Mycosphere 11(1): 305–1059 (2020) www.mycosphere.org
ISSN 2077 7019
Article
Doi 10.5943/mycosphere/11/1/7
Refined families of Sordariomycetes
Hyde KD1, 2, 3, 4, 5, 6, 7, Norphanphoun C2, 3, 6, 15, 19, Maharachchikumbura SSN26,
Bhat DJ21, Jones EBG12, 32, Bundhun D2, 13, Chen YJ2, 6, Bao DF2, 11, 13, Boonmee
S2, Calabon MS2, 6, Chaiwan N2, 6, Chethana KWT2, 23, Dai DQ9, Dayarathne
MC2, 3, 5, 6, Devadatha B2, 18, Dissanayake AJ26, Dissanayake LS29, Doilom M5,
Dong W2, 3, 13, 31, Fan XL30, Goonasekara ID2, 3, 5, 7, Hongsanan S27, 28, Huang SK2,
3, 6, 29
, Jayawardena RS2, Jeewon R14, Karunarathna A2, 5, 13, 15, Konta S2, 3, 5, 6,
Kumar V2, 13, Lin CG2, 3, 6, Liu JK26, Liu NG2, 17, Luangsa-ard J25, Lumyong S4, 8,
10
, Luo ZL11, Marasinghe DS2, 6, 15, McKenzie EHC24, Niego AGT2, 22, Niranjan
M18, Perera RH2, 3, 6, 19, Phukhamsakda C2, Rathnayaka AR2, 6, 16, Samarakoon
MC2, 4, 26, Samarakoon SMBC2, 6, Sarma VV18, Senanayake IC2, 28, Shang QJ2, 6,
19
, Stadler M20, Tibpromma S5, 7, Wanasinghe DN5, Wei DP2, 3, 5, 13,
Wijayawardene NN9, Xiao YP2, 3, 6, 19, Yang J2, 3, 6, 19, Zeng XY2, 6, 19, Zhang SN2, 13,
Xiang MM1
1
Institute of Plant Health, Zhongkai University of Agriculture and Engineering, Haizhu District, Guangzhou 510225,
P.R. China
2
Center of Excellence in Fungal Research, Mae Fah Luang University, Chiang Rai 57100, Thailand
3
Mushroom Research Foundation, 128 M.3 Ban Pa Deng T. Pa Pae, A. Mae Taeng, Chiang Mai 50150, Thailand
4
Department of Biology, Faculty of Science, Chiang Mai University, Chiang Mai 50200, Thailand
5
Key Laboratory for Plant Diversity and Biogeography of East Asia, Kunming Institute of Botany, Chinese Academy of
Science, Kunming 650201, Yunnan, P.R. China
6
School of Science, Mae Fah Luang University, Chiang Rai 57100, Thailand
7
World Agroforestry Centre, East and Central Asia, Kunming 650201, Yunnan, P.R. China
8
Academy of Science, The Royal Society of Thailand, Bangkok 10300, Thailand
9
Center for Yunnan Plateau Biological Resources Protection and Utilization, College of Biological Resource and Food
Engineering, Qujing Normal University, Qujing, Yunnan 655011, P.R. China
10
Center of Excellence in Microbial Diversity and Sustainable Utilization, Faculty of Science, Chiang Mai University,
Chiang Mai 50200, Thailand
11
College of Agriculture and Biological Sciences, Dali University, Dali 671003, Yunnan Province, P.R. China
12
Department of Botany and Microbiology, College of Science, King Saud University, P.O Box 2455, Riyadh 11451,
Kingdom of Saudi Arabia
13
Department of Entomology and Plant Pathology, Faculty of Agriculture, Chiang Mai University, Chiang Mai 50200,
Thailand
14
Department of Health Sciences, Faculty of Science, University of Mauritius, Reduit, Mauritius
15
Department of Plant Medicine, National Chiayi University, 300 Syuefu Road, Chiayi City 60004, Taiwan
16
Engineering Research Center of Southwest Bio-Pharmaceutical Resources, Ministry of Education, Guizhou
University, Guiyang, Guizhou 550025, P.R. China
17
Faculty of Agriculture, Natural Resources and Environment, Naresuan University, Phitsanulok 65000, Thailand
18
Fungal Biotechnology Lab, Department of Biotechnology, School of Life Sciences, Pondicherry University, Kalapet,
Pondicherry-605014, India
19
Guizhou Key Laboratory of Agricultural Biotechnology, Guizhou Academy of Agricultural Sciences, Guiyang,
Guizhou 550006, P.R. China
20
Helmholtz Centre for Infection Research, Department Microbial Drugs, Inhoffenstrasse 7, 38124 Braunschweig,
Germany
21
House No. 128/1-J, Azad Housing Society, Curca, Goa Velha 403108, India
Submitted 15 November 2019, Accepted 13 February 2020, Published 28 February 2020
Corresponding Author: Xiang MM – e-mail – xiang mm_xiang@163.com
305
22
Iloilo Science and Technology University, La Paz, Iloilo, Philippines
Institute of Plant and Environment Protection, Beijing Academy of Agriculture and Forestry Sciences, Beijing
100097, P.R. China
24
Landcare Research Manaaki Whenua, Private Bag 92170, Auckland, New Zealand
25
National Center for Genetic Engineering and Biotechnology (BIOTEC), 113 Thailand Science Park, Phahonyothin
Road, Khlong Nueng, Khlong Luang, Pathum Thani 12120, Thailand
26
School of Life Science and Technology, University of Electronic Science and Technology of China, Chengdu 611731,
P.R. China
27
Shenzhen Key Laboratory of Laser Engineering, College of Optoelectronic Engineering, Shenzhen University,
Shenzhen, P.R. China
28
Shenzhen Key Laboratory of Microbial Genetic Engineering, College of Life Sciences and Oceanography, Shenzhen
University, Shenzhen, P.R. China
29
The Engineering Research Center of Southwest Bio–Pharmaceutical Resource Ministry of Education, Guizhou
University, Guiyang 550025, Guizhou Province, P.R. China
30
The Key Laboratory for Silviculture and Conservation of Ministry of Education, Beijing Forestry University, Beijing
100083, P.R. China
31
Yunnan Institute of Food Safety, Kunming University of Science and Technology, Kunming 650500, P.R. China
32
33B St Edwards Road, Southsea, Hants., PO53DH, UK
Faculty of Agriculture, Natural Resources and Environment, Naresuan University, Phitsanulok 65000, Thailand
23
Hyde KD, Norphanphoun C, Maharachchikumbura SSN, Bhat DJ, Jones EBG, Bundhun D, Chen
YJ, Bao DF, Boonmee S, Calabon MS, Chaiwan N, Chethana KWT, Dai DQ, Dayarathne MC,
Devadatha B, Dissanayake AJ, Dissanayake LS, Doilom M, Dong W, Fan XL, Goonasekara ID,
Hongsanan S, Huang SK, Jayawardena RS, Jeewon R, Karunarathna A, Konta S, Kumar V, Lin
CG, Liu JK, Liu NG, Luangsa-ard J, Lumyong S, Luo ZL, Marasinghe DS, McKenzie EHC, Niego
AGT, Niranjan M, Perera RH, Phukhamsakda C, Rathnayaka AR, Samarakoon MC, Samarakoon
SMBC, Sarma VV, Senanayake IC, Shang QJ, Stadler M, Tibpromma S, Wanasinghe DN, Wei DP,
Wijayawardene NN, Xiao YP, Yang J, Zeng XY, Zhang SN, Xiang MM 2020 – Refined families
of Sordariomycetes. Mycosphere 11(1), 305–1059, Doi 10.5943/mycosphere/11/1/7
Abstract
This is a continuation of the papers “Towards a classification of Sordariomycetes” (2015) and
“Families of Sordariomycetes” (2016) in which we compile a treatment of the class
Sordariomycetes. The present treatment is needed as our knowledge has rapidly increased, from 32
orders, 105 families and 1331 genera in 2016, to 45 orders, 167 families and 1499 genera (with 308
genera incertae sedis) at the time of publication. In this treatment we provide notes on each order,
families and short notes on each genus. We provide up-to-date DNA based phylogenies for 45
orders and 163 families. Three new genera and 16 new species are introduced with illustrations and
descriptions, while 23 new records and three new species combinations are provided. We also list
308 taxa in Sordariomycetes genera incertae sedis. For each family we provide general descriptions
and illustrate the type genus or another genus, the latter where the placement has generally been
confirmed with molecular data. Both the sexual and asexual morphs representative of a family are
illustrated where available. Notes on ecological and economic considerations are also given.
Key words – 19 new taxa – Amphisphaeriales – Amplistromatales – Annulatascales –
Atractosporales – Boliniales – Calosphaeriales – Catabotryales – Cephalothecales –
Chaetosphaeriales – Coniochaetales – Conioscyphales – Coronophorales – Delonicicolales –
Diaporthales – Distoseptisporales – Falcocladiales – Fuscosporellales – Glomerellales –
Hypocreales – Jobellisiales – Koralionastetales – Lulworthiales – Magnaporthales – Meliolales –
Microascales,
Myrmecridiales,
new
records,
Ophiostomatales,
Pararamichloridiales,
Parasympodiellales – Phomatosporales – Phyllachorales – Pisorisporiales – Pleurotheciales –
Pseudodactylariales – Savoryellales – Sordariales – Spathulosporales – Sporidesmiales –
Tirisporellales – Togniniales – Torpedosporales – Tracyllalales – Vermiculariopsiellales –
Xenospadicoidales – Xylariales
306
Contents
Class Sordariomycetes
Subclasses
1. Diaporthomycetidae Senan., Maharachch. & K.D. Hyde, Fungal Divers. 72: 208 (2015) –
contributed by Senanayake IC, Maharachchikumbura SSN
2. Hypocreomycetidae O.E. Erikss. & Winka, Myconet 1: 6 (1997) – contributed by Perera
RH, Maharachchikumbura SSN
3. Lulworthiomycetidae Dayar., E.B.G. Jones & K.D. Hyde, Fungal Divers. 72: 208 (2015)
– contributed by Dayarathne MC, Jones EBG
4. Pisorisporiomycetidae Bundhun, Maharachch. & K.D. Hyde, subclass nov.– contributed
by Bundhun D, Maharachchikumbura SSN, Hyde KD
5. Savoryellomycetidae Hongsanan, K.D. Hyde & Maharachch., Fungal Divers. 84: 35
(2017) – contributed by Dayarathne MC, Jones EBG
6. Sordariomycetidae O.E. Erikss. & Winka, Myconet 1: 10 (1997) – contributed by Huang
SK, Jeewon R
7. Xylariomycetidae O.E. Erikss. & Winka, Myconet 1: 12 (1997) – contributed by
Samarakoon MC, Stadler M
Orders
1. Amphisphaeriales Hawksw. & O.E. Erikss., Syst. Ascom. 5: 177 (1986) – contributed by
Samarakoon MC, Maharachchikumbura SSN
2. Amplistromatales M.J. D'souza, Maharachch. & K.D. Hyde, Fungal Divers. 72: 212
(2015) – contributed by Samarakoon MC, Liu JK
3. Annulatascales M.J. D'souza, Maharachch. & K.D. Hyde, Fungal Divers. 72: 212 (2015)
– contributed by Dong W, Doilom M
4. Atractosporales H. Zhang, K.D. Hyde & Maharachch., Fungal Divers. 85: 88 (2017) –
contributed by Dong W, Doilom M
5. Boliniales P.F. Cannon, Ainsworth & Bisby's Dictionary of the Fungi, Edn 9
(Wallingford): x (2001) – contributed by Huang SK, Jeewon R, Maharachchikumbura
SSN
6. Calosphaeriales M.E. Barr, Mycologia 75: 11 (1983) – contributed by Huang SK, Jeewon
R
7. Catabotryales K.D. Hyde & Senan., ord. nov. – contributed by Hyde KD, Senanayake IC
8. Cephalothecales Maharach. & K.D. Hyde, ord. nov. – contributed by
Maharachchikumbura SSN, Hyde KD
9. Chaetosphaeriales Huhndorf, A.N. Mill. & F.A. Fernández, Mycologia 96: 378 (2004) –
contributed by Perera RH, Lin CG, Liu JK
10. Coniochaetales Huhndorf, A.N. Mill. & F.A. Fernández, Mycologia 96: 378 (2004) –
contributed by Goonasekara ID, Samarakoon MC, Jayawardena RS
11. Conioscyphales Réblová & Seifert, Persoonia 37: 63 (2015) – contributed by Bundhun D,
Liu NG, Dayarathne MC, Jones EBG
12. Coronophorales Nannf., Nova Acta R. Soc. Scient. upsal., Ser. 4 8(no. 2): 54 (1932) –
contributed by Huang SK, Hyde KD
13. Delonicicolales R.H. Perera, Maharachch. & K.D. Hyde, Cryptog. Mycol. 38: 329 (2017)
– contributed by Samarakoon MC, Perera RH
14. Diaporthales Nannf., Nova Acta R. Soc. Scient. upsal., Ser. 4 8(no. 2): 53 (1932) –
contributed by Senanayake IC, Maharachchikumbura SSN
15. Distoseptisporales Z.L. Luo, H.Y. Su & K.D. Hyde, Fungal Divers. 99: 482 (2019) –
contributed by Luo ZL
307
16. Falcocladiales R.H. Perera, Maharachch. & K.D. Hyde, Fungal Divers. 72: 218 (2015) –
contributed by Perera RH
17. Fuscosporellales J. Yang, J. Bhat & K.D. Hyde, Cryptog. Mycol. 37: 457 (2016) –
contributed by Bundhun D, Dayarathne MC, Jones EBG
18. Glomerellales Chadef. ex Réblová, W. Gams & Seifert, Stud. Mycol. 68: 170 (2011) –
contributed by Jayawardena RS, Maharachchikumbura SSN
19. Hypocreales Lindau, Nat. Pflanzenfam., Teil. I (Leipzig) 1: 343 (1897) – contributed by
Perera RH, Xiao YP, Liu JK
20. Jobellisiales M.J. D'souza & K.D. Hyde, Fungal Divers. 72: 219 (2015) – contributed by
Huang SK, Hyde KD, Jeewon R, Maharachchikumbura SSN
21. Koralionastetales Kohlm., Volkm.-Kohlm., J. Campb. & Inderb., Mycol. Res. 113: 377
(2009) – contributed by Bundhun D, Dayarathne MC, Jones EBG
22. Lulworthiales Kohlm., Spatafora & Volkm.-Kohlm., Mycologia 92: 456 (2000) –
contributed by Bundhun D, Dayarathne MC, Jones EBG
23. Magnaporthales Thongk., Vijaykr. & K.D. Hyde, Fungal Divers. 34: 168 (2009) –
contributed by Norphanphoun C, Maharachchikumbura SSN
24. Meliolales Gäum. ex D. Hawksw. & O.E. Erikss., Syst. Ascom. 5: 180 (1986) –
contributed by Hongsanan S
25. Microascales Luttr. ex Benny & R.K. Benj., Mycotaxon 12: 40 (1980) – contributed by
Kumar V, Dayarathne MC, Jones EBG
26. Myrmecridiales Crous, Persoonia 34: 219 (2015) – contributed by Wei DP, Doilom M
27. Ophiostomatales Benny & Kimbr., Mycotaxon 12: 48 (1980) – contributed by
Norphanphoun C, Maharachchikumbura SSN, Doilom M
28. Pararamichloridiales Crous, Persoonia 39: 357 (2017) – contributed by Lin CG,
Maharachchikumbura SSN
29. Parasympodiellales Hern.-Restr., Gené, R.F. Castañeda & Crous, Stud. Mycol. 86: 87
(2017) – contributed by Bundhun D, Dayarathne MC, Jones EBG
30. Phomatosporales Senan., Maharachch. & K.D. Hyde, Mycosphere 7: 631 (2016) –
contributed by Bundhun D, Dayarathne MC, Jones EBG
31. Phyllachorales M.E. Barr, Mycologia 75: 11 (1983) – contributed by Dayarathne MC,
Maharachchikumbura SSN
32. Pisorisporiales Réblová & J. Fourn., Persoonia 34: 43 (2014) – contributed by Bundhun
D, Dayarathne MC, Jones EBG
33. Pleurotheciales Réblová & Seifert, Persoonia 37: 63 (2015) – contributed by Bundhun D,
Dayarathne MC, Jones EBG, Yang J
34. Pseudodactylariales Crous, Persoonia 39: 421 (2017) – contributed by Lin CG, Liu JK
35. Savoryellales Boonyuen, Suetrong, Sivichai, K.L. Pang & E.B.G. Jones, Mycologia 103:
1368 (2011) – contributed by Bundhun D, Dayarathne MC, Jones EBG
36. Sordariales Chadef. ex D. Hawksw. & O.E. Erikss., Syst. Ascom. 5: 182 (1986) –
contributed by Huang SK, Jeewon R, Maharachchikumbura SSN
37. Spathulosporales Kohlm., Mycologia 65: 615 (1973) – contributed by Dayarathne MC,
Jones EBG
38. Sporidesmiales Crous, Persoonia 40: 377 (2018) – contributed by Luo ZL
39. Tirisporellales Suetrong, E.B.G. Jones & K.L. Pang, Fungal Divers. 73: 42 (2015) –
contributed by Dayarathne MC, Jones EBG
40. Togniniales Senan., Maharachch. & K.D. Hyde, Fungal Divers. 72: 220 (2015) –
contributed by Huang SK, Wei DP, Jeewon R
41. Torpedosporales E.B.G. Jones, Abdel-Wahab & K.L. Pang, Fungal Divers. 73: 42 (2015)
– contributed by Dayarathne MC, Jones EBG
42. Tracyllalales Crous, Persoonia 40: 365 (2018) – contributed by Calabon MS
43. Vermiculariopsiellales Hern.-Restr., J. Mena, Gené & Crous, Stud. Mycol. 86: 91 (2017)
– contributed by Lin CG, Liu JK
308
44. Xenospadicoidales Hern.-Restr., J. Mena & Gené, Stud. Mycol. 86: 91 (2017) –
contributed by Dong W, Doilom M
45. Xylariales Nannf., Nova Acta R. Soc. Scient. upsal., Ser. 4 8(no. 2): 66 (1932) –
contributed by Samarakoon MC, Stadler M
Families
1. Acrodictyaceae J.W. Xia & X.G. Zhang, Scientific Reports 7 (no. 7888): 2 (2017) –
contributed by Liu NG
2. Amphisphaeriaceae G. Winter, Rabenh. Krypt.-Fl., Edn 2 (Leipzig) 1.2: 259 (1885) –
