Beitr. Mykol. 2: 62
acervular, with paraphyses. Conidiophores cylindrical, (1852)
hyaline. Conidiogenous cells annellidic. Conidia brown,
cylindrical, clavate to pyriform, eu- or distoseptate, with Other genus included
or without oblique or longitudinal septa. Crinitospora B. Sutton & Alcorn, Trans. Br. mycol. Soc.
Type: Stilbospora Pers. 84(3): 437 (1985)
Notes: The family Stilbosporaceae was introduced by Link Type species: Crinitospora pulchra B. Sutton & Alcorn,
(1826) to accommodate Prosthecium and its asexual morph. Trans. Br. mycol. Soc. 84(3): 439 (1985)
However, it was not a phylogenetically well-supported Natarajania Pratibha & Bhat, Kavaka 33: 129 (2006)
family. Hence Stilbosporaceae was synonymized under [2005]
several families. Voglmayr and Jaklitsch (2014) Type species: Natarajania indica Pratibha & Bhat, Kavaka
resurrected the family Stilbosporaceae in Diaporthales 33: 130 (2006) [2005]
based on phylogenetic analysis of LSU sequence data Stegonsporium Corda, in Opiz, Naturalientausch 11: 458.
and accommodated the genera Stegonsporium and 1827.
Stilbospora within the family synonymizing Type species: Stegonsporium pyriforme (Hoffm.) Corda,
Prosthecium under Stilbospora. Icon. Fungorum (Prague) 3: 23. 1839.
Stilbospora Pers., Neues Mag. Bot. 1: 93 (1794)
Facesoffungi number: FoF 02135; Fig. 139
S a p ro b i c o n d e c a y i n g w o o d . S e x u a l m o r p h : Key to genera of Stilbosporaceae
Pseudostromata not distinct, immersed in substrate, causing
cracks. Ectostroma not obvious, rarely erumpent, limited 1. Coelomycetous. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
to a light grey, orange to brown disc of a gel matrix, 1. Hyphomycetous . . . . . . . . . . . . . . . . . . . . Natarajania
containing numerous, tightly packed periphyses, extend- 2. Conidia hyaline with several tubular, unbranched, fili-
ing from the ostioles. Entostroma a loose network of form apical appendages . . . . . . . . . . . . . . Crinitospora
hyaline to brownish hyphae, circular groups of tightly 2. Conidia brown without any appendages . . . . . . . . . . . 3
packed ascomata, filling the entostromatic area, or ar- 3. Ascospores and conidia with three transverse eusepta,
ranged as a valsoid ring. Ascomata arranged in a single ellipsoid to oblong; asci without a refractive canal in the
layer, immersed, aggregated, depressed globose to len- apex. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Stilbospora
ticular, brownish black, coriaceous, papillate, ostiolate. 3. Ascospores and conidia with more than three transverse
Papilla not obvious, cylindrical, convergent, with pale distosepta, ascospores sometimes and conidia always
brownish walls, ostiolar openings appearing as with additional longitudinal distosepta, ascospores ellip-
subhyaline to brownish circles. Peridium comprising soid to oblong, conidia mostly pyriform; asci with a
dark brown cells of textura angularis. Asci 8-spored, cylindrical, slightly refractive canal in the
unitunicate, ellipsoid to fusoid, short pedicellate, apex apex. . . . . . . . . . . . . . . . . . . . . . . . . . . Stegonsporium
blunt, without a refractive ring. Ascospores uni- or
biseriate, brown, ellipsoid to oblong, 3-euseptate, with Sydowiellaceae Lar.N. Vassiljeva, Pirenomits. Lokuloaskomits.
a gelatinous appendage at each end. Asexual morph: Severa Dal’nego Vostoka (Leningrad): 210 (1987)
Coelomycetous. Conidiomata immersed, acervular, cir- Facesoffungi number: FoF 01412
cular, appearing as dark brown to black spots, contain- Saprobic or parasitic on plant matter. Sexual morph:
ing simple, septate, hyaline paraphyses. Conidiophores Stromata present or absent, if present, scattered, immersed to
hyaline, unbranched cylindrical. Conidiogenous cells erumpent, appear as an aggregation of ostioles, rounded or
annellidic. Conidia brown, ellipsoid or oblong, slightly elliptic in shape, dark brown to black, composed of compact
curved, truncate at the base, 3-euseptate, with a hyaline pseudoparenchymatous tissues, several ascomata in a stroma-
sheath. ta, some species give red colour with 5 % KOH. Ascomata
Fungal Diversity
 Fungal Diversity

ƒFig. 139 Stilbospora macrosperma (Material examined: AUSTRIA, thin-walled, hyaline cells of textura angularis and outer,
Niederösterreich, Rekawinkel, grid square 7862/1, on a trunk of thick-walled, brown cells of textura angularis, papilla com-
Carpinus betulus L. (Betulaceae), 20 October 2001, W. Jaklitsch, W.J.
1840, D25 WU 24708, epitype) a, b Herbarium packet and specimen c
prising brown, elongated cells. Hamathecium comprising cel-
Appearance of ascomata on substrate d Horizontal section of ascomata e, lular, septate, hyaline paraphyses. Asci 8-spored, unitunicate,
f Vertical sections of ascomata g Peridium h Paraphyses i–l Asci m–q cylindrical, short pedicellate, apex blunt with J– apical ring.
Ascospores r Conidia attached to the conidiogenous cells s Conidia. Ascospores overlapping uniseriate, oval to ellipsoid, one me-
Notes: s and r based on Voglmayr and Jaklitsch (2014). Scale bars: c, e,
f = 200 μm, d = 500 μm, g = 50 μm, h = 10 μm, i–r = 20 μm
dian septate, not constricted at the septum, hyaline, smooth-
walled. Asexual morph: Undetermined.
Type species: Sydowiella fenestrans (Duby) Petr., Annls
solitary or aggregated, immersed or erumpent, globose to sub- mycol. 21(1/2): 30 (1923)
globose, coriaceous, central or asymmetrically located ostiolar Notes: Sydowiella was introduced and typified by
canal opens through an individual or combined ostiole, inter- Sydowiella fenestrans. This genus is characterized by the sol-
nally covered by filamentous, hyaline periphyses, sometime itary, erumpent perithecia without stromatic tissue, a
ostiolar opening wider than canal, black to brown. Peridium Melanconis-like hamathecium, and the didymosporous asco-
comprising few layers of brown, thick-walled, textura spores (Kobayashi 1970). Molecular studies have placed the
angularis cells. Hamathecium comprising cellular, septate, g en u s i n S y do w i e l l ac e ae ( R o s s m a n e t a l . 20 0 7;
branched, hyaline paraphyses. Asci 8-spored, unitunicate, cy- Maharachchikumbura et al. 2015).
lindrical to sub-globose, short pedicellate, apex blunt with J–
apical ring. Ascospores uniseriate to multi-seriate, filamen-
tous, ellipsoid or long fusoid-cylindric, 1–11 septa, hyaline, Other genera included
light brown to dark brown, sometime with apical and basal
appendages, wall smooth. Asexual morph: Conidiomata Cainiella E. Müll., Sydowia 10(1–6): 120 (1957) [1956]
pycnidia, superficial, aggregated 3–5 in one group, globose, Type species: Cainiella johansonii (Rehm) E. Müll.,
orange to brown, Conidiomatal wall comprising thick-walled, Sydowia 10(1–6): 121 (1957) [1956]
orange, 5–10 cell layers of cells of textura angularis. Calosporella J. Schröt., in Cohn, Krypt.-Fl. Schlesien
Conidiophores branched, hyaline, short, few conidiogenous (Breslau) 3.2(4): 442 (1897) [1908]
cells arising from one conidiophore, attached to conidiomatal Type species: Calosporella platanoidis (Pers.) Höhn., Ber.
wall. Conidiogenous cells cylindrical, hyaline, bottle-shaped, dt. bot. Ges. 35: 631 (1917)
ends pointed, phialidic. Conidia ellipsoid, one-celled, hyaline, Chapeckia M.E. Barr, Mycol. Mem. 7: 164 (1978)
smooth-walled. Type species: Chapeckia nigrospora (Peck) M.E. Barr,
Type: Sydowiella Petr. Mycol. Mem. 7: 165 (1978)
Notes: The family Sydowiellaceae was established to ac- Hapalocystis Auersw. ex Fuckel, Fungi rhenani exsic.,
commodate the genus Sydowiella, which is typified by fasc. 6: no. 585 (1863)
S. fenestrans and a collection of several fungal taxa that appear Type species: Hapalocystis berkeleyi Auersw. ex Fuckel,
to have somewhat variable morphological characters. Initially, Fungi rhenani exsic., fasc. 6: no. 585 (1863)
most genera and species in this family were placed in Lambro Racib., Parasit. Alg. Pilze Java’s (Jakarta) 2: 13
Diaporthales genera incertae sedis (Rossman et al. 2006). (1900)
However, molecular analysis of different gene regions of taxa Type species: Lambro insignis Racib., Parasit. Alg. Pilze
in family Sydowiellaceae proved it to be a well-supported Java’s (Jakarta) 2: 10 (1900)
family (Rossman et al. 2007, Maharachchikumbura et al. Rossmania Lar.N. Vassiljeva, Mycoscience 42(4): 401 (2001)
2015). Sydowiellaceae includes the genera Chapeckia, Type species: Rossmania ukurunduensis Lar.N.
Hapalocystis, Rossmania, Stegophora, and Sillia. In addition, Vassiljeva, Mycoscience 42(4): 401 (2001)
a few species in family Gnomoniaceae (e.g., Gnomonia Sillia P. Karst., Bidr. Känn. Finl. Nat. Folk 23: 20, 159, 251
rostellata) show an affinity to the Sydowiellaceae. (1873)
Members of this family occur on herbaceous plants, Type species: Sillia ferruginea (Pers.) P. Karst., Bidr.
dicotyledonous and hardwood trees as saprobes, para- Känn. Finl. Nat. Folk 23: 159 (1873)
sites and pathogens. Stegophora Syd. & P. Syd., Annls mycol. 14(5): 364
(1916)
Sydowiella Petr., Annls mycol. 21(1/2): 30 (1923) Type species: Stegophora ulmea (Fr.) Syd. & P. Syd.,
Facesoffungi number: FoF 02136; Fig. 140 Annls mycol. 14(5): 364 (1916)
Saprobic on dead plant matter. Sexual morph: Ascomata Uleoporthe Petr., Annls mycol. 39(4/6): 279 (1941)
solitary, scattered, erumpent to immersed, globose, brown, Type species: Uleoporthe orbiculata (Syd. & P. Syd.) Petr.,
short and wide papillate, ostiolate. Peridium comprising inner, Annls mycol. 39(4/6): 280 (1941)
Fungal Diversity

Fig. 140 Sydowiella fenestrans (Material examined: GERMANY, on host surface f Cross section of ascoma g Peridium h-k Asci l-n
Hessen, Wiesenwald pr. Oestrich (Nassau), on Chamerion Ascospores. Scale bars: d–e = 200 μm, f = 100 μm, g–k = 20 μm, l–
angustifolium (L.) Holub. (Onagraceae) 1894, F109251, holotype) a, b n = 10 μm
Herbarium packets c Herbarium specimen d, e Appearance of ascomata

Winterella (Sacc.) Kuntze, Revis. gen. pl. (Leipzig) 1: 34 2 . A s c o s p o r e s l o n g f u s o i d - c y l i n d r i c a l , 5 – 11

(1891) septate . . . . . . . . . . . . . . . . . . . . . . . . . . . Rossmania
Type species: Winterella tuberculifera (Ellis & Everh.) 2. Ascospores ellipsoid, 1-septate. . . . . . . . . . . . . . . . . . 4
Berl., Icon. fung. (Abellini) 1(3): 94 (1893) 3. Ascospores apiosporous . . . . . . . . . . . . . . . . . . . . . . . 5
3. Ascospores non-apiosporous . . . . . . . . . . . . . . . . . . . 6
4. Stromata well developed, . . . . . . . . . . . . . . . . .Lambro
Key to genera of Sydowiellaceae 4. Stromata poorly developed, . . . . . . . . . . . . Stegophora
5. Ascospores oval to fusoid, not constricted at the
1. Stromata absent; ascomata located in host substrate . . . . 2 septa..……….……………..….Cainiella
1. Stromata present; ascomata located in stromatic tissues3 5. Ascospores fusoid, constricted at the septa . Sydowiella
 Fungal Diversity

6. Ascospores with appendages . . . . . . . . . . . . . . . . . . . 7 Thyridium, Barr (1990) included Valsaria and Valsonectria
6. Ascospores without appendages . . . . . . . . . . . . . . . . . 8 in Thyridiaceae. Valsaria is currently placed in Diaporthales
7. Ascospores ellipsoidal with broadly rounded ends, 1–3 and Valsonectria in Hypocreales (Kirk et al. 2008; Jaklitsch
septate, constricted at the septa. . . . . . . . . Hapalocystis et al. 2015). Rossman et al. (1999) placed Balzania, Mattirolia
7. Ascospores, fusoid to oblong, 3–4 eusepta, not constrict- and Thyronectroidea in Thyridiaceae. Checa et al. (2013) ex-
ed at the septa . . . . . . . . . . . . . . . . . . . . . Calosporella amined the type species of Balzania, Mattirolia, Thyridium
8. Stromata erumpent, circular, with branched splits, yel- and Thyronectroidea, and accepted only Mattirolia and
lowish brown. . . . . . . . . . . . . . . . . . . . . . . Uleoporthe Thyridium within Thyridiaceae. They recognized Balzania
8. Stromata immersed, round or elliptic, dark brown to and Thyronectroidea as synonyms of Mattirolia (Checa et al.
black . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 2013). Jaklitsch and Voglmayr (2014) revised Mattirolia and
9. Perithecia valsoid, stromatic tissues not colouring with synonymised under Thyronectria. Thyronectroidea was con-
10 % KOH . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 sidered as a synonym of Thyronectria, whereas Thyronectria
9. Perithecia diatrypoid, stromatic tissues become dull red is distantly related with Thyridiaceae (Jaklitsch and Voglmayr
with 10 % KOH. . . . . . . . . . . . . . . . . . . . . . . . . . Sillia 2014; Maharachchikumbura et al. 2015). According to the
10. Ascospores oval to fusoid-oval, . . . . . . . . Chapeckia molecular analysis, Thyridiaceae is considered as a member
10. Ascospores cylindrical to fusoid-cylindrical,. . .Winterella of subclass Sordariomycetidae (Spatafora et al. 2006; Réblová
2013; Jaklitsch and Voglmayr 2014). Maharachchikumbura
Thyridiaceae J.Z. Yue & O.E. Erikss., Syst. Ascom. 6(2): 233 et al. (2015) placed Thyridiaceae in the newly introduced
(1987) subclass Diaporthomycetidae, the phylogenetic placement of
Facesoffungi number: FoF 01913 which is still uncertain within the subclass
Saprobic or hemibiotrophic in woody substrates. Sexual (Maharachchikumbura et al. 2015).
morph: Stromata immersed, becoming erumpent to superfi-
cial, soft-textured, reddish brown to brightly pigmented. Thyridium Nitschke, Pyrenomyc. Germ. 1: 110 (1867)
Ascomata immersed in stromata, globose, medium-sized, Facesoffungi number: FoF 02137; Fig. 141
ostiolate. Ostioles periphysate, with short or long papilla or Saprobic or hemibiotrophic in woody substrates. Sexual
necks, sometimes convergent necks merging into one ostiole. morph: Stromata immersed or erumpent, formed by interwo-
Peridium composed of compressed rows of cells, externally ven yellowish brownish to black hyphae and cells of the sub-
brown, internally hyaline. Paraphyses filamentous, septate hy- strate, KOH +/−. Ascomata perithecial, globose, immersed in
aline. Asci unitunicate, usually 8-spored, oblong cylindric, stromata, with short or long necks with convergent or separate
with shallow J-, apical ring. Ascospores uniseriate, shades of ostioles. Ostioles periphysate. Asci 8-spored, cylindrical, with
brown, ellipsoid or biconoid, symmetric, 1-septate or a J-, apical ring. Ascospores uniseriate, muriform, ellipsoid or
muriform. Asexual morph: Coelomycetous. biconoid, symmetric, pale brown to dark brown, or with thick-
Conidiomata stromatic, multi-loculate. Conidiogenous ened and pigmented septa. Asexual morph: Undetermined
cells enteroblastic phialidic. Conidia small, hyaline. (Checa et al. 2013).
Holoblastic sympodial conidia also produced from hy- Type species: Thyridium vestitum (Fr.) Fuckel, Jb. nassau.
phae (Barr 1990). Ver. Naturk. 23–24: 195 (1870) [1869–70]
Type: Thyridium Nitschke Notes: The genus Thyridium was introduced by Nitschke
Notes: The family Thyridiaceae was introduced by Yue and (1867) to accommodate species with uniseriate, muriform,
Eriksson (1987) with Thyridium vestitum (Fr.) Fuckel as the dark coloured ascospores, 8-spored asci and filiform paraph-
type species. Thyridium and Sinosphaeria were included in yses (Nitschke 1867). Later, Thyridium was emended to in-
this family (Yue and Eriksson 1987). Later, Thyridium was corporate Bivonella and Sinosphaeria as synonyms by
expanded to include Bivonella and Sinosphaeria (Eriksson Eriksson and Yue (1989). However, Barr (1990) did not ac-
and Yue 1989). The family Thyridiaceae was placed in cept Bivonella as a synonym of Thyridium since the type spe-
Hypocreales and synonymised under Hypocreaceae by con- cies of Bivonella (= Sinosphaeria) has periphysoids, which
sidering the bright pigmentation of stromatic tissues (in some are not seen in the type species of Thyridium (Checa et al.
genera like Sinosphaeria, pigments are soluble in alcohol and 2013).
5 % KOH) and the presence of both paraphyses and
periphysoids (Eriksson and Yue 1989). Barr (1990) observed
that bright pigments are not a unique character in Hypocreales Other genera included
and believed Thyridium (including Sinosphaeria and
Bivonella) is different from Hypocreales and further main- Mattirolia Berl. & Bres., Microm. Trid.: 55 (1889)
tained this as a separate family Thyridiaceae under Type species: Mattirolia roseovirens Berl. & Bres.,
Xylariales. Considering the morphological similarities to Microm. Trid.: 55 (1889)
Fungal Diversity

morph: Colonies effuse or punctiform, white.