contributed by Samarakoon MC, Senanayake IC, Liu JK
3. Amplistromataceae Huhndorf, A.N. Mill., Greif & Samuels, Mycologia 101: 905 (2009)
– contributed by Samarakoon MC, Liu JK
4. Annulatascaceae S.W. Wong, K.D. Hyde & E.B.G. Jones, Syst. Ascom. 16: 18 (1998) –
contributed by Dong W, Doilom M
5. Apiosporaceae K.D. Hyde, J. Fröhl., Joanne E. Taylor & M.E. Barr, Sydowia 50: 23
(1998) – contributed by Dai DQ, Hyde KD
6. Apiosporopsidaceae Senan., Maharachch. & K.D. Hyde, Stud. Mycol. 86: 234 (2017) –
contributed by Senanayake IC
7. Apoharknessiaceae Senan., Maharachch. & K.D. Hyde, Stud. Mycol. 86: 234 (2017) –
contributed by Senanayake IC
8. Armatellaceae Hosag., Sydowia 55: 165 (2003) – contributed by Zeng XY, Hongsanan S
9. Asterosporiaceae Senan., Maharachch. & K.D. Hyde, Stud. Mycol. 86: 236 (2017) –
contributed by Senanayake IC
10. Atractosporaceae H. Zhang, K.D. Hyde & Maharachch., Fungal Divers. 85: 88 (2017) –
contributed by Dong W, Doilom M
11. Auratiopycnidiellaceae Senan., Maharachch. & K.D. Hyde, Stud. Mycol. 86: 237 (2017)
– contributed by Senanayake IC
12. Australiascaceae Réblová & W. Gams, Stud. Mycol. 68: 171 (2011) – contributed by
Shang QJ, Jayawardena RS, Liu JK
13. Barbatosphaeriaceae H. Zhang, K.D. Hyde & Maharachch., Fungal Divers. 85: 94
(2017) – contributed by Dong W, Doilom M
14. Barrmaeliaceae Voglmayr & Jaklitsch, Mycol. Progr. 17: 162 (2018) – contributed by
Samarakoon MC, Liu JK
15. Batistiaceae Samuels & K.F. Rodrigues, Mycologia 81: 54 (1989) – contributed by Shang
QJ, Liu JK
16. Beltraniaceae Nann., Repert. mic. uomo: 498 (1934) – contributed by Lin CG, Liu JK
17. Bertiaceae Smyk, Ukr. bot. Zh. 38: 47 (1981) – contributed by Huang SK, Jeewon R
18. Bionectriaceae Samuels & Rossman, Stud. Mycol. 42: 15 (1999) – contributed by Perera
RH, Niranjan M, Sarma VV, Jeewon R
19. Boliniaceae Rick, Brotéria, sér. bot. 25: 65 (1931) – contributed by Huang SK, Jeewon R
20. Cainiaceae J.C. Krug, Sydowia 30:123 (1978) – contributed by Samarakoon MC, Liu JK,
Senanayake IC
21. Calcarisporiaceae Jing Z. Sun, X.Z. Liu & K.D. Hyde, Mycol. Prog. 16: 435 (2017) –
contributed by Calabon MS
22. Calosphaeriaceae Munk, Dansk bot. Ark. 17: 278 (1957) – contributed by Huang SK,
Jeewon R
23. Castanediellaceae Hern.-Restr., Guarro & Crous, Stud. Mycol. 86: 93 (2017) –
contributed by Samarakoon MC, Liu JK, Lin CG
24. Catabotryaceae Petr. ex M.E. Barr, Mycotaxon 39: 83 (1990) – contributed by Zhang SN,
Liu JK
25. Cephalothecaceae Höhn., Annls mycol. 15: 362 (1917) – contributed by Shang QJ,
Maharachchikumbura SSN, Jeewon R
309
26. Ceratocystidaceae Locq. ex Réblová, W. Gams & Seifert, Stud. Mycol. 68: 188 (2011) –
contributed by Doilom M
27. Ceratosphaeriaceae Z.L. Luo, H.Y. Su & K.D. Hyde, Fungal Divers. 99: 490 (2019) –
contributed by Luo ZL
28. Ceratostomataceae G. Winter, Rabenh. Krypt.-Fl., Edn 2 (Leipzig) 1.2: 247 (1885) –
contributed by Norphanphoun C, Hongsanan S
29. Chadefaudiellaceae Faurel & Schotter ex Benny & Kimbr., Mycotaxon 12: 46 (1980) –
contributed by Kumar V, Dayarathne MC
30. Chaetomiaceae G. Winter, Rabenh. Krypt.-Fl., Edn 2 (Leipzig) 1.2: 153 (1885) –
contributed by Shang QJ, Niranjan M, Sarma VV, Liu JK
31. Chaetosphaerellaceae Huhndorf, A.N. Mill. & F.A. Fernández, Mycol. Res. 108: 1387
(2004) – contributed by Huang SK, Jeewon R
32. Chaetosphaeriaceae Réblová, M.E. Barr & Samuels, Sydowia 51: 56 (1999) –
contributed by Perera RH, Luo ZL
33. Clavicipitaceae O.E. Erikss., Mycotaxon 15: 224 (1982) – contributed by Xiao YP,
Hongsanan S, Luang-sard J
34. Clypeophysalosporaceae A. Giraldo & Crous, Mycol. Progr. 16: 340 (2017) –
contributed by Chaiwan N
35. Clypeosphaeriaceae G. Winter, Rabenh. Krypt.-Fl., Edn 2 (Leipzig) 1.2: 554 (1886) –
contributed by Konta S, Boonmee S
36. Cocoonihabitaceae W.Y. Zhuang & Z.Q. Zeng – contributed by Bundhun D
37. Coniocessiaceae Asgari & Zare, Mycol. Progr. 10: 195 (2011) – contributed by
Norphanphoun C, Hongsanan S
38. Coniochaetaceae Malloch & Cain, Can. J. Bot. 49: 878 (1971) – contributed by
Samarakoon MC, Liu JK
39. Conioscyphaceae Réblová & Seifert, Persoonia 37: 63 (2015) – contributed by Bundhun
D
40. Conlariaceae H. Zhang, K.D. Hyde & Maharachch., Fungal Divers. 85: 90 (2017) –
contributed by Dong W, Doilom M
41. Cordanaceae (Sacc.) Nann., Repert. mic. uomo: 498 (1934) – contributed by
Goonasekara ID, Jayawardena RS
42. Cordycipitaceae Kreisel ex G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, Stud.
Mycol. 57: 48 (2007) – contributed by Xiao YP, Hongsanan S, Luang-sard J
43. Coronophoraceae Höhn., Sber. Akad. Wiss. Wien, Math.-naturw. Kl., Abt. 1 116: 624
(1907) – contributed by Huang SK, Maharachchikumbura SSN
44. Coryneaceae Corda, Icon. fung. (Prague) 3: 36 (1839) – contributed by Senanayake IC,
Bundhun D
45. Cryphonectriaceae Gryzenh. & M.J. Wingf., Mycologia 98: 246 (2006) – contributed by
Senanayake IC
46. Cylindriaceae Crous & L. Lombard, Fungal Systematics and Evolution 1: 183 (2018) –
contributed by Perera RH, Norphanphoun C
47. Cytosporaceae Fr., Syst. orb. veg. (Lundae) 1: 118 (1825) – contributed by
Norphanphoun C, Hongsanan S
48. Delonicicolaceae R.H. Perera, Maharachch. & K.D. Hyde, Cryptog. Mycol. 38: 334
(2017) – contributed by Perera RH, Jones EBG
49. Diaporthaceae Höhn. ex Wehm., Am. J. Bot. 13: 638 (1926) – contributed by
Dissanayake AJ, Niranjan M
50. Diaporthosporellaceae C.M. Tian & Q. Yang, Mycoscience 59: 230 (2018) – contributed
by Rathnayaka AR, Dissanayake A
51. Diaporthostomataceae X.L. Fan & C.M. Tian, Persoonia 40: 124 (2018) – contributed by
Rathnayaka AR, Dissanayake A
310
52. Diatrypaceae Verh. naturh. Ver. preuss. Rheinl. 26: 73 (1869) – contributed by Shang QJ
S, Niranjan M, Sarma VV, Tibpromma S
53. Distoseptisporaceae K.D. Hyde & McKenzie, Fungal Divers. 80: 402 (2016) –
contributed by Yang J, Liu JK, Luo ZL
54. Dwiroopaceae K.V. Xavier, A.N. KC, J.Z. Groenew., Vallad & Crous, Fungal
Systematics and Evolution 4: 38 (2019) – contributed by Devadatha B, Norphanphoun C
55. Erythrogloeaceae Senan., Maharachch. & K.D. Hyde, Stud. Mycol. 86: 258 (2017) –
contributed by Senanayake IC, Bundhun D
56. Etheirophoraceae Rungjind., Somrith. & Suetrong, Cryptog. Mycol. 35: 134 (2014) –
contributed by Dayarathne MC, Jones EBG
57. Falcocladiaceae Somrith., E.B.G. Jones & K.L. Pang, Cryptog. Mycol. 35: 134 (2014) –
contributed by Kumar V
58. Flammocladiellaceae Crous, L. Lombard & R.K. Schumach., Sydowia 67: 103 (2015) –
contributed by Perera RH, Wei DP, Jones EBG
59. Fuscosporellaceae J. Yang, J. Bhat & K.D. Hyde, Cryptog. Mycol. 37: 457 (2016) –
contributed by Yang J, Liu JK, Luo ZL
60. Glomerellaceae Locq. ex Seifert & W. Gams, Mycologia 98: 1083 (2007) – contributed
by Jayawardena RS, Hyde KD
61. Gnomoniaceae G. Winter, Rabenh. Krypt.-Fl., Edn 2 (Leipzig) 1.2: 570 (1886) –
contributed by Senanayake IC
62. Gondwanamycetaceae Réblová, W. Gams & Seifert, Stud. Mycol. 68: 188 (2011) –
contributed by Dayarathne MC, Jones EBG
63. Graphiaceae Z.W. de Beer, Seifert & M.J. Wingf., CBS Diversity Ser. (Utrecht) 12: 8
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66. Hansfordiaceae Crous, Fungal Systematics and Evolution 3: 84 (2019) – contributed by
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67. Harknessiaceae Crous, Persoonia 28: 55 (2012) – Marasinghe DS, Boonmee S
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69. Hispidicarpomycetaceae Nakagiri, Mycologia 85: 649 (1993) – contributed by
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70. Hypocreaceae De Not., G. bot. ital. 2: 48 (1844) – contributed by Perera RH, Niranjan M,
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71. Hyponectriaceae Petr., Annls mycol. 21: 305 (1923) – contributed by Konta S, Boonmee
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72. Hypoxylaceae DC., Fl. franç., Edn 3 (Paris) 2: 280 (1805) – contributed by Samarakoon
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73. Induratiaceae Samarak., Thongbai, K.D. Hyde & M. Stadler (in press) – contributed by
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74. Iodosphaeriaceae O. Hilber, The Genus Lasiosphaeria and Allied Taxa (Kelheim): 7
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77. Juncigenaceae E.B.G. Jones, Abdel-Wahab & K.L. Pang, Cryptog. Mycol. 35: 133 (2014)
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79. Kathistaceae Malloch & M. Blackw., Can. J. Bot. 68: 1719 (1990) – contributed by
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80. Koralionastetaceae Kohlm. & Volkm.-Kohlm., Mycologia 79: 764 (1987) – contributed
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81. Lamproconiaceae Norph., T.C. Wen & K.D. Hyde, Phytotaxa 270: 94 (2016) –
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82. Lasiosphaeriaceae Nannf., Nova Acta R. Soc. Scient. upsal., Ser. 4 8(no. 2): 50 (1932) –
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83. Lautosporaceae Kohlm., Volkm.-Kohlm. & O.E. Erikss., Bot. Mar. 38: 169 (1995) –
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84. Leptosilliaceae Voglmayr & Jaklitsch, Persoonia 42: 240 (2019) – contributed by
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85. Leptosporellaceae S. Konta & K.D. Hyde, Mycosphere 8: 1956 (2017) – contributed by
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87. Lopadostomataceae Daranag. & K.D. Hyde, Fungal Divers. 73: 129 (2015) – contributed
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88. Lulworthiaceae Kohlm., Spatafora & Volkm.-Kohlm., Mycologia 92: 456 (2000) –
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89. Macrohilaceae Crous, IMA Fungus 6: 180 (2015) – contributed by Wijayawardene N,
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90. Magnaporthaceae P.F. Cannon, Syst. Ascom. 13: 26 (1994) – contributed by
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92. Melanconidaceae G. Winter, Rabenh. Krypt.-Fl., Edn 2 (Leipzig) 1.2: 764 (1886) –
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94. Meliolaceae W. Martin ex Hansf., Mycol. Pap. 15: 23(1946) – contributed by Zeng XY,
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95. Melogrammataceae G. Winter, Rabenh. Krypt.-Fl., Edn 2 (Leipzig) 1.2: 797 (1886) –
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96. Microascaceae Luttr. ex Malloch, Mycologia 62: 734 (1970) – contributed by Kumar V
97. Microdochiaceae Hern.-Restr., Crous & J.Z. Groenew., Persoonia 36: 64 (2015) –
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99. Myrmecridiaceae Crous, Persoonia 34: 219 (2015) – contributed by Wei DP, Wanasinghe
DN
100. Myrotheciomycetaceae Crous, Persoonia 40: 351 (2018) – contributed by Norphanphoun
C
101. Nectriaceae Tul. & C. Tul., Select. fung. carpol. (Paris) 3: 3 (1865) – contributed by
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102. Neomelanconiellaceae Crous, Persoonia 41: 267 (2018) – contributed by Senanayake IC
103. Niessliaceae Kirschst., Annls mycol. 37: 89 (1939) – contributed by Huang SK, Jeewon R,
Hyde KD
104. Nitschkiaceae Nannf., Nova Acta R. Soc. Scient. upsal., Ser. 4 8(no. 2): 56 (1932) –
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105. Obryzaceae Körb., Syst. lich. germ. (Breslau): 427 (1855) – contributed by Bundhun D
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106. Ophioceraceae Klaubauf, E.G. LeBrun & Crous, Studies in Mycology 79: 85–120 (2014)
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107. Ophiocordycipitaceae G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, Stud. Mycol.
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108. Ophiostomataceae Nannf., Nova Acta R. Soc. Scient. upsal., Ser. 4 8(no. 2): 30 (1932) –
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124. Polystigmataceae Höhn. ex Nannf., Nova Acta R. Soc. Scient. upsal., Ser. 4 8(no. 2): 51
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126. Pseudodactylariaceae Crous, Persoonia 39: 421 (2017) – contributed by Lin CG, Liu JK
127. Pseudohalonectriaceae Hongsanan & K.D. Hyde, Fungal Divers. 84: 25–41(2017) –
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129. Pseudomelanconidaceae C.M. Tian & X.L. Fan, Persoonia 40: 128 (2018) – contributed
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131. Pseudoproboscisporaceae H. Zhang, K.D. Hyde & Maharachch., Fungal Divers. 85: 93
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132. Pseudosporidesmiaceae Crous, Persoonia 39: 365 (2017) – contributed by Luo ZL
133. Pseudotruncatellaceae Crous, Persoonia 42: 309 (2019) – contributed by Perera RH
313
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137. Rhamphoriaceae Réblová, Mycologia 110: 750–770 (2018) – contributed by Chen YJ
138. Sarocladiaceae L. Lombard, Persoonia 41: 343 (2018) – contributed by Phukhamsakda C
139. Savoryellaceae Jaklitsch & Réblová, Index Fungorum 209: 1 (2015) – contributed by
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140. Schizoparmaceae Rossman, D.F. Farr & Castl., Mycoscience 48: 137 (2007) –
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141. Scortechiniaceae Huhndorf, A.N. Mill. & F.A. Fernández, Mycol. Res. 108: 1387 (2004)
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142. Sordariaceae G. Winter, Rabenh. Krypt.-Fl., Edn 2 (Leipzig) 1.2: 162 (1885) –
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Introduction
Molecular data based on DNA sequence data is rapidly advancing taxonomic knowledge of
the fungi and recent efforts to provide classification schemes have been published for the basal
fungi (Wijayawardene et al. 2018a), and Basidiomycota (He et al. 2019). The eventual outcome
will be a classification scheme of all fungi, which can be updated periodically (Wijayawardene et
al., in prep). The last 15 years in mycology have seen turbulent times, especially for the
Ascomycota. This began with the AFTOL project which provided molecular data for the main
orders and families of Ascomycota (James et al. 2006). This was followed by the “one fungus one
name” concept to resolve the dual nomenclature problem for pleomorphic genera, a period of great
changes where many genera were combined under an arguably preferred name (Taylor 2011) and
notes on all genera of ascomycetes were provided (Wijayawardene et al. 2017b). Numerous new
higher taxa have been introduced, initially with phylogenetic evidence, followed by support from
molecular dating (Mapook et al. 2016, Hyde et al. 2017a, Hongsanan et al. 2017, Liu et al. 2017,
Zeng et al. 2019).
The first main attempts at classifying all genera of Sordariomycetes were those of Barr (1983,
1987, 1990b) and Eriksson & Hawksworth (1986a, 1993). These were magnificent attempts to
classify often poorly known taxa and were based solely on morphology. We should not forget the
incredible knowledge these authors had and incorporated into these documents. Classification of
the sexual morphs of Sordariomycetes were continued by Lumbsch & Huhndorf (2007, 2010),
which incorporated morphology and available phylogenies. The first classification of
Sordariomycetes in which the asexual and sexual states were included, and was partially based on
molecular data were published by Maharachchimbura et al. (2015b, 2016b). Hongsanan et al.
(2017) used divergence time estimates to provide support for the various orders and families in the
class. These ground-breaking papers provided the background for the present paper, where we
update the classification of Sordariomycetes, with notes on the orders, families and genera.
Sordariomycetes is an important class of ascomycetes, characterized by non-lichenized, flaskshaped fruiting bodies or less frequently cleistothecial ascomata and unitunicate asci (Zhang et al.
2006, Maharachchikumbura et al. 2016b), and can have a varied growth form and colonise diverse
habitats (Hongsanan et al. 2017). It is the second largest class of Ascomycota (Hyde et al. 2013,
Maharachchikumbura et al. 2015, 2016b). Sordariomycetes have a cosmopolitan distribution and
can be found in almost all ecosystems (Pratibha et al. 2014, Jones et al. 2015). Some
Sordariomycetes are phytopathogens causing leaf, stem, root and fruit diseases (e.g.
Colletotrichum, Coniella, Diaporthe), while some are pathogenic to arthropod and mammals (eg.
Sporothrix, Fusarium). Some members are endophytes (e.g. Pestalotiopsis) (Maharachchikumbura
et al. 2014b, Norphanphoun et al. 2019) and some are saprobes involved in decomposition and
nutrient cycling (Chaetomium, Neurospora) (Zhang & Wang 2015). Some taxa are fungicolous
(Sun et al. 2019) and several are opportunistic pathogens of humans (Gostinčar et al. 2018). Some
taxa of Sordariomycetes are economically important as biocontrol agents (Harman et al. 2004,
Hyde et al. 2019a) and some species produce a wide range of important secondary metabolites
(Maharachchikumbura et al. 2016b).
315
The classification of the class Sordariomycetes has changed drastically over the past decade
(Wijayawardene et al. 2018). Characters alone are insufficient to resolve the identification of many
species and even some genera (Tang et al. 2007, 2009). Molecular studies on this class began in the
early 1990s using SSU and LSU sequence data (Berbee & Taylor 1992, Zhang et al. 2006).
However, SSU and LSU sequence data are insufficient to provide a good resolution in most of the
groups in this class. It has been established that the use of protein genes can yield a higher
phylogenetic resolution (Tang et al. 2007, 2009, Schoch et al. 2009, Maharachchikumbura et al.
2016b).
The subclasses of Sordariomycetes have expanded from three to six. Eriksson & Winka
(1997) introduced Hypocreomycetidae, Sordariomycetidae and Xylariomycetidae based on
morphology and nrDNA sequence data. In a revision of Sordariomycetes, Maharachchikumbura et
al. (2016b) used morphology and a combined analysis of LSU, SSU, tef1 and rpb2 sequence data
and introduced another three new subclasses; Diaporthomycetidae, Lulworthiomycetidae, and
Meliolomycetidae. The subclass Meliolomycetidae was found to be a synonym of
Sordariomycetidae, while Savoryellomycetidae was raised to subclass level (Hongsanan et al. 2017,
Wijayawardene et al. 2018a, Dayarathne et al. 2019a).
Kirk et al. (2008) listed 1,119 genera and over 10,000 species in this class. In the outline of
the Sordariomycetes, Maharachchikumbura et al. (2015) included 28 orders, 90 families and 1344
genera. According to the outline by Maharachchikumbura et al. (2016b), Sordariomycetes had six
subclasses, 32 orders, 105 families and 1331 genera. Hongsanan et al. (2017) provided an updated
backbone tree for Sordariomycetes based on divergence times and proposed changes to this class.
In this paper, we deal with the family level classification of Sordariomycetes by updating the
outline given by Maharachchikumbura et al. (2016b).
Layout of the paper
Family descriptions, notes on history and a brief account of the genera including phylogenetic
data if available are provided for each family. The type genus/new species/new host record are
illustrated with a representative plate of features followed by a description and a note for each
family. This is followed by ecological and economic significance of the family and the accepted
genera and their type species. For consistency, authorities and abbreviations follow Index
Fungorum (2020).
Arrangement of Sordariomycetes
The arrangement of this paper follows Maharachchikumbura et al. (2016b), which has been
updated based on recent publications and interpretation of genera from the literature
(Wijayawardene et al. 2018a).
Examination of specimens
Herbarium material
Specimens or slides were obtained from CF, CUP, DAOM, F, FH, G, GZUH, HKAS, IFO,
IFO-H, ILLS, ILLS, IMI, IMS, K(M), KCM, KUN, M, MFLU, MICH, MUCL, NY, O-F, PDD,
PREM, PRM, S-F, TRTC, WU, and ZT MYZ. Fruiting bodies were rehydrated in water and/or 5%
KOH prior to examination and sectioning. Hand sections of the fruiting bodies were mounted in
water for microscopic studies and photomicrography.
Fresh collection
Samples were collected from China, India, Italy, Russia and Thailand. Fungi were isolated by
a modified single spore isolation method (Chomnunti et al. 2014). Cultures were transferred to
different media i.e. 2% potato dextrose agar (PDA) and malt dextrose agar (MEA) or water agar
(WA; 15g/l sterile distilled water), and incubated under different conditions depending on the taxon
to induce sporulation. Extype or representative isolates were deposited in Mae Fah Luang
316
University culture collection (MFLUCC) with duplicates in FU, GZCC, JZB, KUMCC, and NFCC.
The collected specimens were deposited in the Herbarium of Mae Fah Luang University (MFLU),
Thailand, AMH, FU, and HKAS. Vegetative and reproductive structures were mounted in clear
lactic acid or water, either directly from specimens or from colonies sporulating on media. Sections
of fruiting bodies were made by hand and examined. Isolates were grown and incubated on
different culture media and temperatures as required for each genus. Colour notations followed the
colour charts of Rayner (1970). Taxonomic descriptions were deposited in the Facesoffungi (FOF)
database as described in Jayasiri et al. (2015) and Index Fungorum numbers were obtained as
detailed in Index Fungorum database. New species were established based on the guidelines
provided by Jeewon & Hyde (2016).
Both herbarium and fresh collections were examined using a Nikon ECLIPSE 8oi compound
microscope and photographed with a Cannon 450D digital camera fitted to the microscope.
Measurements were made with the Tarosoft (R) Image Frame Work program (v.0.9.0.7) and
images used for figures processed with Adobe Photoshop CS6 software (Adobe Systems, USA).
For the specimens that were in poor condition, hand drawings based on the original publications
were prepared using drawing pens and parchment papers. DNA extraction, PCR amplification,
sequencing and phylogenetic analyses follow previous papers (Hyde et al. 2016a, Tibpromma et al.
2017).
Phylogenetic analysis and divergence estimates
The phylogenetic analysis follows Maharachchimbura et al. (2015b, 2016b). The MCC
analysis was performed in BEAST v1.10.2. The crown age of Sordariomycetes was set with
Normal distribution, mean = 250, SD = 30, with 97.5% of CI = 308.8 MYA, and crown age of
Dothideomycetes with Normal distribution mean = 360, SD = 20, with 97.5% of CI = 399 MYA.
The substitution models were selected based on jModeltest2.1.1; GTR+I+G for LSU, RPB2 and
SSU, and TrN+I+G for TEF (the model TrN is not available in BEAUti 1.10.2, thus we used
TN93). Lognormal distribution of rates was used during the analyses with uncorrelated relaxed
clock model. The Yule process tree prior was used to model the speciation of nodes in the topology
with a randomly generated starting tree. The analyses were performed for 100 million generations,
with sampling parameters every 10000 generations. The effective sample sizes were checked in
Tracer v.1.6 and the acceptable values are higher than 200. The first 20% representing the burn-in
phase were discarded and the remaining trees were combined in LogCombiner 1.10.2., summarized
data and estimated in TreeAnnotator 1.10.2. Bars correspond to the 95% highest posterior density
(HPD) intervals. The scale axis (Fig. 2) shows divergence times as millions of years ago (MYA).
Arrangement of Sordariomycetes with classes, subclasses, orders, families and genera with
numbers of species in brackets
Class Sordariomycetes O.E. Erikss. & Winka
Subclass Diaporthomycetidae Senan. et al.
Annulatascales D’souza et al.
Annulatascaceae S.W. Wong et al.
Annulatascus K.D. Hyde
Annulusmagnus J. Campb. & Shearer
Aqualignicola Ranghoo et al.
Ascitendus J. Campb. & Shearer
Ayria Fryar & K.D. Hyde
Cataractispora K.D. Hyde
Chaetorostrum Zelski et al.
Longicollum Zelski et al.
Submersisphaeria K.D. Hyde
Vertexicola K.D. Hyde et al.
317
Annulatascales genera incertae sedis
Clohiesia K.D. Hyde
Atractosporales H. Zhang et al.
Atractosporaceae H. Zhang et al.
Atractospora Réblová & J. Fourn.
Rubellisphaeria Réblová & J. Fourn.
Conlariaceae H. Zhang et al.
Conlarium F. Liu & L. Cai
Riomyces A. Ferrer et al.
Pseudoproboscisporaceae H. Zhang et al.
Diluviicola K.D. Hyde et al.
Pseudoproboscispora Punith.
Calosphaeriales M.E. Barr
Calosphaeriaceae Munk
Calosphaeria Tul. & C Tul.
Flabellascus Réblová
Jattaea Berl
Togniniella Réblová et al.
Pleurostomataceae Réblová et al.
Pleurostoma Tul. & C. Tul.
Calosphaeriales genus incertae sedis
Calosphaeriopsis Petr.
Enchnoa Fr.
Kacosphaeria Speg.
Sulcatistroma A.W. Ramaley
Diaporthales Nannf.
Apiosporopsidaceae Senan. et al.
Apiosporopsis (Traverso) Mariani.
Apoharknessiaceae Senan. et al.
Apoharknessia Crous & S.J. Lee
Lasmenia Speg.
Asterosporiaceae Senan. et al.
Asterosporium Kunze
Auratiopycnidiellaceae Senan. et al.
Auratiopycnidiella Crous & Summerell
Coryneaceae Corda
Coryneum Nees
Cryphonectriaceae Gryzenh. & M.J. Wingf.
Amphilogia Gryzenh. et al.
318
Aurantioporthe G. Beier & R.A. Blanchette
Aurantiosacculus Dyko & B. Sutton
Aurapex Gryzenh. & M.J. Wingf.
Aurifilum Begoude et al.
Capillaureum M.E.S. Oliveira
Celoporthe Nakab. et al.
Chromendothia Lar.N. Vassiljeva
Chrysofolia Crous & M.J. Wingf.
Chrysomorbus S.F. Chen
Chrysoporthe Gryzenh. & M.J. Wingf.
Corticimorbus S.F. Chen & M.J. Wingf.
Cryphonectria (Sacc.) Sacc. & D. Sacc.
Cryptometrion Gryzenh. & M.J. Wingf.
Diversimorbus S.F. Chen & J. Roux
Endothia Fr.
Foliocryphia Cheew. & Crous
Holocryphia Gryzenh. & M.J. Wingf.
Immersiporthe S.F. Chen et al.
Latruncellus M. Verm. et al.
Luteocirrhus C.F. Crane & T.I. Burgess
Mastigosporella Höhn.
Microthia Gryzenh. & M.J. Wingf.
Myrtonectria Marinc. et al.
Rostraureum Gryzenh. & M.J. Wingf.
Ursicollum Gryzenh. & M.J. Wingf.
Wuestneia Auersw. ex Fuckel
Cytosporaceae Fr.
Cryptascoma Ananthap.
Cytospora Ehrenb.
Pachytrype Berl. ex M.E. Barr et al.
Paravalsa Ananthap.
Waydora B. Sutton
Xenotypa Petr.
Diaporthaceae Höhn. ex Wehm.
Apioporthella Petr.
Apiosphaeria Höhn.
Chaetoconis Clem.
Chiangraiomyces Senan. & K.D. Hyde
Diaporthe Nitschke
Hyaliappendispora Senan. et al.
Leucodiaporthe M.E. Barr & Lar.N. Vassiljeva
Massariothea Syd.
Mazzantia Mont.
Ophiodiaporthe Y.M. Ju et al.
Paradiaporthe Senan. et al.
Phaeocytostroma Petr.
Phaeodiaporthe Petr.
Pustulomyces D.Q. Dai et al.
Stenocarpella Syd. & P. Syd.
319
Diaporthosporellaceae C.M. Tian & Q. Yang
Diaporthosporella C.M. Tian & Q. Yang
Diaporthostomataceae X.L. Fan & C.M. Tian
Diaporthostoma X.L. Fan & C.M. Tian
Dwiroopaceae K.V. Xavier et al.
Dwiroopa Subram. & Muthumary
Erythrogloeaceae Senan. et al.
Chrysocrypta Crous & Summerell
Dendrostoma X.L. Fan & C.M. Tian
Disculoides Crous et al.
Erythrogloeum Petr.
Gnomoniaceae G. Winter
Alnecium Voglmayr & Jaklitsch
Ambarignomonia Sogonov
Amphiporthe Petr.
Anisomyces Theiss. & Syd.
Apiognomonia Höhn.
Apioplagiostoma M.E. Barr
Asteroma DC.
Bagcheea E. Müll. & R. Menon
Chadefaudiomyces Kamat
Clypeoporthe Höhn.
Cryptosporella Sacc.
Dictyoporthe Petr.
Diplacella Syd.
Ditopella De Not.
Ditopellopsis J. Reid & C. Booth
Gloeosporidina Petr.
Gnomonia Ces. & De Not.
Gnomoniella Sacc.
Gnomoniopsis Berl.
Maculatipalma J. Fröhlich & K.D. Hyde
Mamianiella Höhn.
Marsupiomyces Senan. & K.D. Hyde
Millerburtonia Cif .
Occultocarpon L.C. Mejía & Zhu L. Yang
Ophiognomonia (Sacc.) Sacc.
Phragmoporthe Petr.
Phylloporthe Syd.
Plagiostoma Fuckel
Pleuroceras Riess.
Sirococcus Preuss
Spataporthe Bronson et al.
Tenuignomonia Minosh. et al.
Uleoporthe Petr.
Uniseta Ciccar
Valsalnicola D.M. Walker & Rossman
Vismaya V.V. Sarma & K.D. Hyde
320
Harknessiaceae Crous
Harknessia Cooke
Mebarria J. Reid & C. Booth
Juglanconidaceae Voglmayr & Jaklitsch
Juglanconis Voglmayr & Jaklitsch
Lamproconiaceae C. Norphanphoun et al.
Hercospora Fr.
Lamproconium (Grove) Grove
Macrohilaceae Crous
Macrohilum H.J. Swart
Melanconidaceae G. Winter
Melanconis Tul. & C. Tul.
Melanconiellaceae Senan. et al.
Dicarpella Syd. & P. Syd.
Greeneria Scribn. & Viala
Massariovalsa Sacc.
Melanconiella Sacc.
Microascospora Senan. & K.D. Hyde
Septomelanconiella Samarak. & K.D. Hyde
Sheathospora X.L. Fan
Neomelanconiellaceae Crous
Neomelanconiella Crous
Phaeoappendicosporaceae Crous & M.J. Wingf.
Neophaeoappendicospora Crous & M.J. Wingf.
Phaeoappendicospora Senan. et al.
Phaeochorellaceae Guterres et al.
Phaeochorella (Henn.) Theiss. & Syd.
Prosopidicolaceae Senan. & K.D. Hyde
Prosopidicola Crous & C.L. Lennox
Pseudomelanconidaceae C.M. Tian & X.L. Fan
Pseudomelanconis C.M. Tian & X.L. Fan
Neopseudomelanconis C.M. Tian & N. Jiang
Pseudoplagiostomataceae Cheew. et al.
Pseudoplagiostoma Cheew. et al.
Schizoparmaceae Rossman
Coniella Höhn.
Stilbosporaceae Link
Crinitospora B. Sutton & Alcorn
321
Natarajania Pratibha & Bhat
Stegonsporium Corda
Stilbospora Pers.
Sydowiellaceae Lar.N. Vassiljeva
Alborbis Senan. & K.D. Hyde
Breviappendix Senan. & K.D. Hyde
Cainiella E. Müll.
Calosporella J. Schröt
Caudospora Starbäck
Chapeckia M.E. Barr
Hapalocystis Auersw. ex Fuckel
Italiomyces Senan. et al.
Lambro Racib.
Paragnomonia Senan. & K.D. Hyde
Ranulospora Senan. et al.
Rossmania Lar.N. Vassiljeva
Sillia P. Karst.
Sydowiella Petr.
Tenuiappendicula Senan. et al.
Tortilispora Senan. & K.D. Hyde
Synnemasporellaceae X.L. Fan & J.D.P. Bezerra
Synnemasporella X.L. Fan & J.D.P. Bezerra
Tubakiaceae U. Braun et al.