Conidiophores acremonium-like, basitonously verticillate or
synnematous. Synnemata when present, simple or
branched, cylindrical, narrowing towards the apex,
consisting of parallel, longitudinal bundles.
Conidiogenous cells phialidic. Phialides scattered,
cymbiform to cylindrical or allantoid, gradually
narrowing towards the apex, hyaline to yellow, smooth
or becoming verrucose, arranged in groups of 1–4 at the
terminal part of synnema, with obvious collarette, some-
times integrated in septate branches. Conidia hyaline to
yellow, cylindrical, fusiform or ellipsoid to oblong, 0–7
septate, smooth or verrucose to roughened, with distinct
hilum at both ends, with or without a mucoid sheath,
formed in chains or aggregating into large sphaerical or
irregular white masses (description modified from
Lombard et al. 2015).
Type: Tilachlidium Preuss
Notes: The family Tilachlidiaceae was introduced by
Lombard et al. (2015) based on molecular analysis to accom-
modate the asexual genera Septofusidium and Tilachlidium.
Gams (1971) classified Septofusidium in Nectriaceae, while
Tilachlidium was classified as genus incertae sedis in the or-
der Hypocreales (Gams 1971).
Tilachlidium Preuss, Linnaea 24: 126 (1851)
Facesoffungi number: FoF 02138; Fig. 143
Saprobic on dead fungi or hyperparasitic on living
entomogenous fungi. Sexual morph: Undetermined.
Asexual morph: Conidiophores synnematous. Synnemata
simple or branched, cylindrical, narrowing towards the
Fig. 141 Thyridium vestitum (redrawn from Leuchtmann and Müller apex, consisting of bundles of parallel, longitudinally
1986). a Section through the host tissue and stromata b Ascus with
ascospores c Apical part of an ascus d Ascospores e Germinated
arranged, usually closely compacted hyphae.
ascospore with primary conidia Conidiogenous cells phialidic, scattered, subulate, grad-
ually narrowing to an acute apex, hyaline, terminating
hyphal cells of the synnema, or as lateral cells of the
Pleurocytospora Petr., Annls mycol. 21(3/4): 256 (1923) hyphae, arrange singly or in groups. Conidia ellipsoidal
Type species: Pleurocytospora vestita Petr., Annls mycol. to oblong, hyaline, aseptate, smooth, covered by a mu-
21(3/4): 256 (1923); Fig. 142 coid sheath, aggregating into large sphaerical or irregu-
lar masses (description modified from Lombard et al.
2015).
Key to sexual genera of Thyridiaceae Type species: Tilachlidium brachiatum (Batsch) Petch,
Trans. Brit. mycol. Soc. 21: 66. 1937
1. Stromata with short or long necks with convergent or sep- Notes: Tilachlidium was described by Preuss (1851) based
arate ostioles, ascospores very pale brown to dark brown, or on Tilachlidium pinnatum Preuss reported on dried fungi,
with thickened and pigmented septa . . . . . . . . Thyridium chiefly as Agaricus galericulatus (Mains 1951). There are
1. Stromata without convergent ostioles, ascospores hya- 19 epithets in Index Fungorum (2016).
line to greenish yellow when mature . . . . . . Mattirolia

Tilachlidiaceae L. Lombard & Crous, in Lombard et al., Stud.

Mycol. 80: 237 (2015) Other genus included
Facesoffungi number: FoF 01280
Saprobic or parasitic on dead or living entomogenous or Septofusidium W. Gams, Cephalosporium -artige
foliicolous fungi. Sexual morph: Undetermined. Asexual Schimmelpilze (Stuttgart): 147 (1971)
 Fungal Diversity

Fig. 142 Pleurocytospora vestita

(redrawn from Leuchtmann and
Müller 1986) a Section through
the host tissue and stroma b
Conidiophores and conidia c
Conidia forming mycelium and
conidia of 10 days old culture d
Conidiophores and conidia of the
stroma in culture

Type species: Septofusidium elegantulum (Pidopl.) W. Saprobic on decaying submerged petioles of the mangrove
Gams, Cephalosporium-artige Schimmelpilze (Stuttgart): palm (Nypa fruticans). Sexual morph: Ascomata scattered,
147 (1971) solitary or gregarious, semi-immersed to superficial, globose
to subglobose, coriaceous to carbonaceous, black, papillate,
ostiolate. Neck short or very long, central, cylindrical, with
Key to genera of Tilachlidiaceae short hyaline periphyses, black. Peridium comprising inner,
hyaline, few layers of cells of textura angularis, middle,
1. Conidia with mucoid sheath, aggregating into large brown, many layers of cells of textura intricata and outer,
sphaerical or irregular masses . . . . . . . . . . . . . . Tilachlidium small, black to brown cells of textura angularis.
1. Conidia without mucoid sheath, arranged in Hamathecium comprising deliquescent, irregular in width,
chains. . . . . . . . . . . . . . . . . Septofusidium septate, hyaline, paraphyses tapering towards apices and em-
bedded in a mucilaginous matrix. Asci 4–8 spored,
Tirisporellaceae Suetrong et al., in Suetrong et al., Cryptog. unitunicate, cylindrical to clavate, thick-walled, indistinct ped-
Mycol. 36(3): 323 (2015) icellate, apex blunt, J- subapical ring comprising a canal and
Facesoffungi number: FoF 01413 pore. Ascospores overlapping bi-seriate to triseriate, fusoid,
Fungal Diversity

Fig. 143 Tilachlidium

brachiatum (redrawn from CBS
505.67; Lombard et al. 2015) a
Phialides terminating on hyphae
of synnema b Lateral phialides
extending from synnema c
Conidia. Scale bars: a–c = 10 μm

falcate to lunate, straight or curved, hyaline to brown, basal o r d e r Ti r i s p o re l l a l e s ( J o n e s e t a l . 2 0 1 5 ) .

cell present or absent, if present pointed, hyaline to pale Maharachchikumbura et al. (2015) showed that the family
brown, 1–7-septate, smooth or verrucose-walled, with or grouped in the Diaporthales along with a freshwater fungus
without 4–5 large guttules, with or lacking appendages. Thailandiomyces bisetulosus. Suetrong et al. (2015) and Jones
Asexual morph: Hyphomycetous. Mycelium comprising et al. (2015) introduced the order Tirisporellales to accommo-
spreading, septate, superficial, smooth, pale brown, branched date three aquatic ascomycetes, which formed a sister group to
hyphae. Conidiophores reduced to conidiogenous cells or the Diaporthales.
if present macronematous, mononematous, erect, brown, Tirisporella E.B.G. Jones et al., Can. J. Bot. 74(9): 1489
paler toward the apex, straight or flexuous, smooth. (1996)
C o n i d i o g e n o u s c e l l s i n t e r c a l a r y, e n t e r o b l a s t i c , Facesoffungi number: FoF 02139; Fig. 144
monophialidic, integrated on hyphae, terminal, sometime Saprobic on decaying submerged mangrove petioles.
erect on hyphae, pale brown, subcylindrical, narrowly Sexual morph: Ascomata scattered, solitary or gregarious,
ellipsoid, ampulliform to doliiform. Conidia solitary, initially immersed becoming to superficial, globose to
smooth, clavate, fusoid-ellipsoid, obovoid or ellipsoid, subglobose, coriaceous becoming carbonaceous, black,
apex obtuse or broadly rounded, tapering to a truncate papillate, ostiolate. Neck short, central, cylindrical, with
base, hyaline to pale brown, unicellular or 1-septate. short hyaline periphyses, black. Peridium comprising in-
Chlamydospores intercalary, pale brown to brown, ner, hyaline, thin-walled, few layers of cells of textura
smooth, globose to narrowly ellipsoid, 0–1-septate. angularis, middle, light brown, many layers of cells of
Type: Tirisporella E.B.G. Jones et al. textura intricata and outer, small, thick-walled, black to
Notes: Tirisporellaceae was introduced and typified by brown cells of textura angularis. Hamathecium compris-
Tirisporella beccariana and this family comprises marine ing deliquescent, irregular in width, septate, hyaline, un-
and freshwater fungal species. The family is referred to the branched paraphyses tapering towards apices and
 Fungal Diversity

embedded in a mucilaginous matrix. Asci 8 spored, Basionym: Sphaeria beccariana Ces., Atti Accad. Sci. fis.
unitunicate, cylindrical to clavate, thick-walled, indis- mat. Napoli 8(no. 3): 20 (1879)
tinct pedicellate, apex flattened, J- subapical ring com- Notes: Tirisporella was introduced by Jones et al. (1996)
prising with canal and pore. Ascospores biseriate to and placed in Dothideomycetes incertae sedis because of the
triseriate, falcate to lunate, straight or curved, brown, thick ascus wall recognized as two walls. This monotypic
basal cell pointed, hyaline to pale brown, 4–7-septate, genus consists of only the type species Tirisporella
verrucose, appendages only at the apical end. Asexual beccariana. The most obvious characters of this genus are
morph: Hyphomycetous. Mycelium comprising spread- the first basal septum delimiting a hyaline to light-coloured
ing, septate, smooth, branched, pale brown hyphae. basal cell and verrucose cell wall, and the apical ascospore
Conidiophores reduced to conidiogenous cells. appendage.
Conidiogenous cells intercalary monophialidic, integrat-
ed on hyphae, pale brown, subcylindrical, ampulliform,
doliiform to narrowly ellipsoid. Conidia solitary, clavate
to fusoid-ellipsoid, apex obtuse, tapering to a truncate Other genera included
base, unicellular to one septate, hyaline to pale brown,
smooth. Chlamydospores intercalary, globose to narrow- Bacusphaeria Norlailatul et al., Phytotaxa. In press;
ly ellipsoid, unicellular to one septate, pale brown to Type species: Bacusphaeria nypenthi Norlailatul et al.,
brown, smooth. Phytotaxa. In press
Type species: Tirisporella beccariana (Ces.) E.B.G. Jones Thailandiomyces Pinruan et al., Fungal Diversity 29: 91
et al., in Jones et al., Can. J. Bot. 74(9): 1490 (1996) (2008)

Fig. 144 Morphological features of marine Tirisporellaceae: Tirisporella Cylindrical ascus e Cylindrical ascus of Bacusphaeria nypenthi f Ascus
beccariana and Bacusphaeria nypenthi (Material examined: tip of Tirisporella beccariana g Ascus tip of Bacusphaeria nypenthi h, i
THAILAND, Trat Province, Ao Salak Phet, Mu Ko Chang National Ascospores with apical appendage of Tirisporella beccariana (arrowed) j
Park, on frond base of N. fruticans, 29 June 2008, Satinee Suetrong, Ellipsoidal ascospores of Bacusphaeria nypenthi Scale bars: a,
BCC32339) a-c Partially immersed to superficial ascomata of b = 1000 μm, c = 500 μm, d–h = 25 μm, I, j = 10 μm
T. beccariana on the lower base of fronds of Nypa fruticans d
Fungal Diversity

Type species: Thailandiomyces bisetulosus Pinruan et al., analysis of LSU, SSU, TEF and RPB2 sequence data. They
Fungal Diversity 29: 91 (2008) introduced this clade as Togniniales to accommodate
Togniniaceae.

Key to genera of Tirisporellaceae Phaeoacremonium W. Gams, Crous & M.J. Wingf.,

Mycologia 88: 789 (1996)
1. Ascomata, laterally spread on substrate, with long neck; Facesoffungi number: FoF 02140; Figs 145, 146
ascospores fusoid, hyaline, . . . . . . . . . . Thailandiomyces Saprobic on plants, or pathogenic on human and animals or
1. Ascomata, erect, ascospores falcate to lunate, brown soil fungi. Sexual morph: Ascomata perithecial, aggregat-
with pointed, hyaline to pale brown basal cell . . . . . . 2 ed or solitary, superficial, erumpent to immersed,
2. Ascospores 4–7 septate . . . . . . . . . . . . . . . Tirisporella astromatic, globose to subglobose, coriaceous, black,
2. Ascospores predominantly 1–3 septate . . Bacusphaeria ostiolate, with longnecks. Neck straight or curved, inter-
nally lined by periphyses. Peridium comprising two
Togniniaceae Réblová et al., Stud. Mycol. 50: 540 (2004) layers, outer layer of brown to dark brown, small,
Facesoffungi number: FoF 01414 rounded cells of textura angularis, inner layer of hya-
Saprobic on plants, or pathogenic on human and animals, line to pale brown, flattened cells of textura angularis.
or soil fungi. Sexual morph: Ascomata perithecial, aggregat- Hamathecium comprising abundant, cellular, branched,
ed or solitary, superficial, erumpent to immersed, astromatic, septate, hyaline paraphyses, slightly constricted at the
globose to subglobose, coriaceous, black, ostiolate, necks septa and tapering towards the apex. Ascogenous
long with periphyses. Peridium comprising three layers, outer hyphae hyaline, branched, elongate during ascus forma-
layer of a few dark brown to black, compressed cell layers, tion, with remnant bases from which single ascus arises.
middle of brown to dark brown, smaller, rounded or brick-like Asci arising in acropetal succession, 8-spored,
cells of textura angularis, textura prismatica to textura unitunicate, sessile, apically thickened, base bluntly ob-
epidermoidea, and inner layer of hyaline to pale brown, flat- tuse. Ascospores biseriate to uniseriate, hyaline,
tened cells of textura angularis. Hamathecium comprising allantoid, reniform, cylindrical or oblong-ellipsoidal,
abundant, paraphyses. Asci arising in acropetal succession, unicellular. Asexual morph: Hyphomycetous. Mycelium
8-spored, unitunicate, sessile, with broadly rounded to obtuse, branched, septate, with single or bundled hyphae, light
thick-walled apex. Ascospores uniseriate to biseriate, hyaline, brown, are as near to conidia formation becoming pale
unicellular, allantoid, reniform, cylindrical or oblong- brown to hyaline, smooth, verruculose or verrucose.
ellipsoidal or subglobose to ellipsoid, 0–1-septate. Asexual Conidiophores branched or unbranched, arising from ae-
morph: Hyphomycetous. Conidiophores branched or un- rial or submerged hyphae, erect, cylindrical or slightly
branched, erect, cylindrical or slightly tapering, pale brown, tapering, straight or curved, up to 7-septate, pale brown,
with one integrated terminal phialide. Conidiogenous cells paler towards the apex, small warts or verruculose or-
mostly monophialidic, discrete or integrated, terminal or lat- namentation at the base, with one integrated terminal
eral, with an inconspicuous funnel-shaped collarette. Conidia phialide and sometimes one or two additional, discrete
aggregated into globose, slimy heads at the apices of phialides at the upper most septum. Conidiogenous cells
phialides, hyaline, aseptate. phialidic, discrete or integrated, terminal or lateral,
Type: Phaeoacremonium W. Gams, Crous & M.J. Wingf. mostly monophialidic, sometimes polyphialidic, warted,
Notes: Togniniaceae was introduced by Réblová et al. verruculose or smooth, pale brown to hyaline, with an
(2004) based on Togninia minima (Tul. & C. Tul.) Berl., and inconspicuous funnel-shaped collarette. Conidia hyaline,
later Togninia was synonymised under Phaeoacremonium aggregated into globose, slimy heads at the apices of
(Gramaje et al. 2015). The family comprises the genera phialides, oblong-ellipsoidal to obovate, cylindrical,
Conidiotheca and Phaeoacremonium (Réblová and Mostert allantoid or reniform, uncommonly fusiform-ellipsoidal
2007) based on phylogenetic and morphological or globose, aseptate, smooth-walled.
distinctiveness. Mostert et al. (2003) placed Togniniaceae in Type species: Phaeoacremonium parasiticum (Ajello
Calosphaeriales based on its unique morphological characters et al.) W. Gams et al., in Crous et al., Mycologia 88(5): 794
and phylogenetic analysis. Subsequently, Mostert et al. (2006) (1996).
placed this family in Diaporthales based on combined gene Basionym: Phialophora parasitica Ajello, Georg & C.J.K.
analysis of SSU and LSU sequence data. The morphological Wang, Mycologia 66(3): 493 (1974)
characters of Togniniaceae are quite different from other Note: Togninia was introduced and typified by
families in Diaporthales. Maharachchikumbura et al. (2015) T. minima. Togninia species reported as the sexual morph
showed the placement of Togniniaceae to be between of Phaeoacremonium. Phaeoacremonium comprises 44
Diaporthales and Calosphaeriales based on combined gene species (Réblová et al. 2015). The type species of
 Fungal Diversity