Apiognomonioides U. Braun et al.
Involutscutellula U. Braun & C. Nakash.
Oblongisporothyrium U. Braun & C. Nakash.
Paratubakia U. Braun & C. Nakash.
Racheliella Crous & U. Braun
Saprothyrium U. Braun et al.
Sphaerosporithyrium U. Braun et al.
Tubakia B. Sutton
Diaporthales genera incertae sedis
Ceratoporthe Petr.
Cryptoleptosphaeria Petr.
Cytomelanconis Naumov
Diaporthella Petr.
Diatrypoidiella Manohar et al.
Ditopellina J. Reid & C. Booth
Durispora K.D. Hyde
Exormatostoma Gray
Fremineavia Nieuwl.
Gibellia Sacc.
Gyrostroma Naumov
Hyalorostratum Raja & Shearer
Hypophloeda K.D. Hyde & E.B.G. Jones
Hypospilina (Sacc) Traverso
Kapooria J. Reid & C. Booth
Keinstirschia J. Reid & C. Booth
322
Lollipopaia Inderb.
Macrodiaporthe Petr.
Melanamphora Lafl.
Phragmodiaporthe Wehm.
Phruensis Pinruan
Plagiophiale Petr.
Plagiostigme Syd.
Prostratus Sivan. et al.
Pseudocryptosporella J. Reid & C. Booth
Pseudothis Theiss. & Syd.
Pseudovalsella Höhn.
Savulescua Petr.
Sphaerognomoniella Naumov & Kusnezowa
Stioclettia Dennis
Trematovalsa Jacobesco
Wehmeyera J. Reid & C. Booth
Distoseptisporales Z.L. Luo et al.
Distoseptisporaceae K.D. Hyde & McKenzie
Distoseptispora K.D. Hyde et al.
Jobellisiales M.J. D’souza & K.D. Hyde
Jobellisiaceae Réblová
Jobellisia M.E. Barr
Magnaporthales Thongk. et al.
Ceratosphaeriaceae Z.L. Luo et al.
Ceratosphaeria Niessl
Magnaporthaceae P.F. Cannon
Bifusisporella R.M.F. Silva
Budhanggurabania P. Wong et al.
Buergenerula Syd.
Bussabanomyces Klaubauf et al.
Ceratosphaerella Huhndorf et al.
Clasterosporium Schwein
Clavatisporella K.D. Hyde
Falciphora J. Luo & N. Zhang
Falciphoriella M. Hern.-Restr. & Crous
Gaeumannomycella M. Hern.-Restr. & Crous
Gaeumannomyces Arx & D.L. Olivier
Herbampulla Scheuer & Nograsek
Kohlmeyeriopsis Klaubauf et al.
Magnaporthiopsis J. Luo & N. Zhang
Muraeriata Huhndorf et al.
Mycoleptodiscus Ostaz.
Nakataea Hara
Neogaeumannomyces D.Q. Dai & K.D. Hyde
Omnidemptus P.F. Cannon & Alcorn
Pseudophialophora J. Luo & N. Zhang
Pyriculariopsis M.B. Ellis
Slopeiomyces Klaubauf et al.
323
Ophioceraceae Klaubauf et al.
Ophioceras Sacc.
Pseudohalonectriaceae Hongsanan & K.D. Hyde
Pseudohalonectria Minoura & T. Muroi
Pyriculariaceae Klaubauf et al.
Bambusicularia Klaubauf et al.
Barretomyces Klaubauf et al.
Deightoniella S. Hughes
Macgarvieomyces Klaubauf et al.
Neocordana Hern.-Rest. & Crous
Neopyricularia Klaubauf et al.
Proxipyricularia Klaubauf et al.
Pseudopyricularia Klaubauf et al.
Pyricularia Sacc.
Pyriculariomyces Y. Marín et al.
Xenopyricularia Klaubauf et al.
Myrmecridiales Crous
Myrmecridiaceae Crous
Myrmecridium Arzanlou et al.
Neomyrmecridium Crous
Xenodactylariaceae Crous
Xenodactylaria Crous
Ophiostomatales Benny & Kimbr.
Kathistaceae Malloch & M. Blackw.
Kathistes Malloch & M. Blackw.
Mattirolella S. Colla
Termitariopsis M. Blackw. et al.
Ophiostomataceae Nannf.
Afroraffaelea C.C. Bateman et al.
Aureovirgo J.A. van der Linde et al.
Ceratocystiopsis H.P. Upadhyay & W.B. Kendr.
Fragosphaeria Shear
Graphilbum H.P. Upadhyay & W.B. Kendr.
Hawksworthiomyces Z.W. de Beer et al.
Klasterskya Petr.
Leptographium Lagerb. & Melin
Ophiostoma Syd. & P. Syd.
Raffaelea Arx & Hennebert
Sporothrix Hektoen & C.F. Perkins
Spumatoria Massee & E.S. Salmon
Subbaromyces Hesselt.
Pararamichloridiales Crous
Pararamichloridiaceae Crous
Pararamichloridium Crous
324
Phomatosporales Senan. et al.
Phomatosporaceae Senan. & K.D. Hyde
Lanspora K.D. Hyde & E.B.G. Jones
Phomatospora Sacc.
Tenuimurus Senan. et al.
Sporidesmiales Crous
Sporidesmiaceae Fr.
Sporidesmium Link
Tirisporellales Suetrong et al.
Tirisporellaceae Suetrong et al.
Bacusphaeria Norlailatul et al.
Thailandiomyces Pinruan et al.
Tirisporella E.B.G. Jones et al.
Togniniales Senan. et al.
Togniniaceae Réblová et al.
Conidiotheca Réblová & L Mostert
Phaeoacremonium W. Gams et al.
Xenospadicoidales Hern.-Restr., J. Mena & Gené
Xenospadicoidaceae Hern.-Restr., J. Mena & Gené
Calyptosphaeria Réblová & A.N. Mill.
Lentomitella Höhn.
Neospadicoides Z.L. Luo
Spadicoides S. Hughes
Torrentispora K.D. Hyde et al.
Diaporthomycetidae families incertae sedis
Barbatosphaeriaceae H. Zhang et al.
Barbatosphaeria Réblová
Ceratostomella Sacc.
Xylomelasma Réblová
Papulosaceae Winka & O.E. Erikss.
Brunneosporella V.M. Ranghoo & K.D. Hyde
Fluminicola S.W. Wong et al.
Papulosa Kohlm & Volkm-Kohlm.
Wongia Khemmuk et al.
Rhamphoriaceae Réblová
Rhamphoria Niessl
Rhamphoriopsis Réblová & Gardiennet
Rhodoveronaea Arzanlou, W. Gams & Crous
Xylolentia Réblová
Thyridiaceae J.Z. Yue & O.E. Erikss.
Pleurocytospora Petr.
Thyridium Nitschke
Trichosphaeriaceae G. Winter
325
Brachysporium Sacc.
Collematospora Jeng & Cain
Coniobrevicolla Réblová
Eriosphaeria Sacc.
Koorchaloma Subram.
Rizalia Syd. & P. Syd.
Schweinitziella Speg.
Setocampanula Sivan. & W.H. Hsieh
Trichosphaeria Fuckel
Unisetosphaeria Pinnoi et al.
Woswasiaceae H. Zhang et al.
Cyanoannulus Raja et al.
Woswasia Jaklitsch et al.
Xylochrysis Réblová et al.
Diaporthomycetidae genera incertae sedis
Aquapteridospora Jiao Yang et al.
Aquaticola W.H. Ho et al.
Fusoidispora D. Vijaykrishna et al.
Platytrachelon Réblová
Proliferophorum G.N. Wang et al.
Pseudostanjehughesia J. Yang & K.D. Hyde
Subclass Hypocreomycetidae O.E. Erikss. & Winka
Coronophorales Nannf.
Bertiaceae Smyk
Bertia De Not.
Gaillardiella Pat.
Ceratostomataceae G. Winter
Arxiomyces P.F. Cannon & D. Hawksw.
Dactylidispora Y. Marín et al.
Echinusitheca Y. Marín et al.
Erythrocarpon Zukal
Harzia Costantin
Melanospora Corda
Microthecium Corda
Pseudomicrothecium Y. Marín et al.
Pustulipora P.F. Cannon
Rhytidospora Jeng & Cain
Scopinella Lév.
Setiferotheca Matsush.
Syspastospora P.F. Cannon & D. Hawksw.
Vittatispora P. Chaudhary et al.
Chaetosphaerellaceae Huhndorf et al.
Chaetosphaerella E. Müll. & C. Booth
Crassochaeta Réblová
Spinulosphaeria Sivan.
Coronophoraceae Höhn.
326
Coronophora Fuckel
Nitschkiaceae (Fitzp.) Nannf.
Acanthonitschkea Speg.
Biciliosporina Subram. & Sekar
Botryola Bat. & J.L. Bezerra
Fracchiaea Sacc.
Groenhiella Jørg. Koch et al.
Janannfeldtia Subram. & Sekar
Lasiosphaeriopsis D. Hawksw. & Sivan.
Loranitschkia Lar.N. Vassiljeva
Neochaetosphaerella Lar.N. Vassiljeva et al.
Neotrotteria Sacc.
Nitschkia G.H. Otth ex P. Karst.
Rhagadostoma Körb.
Rhagadostomella Etayo
Tortulomyces Lar.N. Vassiljeva et al.
Scortechiniaceae Huhndorf et al.
Biciliospora Petr.
Coronophorella Höhn.
Cryptosphaerella Sacc.
Euacanthe Theiss.
Neofracchiaea Teng
Pseudocatenomycopsis Crous & L.A. Shuttlew.
Scortechinia Sacc.
Scortechiniella Arx & E. Müll.
Scortechiniellopsis Sivan.
Tympanopsis Starbäck
Coronophorales genus incertae sedis
Papulaspora Preuss
Sphaerodes Clem.
Falcocladiales R.H. Perera et al.
Falcocladiaceae Somrithipol et al.
Falcocladium S.F. Silveira et al.
Glomerellales Chadef. ex Réblová et al.
Australiascaceae Réblová & W. Gams
Monilochaetes Halst. ex Harter
Glomerellaceae Locq. ex Seifert & W. Gams
Colletotrichum Corda
Malaysiascaceae Tibpromma & K.D. Hyde
Malaysiasca Crous & M.J. Wingf.
Plectosphaerellaceae W. Gams et al.
Acremoniisimulans Tibpromma & K.D. Hyde
Acrostalagmus Corda
Brunneochlamydosporium Giraldo López & Crous
Brunneomyces A. Giraldo, Gené & Guarro
327
Chlamydosporiella Giraldo López & Crous
Chordomyces Bilanenko et al.
Furcasterigmium Giraldo López & Crous
Fuscohypha Giraldo López & Crous
Gibellulopsis Bat. & H. Maia
Lectera P.F. Cannon
Longitudinalis Tibpromma & K.D. Hyde
Musicillium Zare & W Gams
Musidium Giraldo López & Crous
Nigrocephalum Giraldo López & Crous
Paragibellulopsis Giraldo López & Crous
Paramusicillium Giraldo López & Crous
Phialoparvum Giraldo López & Crous
Plectosphaerella Kleb.
Sayamraella Giraldo López & Crous
Sodiomyces A.A. Grum-Grzhim. et al.
Stachylidium Link
Summerbellia Giraldo López & Crous
Theobromium Giraldo López & Crous
Verticillium Nees
Reticulascaceae Réblová & W. Gams
Blastophorum Matsush.
Cylindrotrichum Bonord.
Kylindria DiCosmo et al.
Sporoschismopsis Hol-Jech. & Hennebert
Glomerellales genera incertae sedis
Ascocodinaea Samuels et al.
Hypocreales Lindau
Bionectriaceae Samuels & Rossman
Acremonium Link
Anthonectria Döbbeler
Aphanotria Döbbeler
Battarrina (Sacc.) Clem. & Shear
Bryocentria Döbbeler
Bryotria Döbbeler & P.G. Davison
Bullanockia Crous
Clibanites (P. Karst.) P. Karst.
Clonostachys Corda
Dimerosporiella Speg.
Fusariella Sacc.
Geonectria Lechat & J. Fourn.
Geosmithia Pitt
Gliomastix Guég.
Globonectria Etayo
Gracilistilbella Seifert
Halonectria E.B.G. Jones
Heleococcum P.M. Jørg.
Hydropisphaera Dumort
Ijuhya Starbäck
328
Kallichroma Kohlm. & Volkm.-Kohlm.
Laniatria Döbbeler & P.G. Davison
Lasionectria (Sacc.) Cooke
Lasionectriella Lechat & J. Fourn.
Mycoarachis Malloch & Cain
Mycocitrus Möller
Nectriella Nitschke ex Fuckel
Nectriopsis Maire
Nigrosabulum Malloch & Cain
Ochronectria Rossman & Samuels
Ovicuculispora Etayo
Paracylindrocarpon Crous et al.
Paranectria Sacc.
Periantria Döbbeler & P.G. Davison
Peristomialis (W. Phillips) Boud.
Pronectria Clem.
Protocreopsis Yoshim Doi
Rhopalocladium Schroers et al.
Roumegueriella Speg.
Selinia P. Karst.
Stephanonectria Schroers & Samuels
Stilbocrea Pat.
Stromatonectria Jaklitsch & H. Voglmayr
Synnemellisia N.K. Rao et al.
Trichonectria Kirschst.
Verrucostoma Hirooka et al.
Xanthonectria Lechat et al.
Calcarisporiaceae Jing Z. Sun et al.
Calcarisporium Preuss
Clavicipitaceae (Lindau) Earle ex Rogerson
Aciculosporium I. Miyake
Aschersonia Mont.
Atkinsonella Diehl
Balansia Speg.
Cavimalum Yoshim. Doi et al.
Claviceps Tul.
Collarina Giraldo et al.
Conoideocrella D. Johnson et al.
Corallocytostroma Y.N. Yu & Z.Y. Zhang
Dussiella Pat.
Ephelis Fr.
Epichloë (Fr.) Tul. & C. Tul.
Epicrea Petr.
Helicocollum Luangsa-ard
Helminthascus Tranzschel
Heteroepichloë E. Tanaka et al.
Konradia Racib.
Loculistroma F. Patt & Charles
Metapochonia Kepler et al.
Metarhiziopsis D.W. Li et al.
329
Metarhizium Sorokīn
Moelleriella Bres.
Mycomalus A. Möller
Mycophilomyces Crous & M.J. Wingf.
Myriogenospora G.F. Atk.
Neobarya Lowen
Neocordyceps Kobayasi
Nigelia Luangsa-ard
Nigrocornus Ryley & Langdon
Orbiocrella D. Johnson et al.
Parepichloë J.F. White & P.V. Reddy
Periglandula U. Steiner et al.
Pochonia Bat. & O.M. Fonseca
Pseudomeria G.L. Barron
Regiocrella Chaverri & K.T. Hodge
Romanoa Thirum.
Rotiferophthora G.L. Barron
Samuelsia Chaverri & K.T. Hodge
Shimizuomyces Kobayasi
Sphaerocordyceps Kobayasi
Tyrannicordyceps Kepler & Spatafora
Ustilaginoidea Bref.
Cocoonihabitaceae W.Y. Zhuang & Z.Q. Zeng
Cocoonihabitus W.Y. Zhuang & Z.Q. Zeng
Cordycipitaceae Kreisel ex G.M. Sung et al.
Akanthomyces Lebert
Amphichorda Fr.
Ascopolyporus Möller
Beauveria Vuill.
Beejasamuha Subram. & Chandrash.
Blackwellomyces Spatafora & Luangsa-ard
Cordyceps (Fr.) Link
Coremiopsis Sizova & Suprun
Engyodontium de Hoog
Gibellula Cavara
Hevansia Luangsa-ard et al.
Hyperdermium J. White et al.
Leptobacillium Zare & W. Gams
Parengyodontium C.C. Tsang
Pseudogibellula Samson & H.C. Evans
Samsoniella Mongkols. et al.
Simplicillium W. Gams & Zare
Flammocladiellaceae Crous et al.
Flammocladiella Crous et al.
Hypocreaceae De Not.
Arachnocrea Z. Moravec.
Dialhypocrea Speg.
Escovopsioides H.C. Evans & J.O. Augustin
Escovopsis J.J. Muchovej & Della Lucia
330
Hypocreopsis P. Karst.
Hypomyces (Fr.) Tul. & C. Tul.
Kiflimonium Summerb. et al.
Lichenobarya Etayo et al.
Mycogone Link
Protocrea Petch
Rogersonia Samuels & Lodge
Sepedonium Link
Sphaerostilbella (Henn.) Sacc. & D. Sacc
Sporophagomyces K. Põldmaa & Samuels
Stephanoma Wallr.
Trichoderma Pers.
Verticimonosporium Matsush.
Myrotheciomycetaceae Crous
Emericellopsis J.F.H. Beym
Leucosphaerina Arx
Myrotheciomyces Crous
Trichothecium Link
Nectriaceae Tul. & C. Tul.
Albonectria Rossman & Samuels
Allantonectria Earle
Allonectella Petr.
Aphanocladium W. Gams
Aquanectria L. Lombard & Crous
Atractium Link
Baipadisphaeria Pinruan
Bisifusarium L. Lombard et al.
Calonectria De Not.
Calostilbe Sacc. & Syd.
Campylocarpon Halleen et al.
Chaetonectrioides Matsush.
Chaetopsina Rambelli
Coccinonectria Lombard & Crous
Corallomycetella Henn.
Corallonectria C. Herrera & P. Chaverri
Corinectria C. González & P. Chaverri
Cosmospora Rabenh.
Cosmosporella S.K. Huang et al.
Curvicladiella Decock & Crous
Cyanochyta Höhn.