Fig. 145 Phaeoacremonium minima - sexual morph (Material Ascomata on substrate f Cross section of ascoma g Three-layered
examined: SOUTH AFRICA, Western Cape Province, Wellington and peridium h–j Asci forming on ascogenous hyphae k Ascogenous
Paarl respectively, stems of Vitis vinifera L. (Vitaceae), 2001, L. Mostert, hyphae l Ascospores. Scale bars: c-d = 200 μm, e = 500 μm, f–
LM 463 CBS H 6580, epitype) a, b Herbarium packet and specimen c–e g = 100 μm, h–l = 10 μm

Phaeoacremonium was linked to the type species of (1900). Togninia presently comprises 26 epithets and 13
Togninia, and sexual-asexual relationships between are not sufficiently reported and described. Nine Togninia
Togninia and Phaeoacremonium have been confirmed species are already linked to Phaeoacremonium species.
by mating experiments in culture and molecular sequence Phaeoacremonium has recently been monographed, and
data (Mostert et al. 2003). The generic name comprises 46 species; all they are known from culture
Phaeoacremonium (1996) was accepted over Togninia and have been subjected to DNA sequence analysis.
Fungal Diversity

Fig. 146 Phaeoacremonium

minima - asexual morph (CBS
110703, ex-epitype) a–e
Conidiophores with attached
conidia f Conidia. Scale bars: a–
f = 10 μm

The genus name Phaeoacremonium is preferred by plant Facesoffungi number: FoF 01099
pathologists and medical mycologists, and it is the more Saprobic on intertidal or mangrove wood and roots,
common name used in recent publications (Gramaje and bark and leaves, in marine habitats. Sexual morph:
Di Marco 2015). Ascomata perithecial, hyaline, immersed or superficial,
subglobose, ostiolate, papillate, subcarbonaceous to cori-
aceous. Paraphyses narrow, irregular, persistent or early
Other genus included deliquescing. Asci 8-spored, unitunicate, thin-walled, cla-
vate to ellipsoidal, short pedicellate, lacking an apical
Conidiotheca Réblová & L. Mostert, Mycol. Res. 111(3): 305 ring, early deliquescing. Ascospores fasciculate, hyaline,
(2007) cylindrical to ellipsoidal, 3–5-septate, with several radiat-
Type species: Conidiotheca tympanoides (M.E. Barr) ing appendages at one or both ends. Asexual morph:
Réblová & L. Mostert, Mycol. Res. 111(3): 305 (2007) Hyphae septate, branched, hyaline. Conidiophores present
or obsolete, cylindrical, clavate, septate or aseptate,
acrogenous or laterally on the hyphae, hyaline to light
Key to genera of Togniniaceae brown. Conidia holoblastic, irregularly helicoid,
muriform, cells of the conidia tightly fused, more or less
1. Ascomata subglobose to broadly conical, ascospores similar in size and colour, acrogenous, solitary, constrict-
subglobose to ellipsoid, 0–1-septate. . . . . . Conidiotheca ed at the septa, yellow to brown. Conidial cells up to 50
1. Ascomata globose to subglobose, ascospores allantoid, in number.
unicellular . . . . . . . . . . . . . . . . . . . Phaeoacremonium Type: Torpedospora Meyers
Notes: Torpedosporaceae was introduced by Jones et al.
Torpedosporaceae E.B.G. Jones & K.L. Pang, in Jones et al., ( 2 0 1 4 ) w i t h t w o To r p e d o s p o r a s p e c i e s a n d
Cryptog. Mycol. 35(2): 135 (2014) Glomerulispora mangrovis Abdel-Wahab & Nagah.
 Fungal Diversity

(asexual morph), based on molecular analysis of partial Torpedosporaceae. However, it remains unresolved
sequences of the nuclear SSU and LSU sequence data. whether Glomerulispora mangrovis is the asexual morph
Jones et al. (2014) showed that four strains of of T. radiata (Abdel-Wahab et al. 2010). The genus
Torpedospora radiata formed a monophyletic clade with Torpedospora is worldwide in distribution and T. radiata
Glomerulispora mangrovis, while Torpedospora a common species.
ambispinosa Kohlm. was placed in a lower sub-clade.
Sexual morphs of the family Torpedosporaceae produce
perithecial ascomata and ascospores that possess 3–5 ra- Other genus included
diating appendages at one or both ends, while
G. mangrovis produces helicoid conidia. The family Glomerulispora Abdel-Wahab & Nagah., in Abdel-Wahab
groups in Hypocreomycetidae, order incertae sedis et al., Mycol. Progr. 9(4): 552 (2010)
(Jones et al. 2014) and this was followed by Type species: Glomerulispora mangrovis Abdel-Wahab &
Maharachchikumbura et al. (2015). Subsequently, Jones Nagah., in Abdel-Wahab et al. (2010) Mycol. Progr. 9(4): 553
et al. (2015) have referred the family to a new order (2010)
Torpedosporales.

Torpedospora Meyers, Mycologia 49: 496 (1957) Key to the genera of Torpedosporaceae
Facesoffungi number: FoF 02141; Fig. 147
Saprobic on wood and roots, bark and leaves. Sexual 1. Hyphomycete with irregularly helicoid, muriform, yel-
morph: Ascomata perithecial, superficial to immersed, hya- low to brown conidia with tightly fused conidial cells,
line, membranous, collapsing on drying, subglobose to glo- more or less similar in size and colour, constricted at
bose, ostiole central, papillate. Peridium 2-layered, with an the septa . . . . . . . . . . . . . . . . . . .Glomerulispora
outer layer of cells of textura angularis, and inner layer of 1. Ascomycete, perithecioid, with fasciculate, hyaline,
elongated cells. Paraphyses present. Asci 8-spored, elongate- cylindrical to ellipsoidal, 3–5-septate ascospores
clavate, short pedicellate, with an apical ring, thin walled, with several radiating appendages at one or both
early deliquescent. Ascospores fasciculate, hyaline, elongate, ends . . . . . . . . . . . . . . . . . . . . Torpedospora
tapering, 3–5-septate, with radiating appendages at one or
both ends. Asexual morph: Undetermined. Trichosphaeriaceae G. Winter [as ‘Trichosphaerieae’],
Type species: Torpedospora radiata Meyers, Mycologia Rabenh. Krypt.-Fl., Edn 2 (Leipzig) 1.2: 191 (1885)
49: 496 (1957) Facesoffungi number: FoF 01809
Notes: The genus Torpedospora was introduced by Saprobic or pathogenic on plant materials, commonly
Meyers (1957) and is distinguished from other genera i s ola t e d f r o m he r b i vo r e d un g. S e x u a l m o r ph:
with appendaged spores by the characteristic torpedo-like Ascomata superficial, immersed or semi-immersed,
shape of the ascospores, together with the radiating ap- ostiolate, subglobose to pyriform, dark brown to black.
pendages at one or both ends. Ascospores of T. radiata Ostiole situated in a small papilla, with bristles, in most
(type species), possess 3–5, radiating, sub-terminal ap- genera setose-like hairs present. Peridium carbonaceous
pendages at one end, that appear to be fibrillar (Jones or sub-carbonaceous, coriaceous or membranaceous,
and Moss 1978), whereas T. ambispinosa has sub- brown to dark brown, cells of textura angularis.
terminal appendages at both ends, that are rigid, straight Hamathecium comprising simple, hyaline, guttulate, sep-
or curved. Ascospores of T. ambispinosa in mass are tate, thin-walled, paraphyses. Asci 4–8-spored,
bright orange, but are hyaline in T. radiata. Molecular unitunicate, cylindrical-clavate, gradually tapering below
studies showed that they group in the same clade with into a short pedicel, with rounded apex, sometimes
high bootstrap support, which suggest that they are con- curved, rounded above, some genera have a distinct
generic (Sakayaroj et al. 2005; Schoch et al. 2007). apical ring. Ascospores uniseriate-overlapping uniseriate,
Schoch et al. (2007) suggested that the monophyly of ellipsoidal-fusiform, hyaline or brown-dark brown,
the genus Torpedospora should be further evaluated with aseptate or 1–3-sepate, some genera (Trichosphaeria)
a wider range of taxa. The marine hyphomycete with rounded apices, smooth-walled, guttulate. Asexual
Glomerulispora mangrovis, with irregularly helicoid co- morph: Hyphomycetous. Conidiophores macronematous
nidia with muriform spores, groups with Torpedospora or some genera (Fluviostroma) synnematous, un-
radiata with variable support (Abdel-Wahab et al. 2010; branched or infrequently branched, flexuous and often
Jones et al. 2014). Depending on molecular and circinate, septate or aseptate, dematiaceous, hyaline to
morphological data, Jones et al. (2014) included the gen- brown-yellow. Conidiogenous cells monophialidic, each
era Torpedospora and Glomerulispora in a new family immediately subtending a septum, forming a chain,
Fungal Diversity

Fig. 147 Torpedospora radiata

(Material examined: TAIWAN,
Keelung City, Chaojing Park,
rocky shore, on driftwood, 03
October 2014, K.L. Pang) a
Section of immersed ascoma b
Clavate ascus c Cylindrical
ascospore with unipolar
appendages and three septa d, e
Ascospore showing four
appendages at one end. Scale
bars: a = 20 μm, b–e = 5 μm

terminal, cylindrical or subcylindrical, solitary, hyaline. Notes: The family Trichosphaeriaceae was established
Conidia ellipsoid to oblong, cylindrical to filiform, invalidly by Winter (1885) to accommodate the type
subglobose, hyaline, brown, aseptate or 1–3-septate, Trichosphaeria pilosa (Pers.) Fuckel. Barr (1983) intro-
some genera (Koorchaloma) producing aggregated co- duced the order Trichosphaeriales to accommodate the
nidia in slimy masses, with mucoid appendages at both family Trichosphaeriaceae. In the multi-gene (LSU,
ends. SSU, TEF and RPB2) phylogenetic analysis by
Type: Trichosphaeria Fuckel Maharachchikumbura et al. (2015) Trichosphaeriaceae
 Fungal Diversity

has affinities with Papulosporaceae and Thyridiaceae. Collematospora Jeng & Cain, Can. J. Bot. 54(21): 2429
However, Maharachchikumbura et al. (2015) maintained (1976)
Trichosphaeriaceae as a separate family until fresh col- Type species: Collematospora venezuelensis Jeng & Cain,
lections and molecular data become available. Certain Can. J. Bot. 54(21): 2430 (1976)
species in this family are coprophilous, while other Coniobrevicolla Réblová, Mycotaxon 70: 422 (1999)
members are known as saprobic or pathogenic on plant Type species: Coniobrevicolla larsenii Réblová,
materials including Chrysopogon zizanioides L., Arenga Mycotaxon 70: 424 (1999)
engleri Becc. and Ulmus minor Mill. (Hudson 1963; Cresporhaphis M.B. Aguirre, in Tibell, Bull. Br. Mus. nat.
Calatayud and Aguirre-Hudson 2001). Hist., Bot. 21(2): 146 (1991)
Type species: Cresporhaphis wienkampii (J. Lahm ex
Trichosphaeria Fuckel, Jb. nassau. Ver. Naturk. 23–24: 144 Hazsl.) M.B. Aguirre, Bull. Br. Mus. nat. Hist., Bot. 21(2):
(1870) [1869–70] 154 (1991)
Facesoffungi number: FoF 01810; Fig. 148 Cryptadelphia Réblová & Seifert, Mycologia 96(2): 348
Saprobic or pathogenic on plant materials. Sexual morph: (2004)
Ascomata perithecial, small, globose to subglobose with a Type species: Cryptadelphia groenendalensis (Sacc. et al.)
flattened base, superficial, amphigenous, dark brown, se- Réblová & Seifert, Mycologia 96(2): 352 (2004)
tose, aggregated in to groups, scattered. Setae numerous, Eriosphaeria Sacc., Atti Soc. Veneto-Trent. Sci. Nat.,
pale brown, septate, smooth, up to 600 μm long and Padova, Sér. 4 4: 86 (1875)
7 μm wide, giving a star-like appearance to the ascoma. Type species: Eriosphaeria vermicularia (Nees) Sacc., Atti
Peridium comprising a single layer of thick-walled, po- Soc. Veneto-Trent. Sci. Nat., Padova, Sér. 4 4: 86 (1875)
lygonal to angular, dark brown, cells of textura Fluviostroma Samuels & E. Müll., Sydowia 33: 283
angularis. Hamathecium comprising simple, hyaline, (1980)
guttulate, septate, thin-walled, paraphyses. Asci 4–8 Type species: Fluviostroma wrightii Samuels & E. Müll.,
spored, unitunicate, cylindrical to clavate, with a non- Sydowia 33: 284 (1980)
amyloid apical ring. Ascospores obliquely uniseriate to Kananascus Nag Raj, Mycotaxon 19: 201 (1984)
biseriate, ellipsoidal, narrowly elliptical to fusiform, hya- Type species: Kananascus verrucisporus Nag Raj,
line, aseptate, smooth, sometimes slightly curved, lacking Mycotaxon 19: 207 (1984)
germ pores. Asexual morph: Undetermined. Khuskia H.J. Huds., Trans. Br. mycol. Soc. 46(3): 358
Type species: Trichosphaeria pilosa (Pers.) Fuckel, Jb. (1963)
nassau. Ver. Naturk. 23–24: 145 (1870) [1869–70] Type species: Khuskia oryzae H.J. Huds., Trans. Br. mycol.
Notes: The genus Trichosphaeria was introduced by Soc. 46(3): 358 (1963)
Fuckel (1870) with T. pilosa (Pers.) Fuckel as the type
Koorchaloma Subram., J. Indian bot. Soc. 32: 124 (1953)
species on rotten wood from Germany. Trichosphaeria
Type species: Koorchaloma madreeya Subram., J. Indian
is characterized with superficial, small, setose perithecia
bot. Soc. 32: 124 (1953)
sitting in a subiculum with cylindrical to oblong, 8-
Neorehmia Höhn., Sber. Akad. Wiss. Wien, Math.-naturw.
spored asci bearing hyaline, aseptate, rarely
Kl., Abt. 1 111: 988 [2 repr.] (1902)
appendiculate ascospores (von Arx and Müller 1954;
Type species: Neorehmia ceratophora Höhn., Sber. Akad.
Réblová and Seifert 2004). Sivanesan and Shivas
Wiss. Wien, Math.-naturw. Kl., Abt. 1 111: 988 [2 repr.]
(2002) accepted 25 species of Trichosphaeria, which
(1902)
were isolated mostly from woody substrates. There are
62 epithets for Trichosphaeria recorded in Index Oplothecium Syd. & P. Syd., Annls mycol. 21(1/2): 97
Fungorum (2016). (1923)
Type species: Oplothecium arecae Syd. & P. Syd., Annls
mycol. 21(1/2): 97 (1923)
Other genera included Rizalia Syd. & P. Syd., Annls mycol. 12(6): 546 (1914)
Type species: Rizalia fasciculata Syd. & P. Syd., Annls
Acanthosphaeria Kirschst., Annls mycol. 37(1/2): 91 mycol. 12(6): 546 (1914)
(1939) Schweinitziella Speg., Anal. Soc. cient. argent. 26(1): 45
Type species: Kirschstein gave no clue for the choice of a (1888)
type species Type species: Schweinitziella styracum Speg., Anal. Soc.
Brachysporium Sacc., Syll. fung. (Abellini) 4: 423 (1886) cient. argent. 26(1): 45 (1888)
Type species: Brachysporium obovatum (Berk.) Sacc., Setocampanula Sivan. & W.H. Hsieh, Mycol. Res. 93(1):
Syll. fung. (Abellini) 4: 427 (1886) 87 (1989)
Fungal Diversity

Fig. 148 Trichosphaeria pilosa (Material examined: SWITZERLAND, Ascomata on substrate d Ascoma e Cross section of ascoma f Peridium
Kt. Graubunden: S of Schiers, Furna, on Alnus (Betulaceae), 3 g, h, i, j Asci k Paraphyses l, m, n, o Ascospores. Scale Bars: c = 400 μm,
Sept. 1971, R.F. Cain, TRTC 48286) a, b Herbarium packets c d, e = 50 μm, f–k = 10 μm, l–o = 5 μm

Type species: Setocampanula taiwanensis Sivan. & W.H. Unisetosphaeria Pinnoi et al., in Pinnoi et al., Mycoscience
Hsieh, Mycol. Res. 93(1): 88 (1989) 44(5): 377 (2003)
 Fungal Diversity

Type species: Unisetosphaeria penguinoides Pinnoi et al., Valsaceae Tul. & C. Tul. [as ‘Valsarum’], Select. fung.
Mycoscience 44(5): 378 (2003) carpol. (Paris) 1: 180 (1861)
Facesoffungi number: FoF 01415
Pathogenic or saprobic on plant tissues. Sexual morph:
Key to the sexual genera of Trichosphaeriaceae Stromata present or absent. If present, Ectostroma circu-
lar or irregular, usually well developed in the upper
1. Ascomata aggregated . . . . . . . . . . . . . . . . . . . . . . . . . . . . .2 regions. Entostroma normally limited to the region near
1. Ascomata solitary. . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 the perithecial walls. Ascomata perithecia, immersed to
2. Ascospores uniseriate, fusiform, 3–5-septa . Fluviostroma erumpent, solitary or 6–10 ascomata aggregated together
2. Ascospores biseriate, granulose, curved, in valsoid configuration, globose to oblong, coriaceous,
aseptate . . . . . . . . . . . . . . . . . . . . . . . . . . . . Khuskia black to brown, with or without long neck which is
3. Ascomata superficial or semi-immersed with/without swollen at the tip, ostiolate. Ostiole periphysate, open
multi-setae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 through the neck. Peridium thin, comprising outer, dark
3. Ascomata immersed or semi-immersed, with a single brown, thick-walled, 4–6 textura angularis cells and in-
long seta arising from the ostiole . . . . Unisetosphaeria ner, small, hyaline, thin-walled, 5–7 textura angularis
4. Ascomata carbonaceous or sub-carbonaceous. . . . . . . 5 cells. Hamathecium comprising few, hyaline paraphyses.
4. Ascomata coriaceous or membranaceous . . . . . . . . . . 6 Asci unitunicate, 8-spored, clavate, short-stalked, apex
5. Asci cylindrical . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 round, with apical ring. Ascospores uniseriate to
5. Asci clavate . . . . . . . . . . . . . . . . . . . . . Schweinitziella biseriate, 0–1-septate, allantoid, fusiform or ellipsoid,
6. Asci cylindrical . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 hyaline, smooth-walled. Asexual morph: Stromata
6. Asci clavate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 uniloculate, black, circular in shape. Locule composed
7. Ascomata superficial . . . . . . . . . . . . . . .Cryptadelphia of numerous interconnecting chambers arranged radially
7. Ascomata semi-immersed, with hispid or irregularly within a continuous mass of ectostromatic
setae . . . . . . . . . . . . . . . . . . . . . . . . . . .Eriosphaeria tissue, one conidiomata per locule. Conidiomata pyri-
8. Ascospores septate . . . . . . . . . . . . . . . . . . . . . . . . . . 10 form in section, brown, divided into compartments by
8. Ascospores aseptate . . . . . . . . . . . . . . . . . . . . . . . . . 11 bending of peridium. Peridium consists of brown, 5–7
9. Perithecial setae with an apical cell . . . . . . . . . . . . . 12 layers of textura angularis cells. Conidiophores reduced
9. Perithecial setae without an apical cell . Cresporhaphis to conidiogenous cells. Conidiogenous cells arising from
10. Ascospores ellipsoidal, narrowly rounded ends, 3- conidiomatal wall, phialidic, simple or branched, hya-
septate . . . . . . . . . . . . . . . . . . . . . . . Coniobrevicolla line, cylindrical. Conidia unicellular, allantoid, hyaline,
10. Ascospores oblong, 0–1-septate . . . . .Setocampanula smooth-walled.
11. Ascospores not fused . . . . . . . . . . . . . . . . . . . . . . . 13 Type: Cytospora Fr. (=Valsa Fr.)
11. Ascospores fusing vertically in groups of two to four Notes: The family Valsaceae was introduced by Tulasne
ascospores . . . . . . . . . . . . . . . . . . . . Collematospora and Tulasne (1861). This family comprises both phytopatho-
12. Ascospores fusiform. . . . . . . . . . . . . . . . Kananascus genic species and saprobes. The type genus Cytospora was
12. Ascospores elongate. . . . . . . . . . . . . . . . . . . . Rizalia typified by Cytospora chrysosperma (Pers.) Fr. and Cytospora
13. Ascospores broadly ellipsoidal . . . . Acanthosphaeria species have been reported as the asexual morph of most taxa
13. Ascospores narrowly ellipsoidal to in Valsaceae. Currently this family comprises 13 genera:
fusiform . . . . . . . . . . . . . . . . . . . . . Trichosphaeria Amphicytostroma, Chadefaudiomyces, Cryptascoma,
Cytospora, Ditopellina, Durispora, Harpostroma,
Hypospilina, Kapooria, Leptosillia, Maculatipalma,
Key to the asexual genera of Trichosphaeriaceae Pachytrype and Paravalsa (Maharachchikumbura et al.
2015).
1. Conidiomata macronematous or synnematous. . . . . . . . . 2
1. Conidiomata pycnidial . . . . . . . . . . . . . Cresporhaphis Cytospora Ehrenb., Sylv. mycol. berol. (Berlin): 2 (1818)
2. Conidiomata macronematous . . . . . . . . . . . . . . . . . . . 3 = Valsa Fr., Summa veg. Scand., Section Post.
2. Conidiomata synnematous . . . . . . . . . . . Fluviostroma (Stockholm): 410 (1849)
3. Conidia aseptate. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 = Valsella Fuckel, Jb. nassau. Ver. Naturk. 23–24: 203
3. Conidia with 1–3 septa. . . . . . . . . . . . .Brachysporium (1870) [1869–70]
4. Conidia fusiform with narrow apex, hyaline, with mu- = Leucostoma (Nitschke) Höhn., Ber. dt. bot. Ges. 35: 637
coid appendages at both ends. . . . . . . . . Koorchaloma (1917)
4. Conidia egg-shaped to flattened-spherical, black, with- = Valseutypella Höhn., Annls mycol. 16(3/6): 224 (1919)
out mucoid appendages . . . . . . . . . . . . . . . . . Khuskia [1918]
Fungal Diversity

Facesoffungi number: FoFxxxxx; Figs 148, 149, 150 Type species: Ditopellina saccardiana (Traverso &
Pathogenic or saprobic on plant tissues. Sexual morph: Spessa) J. Reid & C. Booth [as ‘Ditopellina saccardoana’],
Ectostroma circular or irregular, usually well developed in the (1967), Can. J. Bot. 45: 1481 (1967)
upper regions. Entostroma normally limited to the region near Durispora K.D. Hyde, Sydowia 46(2): 315 (1994)
the perithecial walls. Ascomata perithecia, immersed, aggre- Type species: Durispora elaeidicola K.D. Hyde, Sydowia
gated 6–10 ascomata together in valsoid configuration, glo- 46(2): 316 (1994)
bose to oblong, coriaceous, black, with long neck swollen at Harpostroma Höhn., in Weese, Mitt. bot. Inst. tech.
the tip, ostiolate. Ostiole periphysate, open through the neck. Hochsch. Wien 5: 112 (1928)
Peridium thin, consist of outer, dark brown, thick-walled, 4–6 Type species: Harpostroma notha (Sacc.) Höhn., in
textura angularis cells and inner, small, hyaline, thin-walled, Weese, Mitt. bot. Inst. tech. Hochsch. Wien 5: 112 (1928)
5–7 textura angularis cells. Hamathecium comprising few Hypospilina (Sacc.) Traverso, Fl. ital. crypt.,
paraphyses occurring only in young stage. Asci unitunicate, Pyrenomycetae (Florence): 495 (1913)
8-spored, clavate, short-stalked, apex round, with apical ring. Type species: Hypospilina bifrons (DC.) Traverso, Fl. ital.
Ascospores biseriate, unicellular, allantoid or ellipsoid, hya- crypt. (Florence) 1: 495 (1913)
line, smooth-walled. Asexual morph: Stromata one locule in Kapooria J. Reid & C. Booth, Can. J. Bot. 67(3): 898
a stroma, black, circular in shape. Locule composed of numer- (1989)
ous interconnecting chambers arranged radially or irregularly Type species: Kapooria musarum (J.N. Kapoor) J. Reid &
within a continuous mass of ectostromatic tissue, one C. Booth, Can. J. Bot. 67(3): 898 (1989)
conidiomata per locule. Conidiomata pyriform in section, Leptosillia Höhn., Mitt. bot. Inst. tech. Hochsch. Wien 5:
brown, divided into compartments by bending of peridium. 111 (1928)
Peridium consists of brown, 5–7 layers of textura angularis Type species: Leptosillia notha Höhn., Mitt. bot. Inst. tech.
cells. Conidiophore reduced to conidiogenous cells. Hochsch. Wien 5: 111 (1928)
Conidiogenous cell arising from conidiomatal wall, phialidic, Maculatipalma J. Fröhl. & K.D. Hyde, Mycol. Res. 99(6):
simple or branched, hyaline, cylindrical. Conidia unicellular, 727 (1995)
allantoid, hyaline, smooth-walled. Type species: Maculatipalma frondicola J. Fröhl. & K.D.
Notes: The genus Cytospora was introduced in 1818, typ- Hyde [as ‘fronsicola’], Mycol. Res. 99(6): 727 (1995)
ified by Cytospora chrysosperma. (Adams et al. 2006) Pachytrype Berl. ex M.E. Barr et al., Mycotaxon 48: 530
Maharachchikumbura et al. (2015) and Rossman et al. (1993)
(2015) proposed to conserve the older name Cytospora sup- Type species: Pachytrype princeps (Penz. & Sacc.) M.E.
pressing Leucostoma (Nitschke) Höhn., Valsa Fr., Valsella Barr et al., Mycotaxon 48: 530 (1993)
Fuckel, and Valseutypella Höhn. Cytospora is a plant Paravalsa Ananthap., Mycol. Res. 94(2): 275 (1990)
pathogen forming diseases on various tree species and Type species: Paravalsa indica Ananthap., Mycol. Res.
recently Liu et al. (2015) and Ariyawansa et al. (2015) intro- 94(2): 275 (1990)
duced few new Cytospora species from Russia.
Type species: Cytospora chrysosperma (Pers.) Fr., Syst.
mycol. (Lundae) 2(2): 542 (1823) Key to genera of Valsaceae

1. Asexual morph common in nature; sexual morph

Other genera included undetermined. . . . . . . . . . . . . . . . . . . . Amphicytostroma
1. Asexual morph not common in nature . . . . . . . . . . . . 2
Amphicytostroma Petr., Annls mycol. 19(1/2): 63 (1921) 2. Stromata absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
Type species: Amphicytostroma tiliae (Sacc.) Petr., Annls 2. Stromata present . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
mycol. 19(1/2): 63 (1921) 3. Ascospores apiosporous . . . . . . . . . . . . . . . . . . . . . . . 4
Chadefaudiomyces Kamat et al., in Ullasa et al., Revue 3. Ascospores not apiosporous . . . . . . . . . . . . . . . . . . . . 5
Mycol., Paris 38(1–2): 19 (1974) [1973] 4. Perithecia erect. . . . . . . . . . . . . . . . . . . Maculatipalma
Type species: Chadefaudiomyces indicus Kamat et al., 4. Perithecia horizontal. . . . . . . . . . . . Chadefaudiomyces
Revue Mycol., Paris 38(1–2): 20 (1974) [1973] 5. Ascospores fusiform, with polar spines or appendages6
Cryptascoma Ananthap., Trans. Br. mycol. Soc. 90(3): 479 5. Ascospores allantoid, without polar spines . . Paravalsa
(1988) 6. Ascospores with apical and basal appendages Durispora
Type species: Cryptascoma bisetula Ananthap., Trans. Br. 6. Ascospores without apical and basal appendages
mycol. Soc. 90(3): 479 (1988) Cryptascoma
Ditopellina J. Reid & C. Booth, Can. J. Bot. 45: 1481 7. Stromatic capsules pustuliform (composed of 1, rarely 2
(1967) ascomata). . . . . . . . . . . . . . . . . . . . . . . . . Hypospilina
 Fungal Diversity

Fig. 149 Cytospora chrysosperma – sexual morph (Material examined: substrate e Cross section of ascomata f Peridium g–j Asci k-p
SOUTHERN MORAVIA, near Lednice, Dolniles, 48.8000736N, Ascospores. Scale bars: c, d = 500 μm, e = 100 μm, f–j = 20 μm, k–
16.8445072E, on twigs of Tilia sp., 16 October 1912, H. Zimmermann, p = 10 μm
PRM: 163781, neotype) a, b Herbarium specimen c, d Ascomata on

7. Stromatic capsules non-pustuliform (composed of more 8. Stromata pseudoparenchymatous . . . . . . . . . . . . . . . 10

than 2 ascomata) . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 9. Ascospores apiosporous . . . . . . . . . . . . . Harpostroma
8. Stromata prosenchymatous. . . . . . . . . . . . . . . . . . . . . 9 9. Ascospores not apiosporous . . . . . . . . . . . . .Leptosillia
Fungal Diversity

Fig. 150 Cytospora chrysosperma – asexual morph (Material examined: PRM: 163781, neotype) a Conidiomata on substrate b Cross section of
SOUTHERN MORAVIA, near Lednice, Dolniles, 48.8000736N, conidiomata c Peridium d-h Conidiophore with conidia i Conidia. Scale
16.8445072E, on twigs of Tilia sp., 16 October 1912, H. Zimmermann, bars: a = 500 μm, b = 100 μm, c = 50 μm, d–h = 10 μm, i = 5 μm