Cyanonectria Samuels & Chaverri
Cyanophomella Höhn.
Cylindrocladiella Boesew.
Cylindrodendrum Bonord.
Dacryoma Samuels
Dactylonectria L. Lombard & Crous
Dematiocladium Allegr. et al.
Fusarium Link
Fusicolla Bonord
Geejayessia Schroers et al.
331
Gliocephalotrichum J.J. Ellis & Hesselt.
Gliocladiopsis S.B. Saksena
Ilyonectria P. Chaverri & C. Salgado
Macroconia (Wollenw.) Gräfenhan et al.
Mariannaea G. Arnaud ex Samson
Microcera Desm.
Murinectria M. Niranjan & V.V. Sarma
Nalanthamala Subram.
Nectria (Fr.) Fr.
Neocosmospora E.F. Sm.
Neonectria Wollenw.
Neothyronectria Crous & Thangavel
Ophionectria Sacc.
Pandanaceomyces Tibpromma & K.D. Hyde
Paracremonium L. Lombard & Crous
Payosphaeria W.F. Leong
Penicillifer Emden
Persiciospora P.F. Cannon & D. Hawksw.
Pleiocarpon L. Lombard & D. Aiello
Pleogibberella Sacc.
Pleurocolla Petr.
Pseudoachroiostachys Tibpromma & K.D. Hyde
Pseudocosmospora C. Herrera & P. Chaverri
Pseudonectria Seaver
Rectifusarium Lombard et al.
Rugonectria P. Chaverri & Samuels
Sarcopodium Ehrenb.
Stylonectria Höhn.
Thelonectria P. Chaverri & C.G. Salgado
Thyronectria Sacc.
Varicosporella Lechat & J. Fourn.
Varicosporellopsis Lechat & J. Fourn.
Volutella Fr.
Xenoacremonium Lombard & Crous
Xenocylindrocladium Decock et al.
Xenogliocladiopsis Crous & W.B. Kendr.
Xenoleptographium Marinc. et al.
Xenonectriella Weese
Niessliaceae Kirschst.
Atronectria Etayo
Circinoniesslia Samuels & M.E. Barr
Cryptoniesslia Scheuer
Eucasphaeria Crous
Hyaloseta A.W. Ramaley
Malmeomyces Starb.
Melchioria Penz. & Sacc.
Miyakeomyces Hara
Myrmaeciella Lindau
Myrtacremonium Crous
Neoeucasphaeria Crous
Niesslia Auersw.
332
Paraniesslia K.M. Tsui et al.
Pseudohyaloseta Tibpromma & K.D. Hyde
Pseudonectriella Petr.
Pseudorhynchia Höhn.
Rosasphaeria Jaklitsch & Voglmayr
Taiwanascus Sivan & H.S. Chang
Trichosphaerella E. Bommer et al.
Valetoniella Höhn.
Valetoniellopsis Samuels & M.E. Barr
Ophiocordycipitaceae G.H. Sung et al.
Drechmeria W. Gams & H.B. Jansson
Harposporium Lohde
Hirsutella Pat.
Hymenostilbe Petch
Ophiocordyceps Petch
Paraisaria Samson & B.L. Brady
Perennicordyceps Matočec & I. Kušan
Polycephalomyces Kobayasi
Purpureocillium Luangsa-ard et al.
Tolypocladium W. Gams
Sarocladiaceae L. Lombard
Parasarocladium Summerb. et al.
Sarocladium W. Gams & D. Hawksw.
Stachybotryaceae L. Lombard & Crous
Achroiostachys L. Lombard & Crous
Albifimbria L. Lombard & Crous
Albosynnema E.F. Morris
Alfaria Crous et al.
Alfariacladiella Crous & R.K. Schumach.
Brevistachys L. Lombard & Crous
Capitofimbria L. Lombard & Crous
Cymostachys L. Lombard & Crous
Didymostilbe Henn.
Digitiseta Gordillo & Decock
Dimorphiseta L. Lombard & Crous
Globobotrys L. Lombard & Crous
Grandibotrys L. Lombard & Crous
Gregatothecium L. Lombard & Crous
Hyalinostachys C.G. Lin & K.D. Hyde
Inaequalispora L. Lombard & Crous
Kastanostachys L. Lombard & Crous
Koorchalomella Chona et al.
Melanopsamma Niessl
Memnoniella Höhn.
Myrothecium Tode
Myxospora L. Lombard & Crous
Neomyrothecium L. Lombard & Crous
Paramyrothecium L. Lombard & Crous
Parasarcopodium Mel’nik et al.
333
Parvothecium L. Lombard & Crous
Peethambara Subram. & Bhat
Pseudoornatispora Tibpromma & K.D. Hyde
Septomyrothecium Matsush.
Sirastachys L. Lombard & Crous
Smaragdiniseta L. Lombard & Crous
Stachybotrys Corda
Striatibotrys L. Lombard & Crous
Striaticonidium L. Lombard & Crous
Tangerinosporium L. Lombard & Crous
Virgatospora Finley
Xenomyrothecium L. Lombard & Crous
Xepicula Nag Raj
Xepiculopsis Nag Raj
Tilachlidiaceae Lombard & Crous
Psychronectria J. Pawłowska et al.
Septofusidium W. Gams
Tilachlidium Preuss
Hypocreales genera incertae sedis
Acremoniopsis Giraldo et al.
Berkelella (Sacc.) Sacc.
Bulbithecium Udagawa & T Muroi
Cephalosporiopsis Peyronel
Chondronectria Etayo et al.
Cylindronectria Etayo
Diploöspora Grove
Gynonectria Döbbeler
Hapsidospora Malloch & Cain
Haptospora G.L. Barron
Illosporiopsis D. Hawksw.
Illosporium Mart.
Leptobarya Etayo
Lichenopenicillus Etayo
Metadothella Henn.
Munkia Speg.
Neomunkia Petr.
Peloronectria Möller
Pseudoacremonium Crous
Pseudoidriella Crous & R.G. Shivas
Pseudomeliola Speg.
Rodentomyces Doveri et al.
Roselliniella Vain
Saksenamyces A.N. Rai & P.N. Singh
Sedecimiella K.L. Pang et al.
Stanjemonium W. Gams et al.
Stilbella Lindau
Ticonectria Döbbeler
Tilakidium Vaidya et al.
Microascales Luttr. ex Benny & Kimbr.
334
Ceratocystidaceae Locq. ex Réblová et al.
Ambrosiella Brader ex Arx & Hennebert
Berkeleyomyces W.J. Nel et al.
Bretziella Z.W. de Beer et al.
Ceratocystis Ellis & Halst.
Chalaropsis Peyronel
Davidsoniella Z.W. de Beer et al.
Endoconidiophora Münch
Huntiella Z.W. de Beer et al.
Meredithiella McNew et al.
Phialophoropsis L.R. Batra emend. T.C. Harr.
Thielaviopsis Went.
Chadefaudiellaceae Faurel & Schotter ex Benny & Kimbr.
Chadefaudiella Faurel & Schotter
Faurelina Locq-Lin.
Gondwanamycetaceae Réblová et al.
Custingophora Stolk
Knoxdaviesia M.J. Wingf et al.
Graphiaceae De Beer
Graphium Corda
Halosphaeriaceae E. Müll & Arx ex Kohlm.
Alisea J. Dupont & E.B.G. Jones
Amphitrite S. Tibell
Aniptodera Shearer & M. Miller
Aniptosporopsis (K.D. Hyde) K.L. Pang
Anisostagma K.R.L. Petersen & Jørg. Koch
Antennospora Meyers
Appendichordella R.G. Johnson et al.
Arenariomyces Höhnk
Bathyascus Kohlm.
Carbosphaerella I. Schmidt
Ceriosporopsis Linder
Chadefaudia Feldm.-Maz.
Corallicola Volkm.-Kohlm. & Kohlm.
Corollospora Werderm
Cucullosporella K.D. Hyde & E.B.G. Jones
Ebullia K.L. Pang
Gesasha Abdel-Wahab & Nagah.
Haiyanga K.L. Pang & E.B.G. Jones
Haligena Kohlm.
Halosarpheia Kohlm. & E. Kohlm.
Halosphaeria Linder
Halosphaeriopsis T.W. Johnson
Havispora K.L. Pang & Vrijmoed
Iwilsoniella E.B.G. Jones
Kitesporella Jheng & K.L. Pang
Kochiella Sakay. et al.
Lautisporopsis E.B.G. Jones et al.
335
Lignincola Höhnk
Limacospora Jørg. Koch & E.B.G. Jones
Luttrellia Shearer
Magnisphaera J. Campb. et al.
Marinospora A.R. Caval.
Moana Kohlm. & Volkm.-Kohlm.
Morakotiella Sakay.
Naïs Kohlm.
Natantispora J. Campb. et al.
Nautosphaeria E.B.G. Jones
Neptunella K.L. Pang & E.B.G. Jones
Nereiospora E.B.G. Jones et al.
Nimbospora Jørg. Koch
Nohea Kohlm. & Volkm.-Kohlm.
Oceanitis Kohlm.
Ocostaspora E.B.G. Jones et al.
Okeanomyces K.L. Pang & E.B.G. Jones
Ondiniella E.B.G. Jones et al.
Ophiodeira Kohlm. & Volkm.-Kohlm.
Paraaniptodera K.L. Pang et al.
Phaeonectriella R.A. Eaton & E.B.G. Jones
Praelongicaulis Jones et al.
Panorbis J. Campb. et al.
Pileomyces K.L. Pang & Jheng
Pseudolignincola Chatmala & E.B.G. Jones
Remispora Linder
Saagaromyces K.L. Pang & E.B.G. Jones
Sablicola E B.G. Jones et al.
Thalassogena Kohlm. & Volkm.-Kohlm.
Thalespora Chatmala & E.B.G. Jones
Tinhaudeus K.L. Pang et al.
Tirispora E.B.G. Jones & Vrijmoed
Toriella Sakay. et al.
Trailia G.K. Sutherl.
Trichomaris Hibbits et al.
Tubakiella Sakay. et al.
Tunicatispora K.D. Hyde
Microascaceae Luttr. ex Malloch
Acaulium Sopp
Brachyconidiellopsis Decock et al.
Canariomyces Arx
Cephalotrichum Link
Doratomyces Corda
Echinobotryum Corda
Enterocarpus Locq.-Lin.
Fairmania Sacc.
Gamsia M. Morelet
Kernia Nieuwl.
Lomentospora Hennebert & B.G. Desai
Lophotrichus R.K. Benj.
Microascus Zukal
336
Parascedosporium Gilgado et al.
Petriella Curzi
Pseudallescheria Negroni & I. Fisch.
Pseudoscopulariopsis M. Sandoval-Denis et al.
Rhinocladium Sacc. & Marchal
Scedosporium Sacc. ex Castell. & Chalm.
Scopulariopsis Bainier
Wardomyces F.T. Brooks & Hansf.
Wardomycopsis Udagawa & Furuya
Yunnania H.Z. Kong
Triadelphiaceae Y.Z. Lu et al.
Synnematotriadelphia Chuaseehar et al.
Triadelphia Shearer & J.L. Crane
Microascales genera incertae sedis
Bisporostilbella Brandsb. & E.F. Morris
Cephalotrichiella Crous
Cornuvesica C.D. Viljoen et al.
Gabarnaudia Samson & W. Gams
Sporendocladia G. Arnaud ex Nag Raj & W.B. Kendr.
Parasympodiellales Hern.-Restr. et al.
Parasympodiellaceae Hern.-Restr. et al.
Parasympodiella Ponnappa
Torpedosporales E.B.G. Jones et al.
Etheirophoraceae Rungjindamai et al.
Etheirophora Kohlm. & Volkm.-Kohlm.
Swampomyces Kohlm. & Volkm.
Juncigenaceae E.B.G. Jones et al.
Elbamycella A. Poli
Fulvocentrum E.B.G. Jones & Abdel-Wahab
Juncigena Kohlm Kohlm. et al.
Khaleijomyces Abdel-Wahab
Marinokulati E.B.G. Jones & K.L. Pang
Torpedosporaceae E.B.G. Jones & K.L. Pang
Torpedospora Meyer
Hypocreomycetidae genera incertae sedis
Campylospora Ranzoni
Dendroclathra Voglmayr & G. Delgado
Subclass Lulworthiomycetidae Dayar. et al.
Koralionastetales Kohlm. et al.
Koralionastetaceae Kohlm. & Volkm.-Kohlm.
Koralionastes Kohlm. & Volkm.-Kohlm.
Pontogeneia Kohlm.
Lulworthiales Kohlm. et al.
337
Lulworthiaceae Kohlm. et al.
Cumulospora I. Schmidt
Halazoon Abdel-Aziz et al.
Haloguignardia A. Cribb & J. Cribb
Hydea K.L. Pang & E.B.G Jones
Kohlmeyeriella E.B.G. Jones et al.
Lindra I. Wilson
Lulwoana Kohlm. et al.
Lulwoidea Kohlm. et al.
Lulworthia G.K. Sutherl
Matsusporium E.B.G. Jones & K.L. Pang
Moleospora Abdel-Wahab et al.
Moromyces Abdel-Wahab et al.
Orbimyces Linder
Rostrupiella Jørg Koch et al.
Sammeyersia S.Y. Guo et al.
Subclass Pisorisporiomycetidae Bundhun et al.
Pisorisporiales Réblová & J. Fourn.
Pisorisporiaceae Réblová & J. Fourn.
Achroceratosphaeria Réblová et al.
Pisorisporium Réblová & J. Fourn.
Subclass Savoryellomycetidae Hongsanan et al.
Conioscyphales Réblová & Seifert
Conioscyphaceae Réblová & Seifert
Conioscypha Höhn.
Fuscosporellales J. Yang et al.
Fuscosporellaceae J. Yang et al.
Bactrodesmiastrum Hol.-Jech.
Fuscosporella J. Yang et al.
Mucispora J. Yang et al.
Parafuscosporella J. Yang et al.
Plagiascoma Réblová & J. Fourn.
Pseudoascotaiwania J. Yang et al.
Pleurotheciales Réblová & Seifert
Pleurotheciaceae Réblová & Seifert
Adelosphaeria Réblová
Anapleurothecium Hern.-Restr. et al.
Helicoascotaiwania Dayar. et al.
Melanotrigonum Réblová
Monotosporella S. Hughes
Phaeoisaria Höhn.
Phragmocephala E.W. Mason & S. Hughes
Pleurotheciella Réblová
Pleurothecium Höhn.
Sterigmatobotrys Oudem.
Savoryellales Boonyuen et al.
Savoryellaceae Jaklitsch & Réblová
338
Ascotaiwania Sivan. & H.S. Chang
Canalisporium Nawawi & Kuthub.
Dermatiosporium Z.L. Luo et al.
Savoryella E.B.G. Jones & R.A. Eaton
Subclass Sordariomycetidae O.E. Erikss & Winka
Boliniales P.F. Cannon
Boliniaceae Rick
Apiocamarops Samuels & J.D. Rogers
Apiorhynchostoma Petr.
Camaropella Lar.N. Vassiljeva
Camarops P. Karst.
Cornipulvina Huhndorf et al.
Endoxyla Fuckel
Mollicamarops Lar.N. Vassiljeva
Neohypodiscus J.D. Rogers et al.
Pseudovalsaria Spooner
Cephalothecales Maharachch. & K.D. Hyde
Cephalothecaceae Höhn.
Albertiniella Kirschst.
Cephalotheca Fuckel
Cryptendoxyla Malloch & Cain
Phialemonium W. Gams & McGinnis
Victoriomyces D Davolos et al.
Chaetosphaeriales Huhndorf et al.
Chaetosphaeriaceae Réblová et al.
Adautomilanezia Gusmão et al.
Anacacumisporium Y.R. Ma & X.G. Zhang
Ascochalara Réblová
Brunneodinemasporium Crous & R.F. Castañeda
Catenularia Grove
Chaetosphaeria Tul. & C. Tul.
Chloridium Link
Codinaea Maire
Conicomyces R.C. Sinclair et al.
Craspedodidymum Hol.-Jech.
Cryptophiale Piroz.
Cryptophialoidea Kuthub. & Nawawi
Dendrophoma Sacc.
Dictyochaeta Speg.
Dictyochaetopsis Aramb. & Cabello
Dinemasporium Lév.
Eucalyptostroma Crous & M.J. Wingf.
Exserticlava S. Hughes
Hemicorynespora M.B. Ellis
Infundibulomyces Plaingam et al.
Kionochaeta P.M. Kirk & B. Sutton
Lecythothecium Réblová & Winka
Menispora Pers.
Menisporopsis S. Hughes
339
Miyoshiella Kawam.
Morrisiella Saikia & A.K. Sarbhoy
Nawawia Marvanová
Neopseudolachnella A. Hashim. & Kaz. Tanaka
Paliphora Sivan. & B. Sutton
Phialosporostilbe Mercado & J. Mena
Polynema Lév.
Pseudodinemasporium A. Hashim. & Kaz. Tanaka
Pseudolachnea Ranoj.
Pseudolachnella Teng
Pyrigemmula D. Magyar & R. Shoemaker
Rattania Prabhug. & Bhat
Sporoschisma Berk. & Broome
Striatosphaeria Samuels & E. Müll.
Tainosphaeria F.A. Fernández & Huhndorf
Thozetella Kuntze
Umbrinosphaeria Réblová
Verhulstia Hern.-Rest.
Zanclospora S. Hughes & W.B. Kendr.
Helminthosphaeriaceae Samuels et al.
Echinosphaeria A.N. Mill. & Huhndorf
Endophragmiella B. Sutton
Helminthosphaeria Fuckel
Hilberina Huhndorf & A.N. Mill.