10. Ascospores ellipsoid to more or less allantoid . . . . 11 sometimes tapering towards the apex or base, sessile,
10. Ascospores fusoid but with sharply pointed thin-walled except at the apex, apex obtuse or truncate,
ends . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Kapooria with subconical or subapical, J+ apical ring. Ascospores
11. Perithecia in groups with convergent beaks; asci clavate biseriate, triseriate to fasciculate, sometimes weakly heli-
to fusoid . . . . . . . . . . . . . . . . . . . . . . . . . . . Cytospora cally coiled, strongly isthmoid, ends fusiform to rhombic,
11. Perithecia in groups with non-convergent beaks; asci apical portion sometimes slightly larger than the basal
more or less rectangular . . . . . . . . . . . . . . Pachytrype part, 1–3-septate, hyaline, smooth-walled. Asexual
morph: Coelomycetous. Conidiomata pycnidia, superfi-
Vialaeaceae P.F. Cannon, Mycol. Res. 99(3): 368 (1995) cial, solitary, scattered, globose, with slimy, shining spore
Facesoffungi number: FoF 01914 mass and basal mycelium forming thick, black strands.
Parasitic, endophytic or saprobic on dead plant matter. Conidiophores erect, branched, septate, hyaline.
Sexual morph: Pseudostromata appear as ellipsoidal, Conidiogenous cells phialidic, discrete or in small
black, slightly raised dots on substrate. Ascomata perithe- whorls, lageniform to cylindrical, hyaline. Conidia ob-
cial, solitary or aggregated, immersed, globose, sub- long to ellipsoidal, one-celled, hyaline, smooth, with
globose to ellipsoidal, coriaceous, black to brown, papil- truncate abscission scar.
late, ostiolate. Papilla long, straight or curved, Type genus: Vialaea Sacc.
periphysate. Periphyses hyaline, filamentous. Peridium Notes: Vialaeaceae was formally established by Cannon
thick, composed of outer, dark brown, thick-walled cells (1995). Vialaea is the type genus of this family and is typified
of textura angularis and inner, hyaline, thick-walled cells by Vialaea insculpta (Fr.) Sacc. The phylogenetic placement
of textura angularis. Hamathecium comprising filiform, of this monotypic family was confirmed by LSU sequence
rarely branched, septate paraphyses, sometimes apex data (Shoemaker et al. 2013), combined LSU and ITS se-
slightly swollen. Asci 8-spored, unitunicate, cylindrical, quence data (Senanayake et al. 2014) and combined LSU,
 Fungal Diversity

SSU, TEF1 and RPB2 sequence data (Maharachchikumbura Parasitic, endophytic or saprobic on twigs of trees. Sexual
et al. 2015). morph: Pseudostromata (pseudoclypeus) absent or if present,
immersed in host tissues, with brown to black tissues around
Vialaea Sacc., Bull. Soc. mycol. Fr. 12: 66 (1896) immersed neck, containing 1–9 ascomata. Ascomata perithe-
Facesoffungi number: FoF 000011; Figs 151, 152 cial, immersed solitary or aggregated in circular groups, with

Fig. 151 Vialaea insculpta (Material examined: GERMANY, North section of ascoma e Peridium f Periphyses in ostiolar canal g, h Asci i
Rhine-Westphalia, Westfalen, Bottrop, at the zerzogstrasse, on twigs of Apical ring bluing in Melzer’s reagent j-l Ascospores m Ascospores in
Ilex aquifolium L. (Aquifoliaceae), 2 August 1922, H. Rupprecht, S- cotton blue reagent. Scale bars: b = 1 cm, c = 500 μm, d, f = 100 μm,
F121733). a Herbarium packet b, c Pseudostroma on host d Vertical e = 20 μm, g–I = 50 μm, j–m = 20 μm
Fungal Diversity

Fig. 152 Asexual morph of

Vialaea mangiferae (Material
examined: THAILAND, Chiang
Rai Province, Muang District,
near Bandu, Baan Khuakhae, at
31 M. 17, (19° 59′ 52.05″ N; 99°
49′ 25.15″ E), on twigs of
Mangifera indica L.
(Anacardiaceae), 15 November
2012, K.D. Hyde, CHUNI001,
MFLU13-0342, holotype). a
Culture from above b, c Culture
from below d-f Conidiomata
forming on water agar g Conidia
forming on phialides attached to
conidiophores h, i Conidia
attached to lageniform intercalary
cells j Conidia. Note: Conidia,
conidiogenous cells,
conidiophores are arrowed in g
from up to down. Scale bars:
d = 1 mm, e–f = 500 μm, g–j
=10 μm

ostioles converging in their center, globose or subglobose, smooth-walled. Asexual morph: Coelomycetous.
coriaceous, black to brown, papillate, ostiolate. Papilla elon- Conidiomata pycnidial, superficial, solitary, globose.
gate, dark brown to black, internal wall covered by Conidiophores erect, branched, septate, hyaline.
periphyses. Peridium comprising several layers of slightly Conidiogenous cells proliferating percurrently conidia.
flattened, strongly melanized, thick-walled cells of textura Conidia oblong to fusiform, one-celled, hyaline, smooth-
globulosa or textura angularis with peridial pores between walled.
cells. Hamathecium comprising poorly developed, thin- Type species: Vialaea insculpta (Fr.) Sacc., Bull. Soc.
walled, septate or aseptate, sparse paraphyses, sometimes del- mycol. Fr. 12: 66 (1896).
iquescent at maturity, much shorter than the asci. Asci 8- Notes: Vialaea is typified by Vialaea insculpta (Fr.) Sacc.
spored, unitunicate, cylindrical, sometimes tapering towards (Saccardo 1896) and was placed in Amphisphaeriaceae
the apex or base, short pedicellate or sessile, thin-walled ex- (Xylariales) based on the J+ apical ring, and Cannon (1995)
cept at the apex, apex obtuse or truncate, with subconical or placed the genus in Vialaeaceae. LSU sequence analysis
subapical, J+ apical ring. Ascospores biseriate, triseriate to showed that Vialaeaceae is a distinct monotypic family in
fasciculate, sometimes weakly helically coiled, strongly the order Xylariales (Shoemaker et al. 2013). A new species,
isthmoid, ends fusiform to rhombic, apical portion sometimes Vialaea mangiferae Senan. & K. D. Hyde was introduced in
slightly larger than the basal part, 1–3-septate, hyaline, the genus by Senanayake et al. (2014).
 Fungal Diversity

Xylariaceae Tul. & C. Tul. [as ‘Xylariei’], Select. fung. subfamilies (Chesters and Greenhalgh 1964; Ju and Rogers
carpol. (Paris) 2: 3 (1863) 1996; Bitzer et al. 2008; Stadler et al. 2010). However this
= Graphostromataceae M.E. Barr et al., Mycotaxon 48: grouping of taxa with geniculosporium-like (Xylarioideae)
533 (1993) and nodulisporium-like (Hypoxyloideae) conidiophores, was
Facesoffungi number: FoF 00070 not formerly introduced by the authors. Most subsequent stud-
Saprobic, pathogenic, or endophytic in wood, leaves and ies based on phylogenetic analyses have strongly supported
fruits, dung inhabitants or associated with insect vectors, in this separation of subfamilies within the Xylariaceae.
terrestrial and aquatic habitats. Sexual morph: Stromata if Pirozynski (1974) found that Sphaeria platystoma
present extremely variable in size, shape and colour, erect or Schwein. resembled Diatrype stigma (Hoffm.) Fr. and placed
applanate or effuse-pulvinate or sometimes rudimentary, aris- it in a new monotypic genus Graphostroma. Graphostroma
ing singly or aggregated, with 1 to many ascomata, ostiolate, platystomum (Schwein.) Piroz. 1974 was associated with a
with or without extractable stromatal pigments, bipartite or nodulisporium-like asexual morph, which was identical to
unipartite. Ascomata variable in size and shape, globose-pyr- most of the hypoxyloid Xylariaceae. Therefore, Pirozynski
iform, embedded in the stroma, single or multi-layered. (1974) placed Graphostroma platystomum in Xylariaceae.
Hamathecium comprising hyphoid, filamentous, septate pa- Later, Barr et al. (1993) introduced a new monotypic family
raphyses, embedded in a gelatinous matrix. Asci 4–8-spored, Grap hostromata ceae to accommodate the genus
unitunicate, cylindrical to clavate, pedicellate, apically round- Graphostroma. This segregation was based on the combina-
ed, with or without a J+, or J–, apical ring, or with apical tion of its nodulisporium-like conidial morph and a sexual
thickenings. Ascospores uniseriate-biseriate (except for the morph similar to that of the Diatrypaceae, in having allantoid
genera Phylacia and Pyrenomyxa which are produced in ascospores (Barr et al. 1993). Phylogenetic studies (Stadler
spherical or oval clusters), brown to black, rarely hyaline, 1– et al. 2013; Maharachchikumbura et al. 2015) placed
2-celled, variously-shaped, mostly with a germ slit. Perispore Graphostromataceae close to Xylariaceae, while in
dehiscent or lacking, smooth or with patterns. Asexual Daranagama et al. (2015) and Senanayake et al. (2015)
morph: Hyphomycetous. Two major types of asexual morph Graphostroma platystomum clusters with xylariaceous genera
reported are nodulisporium-like and geniculosporium-like as with bipartite stromata, such as Biscogniauxia and Camillea
defined by Bitzer et al. (2008), featuring hyaline to light within the Xylariaceae clade. Thus the position of the
brown, smooth, branched conidiophores, bearing hyaline, Graphostromataceae as a separate family has been shown to
roughened or smooth, ellipsoidal conidia. Several other hy- be doubtful with the available molecular data. Considering
phomycetous and coelomycetous genera have been linked to these molecular studies, as well as the morphological charac-
Xylariaceae (Stadler et al. 2013, Senanayake et al. 2015). ters, especially the asexual morph which is nodulisporium-
Type: Xylaria Hill ex Schrank like, we believe that Graphostroma should be considered as
Notes: The family Xylariaceae was introduced by Tulasne a genus in Xylariaceae. Therefore, we have synonymized
and Tulasne (1863b) using the term “Xylariei” and the rank of Graphostromataceae and consider Graphostroma as a genus
the name was not certain as it did not address the family concept of Xylariaceae.
(Stadler et al. 2013). Even though the exact number of currently Fassia was introduced by Dennis (1964) with Fassia
accepted taxa varies, according to different authors, the recent scabrosa Dennis as the type species. This monotypic genus
outline of Xylariaceae (Maharachchikumbura et al. 2015) ac- is characterized by superficial stromata with long, slender,
cepted 85 genera. The segregation of Xylariaceae into the sub- protruding, ostiolar necks (Dennis 1964). According to
families Hypoxyloideae and Xylarioideae has been supported by Dennis (1964), Müller observed part of the type species col-
molecular data (Stadler et al. 2013; Daranagama et al. 2015; lection and suggested a relationship with Wawelia. Læssøe
Maharachchikumbura et al. 2015). Xylariaceous members are (1994), however, suggested that Fassia scabrosa should be
well-known for their secondary metabolite production (Stadler placed in Diatrypaceae, based on its morphological similarity
2011). Several studies have been carried out to determine the to Echinomyces, which is accepted in Diatrypaceae. Rappaz
chemical composition of these secondary metabolites (Whalley (1987) had not compared Echinomyces with Fassia when de-
and Edwards 1995; Mühlbauer et al. 2002; Quang et al. 2002, scribing the new genus. The clavate asci with allantoid asco-
2004, 2006; Hellwig et al. 2005). Chemotaxonomic data also spores are characters of diatrypaceous taxa, rather than
play an important role in the identification and characterization xylariaceous taxa, therefore we prefer to exclude Fassia from
of these stromatic Xylariaceae (Whalley and Edwards 1995; Xylariaceae and place it in Xylariales genera incertae sedis
Stadler et al. 2001; Bitzer et al. 2008; Kuhnert et al. 2014). For until the phylogenetic position is established by fresh
example, Stadler et al. (2014a) used a polyphasic taxonomic collections and molecular data.
approach in their world monograph of Daldinia. The genus Muscodor was introduced by Worapong et al.
Based on the asexual morphs and their characters, (2001) for an endophytic fungus, Muscodor albus Worapong
Xylariaceae has been traditionally classified into two et al., which produces volatile antibiotics. Since then 16
Fungal Diversity

species of Muscodor have been described by various authors 2014b). Most Xylaria species are saprobic, growing on moist,
(Kirk et al. 2008), based on the differences in their chemical decayed dicotyledonous wood, less on monocotyledons,
profiles of volatile compounds. However, when describing fruits, seeds and fallen leaves. Interestingly the subgenus
this genus, Worapong et al. (2001) used only a few reference Pseudoxylaria, which is associated with termite nests (Visser
DNA sequences representing only four genera of Xylariaceae, et al. 2009; Hsieh et al. 2010) are also commonly endophytes
which were not derived from type species strains. mainly from tropical plants (Petrini and Petrini 1985; Bills
Furthermore, they did not compare the morphology when et al. 2012).
discussing the current status of Muscodor and presented argu-
ments as to why it cannot be recognized as a typical
Xylariaceae member. The concept of this genus is incongruent
with other concepts of Xylariaceae, thus in this study Other genera included
Muscodor is placed in Xylariales genera incertae sedis, until
it can be studied in relation to xylarioid Xylariaceae with Amphirosellinia Y.M. Ju et al., Mycologia 96(6): 1393 (2004)
molecular data. Type species: Amphirosellinia nigrospora Y.M. Ju et al.,
Xylaria Hill ex Schrank, Baier. Fl. (München) 1: 200 Mycologia 96(6): 1397 (2004)
(1789) Annulohypoxylon Y.M. Ju et al., in Hsieh et al., Mycologia
Facesoffungi number: FoF 00696; Figs 153, 154 97(4): 855 (2005)
Saprobic on deciduous dead wood or endophytes of vari- Type species: Annulohypoxylon truncatum (Schwein.)
ous plants. Sexual morph: Stromata extremely variable in Y.M. Ju et al. Mycologia 97(4): 861 (2005)
size and shape, simple to branched from the base, nearly ses- Anthocanallis Daranagama et al., Fungal Diversity
sile or arising from long rooting stipes, always with flattened 73:203–238 (2015)
to mucronate sterile apices, whitish when immature, gradually Type species: Anthocanalis sparti Daranagama et al.,
turning silvery grey and eventually black, longitudinally Fungal Diversity 73:203–238 (2015)
furrowed delimiting narrow strips, perithecial contours most Anthostomella Sacc., Atti Soc. Veneto-Trent. Sci. Nat.,
often inconspicuous in well-developed stromata; leathery, Padova, Sér. 4 4: 84 (1875)
black, homogeneous, white to cream, with a slightly darker Type species: Anthostomella tomicoides Sacc., Atti Soc.
core, stipes cylindrical to strap-like, longitudinally puckered, Veneto-Trent. Sci. Nat., Padova, Sér. 4 4: 101 (1875)
black, with a hairy-tomentose, broadened base. Perithecia Appendixia B.S. Lu & K.D. Hyde, Fungal Diversity Res.
subglobose, immersed to slightly exposed. Ostioles if present Ser. 4: 224 (2000)
raised-discoid, grey brown to black, with a low conical papilla Type species: Appendixia closterium (Berk. & M.A.
at the center. Asci (6–)8-spored, unitunicate, cylindrical, long- Curtis) B.S. Lu & K.D. Hyde, Fungal Diversity Res. Ser. 4:
pedicellate, with tubular, apical ring, bluing in Melzer’s re- 225 (2000)
agent. Paraphyses sparse, hypha-like, hyaline, septate. Areolospora S.C. Jong & E.E. Davis, Norw. Jl Bot. 21(1):
Ascospores overlapping uniseriate in the ascus, medium 28 (1974)
brown, most often ellipsoid-inequilateral, with narrow to Type species: Areolospora terrophila S.C. Jong & E.E.
broadly rounded ends, smooth-walled, with a fugacious cellu- Davis, Norw. Jl Bot. 21(1): 28 (1974)
lar appendage, usually disappearing at maturity, containing Arthroxylaria Seifert & W. Gams, in Seifert, Gams &
two large guttules, with a conspicuous straight germ slit, Louis-Seize, Czech Mycol. 53(4): 299 (2002)
1/2–4/5 spore-length on the flattened side. Atypical asco- Type species: Arthroxylaria elegans Seifert & W. Gams, in
spores may have acute ends, slightly oblique or sinuous germ Seifert, Gams & Louis-Seize, Czech Mycol. 53(4): 299 (2002)
slits located next to one end, or on the convex side. Asexual Ascotricha Berk., Ann. nat. Hist., Mag. Zool. Bot. Geol. 1:
morph: Where known geniculosporium–like, hyaline–light 257 (1838)
brown, smooth, branched conidiophores bearing hyaline, Type species: Ascotricha chartarum Berk., Ann. nat. Hist.,
roughened or smooth, ellipsoidal conidia (Ju and Rogers Mag. Zool. Bot. Geol. 1: 257 (1838)
1996; Stadler et al. 2014b). Astrocystis Berk. & Broome, J. Linn. Soc., Bot. 14(no. 74):
Type species: Xylaria hypoxylon (L.) Grev., Fl. Edin.: 355 123 (1873) [1875]
(1824) Type species: Astrocystis mirabilis Berk. & Broome, J.
Notes: Xylaria is the type genus in the family Xylariaceae Linn. Soc., Bot. 14(no. 74): 123 (1873) [1875]
with more than 700 epithets listed in Index Fungorum (2016). Barrmaelia Rappaz, Mycol. helv. 7(1): 130 (1995)
Many Xylaria species possess massive stromata, which can Type species: Barrmaelia rhamnicola Rappaz, Mycol.
vary in colour, shape and size, cylindrical asci, a J+ apical ring helv. 7(1): 130 (1995)
and ellipsoidal, dark ascospores, as well as geniculosporium- Biscogniauxia Kuntze, Revis. gen. pl. (Leipzig) 2: 398
like asexual morphs (Ju and Rogers 1996; Stadler et al. 2013, (1891)
 Fungal Diversity