Ruzenia O. Hilber
Synaptospora Cain
Tengiomyces Réblová
Leptosporellaceae Konta & K.D. Hyde
Leptosporella Penz. & Sacc.
Linocarpaceae Konta & K.D. Hyde
Linocarpon Syd. & P. Syd.
Neolinocarpon K.D. Hyde
Chaetosphaeriales genera incertae sedis
Calvolachnella Marinc. et al.
Caudatispora J. Fröhl. & K.D. Hyde
Erythromada Huhndorf et al.
Lasiosphaeriella Sivan.
Neoleptosporella Phukhums. et al.
Rimaconus Huhndorf et al.
Coniochaetales Huhndorf et al.
Coniochaetaceae Malloch & Cain
Barrina A.W. Ramaley
Coniochaeta (Sacc.) Cooke
Cordanaceae Nann.
Cordana Preuss
340
Coniochaetales genera incertae sedis
Cannonia J.E. Taylor & K.D. Hyde
Pseudogliomastix W. Gams
Meliolales Gäum. ex D. Hawksw. & O.E. Erikss.
Armatellaceae Hosag.
Armatella Theiss. & Syd.
Meliolaceae G.W. Martin ex Hansf.
Amazonia Theiss.
Appendiculella Höhn.
Asteridiella McAlpine
Cryptomeliola S. Hughes & Piroz.
Endomeliola S. Hughes & Piroz.
Irenopsis F. Stevens
Meliola Fr.
Setameliola D.R. Reynolds
Phyllachorales M.E. Barr
Phaeochoraceae K.D. Hyde et al.
Cocoicola K.D. Hyde
Phaeochora Höhn.
Phaeochoropsis K.D. Hyde & P.F. Cannon
Serenomyces Petr.
Phyllachoraceae Theiss. & H. Syd.
Ascovaginospora Fallah et al.
Brobdingnagia K.D. Hyde & P.F. Cannon
Camarotella Theiss. & Syd.
Coccodiella Hara
Cyclodomus Höhn.
Deshpandiella Kamat & Ullasa
Diachora Müll. Arg.
Diatractium Syd. & P. Syd.
Erikssonia Penz. & Sacc.
Fremitomyces P.F. Cannon & H.C. Evans
Geminispora Pat.
Gibellina Pass. Ex Roum.
Imazekia Tak. Kobay. & Y. Kawabe
Isothea Fr.
Lichenochora Hafellner
Lindauella Rehm
Linochora Höhn.
Lohwagia Petr.
Maculatifrondes K.D. Hyde
Malthomyces K.D. Hyde & P.F. Cannon
Muelleromyces Kamat & Anahosur
Neoflageoletia J. Reid & C. Booth
Neophyllachora Dayar. & K.D. Hyde
Ophiodothis Sacc.
Ophiodothella (Henn.). Höhn.
Orphnodactylis Malloch & Mallik
341
Oxodeora K.D. Hyde & P.F. Cannon
Parberya C.A. Pearce & K.D. Hyde
Petrakiella Syd.
Phycomelaina Kohlm.
Phyllachora Nitschke ex Fuckel
Phylleutypa Petr.
Phyllocrea Höhn.
Pseudothiella Petr.
Pseudothiopsella Petr.
Pterosporidium W.H. Ho & K.D. Hyde
Rehmiodothis Theiss. & Syd.
Retroa P.F. Cannon
Rhodosticta Woron.
Rikatlia P.F. Cannon
Schizochora Syd. & P. Syd.
Sphaerodothella C.A. Pearce & K.D. Hyde
Sphaerodothis (Sacc. & P. Syd.) Shear
Stigmatula (Sacc.) Syd. & P. Syd.
Stigmochora Theiss. & Syd.
Stromaster Höhn.
Tamsiniella S.W. Wong et al.
Telimenella Petr.
Telimenochora Sivan.
Trabutia Sacc. & Roum.
Tribulatia J.E. Taylor et al.
Uropolystigma Maubl.
Vitreostroma P.F. Cannon
Zimmermanniella Henn.
Telimenaceae Mardones et al.
Telimena Racib.
Phyllachorales genus incertae sedis
Marinosphaera K.D. Hyde
Pseudodactylariales Crous
Pseudodactylariaceae Crous
Pseudodactylaria Crous
Sordariales Chad. ex D. Hawksw. & O.E. Erikss.
Chaetomiaceae G. Winter
Achaetomium J.N. Rai et al.
Acrophialophora Edward
Allobotryotrichum M. Raza & L. Cai
Amesia X. Wei Wang et al.
Arcopilus X. Wei Wang et al.
Arxotrichum A. Nováková & M. Kolařik
Botryotrichum Sacc. & Marchal
Chaetomium Kunze
Collariella X. Wei Wang et al.
Corynascella Arx & Hodges
Crassicarpon Y. Marín et al.
342
Dichotomopilus X. Wei Wang et al.
Guanomyces M.C. Gonzáles et al.
Humicola Traaen
Madurella Brumpt
Melanocarpus Arx
Myceliophthora Costantin
Mycothermus D.O. Natvig et al.
Ovatospora X. Wei Wang et al.
Remersonia Samson & Seifert
Staphylotrichum J.A. Mey. & Nicot
Subramaniula Arx
Thermothelomyces Y. Marín et al.
Thielavia Zopf
Trichocladium Harz
Lasiosphaeriaceae Nannf.
Anopodium Lundq.
Apiosordaria Arx & W. Gams
Apodospora Cain & J.H. Mirza
Apodus Malloch & Cain
Arnium Nitschke ex G. Winter
Bellojisia Réblová
Biconiosporella Schaumann
Bombardia (Fr.) P. Karst.
Bombardioidea C. Moreau ex N. Lundqv.
Camptosphaeria Fuckel
Cercophora Fuckel
Corylomyces Stchigel et al.
Diffractella Guarro et al.
Diplogelasinospora Cain
Emblemospora Jeng & J.C. Krug
Eosphaeria Höhn.
Episternus Górz & Boroń
Fimetariella N. Lundq.
Immersiella A.N. Mill. & Huhndorf
Jugulospora N. Lundq.
Lasiosphaeria Ces. & De Not.
Mammaria Ces. ex Rabenh.
Periamphispora J.C. Krug
Ramophialophora M. Calduch et al.
Rinaldiella Deanna A. Sutton et al.
Schizothecium Corda
Strattonia Cif.
Thaxteria Sacc.
Tripterosporella Subram. & Lodha
Zopfiella G. Winter
Zygopleurage Boedijn
Zygospermella Cain
Podosporaceae X. Wei Wang & Houbraken
Cladorrhinum Sacc. & Marchal
Podospora Ces.
343
Triangularia Boedijn
Sordariaceae G. Winter
Copromyces N. Lundq.
Effetia Bartoli et al.
Guilliermondia Boud.
Neurospora Shear & B.O. Dodge
Pseudoneurospora Dania García et al.
Sordaria Ces. & De Not.
Stellatospora T. Ito & A. Nakagiri
Sordariales genera incertae sedis
Abyssomyces Kohlm
Acanthotheciella Höhn.
Ascolacicola Ranghoo & K.D. Hyde
Bombardiella Höhn.
Coronatomyces Dania García et al.
Cuspidatispora Shearer & Bartolata
Globosphaeria D. Hawksw.
Isia D. Hawksw & Manohar
Lasiosphaeris Clem.
Lunulospora Ingold
Lockerbia K.D. Hyde
Nitschkiopsis Nannf. & R. Sant.
Onygenopsis Henn.
Phaeosporis Clem.
Reconditella Matzer & Hafellner
Rhexodenticula W.A. Baker & Morgan-Jones
Rhexosporium Udagawa & Furuya
Roselliniomyces Matzer & Hafellner
Roselliniopsis Matzer & Hafellner
Stromatographium Höhn.
Utriascus Réblová
Ypsilonia Lév.
Sordariomycetidae families incertae sedis
Batistiaceae Samuels & K.F. Rodrigues
Batistia Cif.
Sordariomycetidae genera incertae sedis
Arecacicola Joanne E. Taylor et al.
Bullimyces A. Ferrer et al.
Cancellidium Tubaki
Ceratolenta Réblová
Chaetosphaerides Matsush.
Cryptophyllachora L. Kiss et al.
Hanliniomyces Raja & Shearer
Hydromelitis A. Ferrer et al.
Merugia Rogerson & Samuels
Mycomedusiospora G.C. Carroll & Munk
Myxocephala G. Weber et al.
Nigromammilla K.D. Hyde & J. Fröhl.
344
Phaeotrichosphaeria Sivan.
Phragmodiscus Hansf.
Plagiosphaera Petr.
Pseudobotrytis Krzemien. & Badura
Subclass Xylariomycetidae O.E. Erikss & Winka
Amphisphaeriales D. Hawksw. & O.E. Erikss.
Amphisphaeriaceae G. Winter
Amphisphaeria Ces. & De Not.
Griphosphaerioma Höhn.
Lepteutypa Petr.
Apiosporaceae K.D. Hyde et al.
Appendicospora K.D. Hyde
Arthrinium Kunze
Dictyoarthrinium S. Hughes
Endocalyx Berk. & Broome
Nigrospora Zimm.
Beltraniaceae Nann.
Beltrania Penz.
Beltraniella Subram.
Beltraniopsis Bat. & J.L. Bezerra
Hemibeltrania Piroz.
Parapleurotheciopsis P.M. Kirk
Porobeltraniella Gusmão
Pseudobeltrania Henn.
Pseudosubramaniomyces Crous
Subsessila C.G. Lin & K.D. Hyde
Castanediellaceae Hern.-Restr. et al.
Castanediella Hern.-Restr. et al.
Clypeophysalosporaceae Giraldo & Crous
Bagadiella Cheew. & Crous
Clypeophysalospora H.J. Swart
Neophysalospora Crous & M.J. Wingf.
Plectosphaera Theiss.
Cylindriaceae Crous & L. Lombard
Cylindrium Bonord
Hyponectriaceae Petr.
Apiothyrium Petr.
Arecomyces K.D. Hyde
Arwidssonia B. Erikss.
Cesatiella Sacc.
Chamaeascus L. Holm et al.
Discosphaerina Höhn.
Exarmidium P. Karst.
Frondicola K.D. Hyde
Hyponectria Sacc.
345
Lichenoverruculina Etayo
Micronectria Speg.
Papilionovela Aptroot
Pellucida Dulym. et al.
Phragmitensis M.K.M. Wong et al.
Physalospora Niessl
Rachidicola K.D. Hyde & J. Fröhl.
Xenothecium Höhn.
Iodosphaeriaceae O. Hilber
Iodosphaeria Samuels
Melogrammataceae G. Winter
Melogramma Fr.
Oxydothidaceae Konta & K.D. Hyde
Oxydothis Penz. & Sacc.
Phlogicylindriaceae Senan. & K.D. Hyde
Ciferriascosea Senan. et al.
Idriellomyces Crous
Phlogicylindrium Crous et al.
Pseudomassariaceae Senan. & K.D. Hyde
Leiosphaerella Höhn.
Pseudapiospora Petr.
Pseudomassaria Jacz.
Pseudomassariella Petr.
Pseudosporidesmiaceae Crous
Pseudosporidesmium K.D. Hyde & McKenzie
Pseudotruncatellaceae Crous
Pseudotruncatella R.H. Perera et al.
Sporocadaceae Corda
Allelochaeta Petr.
Annellolacinia B. Sutton
Bartalinia Tassi
Broomella Sacc.
Ciliochorella Syd.
Diploceras (Sacc.) Died.
Disaeta Bonar
Discosia Lib.
Distononappendiculata F. Liu et al.
Diversimediispora F. Liu et al.
Doliomyces Steyaert
Heterotruncatella F. Liu et al.
Hyalotiella Papendorf
Hymenopleella Munk
Immersidiscosia Kaz. Tanaka et al.
Monochaetia (Sacc.) Allesch.
346
Morinia Berl. & Bres.
Neopestalotiopsis Maharachch. et al.
Nonappendiculata F. Liu et al.
Parabartalinia F. Liu et al.
Pestalotiopsis Steyaert
Pseudopestalotiopsis Maharachch. et al.
Pseudosarcostroma F. Liu et al.
Robillarda Sacc.
Sarcostroma Cooke
Seimatosporium Corda
Seiridium Nees
Sporocadus Corda
Strickeria Körb.
Synnemapestaloides T. Handa & Y. Harada
Truncatella Steyaert
Xenoseimatosporium F. Liu et al.
Vialaeaceae P.F. Cannon
Vialaea Sacc.
Xyladictyochaetaceae Crous & Hern.-Restr
Xyladictyochaeta Hern.-Restr. et al.
Amphisphaeriales genera incertae sedis
Chitonospora E. Bommer et al.
Fasciatispora K.D. Hyde
Delonicicolales R.H. Perera et al.
Delonicicolaceae R.H. Perera et al.
Delonicicola R.H. Perera et al.
Furfurella Voglmayr & Jaklitsch
Leptosilliaceae Voglmayr & Jaklitsch
Leptosillia Höhn.
Xylariales Nannf.
Barrmaeliaceae Voglmayr & Jaklitsch
Barrmaelia Rappaz
Entosordaria (Sacc.) Höhn.
Cainiaceae J.C. Krug
Alishanica Karun. et al.
Amphibambusa D.Q. Dai & K.D. Hyde
Arecophila K.D. Hyde
Atrotorquata Kohlm. & Volkm.-Kohlm.
Cainia Arx & E. Müll.
Seynesia Sacc.
Clypeosphaeriaceae G. Winter
Aquasphaeria K.D. Hyde
Apioclypea K.D. Hyde
Brunneiapiospora K.D. Hyde et al.
Clypeosphaeria Fuckel
347
Crassoascus Checa et al.
Palmaria K.D. Hyde et al.
Coniocessiaceae Asgari & Zare
Coniocessia Dania García et al.
Paraxylaria Wanas. et al.
Diatrypaceae Nitschke
Allocryptovalsa Senwanna et al.
Anthostoma Nitschke
Cryptosphaeria Ces & De Not.
Cryptovalsa Ces. & De Not. ex Fuckel
Diatrypasimilis J.J. Zhou & Kohlm.
Diatrype Fr.
Diatrypella (Ces. & De Not.) De Not.
Echinomyces Rappaz
Endoxylina Romell
Eutypa Tul. & C. Tul.
Eutypella (Nitschke) Sacc.
Halocryptovalsa Dayar. & K.D. Hyde
Halodiatrype Dayar. & K.D. Hyde
Leptoperidia Rappaz
Libertella Desm.
Monosporascus Pollack & Uecker
Neoeutypella M. Raza et al.
Pedumispora K.D. Hyde & E.B.G. Jones
Peroneutypa Berl.
Quaternaria Tul. & C. Tul.
Graphostromataceae M.E. Barr et al.
Biscogniauxia Kuntze
Camillea Fr.
Graphostroma Piroz.
Obolarina Pouzar
Vivantia J.D. Rogers et al.
Hansfordiaceae Crous
Hansfordia S. Hughes
Hypoxylaceae DC.
Annulohypoxylon Y.M. Ju et al.
Anthocanalis Daranag. et al.
Chlorostroma A.N. Mill. et al.
Daldinia Ces. & De Not.
Durotheca Læssøe et al.
Entonaema Möller
Hypomontagnella Sir et al.
Hypoxylon Bull.
Jackrogersella L. Wendt et al.
Natonodosa Heredia et al.
Phylacia Lév.
Pyrenomyxa Morgan
348
Pyrenopolyporus Lloyd
Rhopalostroma D. Hawksw.
Rostrohypoxylon J. Fourn. & M. Stadler
Ruwenzoria J. Fourn. et al.
Thamnomyces Ehrenb.
Theissenia Maubl.
Thuemenella Penz. & Sacc.
Induratiaceae Samarak et al.
Induratia Samuels et al.
Emarcea Duong et al.
Lopadostomataceae Daranag. & K.D. Hyde
Creosphaeria Theiss.
Jumillera J.D. Rogers et al.
Lopadostoma (Nitschke) Traverso
Whalleya J.D. Rogers et al.
Microdochiaceae Hern.-Restr. et al.
Idriella P.E. Nelson & S. Wilh.
Microdochium Syd.
Selenodriella R.F. Castañeda & W.B. Kendr.
Polystigmataceae Höhn. ex Nannf.
Polystigma DC.
Requienellaceae Boise
Acrocordiella O.E. Erikss.
Lacrymospora Aptroot
Parapyrenis Aptroot
Requienella Fabre
Xylariaceae Tul. & C. Tul.
Abieticola Hyang B. Lee
Amphirosellinia Y.M. Ju et al.
Anthostomella Sacc.
Anthostomelloides Tibpromma & K.D. Hyde
Ascotricha Berk.
Astrocystis Berk. & Broome
Brunneiperidium Daranag. et al.
Collodiscula I. Hino & Katum.
Coniolariella Dania García et al.
Engleromyces Henn.
Entalbostroma J.D. Rogers & P.R. Johnst.
Entoleuca Syd.
Euepixylon Füisting
Halorosellinia Whalley et al.
Helicogermslita Lodha & D. Hawksw.
Hypocopra (Fr) J. Kickx f .
Hypocreodendron Henn.
Kretzschmaria Fr.
Kretzschmariella Viégas
349
Leprieuria Laessøe et al.
Lunatiannulus Daranag. et al.
Nemania Gray
Podosordaria Ellis & Holw.
Poronia Willd.
Rosellinia De Not.
Sarcoxylon Cooke
Squamotubera Henn.
Stilbohypoxylon Henn.
Vamsapriya Gawas & Bhat
Virgaria Nees
Wawelia Namysl.