Fig. 153 Xylaria hypoxylon (SWEDEN, Uppland, Stockholm, stromata showing perithecia encased in stromatal tissue f Asci with J+,
Vällingby, Grimsta Nature Reserve, near Lake Mälaren, on Sorbus apical ring, bluing in Melzer’s reagent g Cluster of asci and paraphyses h,
aucuparia L. (Rosaceae), 21 October 2007, A.-L. & A. Anderberg i Mature asci j–m Ascospores. Scale bars: c = 5 mm d = 150 μm,
G07-1 (S-F72430, epitype) a Herbarium packet b Stromata c Furrowed e = 0.5 mm, f, g, h = 20 μm, g = 30 μm, j–m = 10 μm
stromatal surface d Black ostioles in stromatal surface e Cross section of

Type species: Biscogniauxia nummularia (Bull.) Kuntze, Camillea Fr., Summa veg. Scand., Section Post.
Revis. gen. pl. (Leipzig) 2: 398 (1891) (Stockholm): 382 (1849)
Brunneiperidium Daranagama et al., Fungal Diversity Type species: Camillea leprieurii (Mont.) Mont., Summa
73:203–238 (2015) veg. Scand., Section Post. (Stockholm): 382 (1849)
Type species: Brunneiperidium gracilentum Daranagama Cannonia Joanne E. Taylor & K.D. Hyde, Mycol. Res.
et al., Fungal Diversity 73:203–238 (2015) 103(11): 1398 (1999)
Calceomyces Udagawa & S. Ueda, Mycotaxon 32: 448 Type species: Cannonia australis (Speg.) Joanne E. Taylor
(1988) & K.D. Hyde, Mycol. Res. 103(11): 1398 (1999)
Type species: Calceomyces lacunosus Udagawa & S. Chaenocarpus Rebent., Prodr. fl. neomarch. (Berolini):
Ueda, Mycotaxon 32(1): 448 (1988) 350 (1804)
Fungal Diversity

Fig. 154 Xylaria sp. with

geniculosporium–like asexual
morph a Macroscopic features on
natural habitat b, c Conidiophores
d, e Conidiophores with spores f
Conidia g Conidiogenous cells at
the apex of conidiophores. Scale
bars: b–e, g = 10 μm, e = 5 μm

Type species: Chaenocarpus setosus (Roth) Rebent., Creosphaeria Theiss., Beih. bot. Zbl., Abt. 2 27: 396 (1910)
Prodr. fl. neomarch. (Berolini): 350, tab. 3:12a-b (1804) Type species: Creosphaeria sassafras (Schwein.) Y.M. Ju
Chlorostroma A.N. Mill. et al., Sydowia 59(1): 142 (2007) et al. Mycotaxon 47: 223 (1993)
Type species: Chlorostroma subcubisporum A.N. Mill Cyanopulvis J. Fröhl. & K.D. Hyde, Fungal Diversity Res.
et al., Sydowia 59(1): 142 (2007) Ser. 3: 308 (2000)
Collodiscula I. Hino & Katum., Bull. Faculty of Type species: Cyanopulvis australiensis J. Fröhl. & K.D.
Agriculture, Yamaguchi University 6: 55 (1955) Hyde, Fungal Diversity Res. Ser. 3: 308 (2000)
Type species: Collodiscula japonica I. Hino & Katum., Daldinia Ces. & De Not., Comm. Soc. crittog. Ital. 1(4):
Bull. Faculty of Agriculture, Yamaguchi University 6: 55 (1955) 197 (1863)
Coniolariella Dania García et al., in García et al., Mycol. Type species: Daldinia concentrica (Bolton) Ces. & De
Res. 110(11): 1285 (2006) Not., Comm. Soc. crittog. Ital. 1(4): 197 (1863)
Type species: Coniolariella gamsii (Asgari & Zare) Dania Durotheca Læssøe et al. IMA Fungus 4(1): 62 (2013)
García et al., in García et al., Mycol. Res. 110(11): 1285 Type species: Durotheca depressa Læssøe & Srikitik.,
(2006) IMA Fungus 4(1): 62 (2013)
 Fungal Diversity

Emarcea Duong et al. Stud. Mycol. 50(1): 255 (2004) Type species: Induratia apiospora Samuels et al.,
Type species: Emarcea castanopsidicola Duong et al., in Mycotaxon 28(2): 484 (1987)
Duong et al., Stud. Mycol. 50(1): 255 (2004) *Jumillera J.D. Rogers et al. Mycotaxon 64: 41 (1997)
Engleromyces Henn., Bot. Jb. 28(3): 327 (1900) Type species: Jumillera mexicana J.D. Rogers et al.,
Type species: Engleromyces goetzei Henn., Bot. Jb. 28(3): Mycotaxon 64: 46 (1997)
327 (1900) Kretzschmaria Fr., Summa veg. Scand., Section Post.
Entoleuca Syd., Annls mycol. 20(3/4): 186 (1922) (Stockholm): 409 (1849)
Type species: Entoleuca callimorpha Syd., in Sydow & Type species: Kretzschmaria clavus (Fr.) Sacc., Syll. fung.
Petrak, Annls mycol. 20(3/4): 186 (1922) (Abellini) 2: XXIX (1883)
Entonaema A. Möller, Bot. Mitt. Trop. 9: 306. (1901) Kretzschmariella Viégas, Bragantia 4(1–6): 105 (1944)
emend. in Stadler et al. Mycol Prog 7:53–73 (2008) Type species: Kretzschmariella culmorum (Cooke) Y.M.
Type species: Entonaema liquescens Möller, Bot. Mitt. Ju & J.D. Rogers, Mycotaxon 51: 242 (1994)
Trop. 9: 307 (1901) Leprieuria Læssøe et al., Mycol. Res. 93(2): 152 (1989)
Euepixylon Füisting, Bot. Ztg. 25(no. 39): 309 (1867) Type species: Leprieuria bacillum (Mont.) Læssøe et al.,
Type species: Euepixylon udum (Pers.) Füisting, Bot. Ztg. Mycol. Res. 93(2): 152 (1989)
25(no. 39): 305, 309 (1867) Leptomassaria Petr., Annls mycol. 12(5): 474 (1914)
Fasciatispora K.D. Hyde, Trans. Mycol. Soc. Japan 32(2): Type species: Leptomassaria simplex (Nitschke ex G.H.
265 (1991) Otth) Petr., Annls mycol. 12(5): 474 (1914)
Type species: Fasciatispora nypae K.D. Hyde, Trans. Libertella Desm., Annls Sci. Nat., Bot., sér. 1 19: 275
Mycol. Soc. Japan 32(2): 267 (1991) (1830)
Gigantospora B.S. Lu & K.D. Hyde, Nova Hedwigia Type species: Libertella betulina Desm., Annls Sci. Nat.,
76(1–2): 202 (2003) Bot., sér. 1 19: 276 (1830)
Type species: Gigantospora gigaspora B.S. Lu & K.D. Lunatiannulus Daranagama et al., Fungal Diversity
Hyde, Nova Hedwigia 76(1–2): 202 (2003) 73:203–238 (2015)
Graphostroma Piroz., Can. J. Bot. 52(10): 2131 (1974) Type species: Lunatiannulus irregularis Daranagama
Type species: Graphostroma platystoma (Schwein.) Piroz., et al., Fungal Diversity 73:203–238 (2015)
Can. J. Bot. 52(10): 2131 (1974) Myconeesia Kirschst., Annls mycol. 34(3): 200 (1936)
Guestia G.J.D. Sm. & K.D. Hyde, Fungal Diversity 7: 107 Type species: Myconeesia formosa (Kirschst.) Kirschst.,
(2001) Annls mycol. 34(3): 200 (1936)
Type species: Guestia gonetropospora G.J.D. Sm. & K.D. Nemania Gray, Nat. Arr. Brit. Pl. (London) 1: 508, 516
Hyde, Fungal Diversity 7: 109 (2001) (1821)
Halorosellinia Whalley et al., in Whalley et al., Mycol. Type species: Nemania serpens (Pers.) Gray, Nat. Arr. Brit.
Res. 104(3): 368 (2000) Pl. (London) 1: 508, 516 (1821)
Type species: Halorosellinia oceanica (S. Schatz) Whalley Nipicola K.D. Hyde, Cryptog. bot. 2(4): 330 (1992)
et al., Mycol. Res. 104(3): 370 (2000) Type species: Nipicola carbospora K.D. Hyde, Cryptog.
Helicogermslita Lodha & D. Hawksw., in Hawksworth & bot. 2(4): 330 (1992)
Lodha, Trans. Br. mycol. Soc. 81(1): 91 (1983) Nodulisporium Preuss, in Rabenhorst, Klotzschii Herb.
Type species: Helicogermslita celastri (S.B. Kale & S.V.S. Viv. Mycol.: no. 1272 (1849)
Kale) Lodha & D. Hawksw., in Hawksworth & Lodha, Trans. Type species: Nodulisporium ochraceum Preuss [as
Br. mycol. Soc. 81(1): 91 (1983) ‘ocheraceum’], Klotzschii Herb. Viv. Mycol.: no. 1272
Hypocopra (Fr.) J. Kickx f., Fl. Crypt. Flandres (Paris) 1: (1849)
362 (1867) Obolarina Pouzar, Česká Mykol. 40(1): 7 (1986)
Type species: Hypocopra merdaria (Fr.) J. Kickx f., Fl. Type species: Obolarina dryophila (Tul. & C. Tul.) Pouzar,
Crypt. Flandres (Paris) 1: 362 (1867) Česká Mykol. 40(1): 7 (1986)
Occultitheca J.D. Rogers & Y.M. Ju, Sydowia 55(2): 359
Hypocreodendron Henn., Hedwigia 36: 223 (1897)
(2003)
Type species: Hypocreodendron sanguineum Henn.,
Type species: Occultitheca costaricensis J.D. Rogers &
Hedwigia 36(4): 223 (1897)
Y.M. Ju, Sydowia 55(2): 360 (2003)
Hypoxylon Bull., Hist. Champ. Fr. (Paris) 1: 168 (1791); Ophiorosellinia J.D. Rogers et al., in Rogers et al.,
Fig. 155 Mycologia 96(1): 172 (2004)
Type species: Hypoxylon fragiforme (Pers.) J. Kickx f., FI. Type species: Ophiorosellinia costaricensis J.D. Rogers
crypt. Louvain (Bruxelles): 116 (1835) et al., in Rogers et al., Mycologia 96(1): 172 (2004)
Induratia Samuels et al. Mycotaxon 28(2): 482 (1987) Pandanicola K.D. Hyde, Sydowia 46(1): 35 (1994)
Fungal Diversity

Fig. 155 Hypoxylon fendleri with nodulisporium–like asexual morph 0104, ICMP). asexual morph a Culture from above b From below c
(Material examined: THAILAND, Chiang Rai, Doi Mae Saloung on Conidia d-f Conidiophores with spores. Scale bars: c = 20 μm, d–
dead bamboo clumps, 12 December 2012, D. A. Daranagama and K. f = 40 μm
D. Hyde, AXL 055 (MFLU 12-0823, living cultures, MFLUCC 13-

Type species: Pandanicola calocarpa (Syd. & P. Syd.) Poroleprieuria M.C. González et al. Aguirre, Mycologia
K.D. Hyde, Sydowia 46(1): 36 (1994) 96(3): 676 (2004)
Paramphisphaeria F.A. Fernández et al., Mycologia Type species: Poroleprieuria rogersii M.C. González et al.,
96(1): 175 (2004) Mycologia 96(3): 676 (2004)
Type species: Paramphisphaeria costaricensis F.A. Poronia Willd., Fl. berol. prodr.: 400 (1787)
Fernández et al., Mycologia 96(1): 175 (2004) Type species: Poronia gleditschii Willd., Fl. berol. prodr.:
Paucithecium Lloyd, Mycol. Notes (Cincinnati) 7(4): 400 (1787)
1200 (1923) Pyrenomyxa Morgan, J. Cincinnati Soc. Nat. Hist. 18: 42
Type species: Paucithecium rickii Lloyd, Mycol. Notes (1895)
(Cincinnati) 7(4): 1200 (1923) Type species: Pyrenomyxa invocans Morgan, J. Cincinnati
Phylacia Lév., Annls Sci. Nat., Bot., sér. 3 3: 61 (1845) Soc. Nat. Hist. 18: 42 (1895)
Type species: Phylacia globosa Lév., Annls Sci. Nat., Bot., Pyriformiascoma Daranagama et al., Fungal Diversity
sér. 3 3: 61 (1845) 73:203–238 (2015)
Pidoplitchkoviella Kiril., Mikrobiol. Zh. 37(5): 603 (1975) Type species: Pyriformiascoma trilobatum Daranagama
Type species: Pidoplitchkoviella terricola Kiril., et al., Fungal Diversity 73:203–238 (2015)
Mikrobiol. Zh. 37(5): 603 (1975) Rhopalostroma D. Hawksw., Kew Bull. 31(3): 422 (1977)
Podosordaria Ellis & Holw., Bot. Gaz. 24: 37 (1897) Type species: Rhopalostroma indicum D. Hawksw. &
Type species: Podosordaria mexicana Ellis & Holw., Bot. Muthappa, in Hawksworth, Kew Bull. 31(3): 426 (1977)
Gaz. 24: 37 (1897) Rosellinia De Not., G. bot. ital. 1(1): 334 (1844)
 Fungal Diversity