Xylaria Hill ex Schrank
Zygosporiaceae Li et al.
Zygosporium Mont.
Xylariales genera incertae sedis
Adomia S. Schatz
Alloanthostomella Daranag. et al.
Anungitea B. Sutton
Ascotrichella Valldos. & Guarro
Basifimbria Subram. & Lodha
Biporispora J.D. Rogers et al.
Castellaniomyces Senan. et al.
Chaenocarpus Rebent.
Circinotrichum Nees
Cryptostroma P.H. Greg. & S. Waller
Cyanopulvis J. Fröhl. & K.D. Hyde
Diamantinia A.N. Mill. et al.
Gigantospora B.S. Lu & K.D. Hyde
Guestia G.J.D. Sm. & K.D. Hyde
Gyrothrix (Corda) Corda
Hadrotrichum Fuckel
Idriellopsis Hern.-Restr. & Crous
Kirstenboschia Quaedvl. et al.
Lanceispora Nakagiri et al.
Lasiobertia Sivan.
Leptomassaria Petr.
Neoanthostomella D.Q. Dai & K.D. Hyde
Neoidriella Hern.-Restr. & Crous
Nipicola K.D. Hyde
Occultitheca J.D. Rogers & Y.M. Ju
Ophiorosellinia J.D. Rogers et al.
Palmicola K.D. Hyde
Pandanicola K.D. Hyde
Paraidriella Hern.-Restr. & Crous
Paramphisphaeria F.A. Fernández et al.
Paraphysalospora Crous
Paucithecium Lloyd
Pidoplitchkoviella Kiril.
Polyancora Voglmayr & Yule
350
Polyscytalum Riess
Poroleprieuria M.C. González et al.
Pseudoanthostomella Daranag. et al.
Pseudophloeospora Crous & R.G. Shivas
Pulmosphaeria Joanne E. Taylor et al.
Pyriformiascoma Daranag. et al.
Roselymyces Fiuza et al.
Sabalicola K.D. Hyde
Spirodecospora B.S. Lu et al.
Sporidesmina Subram. & Bhat
Striatodecospora D.Q. Zhou et al.
Stromatoneurospora S.C. Jong & E.E. Davis
Surculiseries Okane
Synnemadiella Crous & M.J. Wingf.
Tristratiperidium Daranag. et al.
Xylocrea Möller
Xylotumulus J.D. Rogers et al.
Yuea O.E. Erikss.
Xylariomycetidae families incertae sedis
Myelospermataceae K.D. Hyde & S.W. Wong
Myelosperma Syd. & P. Syd.
Xylariomycetidae genera incertae sedis
Calceomyces Udagawa & S. Ueda
Sordariomycetes orders incertae sedis
Amplistromatales D’souza et al.
Amplistromataceae Huhndorf et al.
Acidothrix Hujslová & M. Kolařík
Amplistroma Huhndorf et al.
Wallrothiella Sacc.
Catabotryales K.D. Hyde & Senan.
Catabotryaceae Petr. ex M.E. Barr
Catabotrys Theiss. & Syd.
Spathulosporales Kohlm.
Hispidicarpomycetaceae Nakagiri
Hispidicarpomyces Nakagiri
Spathulosporaceae Kohlm.
Retrostium Nakagiri & Tad Ito
Spathulospora A.R. Caval. & T.W. Johnson
Tracyllalales Crous
Tracyllaceae Crous
Tracylla (Sacc.) Tassi
Vermiculariopsiellales Hern.-Restr. et al.
Vermiculariopsiellaceae Hern.-Restr. et al.
Vermiculariopsiella Bender
351
Sordariomycetes families incertae sedis
Acrodictyaceae J.W. Xia & X.G. Zhang
Acrodictys M.B. Ellis
Junewangiaceae J.W. Xia & X.G. Zhang
Dictyosporella Abdel-Aziz
Junewangia W.A. Baker & Morgan-Jones
Lautosporaceae Kohlm. et al.
Lautospora K.D. Hyde & E.B.G. Jones
Obryzaceae Körb.
Obryzum Wallr.
Sordariomycetes genera incertae sedis
Acerbiella Sacc.
Acrospermoides Miller & G.E. Thomps.
Ameromassaria Hara
Amphisphaerellula Gucevič
Amphisphaerina Höhn.
Amphorulopsis Petr.
Amylis Speg.
Anisomycopsis I. Hino & Katum.
Anthostomaria (Sacc.) Theiss. & Syd.
Anthostomellina L.A. Kantsch.
Apodothina Petr.
Apogaeumannomyces Matsush.
Aquadulciospora Fallah & Shearer
Areolospora S.C. Jong & E.E. Davis
Aropsiclus Kohlm. & Volkm.-Kohlm.
Ascorhiza Lecht.-Trinka
Ascoyunnania L. Cai & K.D. Hyde
Atrogeniculata J.S. Monteiro et al.
Aulospora Speg.
Azbukinia Lar.N. Vassiljeva
Bactrosphaeria Penz. & Sacc.
Basidiobotrys Höhn.
Biciliopsis Diederich
Bombardiastrum Pat.
Botryosporium Corda
Brenesiella Syd.
Byrsomyces Cavalc.
Byssotheciella Petr.
Caleutypa Petr.
Caproniella Berl.
Chaetoamphisphaeria Hara
Ciliofusospora Bat. & J.L. Bezerra
Clypeoceriospora Sousa da Câmara
Clypeosphaerulina Sousa da Câmara
Cryptoascus Petri
Cryptomycella Höhn.
352
Cryptomycina Höhn.
Cucurbitopsis Bat. & Cif.
Curvatispora V.V. Sarma & K.D. Hyde
Dasysphaeria Speg.
Delpinoëlla Sacc.
Diacrochordon Petr.
Didymobotryum Sacc.
Duradens Samuels & Rogerson
Ellisembia Subram.
Esfandiariomyces Ershad
Fantasmomyces D.Hyeon Lee et al.
Fassia Dennis
Flammispora Pinruan et al.
Frondisphaeria K.D. Hyde
Hapsidascus Kohlm. & Volkm.-Kohlm.
Heliastrum Petr.
Hyaloderma Speg.
Hyalotiopsis Punith.
Hydronectria Kirschst.
Immersisphaeria Jaklitsch
Iraniella Petr.
Konenia Hara
Kravtzevia Schwartzman
Kurssanovia Kravtzev
Lecythiomyces Doweld
Leptosacca Syd.
Leptosphaerella Speg.
Mangrovispora K.D. Hyde & Nakagiri
Marisolaris Jørg. Koch & E.B.G. Jones
Melanographium Sacc.
Microcyclephaeria Bat.
Mirannulata Huhndorf et al.
Neonawawia J. Yang et al.
Natantiella Réblová
Naumovela Kravtzev
Neocryptospora Petr.
Neoeriomycopsis Crous & M.J. Wingf.
Neolamya Theiss. & Syd.
Neothyridaria Petr.
Ophiomassaria Jacz.
Ophiomeliola Starbäck
Paoayensis Cabanela et al.
Paradiplococcium Hern.-Restr. et al.
Paramicrodochium Hern.-Restr. & Crous
Pareutypella Y.M. Ju & J.D. Rogers
Phialemoniopsis Perdomo et al.
Phragmeriella Hansf.
Phyllocelis Syd.
Pleocryptospora J. Reid & C. Booth
Pleosphaeria Speg.
Pleurophragmium Costantin
Protocucurbitaria Naumov
353
Pulvinaria Bon.
Pumilus Viala & Marsais
Rehmiomycella E. Müll.
Rhamphosphaeria Kirschst.
Rhizophila K.D. Hyde & E.B.G. Jones
Rhopographella (Henn.) Sacc. & Trotter
Rhynchosphaeria (Sacc.) Berl.
Rivulicola K.D. Hyde
Romellina Petr.
Saccardoëlla Speg.
Sartorya Vuill.
Scharifia Petr.
Scoliocarpon Nyl.
Scotiosphaeria Sivan.
Selenosporella G. Arnaud ex MacGarvie
Servazziella J. Reid & C. Booth
Sporoctomorpha J.V. Almeida & Sousa da Câmara
Stanjehughesia Subram.
Stearophora L. Mangin & Viala
Steganopycnis Syd. & P. Syd.
Stegophorella Petr.
Stellosetifera Matsush.
Stereosphaeria Kirschst.
Stomatogenella Petr.
Sungaiicola Fryar & K.D. Hyde
Synsphaeria Bon.
Teracosphaeria Réblová & Seifert
Thelidiella Fink
Thyridella (Sacc.) Sacc.
Thyrotheca Kirschst.
Trichospermella Speg.
Trichosphaeropsis Bat. & Nasc.
Tunstallia Agnihothr.
Tulipispora Révay & Gönczöl
Urosporella G.F. Atk.
Urupe Viégas
Vleugelia J. Reid & C. Booth
Xenodium Syd.
Zalerion R.T. Moore & Meyers
Results
The combined LSU, SSU, tef1 and rpb2 gene data set comprised 298 taxa, with Botryotinia
fuckeliana (AFTOL ID-59), Dothidea sambuci (DAOM 231303), and Pyxidiophora arvernensis
(AFTOL-ID 2197) as outgroup taxa. The combined dataset comprised 4303 characters including
gaps. The best scoring RAxML tree is shown in Fig 1. In the phylogenetic tree, the
Sordariomycetes strains included in the analysis clustered into seven subclasses including
Sordariomycetidae, Hypocreomycetidae and Xylariomycetidae as in the previous treatment of
Lumbsch & Huhndorf (2010); Diaporthomycetidae and Lulworthiomycetidae as suggested by
Maharachchikumbura et al. (2015), as well as Savoryellomycetidae suggested by Hongsanan et al.
(2017) and the addition of a new subclass, the Pisorisporiomycetidae in this paper. The seven
subclasses separate into 45 orders including the newly introduced Cephalothecales. Furthermore,
sequence data of 153 families are included in the phylogenetic analysis.
354
The subclasses
There are seven subclasses accepted in Sordariomycetes with the addition of
Pisorisporiomycetidae in this paper.
Diaporthomycetidae Senan., Maharachch. & K.D. Hyde, Fungal Divers. 72: 208 (2015)
The subclass Diaporthomycetidae was introduced by Maharachchikumbura et al. (2015) for
some taxa already placed in Sordariomycetidae, but that were phylogenetically and
morphologically distinct from Sordariomycetidae. Members of Diaporthomycetidae occur in both
aquatic and terrestrial habitats as saprobes, pathogens, or endophytes. Previously there were ten
orders in this subclass (Hongsanan et al. 2017). Crous et al. (2017a) introduced
Pararamichloridiales and Crous et al. (2019a) introduced Sporidesmiales. Hyde et al. (2017a)
proposed Catabotryales based on evolutionary data and here we formally introduce it. Currently
there are 15 orders and 65 families in this subclass (Hyde et al. 2017a, this paper). The divergence
time for Diaporthomycetidae is estimated as 247 MYA (Fig. 2). The orders and families in this
subclass are mostly well-supported in our phylogenetic analysis (Figs 6, 8, 13, 14, 18).
Hypocreomycetidae O.E. Erikss. & Winka, Myconet 1(1): 6 (1997)
Maharachchikumbura et al. (2016b) accepted Conioscyphales, Coronophorales,
Falcocladiales, Glomerellales, Hypocreales, Melanosporales, Microascales, Pleurotheciales and
Savoryellales in Hypocreomycetidae. Yang et al. (2016b) introduced Fuscosporellales to the
subclass based on LSU, SSU and rpb2 sequence data. Hongsanan et al. (2017) placed
Fuscosporellales and Pleurotheciales in the newly introduced subclass Savoryellomycetidae. The
families Etheirophoraceae, Juncigenaceae and Torpedosporaceae have previously been treated in
Torpedosporales by Jones et al. (2015), but Torpedosporales was maintained in
Hypocreomycetidae, incertae sedis by Maharachchikumbura et al. (2015). Maharachchikumbura et
al. (2016b) accepted Torpedosporales in Hypocreomycetidae and this was confirmed by Hongsanan
et al. (2017) based on the divergent time estimates. Later, Parasympodiellales was added to the
subclass by Hernández-Restrepo et al. (2017). Our phylogenetic analysis of a combined LSU, SSU,
tef1, rpb2 sequence data (Figs. 1, 24) also supports the placement of Torpedosporales in
Hypocreomycetidae. Currently there are seven orders (Coronophorales, Falcocladiales,
Glomerellales, Hypocreales, Microascales, Parasympodiellales and Torpedosporales) and 37
families in this subclass (Hyde et al. 2017a, this paper). The divergence time for
Hypocreomycetidae has been estimated as 256.5 MYA (Fig. 2). The orders in this subclass are
mostly well-supported in our analysis, except Torpedosporales (Fig. 1).
Lulworthiomycetidae Dayar., E.B.G. Jones & K.D. Hyde, Fungal Divers. 72: 208 (2015)
Maharachchkumbura et al. (2016b) established the subclass Lulworthiomycetidae to
accommodate Koralionastetales, Lulworthiales and Pisorisporiales as proposed by
Maharachchikumbura et al. (2015). Taxa related to Lulworthiomycetidae are saprobic on wood, sea
grasses and marsh plants or parasites on algae. The divergence time for Lulworthiomycetidae has
been estimated as 257 MYA (Fig. 2). Currently there are two orders and two families in this
subclass and Pisorisporiales is transferred to the introduced subclass Pisorisporiomycetidae in this
study.
Pisorisporiomycetidae Bundhun, Maharachch. & K.D. Hyde, subclass nov.
Index Fungorum number: IF556880; Facesoffungi number: FoF06688
Saprobic on submerged wood or driftwood. Sexual morph: Ascomata astromatic, perithecial,
solitary or aggregated in small groups, immersed, semi-immersed to superficial, ostiolate. Ostiole
periphysate. Peridium 2-layered, leathery to fragile, partly carbonaceous. Paraphyses abundant,-
355
Figure 1 – Maximum likelihood (ML) majority rule combined LSU, SSU, tef1 and rpb2 consensus
tree for the analyzed Sordariomycetes isolates. Families are indicated in yellow and green coloured
blocks and orders are indicated in dark and light grey coloured blocks. RAxML bootstrap support
values (MLB above 50 %) are given at the nodes. The scale bar represents the expected number of
changes per site. The tree is rooted with Botryotinia fuckeliana (AFTOL ID-59), Dothidea sambuci
(DAOM 231303), and Pyxidiophora arvernensis (AFTOL-ID 2197).
356
Figure 1 – Continued.
357
Figure 1 – Continued.
-hyaline, persistent. Asci 8-spored, unitunicate, pedicellate, persistent, with a J+ or J-, apical ring.
Ascospores hyaline, multi-septate, often guttulate, lacking any mucilaginous sheath or appendages.
Asexual morph: Undetermined.
Type order – Pisorisporiales Réblová & J. Fourn.
Notes – Pisorisporiales was reported to form a sister clade with Lulworthiales and
Koralionastetales in Lulworthiomycetidae with a stem age of 266 MYA (Hongsanan et al. 2017,
Hyde et al. 2017a). This is congruent to the results obtained in the present study (257 MYA). Since
this stem age falls within the subclass status range (250–300 MYA), Pisorisporiomycetidae is
proposed here as a new subclass. This subclass contains Pisorisporiales, family Pisorisporiaceae
and the two genera Achroceratosphaeria and Pisorisporium (Réblová et al. 2015a).
358
Figure 1 – Continued.
359
Divergence times of lineages were used to analyze the status of higher ranks of fungi
(Hongsanan et al. 2017, 2018, Hyde et al. 2017, Liu et al. 2017). Hyde et al. (2017) provided a
maximum clade credibility (MCC) tree of families in Sordariomycetes and also proposed a series of
evolutionary periods that could be used as a guide to define ranking of fungi in Sordariomycetes. In
our paper, we provide the MCC tree base on updated classification of Sordariomycetes (Fig. 2).
The same dataset as in Fig. 1 was used in our molecular clock analysis to compare the phylogenetic
placement generated from both approaches. We use divergence time to confirm familial status of
families in Sordariomycetes according to the guidance of Hyde et al. (2017). Some families and
orders are not supported by divergence times, but their status is retained due to their unique
characters or lack of species (notes are provided in each family).
Figure 2 – The maximum clade credibility (MCC) tree, using the same dataset from Fig. 1. This
analysis was performed in BEAST v1.10.2. The crown age of Sordariomycetes was set with
Normal distribution, mean = 250, SD = 30, with 97.5% of CI = 308.8 MYA, and crown age of
Dothideomycetes with Normal distribution mean = 360, SD = 20, with 97.5% of CI = 399 MYA.
The substitution models were selected based on jModeltest2.1.1; GTR+I+G for LSU, rpb2 and
SSU, and TrN+I+G for tef1 (the model TrN is not available in BEAUti 1.10.2, thus we used TN93).
Lognormal distribution of rates was used during the analyses with uncorrelated relaxed clock
model. The Yule process tree prior was used to model the speciation of nodes in the topology with
a randomly generated starting tree. The analyses were performed for 100 million generations, with
sampling parameters every 10000 generations. The effective sample sizes were checked in Tracer
v.1.6 and the acceptable values are higher than 200. The first 20% representing the burn-in phase
were discarded and the remaining trees were combined in LogCombiner 1.10.2., summarized data
and estimated in TreeAnnotator 1.10.2. Bars correspond to the 95% highest posterior density
(HPD) intervals. The scale axis shows divergence times as millions of years ago (MYA).
360
Figure 2 – Continued.
361
Figure 2 – Continued.