Type species: Rosellinia aquila (Fr.) Ces. & De Not., G. Thuemenella Penz. & Sacc., Malpighia 11(11–12): 518
bot. ital. 1(1): 334 (1844) (1898) [1897]
Rostrohypoxylon J. Fourn. et al. Fungal Diversity 40: 24 Type species: Thuemenella javanica Penz. & Sacc.,
(2010) Malpighia 11(11–12): 518 (1898) [1897]
Type species: Rostrohypoxylon terebratum J. Fourn. & M. Versiomyces Whalley & Watling, Notes R. bot. Gdn Edinb.
Stadler, in Fournier, Stadler, Hyde & Duong, Fungal Diversity 45(2): 401 (1989) [1988]
40: 24 (2009) Type species: Versiomyces cahuchucosus Whalley &
Ruwenzoria J. Fourn. et al., in Stadler et al., Mycol. Progr. Watling, Notes R. bot. Gdn Edinb. 45(2): 401 (1989)
9(2): 171 (2010) [1988]
Type species: Ruwenzoria pseudoannulata J. Fourn. et al., Virgaria Nees, Syst. Pilze (Würzburg): 54 (1816) [1816–
in Stadler et al., Mycol. Progr. 9(2): 171 (2010) 17]
Sabalicola K.D. Hyde, Nova Hedwigia 60(3–4): 596 Type species: Virgaria nigra (Link) Gray, Nat. Arr. Brit. Pl.
(1995) (London) 1: 553 (1817) [1816–17]
Type species: Sabalicola sabalensioides (Ellis & G. Vivantia J.D. Rogers, Y.M. Ju & Cand., Mycol. Res.
Martin) K.D. Hyde, Nova Hedwigia 60(3–4): 597 (1995) 100(6): 672 (1996)
Sarcoxylon Cooke, Grevillea 12(no. 62): 50 (1883) Type species: Vivantia guadalupensis J.D. Rogers, Y.M. Ju
Type species: Sarcoxylon compunctum (Jungh.) Cooke, & Cand., Mycol. Res. 100(6): 672 (1996)
Grevillea 13(no. 68): 107 (1885) Wawelia Namysł., Bull. int. Acad. Sci. Lett. Cracovie, Cl.
Seynesia Sacc., Syll. fung. (Abellini) 2: 668 (1883) sci. math. nat. Sér. B, sci. nat. 2: 602 (1908)
Type species: Seynesia nobilis (Welw. & Curr.) Sacc., Syll. Type species: Wawelia regia Namysł., Bull. int. Acad.
fung. (Abellini) 2: 668 (1883) Sci. Lett. Cracovie, Cl. sci. math. nat. Sér. B, sci. nat.
Spirodecospora B.S. Lu et al., Fungal Diversity Res. Ser. 2: 597 (1908)
1: 170 (1998) *Whalleya J.D. Rogers et al. Mycotaxon 64: 48
Type species: Spirodecospora bambusicola B.S. Lu et al., (1997)
Fungal Diversity 1: 172 (1998) Type species: Whalleya microplaca (Berk. & M.A. Curtis)
Squamotubera Henn., Hedwigia 42(Beibl.): (308) (1903) J.D. Rogers et al. Mycotaxon 64: 48 (1997)
Type species: Squamotubera le-ratii Henn., Hedwigia Xylotumulus J.D. Rogers et al. Sydowia 58(2): 291 (2006)
42(Beibl.): (308) (1903) Type species: Xylotumulus gibbisporus J.D. Rogers et al.
Steganopycnis Syd. & P. Syd., Annls mycol. 14(5): 370 Sydowia 58(2): 291 (2006)
(1916) Xylocrea Möller, Bot. Mitt. Trop. 9: 397 (1901)
Type species: Steganopycnis oncospermatis Syd. & P. Type species: Xylocrea piriformis Möller, Bot. Mitt. Trop.
Syd., Annls mycol. 14(5): 337 (1916) 9: 397 (1901)
Stereosphaeria Kirschst., Annls mycol. 37(1/2): 96 (1939) * Jumillera J.D. Rogers et al. and Whalleya J.D.
Type species: Stereosphaeria phloeophila Kirschst., Annls Rogers et al. were introduced to encompass taxa re-
mycol. 37(1/2): 96 (1939) moved from Biscogniauxia, based on morphological dif-
Stilbohypoxylon Henn., Hedwigia 41: 16 (1902) ferences of their asexual morphs (Rogers et al. 1997).
Type species: Stilbohypoxylon moelleri Henn., Hedwigia Both Jumillera and Whalleya have xylariaceous sexual
41: 16 (1902) morphs with conspicuous, bipartite stromata and asci
Striatodecospora D.Q. Zhou et al., Mycotaxon 76: 142 with an amyloid apical ring, containing dark, ellipsoidal
(2000) ascospores (Rogers et al. 1997). However, both genera
Type species: Striatodecospora bambusae D.Q. Zhou produce scolecosporous conidia, which is a characteris-
et al., Mycotaxon 76: 143 (2000) tic feature of Lopadostomataceae. Some Jumillera spe-
Stromatoneurospora S.C. Jong & E.E. Davis, Mycologia cies produce libertella-like asexual morphs and species
65(2): 459 (1973) such as J. cinerea (Ellis & Everh.) J.D. Rogers et al.
Type species: Stromatoneurospora phoenix (Kunze) S.C. produce geniculosporium-like synanamorphs in culture
Jong & E.E. Davis, Mycologia 65(2): 459 (1973) (Rogers et al. 1997). Whalleya microplaca (Berk. &
Thamnomyces Ehrenb., in Nees von Esenbeck (ed.), M.A. Curtis) J.D. Rogers et al. produced a similar asex-
Horae Phys. Berol.: 79 (1820) ual morph to Lopadostoma pouzarii Granmo & L.E.
Type species: Thamnomyces chamissonis Ehrenb., in Petrini (Granmo and Petrini 1996). Based on these facts
Nees von Esenbeck (ed.), Horae Phys. Berol.: 79 (1820) Jumillera and Whalleya have similarities with both
Theissenia Maubl., Bull. Soc. mycol. Fr. 30(1): 52 (1914) Xylariaceae and Lopadostomataceae. Therefore their
Type species: Theissenia pyrenocrata (Theiss.) Maubl., phylogenetic placement remains to be established as
Bull. Soc. mycol. Fr. 30(1): 52 (1914) fresh material and molecular data become available.
Fungal Diversity

Key for asexual genera in Xylariaceae 12. Stromata hemisphaerical with gelatinous, soft, leathery
interior, without concentric zones . . . . . . .Entonaema
1. Producing single, asexual morph in culture 2 13. Stromata uniperitheciate . . . . . . . . . . . . . . . Induratia
1. Producing multiple, distinct, asexual morphs in 13. Stromata multi-peritheciate. . . . . . . . . . . . . . . . . . . 14
culture. . . . . . . . . . . . . . . . . . . . . . . . . . Arthroxylaria 14. Ostioles with truncatum-type disc, with carbonaceous
2. Conidiogenous cells produced in digitate tissue encasing each individual perithecium, perispore
configurations or false whorls, proliferating with a thickened area, visible at the position of ca. 1/3
percurrently . . . . . . . . . . . . . . . . . .Hypocreodendron ascospore length . . . . . . . . . . . . . .Annulohypoxylon
2. Conidiogenous cells sympodial, with apically aggregat- 14. Ostioles with bovei-type disc, with carbonaceous tis-
ed scars . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 sue, beneath surface and between perithecia, perispore
3. Conidia hyaline . . . . . . . . . . . . . . . . . . Nodulisporium without a thickened area . . . . . . . . . . . . . Hypoxylon
3. Conidia brown Virgaria 15. Stromata in upright position to the substrate . . . . . . 16
15. Stromata in parallel position to the substrate. . . . . . 18
16. With KOH-extractable pigments. . . . . . . . . . . . . . . 17
Key to sexual genera of Xylariaceae 16. Lacking KOH-extractable pigments . . . . . . . . . . . . 20
17. Stromata hemisphaerical to peltate, sessile, with annu-
1. With nodulisporium-like asexual morphs . . . . . Section A late disks surrounding the ostioles . . . . . Ruwenzoria
1. With geniculosporium-like asexual morphs . Section B 17. Stromata stipitate, head part subglobose to clavate,
1. With libertella-like or other unnamed asexual without conspicuous disks surrounding the
morphs . . . . . . . . . . . . . . . . . . . . . . . . . . . . Section C ostioles . . . . . . . . . . . . . . . . . . . . . . . Rhopalostroma
18. Asci globose to subglobose . . . . . . . . . . Pyrenomyxa
18. Asci cylindrical to clavate . . . . . . . . . . . . . . . . . . . 19
Section A – Hypoxyloideae group 19. Stromata pulvinate, with stout ostiolar necks and deep
cylindrical holes . . . . . . . . . . . . . . . Rostrohypoxylon
1. Stromata reduced or absent. . . . . . . . . . . . . . . . . . . . . . . 2 19. Stromata subglobose to cushion-like, with stromatal
1. Stromata present and well-developed . . . . . . . . . . . . . 3 tissues with green tints. . . . . . . . . . . . . . . Thumnella
2. Ascospores “shoe”- shaped, tapered in side view, brown, 20. Ascospores with germ slit . . . . . . . . . Thamnomyces
reticulate with irregular ridges and shallow 20. Ascospores lacking germ slit . . . . . . . . . . . . Phylacia
depressions. . . . . . . . . . . . . . . . . . . . . . . .Calceomyces
2. Ascospores ellipsoidal, olivaceous green, with hyaline
appendage at one end . . . . . . . . . . . . . . Anthocanalis
3. Stromata bipartite. . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 Section B – Xylarioideae group
3. Stromata unipartite. . . . . . . . . . . . . . . . . . . . . . . . . . . 9
4. Stromata in vertical position. . . . . . . . . . . . . . Camellia 1. Stromata reduced or absent, immersed in substratum . . . 2
4. Stromata in horizontal position. . . . . . . . . . . . . . . . . . 5 1. Stromata erumpent or superficial . . . . . . . . . . . . . . . . 8
5. Apical ring present. . . . . . . . . . . . . . . . . . . . . . . . . . . 6 2. Stromata associated with dung . . . . . . . . . . Hypocopra
5. Apical ring lacking. . . . . . . . . . . . . . . . . . . . . . . . . . . 8 2. Stromata associated with substrates other than dung 3
6. Germ slit present . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 3. Apical ring present in asci . . . . . . . . . . . . . . . . . . . . . 4
6. Germ slit lacking . . . . . . . . . . . . . . . . . . . . . . Vivantia 3. Apical ring lacking in asci . . . . . . . . . . . . . . . . . . . . . 6
7. Ascospores unicellular . . . . . . . . . . . . . . . . . . . . . . . . 8 4. Apical ring cylindrical to barrel-shaped (length>width) 5
7. Ascospores 2-celled . . . . . . . . . . . . . . . Biscogniauxia 4. Apical ring discoid (length<width) . . . . . . . Euepixylon
8. Ascospores sub-allantoid and hyaline. . Graphostroma 5. Ascospores brown, unicellular . . . . . . . .Halorosellinia
8. Ascospores ellipsoidal and brown . . . . . . . . Obolarina 5. Ascospores hyaline, 2-celled . . . . . . . . . . . . .Emarcea
9. Ascospores thick-walled, oblong to allantoid Durotheca 6. Ascomatal surface with dark brown to black
9. A s c o s p o r e s t h i n - w a l l e d , s u b g l o b o s e t o hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ascotricha
ellipsoidal . . . . . . . . . . . . . . . . Thisseina 6. Ascomatal surface not hairy . . . . . . . . . . . . . . . . . . . . 7
10. Apical ring present . . . . . . . . . . . . . . . . . . . . . . . . . 11 7. Ascospores with longitudinal germ slit. . Coniolariella
10. Apical ring lacking. . . . . . . . . . . . . . . . . . . . . . . . . 15 7. Ascospores with helical germ slit . . . . Helicogermslita
11. Stromata erumpent, in upright position. . . . . . . . . . 12 8. Stromata erect, usually several times higher than broad 9
11. Stromata effuse-pulvinate, spreading parallel to the host 8. Stromata effuse, pulvinate, conical, always broader than
surface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13 high . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
12. Stromata hemisphaerical with hard, woody interior, 9. Apical ring present. . . . . . . . . . . . . . . . . . . . . . . . . . 10
with concentric zones . . . . . . . . . . . . . . . . . Daldinia 9. Apical ring lacking. . . . . . . . . . . . . . . . . . . . . . . . . . 11
 Fungal Diversity

10. Stromata reaching 3.5 cm, apex begins to expand into a 5. Mucilaginous sheath present and surrounding entire
disc-shaped head, associated with dung . . . . Poronia ascospores . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
10. Stromata with clavate-globose heads, associated with 5. Mucilaginous sheath lacking or restricted to ends, be-
various substrates. . . . . . . . . . . . . . . . . . . . . . Xylaria coming appendages . . . . . . . . . . . . . . . . . . . . . . . . . . 8
11. Ascospores reniform, with prominent germ 6. Ascospores usually with a hyaline dwarf
slit . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Leprieuria cell. . . . . . . . . . . . . . . . . . . . . . . . . . . Anthostomella
11. Ascospores reniform, with prominent terminal germ 6. Ascospores lacking dwarf cells . . . . . . . . . . . . . . . . . 7
pores . . . . . . . . . . . . . . . . . . . . . . . . . Poroleprieuria 7. Stromata multi-peritheciate. . . . . . . . .Kretzschmariella
12. Amyloid apical ring present . . . . . . . . . . . . . . . . . . 13 7. Stromata uni-peritheciate . . . . . . . . . Brunneiperidium
12. Inamyloid apical ring present . . . . . . . . . Xylotumulus 8. Ascospores with polar, pad-like
13. Ascospores with germ slit . . . . . . . . . . . . . . . . . . . 15 appendages. . . . . . . . . . . . . . . . . . . . . Gigantospora
13. Ascospores lacking germ slit . . . . . . . . . . . . . . . . . 14 8. Ascospores with a hyaline, cellular
14. Ascospores scolecosporous with 6–7 appendage . . . . . . . . . . . . . . . . . . . . . . . Occultitheca
septa . . . . . . . . . . . . . . . . . . . . . . . .Ophiorosellinia 9. Ascospores with polar appendages . . . . . . . . . . . . . . 10
14. Ascospores inequilaterally fusoid with one median 9. Ascospores lacking polar appendages. . . . . . . . . . . . 12
slightly constricted septum . . . . . . . . . ..Collodiscula 10. Ascospores with central pallid band . . . Fasciatispora
15. Stromata multi-peritheciate. . . . . . . . . . . . . . . . . . . 16 10. Ascospores lacking central pallid band. . . . . . . . . . 11
15. Stromata uniperitheciate . . . . . . . . . . . . . . . . . . . . . 19 11. Ascospores brown . . . . . . . . . . . . . . . . . . Appendixia
16. Stromata effused-pulvinate, attached to the substrate by 11. Ascospores hyaline to yellowish. . . . . . . . .Sabalicola
entire base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17 12. Ascospores with conspicuous, spirally arranged
16. Stromata pulvinate, attached to the substrate by simple warts. . . . . . . . . . . . . . . . . . . . . . . . .Spirodecospora
or branched stipes by narrow connectives. . . . . . . . 18 1 2 . A s c o s p o r e s w i t h s t r i a t e
17. Stromata with coarsely papillate ostioles, containing ornamentations . . . . . . . . . . . . .Striatodecospora
few perithecia . . . . . . . . . . . . . . . . . . . . . . Entoleuca 13. Ascospores with hyaline dwarf cells. . . . . . . . . . . . 14
17. Stromata with finely papillate ostioles, containing sev- 13. Ascospores lacking hyaline dwarf cells . . . . . . . . . 15
eral perithecia. . . . . . . . . . . . . . . . . . . . . . . .Nemania 14. Ascospores with straight germ slit . . . . . Myconeesia
18. Stromata associated with dung . . . . . . . Podosordaria 14. Ascospores lacking germ slit . . . . . Pyriformiascoma
18. Stromata associated with wood . . . . . Kretzschmaria 15. Asci clavate, with ascospores always aggregated at the
19. Mature stromata completely surrounded or covered by apex . . . . . . . . . . . . . . . . . . . . . . . . . . Lunatiannulus
epidermal host tissue . . . . . . . . . . . ..Amphirosellinia 15. Asci broadly cylindrical, with ascospores uniseriately
19. Mature stromata loosely covered by epidermal tissue and arranged. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
mostly epidermal tissue residues at base of stromata. 20 16. Ascospores lunate, black, with straight germ
20. Stromata associated with ochraceous, short peg-like slit . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nipicola
protuberances . . . . . . . . . . . . . . . . . Stilbohypoxylon 16. Ascospores elliptic-fusiform, at first hyaline, becoming
20. Stromata lacking protuberances . . . . . . . . . . . . . . . 21 yellowish brown, with polar germ pores Pandanicola
21. Stromata usually surrounded by a subiculum, superfi- 17. Apical ring present . . . . . . . . . . . . . . . . . . . . . . . . . 18
cial, never splitting, asci with long stipes . . Rosellinia 17. Apical ring lacking. . . . . . . . . . . . . . . . . . . . . . . . . 32
21. Stromata not associated with subiculum, often splitting 18. Apical ring amyloid . . . . . . . . . . . . . . . . . . . . . . . . 19
into an outer stromatic layer, developing beneath host 18. Apical ring not amyloid 27
epidermis, asci with short stipes . . . . . . . . Astrocystis 1 9. S tr omata ere ct, sev eral times h ig her th an
broad . . . . . . . . . . . . . . . . . . . . . . . . Squamotubera
19. Stromata pulvinate, hemisphaerical, globose, broader
Section C: Genera with libertella-like or unnamed than high . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
asexual morphs 20. Stromata hemisphaerical, soft interior, interior and ex-
terior mostly light coloured . . . . . . . . . . . . . . . . . . 21
1. Stromata greatly reduced and immersed in substratum 2 20. Stromata pulvinate, effuse, woody to hard, bright
1. Stromata erumpent or superficial . . . . . . . . . . . . . . . 18 coloured to black. . . . . . . . . . . . . . . . . . . . . . . . . . 22
2. Apical ring amyloid . . . . . . . . . . . . . . . . . . . . . . . . . . 3 21. Perithecia with long ostiolar necks; Ostioles brown,
2. Apical ring not amyloid . . . . . . . . . . . . . . . . . . . . . . 14 punctuate scattered at the stromatal
3. Ascospores with germ slits or germ pores . . . . . . . . . 4 surface. . . . . . . . . . . . . . . . . . . . . . . . . Engleromyces
3. Ascospores lacking germ slits or germ pores . . . . . . . 9 21. Perithecia with gelatinous zones beneath perithecial
4. Germ slit straight . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 layer, covering with a whitish membrane, without
4. Germ slit spiral . . . . . . . . . . . . . . . . . . Leptomassaria long necks or visible ostioles . . . . . .Sarcoxylon
Fungal Diversity

22. Apical ring higher than wide . . . . . . . . . . . . . . . . . 23 coincident with Gnomoniaceae G. Winter, which is consid-
22. Apical ring wider than high . . . . . . . . . . . . . . . . . . 24 ered to be a well-known family name in plant pathology, while
23. Ascomata with papillate ostioles, protruding through Obryzaceae was only recently reported in lichenological stud-
star-like cracks on host surface, hemisphaerical, with ies (McNeill et al. 2006). Hawksworth and Eriksson (1988)
wart-like ornamentations . . . . . . . . . . . . . . . . Guestia proposed that the name Obryzaceae should be rejected to con-
23. Ascospores with a central papillate ostiole, often cov- serve Gnomoniaceae and the proposal was accepted (McNeill
ered with fine blue powder. . . . . . . . . . . Cyanopulvis et al. 2006). Obryzaceae was listed in Sordariomycetes, fam-
24. Ascospores unicellular . . . . . . . . . . . . . . . . . . . . . . 25 ilies incertae sedis by Lumbsch and Huhndorf (2010) and this
24. Ascospores 2-celled . . . . . . . . . . . . . . Steganopycnis was followed in Maharachchikumbura et al. (2015) and Index
25. Ascospores smooth-walled . . . . . . . . . . . . . . . . . . . 26 Fungorum (2016), indicative of the problems in picking up
25. Ascospores roughened by small nodular-like or irregu- obscure literature.
lar warts . . . . . . . . . . . . . . . . . . . . . . . . . Areolospora It is obvious that Obryzaceae is distinct from
26. Ascospores with mucilaginous sheath . . . . Jumillera Gnomoniaceae by their mode of life and morphological char-
26. Ascospores lacking mucilaginous sheath . . Whalleya acters. Species of Gnomoniaceae are found in herbaceous
27. Stromata dark coloured, lacking KOH extractable plant material, in leaves, twigs or stems, and sometimes in
pigments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28 bark or wood (Sogonov et al. 2008), have solitary, immersed
27. Stromata green, with KOH extractable astromatic ascomata, or ascomata aggregated with a rudimen-
pigments . . . . . . . . . . . . . . . . . . . . . . Chlorostroma tary stroma and small, mostly less than 25 μm long, mainly
28. Stromata thread-like . . . . . . . . . . . . . . . . . . . Wawelia aseptate to 1-septate, rarely multi-septate ascospores
28. Stromata effuse . . . . . . . . . . . . . . . . . . . . . . . . . . . 29 (Sogonov et al. 2008).
29. Ascospores lacking germ slit . . . . . . . . . . . . . . . . . 30 Type: Obryzum Wallr., Naturgesch. Flecht. 1: 253 (1825);
29. Ascospores with germ slit . . . . . . . . . . Paucithecium Fig. 156
30. Ascospores elongated fusiform, with longitudinal, Lichenicolous on rocks or bark, rarely on mosses or para-
parallel to convergent ridges on the sitic on cyanobacterial lichens of the genus Leptogium. Sexual
wall. . . . . . . . . . . . . . . . . . . . . .Stromatoneurospora morph: Ascomata astromatic, perithecial, immersed, pyri-
30. Ascospores not fusiform, with smooth walls. . . . . . 31 form, ostiolate. Ostiole periphysate. Peridium colourless, but
31. Ascospores unicellular . . . . . . . . . . . . . . . Barrmaelia sometimes the ostiolar region lightly brown. Asci 4–8-spored,
31. Ascospores bi-celled . . . . . . . . . . Paramphisphaeria unitunicate, clavate, narrowly ellipsoid to ovoid, thin-walled,
32. Ascospores ellipsoidal . . . . . . . . . . . . . . . . . . . . . . 33 short-pedicellate, without an apical ring, evanescent.
32. Ascospores cylindrical . . . . . . . . . . . . Chaenocarpus Ascospores aseptate, fusiform to limoniform or ellipsoid, hy-
33. Ascomata embedded in host tissue with long cylindri- aline, smooth-walled, lacking appendages, without a gelati-
cal, slender necks erumpent from substrate nous sheath. Asexual morph: Undetermined.
surface . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cannonia Type species: Obryzum corniculatum (Hoffm.) Wallr.,
33. Ascomata immersed in more or less solid, but gelati- Naturgesch. Flecht. 1: 253 (1825)
nous stromata, with short necks . . . . . . . . . . Xylocrea Notes: This genus is poorly studied and lacks molecular
data in GenBank. Six epithets are currently listed under
Obryzum (Index Fungorum 2016).
Excluded, doubtful and rejected families Pleurotremataceae Walt. Watson, New Phytol. 28: 113
(1929)
Obryzaceae Körb. [as ‘Obryzeae’], Syst. lich. germ. = Dyfrolomycetaceae K.D. Hyde et al., in Pang et al.,
(Breslau): 427 (1855) Cryptog. Mycol. 34(3): 227 (2013)
Lichenicolous with green algae on rocks or bark, rarely on Facesoffungi number: FoF 01911
mosses or parasitic on cyanobacterial lichens, in temperate Saprobic on wood in terrestrial and aquatic habitat. Sexual
regions. Sexual morph: Ascomata astromatic, perithecial, im- morph: Ascomata perithecial, gregarious or solitary, im-
mersed, pyriform, ostiolate, with a periphysate ostiolar region. mersed or erumpent through bark of host, black, ovoid to
Asci 4–8-spored, unitunicate, clavate, narrowly ellipsoid to subglobose, carbonaceous to membranaceous, smooth, with
ovoid, thin-walled, short- pedicellate, apical ring not apparent, ostioles. Peridium thick (60–120 μm), comprising of dark
evanescence. Ascospores aseptate, hyaline, smooth-walled, cells of textura angularis. Hamathecium composed of numer-
lacking appendages, without a gelatinous sheath. Asexual ous, hyaline, septate, branched paraphyses. Asci 8-spored,
morph: Undetermined. unitunicate, clavate to cylindrical, short pedicellate, apical
Type: Obryzum Wallr. ring present, non-amyloid. Ascospores uni-seriate, hyaline,
Notes: This is a poorly studied, monotypic family, for ellipsoidal to cylindrical, septate, smooth-walled, mostly with
which no molecular data are available. Obryzaceae is guttules. Asexual morph: Undetermined.
 Fungal Diversity

Fig. 156 Obryzum corniculatum (Material examined: PC 002061) a through ascoma e Transverse section through ostiole f Peridium g, h
Herbarium specimen with the label b, c Appearance of ascomata on Mature asci i–k Ascospores. Scale bars: b = 500 μm, c = 1 mm,
host substrate (see arrows, c in 5 % KOH). d Transverse section d = 100 μm, e–h = 20 μm, i–k = 10 μm

Pleurotrema Müll. Arg. was not lichenized and with unitunicate asci (Barr 1994).
Notes: The family Pleurotremataceae was introduced by However, Barr (1994) transferred Pleurotremataceae to
Watson (1929) based on Pleurotrema polysemum (Nyl.) Xylariales based on its cylindrical asci, with a non amyloid
Mull. Arg., the sole species. It is characterized by black apical ring and hyaline, distoseptate ascospores. Five genera,
ascomata and cylindrical asci, with septate, hyaline asco- Phomatospora, Melomastia, Pleurotrema, Saccardoella,
spores. This family was introduced as lichenized with Daruvedia, were included in Pleurotremaceae on account of
bitunicate asci (Eriksson 1981). Eriksson and Hawksworth their non-fissitunicate asci (Barr 1994). Hawksworth et al.
(1993) suspected Pleurotremataceae to be synonym of (1995) disagreed with this justification and retained
Pyrenulaceae. Pleurotrema was synonymized under Pleurotrema in the order Pyrenulales. Saccardoella and
Lithothelium as a member of Pyrenulaceae by Aptroot Melomastia were placed in Ascomycota genera incertae sedis
(1991). Later, Harris re-examined the isotype of (Kirk et al. 2001; Lumbsch and Huhndorf 2010), but there are
P. polysemum, determined that it is closely related to no phylogenic datasets. Phomatospora was found in freshwa-
Melomastia and Saccardoella (Barr 1994), and this species ter and marine habitats and positioned in Ascomycota genera
Fungal Diversity
 Fungal Diversity

ƒFig. 157 Pleurotrema polysemum (Material examined: COLOMBIA, pathogens of trees in South Africa. Australas Plant Pathol 35:521–
Nov. Granada, on dead wood, Lindig A, S L1236, isotype). a Material 548
label b Material c Ascomata d Ascoma cross section e Peridium f Ahmed SA, Khan Z, Wang XW, Moussa TA, Al-Zahrani HS, Almaghrabi
Paraphyses g–j Asci (j. ascus with ejected spores) k Apical part of ascus OA, de Hoog GS (2015) Chaetomium-like fungi causing opportu-
l–o Ascospores. Notes: Figs f-o soaked in 3 % KOH. Scale bars: nistic infections in humans: a possible role for extremotolerance.
c = 500 μm, d = 100 μm, g–j = 20 μm, e–f, k = 10 μm, l–o = 3 μm Fungal Divers 76:1–16
Ainsworth GC (ed) (1971) Ainsworth & Bisby’s dictionary of the fungi,
6th edn. CABI, Kew
incertae sedis based on phylogenic analysis (Vijaykrishna Ainsworth GC, Bisby GR (1954) A dictionary of the fungi, 4th edn. The
Commonwealth Mycological Institute, Kew
et al. 2006). Pleurotremataceae was accepted as monotypic
Alexopoulos CJ (1962) Introductory mycology, 2nd edn. Wiley, New
via Pleurotrema and placed under Chaetosphaeriales without York, 613 p
giving a reasons (Maharachchikumbura et al. 2015). Alvarez E, Llano GA, Loke JB, Chacon MI (2012) Characterization of
Pleurotrema Müll. Arg., Bot. Jb. 6: 388 (1885); Fig. 157 Thielaviopsis paradoxa isolates from Oil Palms in Colombia,
Type species: Pleurotrema polysemum (Nyl.) Müll. Arg., Ecuador and Brazil. J Phytopathol 160:690–700
Ames LM (1961) A monograph of the Chaetomiaceae. US Army Res
Bot. Jb. 6: 389 (1885) Dev Ser, no. 2. Washington, D.C
We re-examined the isotype from S of Pleurotrema Anagnostakis SL (1987) Chestnut blight: the classical problem of an
polysemum. It has what appears to be unitunicate asci, and introduced pathogen. Mycologia 79:23–37
septate ascospores and is similar to species of Saccardoella Anderson F, Bianchinotti VA (1996) New species of Bartalinia Tassi
(Deuteromycotina, Coelomycetes). Mycotaxon 60:191–200
and Dyfrolomyces (Hyde et al. 2013). Thus this is an earlier
Andersson K, Eriksson OE, Landvik S (1995) Boliniaceae transferred to
family name for Dyfrolomycetaceae (Dothideomycetes). Sordariales (Ascomycota). Syst Ascomycetum 14:1–16
Fresh collections are needed to substantiate this. Andrianova TV, Minter DW (2007) New species of Bartalinia and
Septoriella from the Altai Mountains (Russia). Mycotaxon 101:
Acknowledgments This work was supported by the International 297–313
Research Group Program (IRG-14-27), Deanship of Scientific Aptroot A (1991) A monograph of the Pyrenulaceae (excluding
Research, King Saud University, Saudi Arabia. The curators of the her- Anthracothecium and Pyrenula) and the Requienellaceae, with notes
barium ANM, AR, B, BBH, BCC, BISH, BP, BPI, BR, BRIP, CBS, on the Pleomassariaceae, the Trypetheliaceae, and Mycomicrothelia
DAOM, FH, G, GZU, GZU, H, HAL, HHUF, HKU (M), IFRD, ILLS, (lichenized and non-lichenized ascomycetes). Bibl Lichenologica
IMI, K, L, LPS, MFU, NY, P, PC, PREM, S, TRTC, UBC, UC, UCR, 44:1–178
UPS, URM, URM and W are thanked for loaning herbarium specimens Ariyawansa HA, Hyde KD, Jayasiri SC, Buyck B, Chethana KWT, Dai
and for being very helpful in locating specimens. Without their help this DQ, Dai YC, Daranagama DA, Jayawardena RS, Lücking R,
work would not have been possible. The abbreviations of herbarium are Ghobad-Nejhad M, Niskanen T, Thambugala KM, Voigt K, Zhao
those listed in Index Herbariorum (2015). We also thank the curator of RL, Li GJ, Doilom M, Boonmee S, Yang ZL, Cai Q, Cui YY,
MFLU for arranging the loan of specimens from various herbaria. The Bahkali AH, Chen J, Cui BK, Chen YY, Monika CD,
authors would like to thank the featured microbial resources and diversity Dissanayake AJ, Ekanayaka AH, Hashimoto A, Hongsanan S,
investigation in Southwest Karst area (2014FY120100). This research Jones EBG, Larsson E, Li WJ, Li QR, Liu JK, Luo ZL,
was also financially supported by The Biodiversity and Training Maharachchikumbura SSN, Mapook A, McKenzie EHC,
Program (BRT R_251006, BRT R_351004, BRT_R352015) and Norphanphoun C, Konta S, Pang KL, Perera RH, Phookamsak R,
National Center for Genetic Engineering and Biotechnology (BIOTEC) Phukhamsakda C, Pinruan U, Randrianjohany E, Singtripop C,
Khlong Luang, Pathum Thani 12120, Thailand. Shi-Ke Huang’ s work Tanaka K, Tian CM, Tibpromma S, Abdel-Wahab MA,
was funded by the grants from the National Natural Science Foundation Wanasinghe DN, Wijayawardene NN, Zhang JF, Zhang H, Abdel-
of China (NSFC, No. 30870009 & 31460011), the International Aziz FA, Wedin M, Westberg M, Ammirati JF, Bulgakov TS, Lima
Collaboration Plan of Guizhou Province (No. G [2012] 7006). Ka-Lai DX, Callaghan TM, Callac P, Chang CH, Coca LF, Dal-Forno M,
Pang thanks the Ministry of Science and Technology, Taiwan (grant no. Dollhofer V, Fliegerová K, Greiner K, Griffith GW, Ho HM,
101–2621-B-019–001-MY3) for financial support. We are also grateful Hofstetter V, Jeewon R, Kang JC, Wen TC, Kirk PM, Kytövuori I,
to Bita Asgari and Rasoul Zare, for images of Coniocessia spp., and Lawrey JD, Xing J, Li H, Liu ZY, Liu XZ, Liimatainen K, Lumbsch
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Asanka Bandara, Kasun Thambugala, Mingkwan Doilom and Wen-Jing Azevedo Santiago ALCM, Sommai S, Song Y, de Souza CAF, de
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