362
Savoryellomycetidae Hongsanan, K.D. Hyde & Maharachch., Fungal Divers. 84: 35 (2017)
According to Maharachchikumbura et al. (2016b) and Yang et al. (2016b) Conioscyphales
was assigned to Diaporthomycetidae, genera incertae sedis, while Fuscosporellales,
Pleurotheciales, and Savoryellales were included in the subclass Hypocreomycetidae. In the
phylogenetic and molecular clock analyses of (Hongsanan et al. 2017, Hyde et al. 2017a),
Conioscyphales, Fuscosporellales, Pleurotheciales, and Savoryellales clustered together as a
distinct clade, with a stem age of 268 MYA. Hence, they were referred to a new subclass
Savoryellomycetidae by Hongsanan et al. (2017) and this was reinforced in the paper by
Dayarathne et al. (2019a). Our phylogenetic analyses with combined LSU, SSU, ITS and rpb2
sequence data also showed Conioscyphales, Fuscosporellales, Pleurotheciales, and Savoryellales
formed well-supported distinct clades (100% ML, 1.00 PP, 100% ML, 1.00 PP, 98% ML, 1.00 PP
and 100% ML, 1.00 PP, respectively) within the subclass Savoryellomycetidae (Fig. 10). Currently
there are four orders and four families in this subclass (this paper).
Sordariomycetidae O.E. Erikss. & Winka, Myconet 1(1): 10 (1997)
The subclass Sordariomycetidae was established by Eriksson & Winka (1997) and comprised
six orders, 12 families and two families incertae sedis. Members of this subclass are mainly
characterized by dark ascomata with inoperculate, unitunicate asci and occur in terrestrial, aquatic
and marine habitats and are widely distributed as plant and animal pathogens, endophytes, saprobes
as well as coprophilous and lichenicolous taxa (Maharachchikumbura et al. 2015, 2016b, Huang et
al. 2019). They are mycophilic, rich in coprophilous taxa and associated with invertebrates and
their ecological aspects and biotechnological potential have been researched (Zhang et al. 2006,
Raghukumar 2008, Bovio et al. 2018). An MCC tree based on a combined SSU, LSU, tef1 and rpb2
sequence data revealed that this subclass evolved around 145–216 MYA (Hongsanan et al. 2017,
Hyde et al. 2017a). The divergence time for Sordariomycetidae is estimated as 247 MYA (Fig. 2).
Currently there are eight orders and 19 families in this subclass (this paper).
Xylariomycetidae O.E. Erikss & Winka, Myconet 1: 12 (1997)
Different outlines of Xylariomycetidae have been published by Maharachchikumbura et al.
(2016b), Samarakoon et al. (2016b) and Hongsanan et al. (2017). However, in the present study, we
have revised the subclass. Concatenated LSU, ITS, rpb2 and tub2 based maximum likelihood
phylogeny resulted in a well-supported backbone tree for 34 families in Xylariomycetidae (Fig. 4).
The divergence time for Xylariomycetidae is estimated as 278 MYA (Fig. 2). There are three
distinct clades in the tree representing the orders discussed in previous studies: Xylariales,
Amphisphaeriales and Delonicicolales. The sister orders Xylariales (15 families) and
Amphisphaeriales (17 families) have moderate statistical support (55% ML) and basal to these is
the highly supported clade Delonicicolales (100% ML). Samarakoon et al. (2016b) and Hongsanan
et al. (2017) provided divergence time estimations as additional information for Amphisphaeriales,
which is estimated to have diverged from Xylariales around 152–187 Mya and provides evidence
for these as distinct orders. Families accepted in Amphisphaeriales in this paper are similar to
Hongsanan et al. (2017). In this study, we accept Cainiaceae as placed in Xylariales (Figs 1, 4),
while Iodosphaeriaceae (Figs 1, 4) which was previously referred to the Xylariomycetidae incertae
sedis, (Hongsanan et al. 2017) is placed in Amphisphaeriales. Xyladictyochaetaceae (Crous et al.
2018b) is accepted in Amphisphaeriales and clusters with Phlogicylindriaceae with high statistical
support (95% ML; Fig. 4). Hansfordiaceae (Crous et al. 2019b) is sister to Coniocessiaceae in
Xylariales with strong statistical support (82% ML; Fig. 4). Cylindriaceae (Crous et al. 2018b) and
Pseudotruncatellaceae (Crous et al. 2019b) are placed in Amphisphaeriales with poor statistical
support (Fig. 4). Induratiaceae will be introduced by Samarakoon et al. (2020) and is placed in
Xylariales (Fig. 4). Voglmayr et al. (2019a) introduced Leptosilliaceae as a new family which is
sister to Delonicicolaceae, while rejecting Delonicicolales. However, with high statistical support
(Fig. 4), we accept Delonicicolales in this study. Currently there are three orders and 35 families in
this subclass (this paper).
363
Figure 3 – Phylogram generated from maximum likelihood analysis based on combined LSU,
SSU, ITS and rpb2 sequence data of Diaporthomycetidae. One hundred and ninety-three strains are
included in the combined analyses which comprised 3545 characters (859 characters for LSU, 972
characters for SSU, 659 characters for ITS) after alignment. Single gene analyses were carried out
and the topology of each tree had clade stability. Tree topology of the maximum likelihood analysis
is similar to the Bayesian analysis. The best RaxML tree with a final likelihood value of 68207.368884 is presented. Estimated base frequencies were as follows: A = 0.248206, C =
0.241993, G = 0.285500, T = 0.224301; substitution rates AC = 1.369088, AG = 2.887040, AT =
1.413053, CG = 1.152137, CT = 6.303994, GT = 1.000000; gamma distribution shape parameter a
= 0.315782. Bootstrap support values for ML greater than 75% and Bayesian posterior probabilities
greater than 0.95 are given near the nodes. The tree is rooted with Diatrype disciformis (AFTOL-ID
927). Ex-type strains are in bold. The newly generated sequences are indicated in blue.
364
Figure 3 – Continued.
365
Figure 3 – Continued.
366
Figure 3 – Continued.
The orders
The orders are listed alphabetically with notes on new data since Maharachchimbura et al.
(2016b). The divergence time for each order has been estimated, and stem ages are used to consider
their ordinal status.
Amphisphaeriales Hawksw. & O.E. Erikss., Syst. Ascom. 5(1): 177 (1986)
Amphisphaeriales has been accepted in consecutive studies using multigene phylogenies in
the subclass Xylariomycetidae (Senanayake et al. 2015, Samarakoon et al. 2016b, Hongsanan et al.
367
2017, Crous et al. 2018b). However, based on previous studies, Wijayawardene et al. (2018a)
accepted 11 families in Amphisphaeriales, i.e. Amphisphaeriaceae, Apiosporaceae, Beltraniaceae,
Clypeophysalosporaceae,
Hyponectriaceae,
Melogrammataceae,
Oxydothidaceae,
Phlogicylindriaceae, Pseudomassariaceae, Sporocadaceae and Vialaeaceae, while Cainiaceae,
Coniocessiaceae, and Iodosphaeriaceae were accepted in Xylariomycetidae families incertae sedis.
In this study, Amphisphaeriales is a moderately supported clade sister to Xylariales and comprises
17 families viz. Amphisphaeriaceae, Apiosporaceae, Beltraniaceae, Castanediellaceae,
Clypeophysalosporaceae, Cylindriaceae, Hyponectriaceae, Iodosphaeriaceae, Melogrammataceae,
Oxydothidaceae,
Phlogicylindriaceae,
Pseudomassariaceae,
Pseudosporidesmiaceae,
Pseudotruncatellaceae, Sporocadaceae, Vialaeaceae, and Xyladictyochaetaceae. The divergence
time for Amphisphaeriales has been estimated as 133 MYA. This number did not include
Vialaeaceae because this family goes to Xylariales in MCC tree (Fig. 2). Currently there are 17
families and 88 genera in this order (this paper).
Amplistromatales D’souza, Maharachch. & K.D. Hyde, Fungal Divers. 72: 212 (2015)
Maharachchikumbura et al. (2015) introduced Amplistromatales (in Sordariomycetes order
incertae sedis) comprising two families, Amplistromataceae and Catabotryaceae. Subsequent
studies by Daranagama et al. (2017) and Hongsanan et al. (2017) accepted Amplistromatales in
Sordariomycetes incertae sedis with a stem age of 165 MYA. Hyde et al. (2017a) proposed
Catabotryaceae as a separate order based on stem age at 165 MYA. In this study, Catabotryaceae is
placed in Catabotryales, and Amplistromataceae is placed in Amplistromatales with high statistical
support (100% MP/100% ML/1.00 PP; Fig. 5). Catabotryales is introduced as a new order based on
its stem age (172 MYA). Currently there is one family and three genera in Amplistromatales (this
paper).
Annulatascales M.J. D’souza, Maharachch. & K.D. Hyde, Fungal Divers. 72(1): 212 (2015)
Annulatascales comprises a single family Annulatascaceae and currently five genera are
confirmed with molecular data (Zhang et al. 2017a; Fig. 3). The phylogenetic relationship of this
family is inferred mainly using LSU gene data as ITS and SSU genes are available for only a few
taxa. More genes generated from fresh collections are needed to clarify the phylogenetic
relationships of Annulatascales species. Paoayensis clustered in Annulatascaceae in our
phylogenetic analyses, but we consider that it should not belong to this family because of its 2–6spored asci and absence of large ring, which is different from familial concept of Annulatascaceae,
and its weak bootstrap support in Annulatascaceae. The placement of Annulatascus nilensis
(phylogenetically distant from other Annulatascus species) also warrant further investigations (Fig.
3). Fresh collections of this taxon are also needed. The divergence time for Annulatascales has been
estimated as 112 MYA (Fig. 2). Currently there is one family and 11 genera in this order (this
paper).
Atractosporales H. Zhang, K.D. Hyde & Maharachch., Fungal Divers. 85: 88 (2017)
Atractosporales was introduced to resolve the taxonomic problems of annulatascaceae-like
taxa previously placed in Annulatascaceae and Sordariomycetes genera incertae sedis. Three
families (viz. Atractosporaceae, Conlariaceae, and Pseudoproboscisporaceae) were included in the
order by Zhang et al. (2017a). The monophyly of this order is not well-supported in previous
studies (Luo et al. 2019) and in this study. Luo et al. (2019) showed that Junewangiaceae and
Cancellidium also clustered in Atractosporales and this agrees with our phylogenetic analyses, but
with low bootstrap support (Fig. 3). We therefore retain Cancellidium (Tubaki 1975) as
Sordariomycetes genera incertae sedis. This ambiguous group may be included in this order with
larger taxon sampling in future phylogenetic analyses. The divergence time for Atractosporales has
been estimated as 106 MYA (Fig. 2), which falls in the range for family status. The status of
Atractosporales members may therefore need revision following further study. Currently there are
three families and six genera in this order (this paper).
368
Figure 4 – Phylogram generated from maximum likelihood analysis based on combined ITS, LSU,
rpb2 and tub2 sequence data for Xylariomycetidae. Two hundred and seventy-two strains are
included in the combined analyses which comprised 4211 characters (1168 characters for ITS, 937
characters for LSU, 1128 characters for rpb2, 978 characters for tub2) after alignment.
Achaetomium macrosporum (CBS 532.94), Chaetomium elatum (CBS 374.66) and Sordaria
fimicola (CBS 723.96) are outgroup taxa. Single gene analyses were carried out and the topology of
each tree had clade stability. The best RaxML tree with a final likelihood value of - 132297.706952
is presented. Estimated base frequencies were as follows: A = 0.241914, C = 0.251908, G =
0.265558, T = 0.240620; substitution rates AC = 1.281946, AG = 3.512297, AT = 1.499895, CG =
1.121065, CT = 6.472834, GT = 1.000000; gamma distribution shape parameter a = 0.678614.
Bootstrap support values for ML greater than 75% are given near the nodes. Ex-type strains are in
bold. The newly generated sequences are indicated in blue.
369
Figure – 4 Continued.
370
Figure – 4 Continued.
371
Figure 4 – Continued.
Boliniales P.F. Cannon, Ainsworth & Bisby's Dictionary of the Fungi, Edn 9 (Wallingford): x
(2001)
Boliniales was established by Kirk et al. (2001) and comprises a single family Boliniaceae
based on perithecial, brown to black ascomata and cylindrical asci with smooth, ellipsoid
ascospores with apical germ pores (Rick 1931, Huang et al. 2019). Most members in Boliniales
have been found in America and Europe and primarily occur on wood and no asexual morph is
372
known (Zhang et al. 2006, Huang et al. 2019). Multi-gene analyses have shown that the order is
closely related to Meliolales, Phyllachorales and Chaetosphaeriales within Sordariomycetidae
(Lumbsch & Huhndorf 2010, Hongsanan et al. 2017, Hyde et al. 2017a, Huang et al. 2019). The
divergence time for Boliniales has been estimated as 158 MYA (Fig. 2). Currently there is one
family and nine genera in this order (this paper).
Figure 5 – Phylogram generated from maximum likelihood analysis based on combined LSU and
ITS sequence data of Amplistromatales. Eighteen strains are included in the combined analyses
which comprised 1661 characters (908 characters for LSU, 753 characters for ITS) after alignment.
Ophioceras dolichostomum (CBS 114926) and O. leptosporum (CBS 894.70) are used as outgroup
taxa. Single gene analyses were also carried out and the phylogenies were similar in topology and
clade stability. Tree topology of the maximum likelihood analysis is similar to the Bayesian
analysis. The best RaxML tree with a final likelihood value of - 6986.920872 is presented.
Estimated base frequencies were as follows: A = 0.226, C = 0.264, G = 0.321, T = 0.190;
substitution rates AC = 0.643764, AG = 1.471407, AT = 1.612417, CG = 0.654293, CT =
5.330856, GT = 1.000000; gamma distribution shape parameter a = 0.653797. Bootstrap support
values for MP and ML greater than 55% and Bayesian posterior probabilities greater than 0.95 are
given near the nodes. Ex-type strains are in bold. The newly generated sequences are indicated in
blue.
Calosphaeriales M.E. Barr, Mycologia 75(1): 11 (1983)
Members of Calosphaeriaceae and Pleurostomataceae are accommodated in Calosphaeriales
and are characterized by brown to black ascomata with reniform to clavate asci, phialidic
conidiophores and ellipsoid to allantoid conidia (Maharachchikumbura et al. 2016b, Huang et al.
2019). Some species are plant pathogens (Barr 1983, Damm et al. 2008, Trouillas et al. 2012). In
previous phylogenetic analyses, species of Calosphaeriales were closely related to Jobellisiales and
Togniniales (Maharachchikumbura et al. 2015, 2016b, Hongsanan et al. 2017). Calosphaeriales
comprises seven genera within the two families Calosphaeriaceae and Pleurostomataceae (Huang et
al. 2019). The monotypic genera, Kacosphaeria and Sulcatistroma are accepted in Calosphaeriales
based on the small, clavate asci with allantoid to ellipsoid, hyaline ascospores similar to
Calosphaeria (Huang et al. 2019). The divergence time for Calosphaeriales has been estimated as
160 MYA (Fig. 2). Currently there are two families and nine genera in this order (this paper).
373
Figure 6 – Phylogram generated from maximum likelihood analysis based on combined LSU, ITS
and tub2 sequence data of Boliniales. Related sequences are taken from Læssøe et al. (2013).
Seventy-three strains are included in the combined analyses which comprised 2081 characters (867
characters for LSU, 561 characters for ITS, 653 characters for tub2) after alignment. Members of
Diaporthales are used as outgroup taxa. Single gene analyses were also carried out and the
phylogenies were similar in topology and clade stability. The best RAxML tree with a final
likelihood value of -34809.341208 is presented. Estimated base frequencies were as follows: A =
0.209463, C = 0.298053, G = 0.289389, T = 0.203095; substitution rates AC = 1.086242, AG =
2.378803, AT = 1.369068, CG = 1.213916, CT = 6.024342, GT = 1.000000; gamma distribution
shape parameter a = 0.567627. Bootstrap support values for ML greater than 60% are given near
the nodes. Ex-type strains are in bold. The newly generated sequences are indicated in blue.
Catabotryales K.D. Hyde & Senan., ord. nov.
Index Fungorum number: IF556718; Facesoffungi number: FoF06864
Saprobic on dead plant parts. Sexual morph: Stromata irregularly scattered, conspicuous,
multi-loculate, superficial, with base slightly penetrating the epidermis, discoid to pulvinate,
reddish brown to black, scurfy surface, flat or slightly convex, composed of rather thin-walled,
reddish yellow cells of textura angularis, textura globosa or textura epidermoidea. Ascomata
perithecial, deeply imbedded in stromatic tissues, globose, with a long, periphysate, ostiolar neck.
Peridium thick, composed of several layers of reddish brown cells of textura angularis. Paraphyses
hypha-like, numerous, tapering towards the apex, not embedded in a gelatinous matrix. Asci 8spored, unitunicate, broad cylindrical, short pedicellate, apically rounded or truncate, with a J-,
discoid, refractive, apical ring. Ascospores bi-seriate, hyaline, 1-celled, ellipsoidal to cylindrical,
smooth-walled, lacking a mucilaginous sheath. Asexual morph: Undetermined.
Type family – Catabotryaceae Petr. ex M.E. Barr
Notes – Catabotryaceae was placed in Amplistromatales based on DNA sequence analyses
(Maharachchikumbura et al. 2015, 2016b). However, this family was later placed in
Sordariomycetes families incertae sedis based on phylogeny and molecular clock evidence
(Hongsanan et al. 2017). Catabotryaceae has only one strain representing its taxonomic placement
and more samples are needed to provide evidence for its ordinal status. Hyde et al. (2017a)
however suggested that a new order Catabotryales was needed to accommodate the monotypic
family Catabotryaceae based on divergent time estimations. Catabotryales is formally introduced
374
here for the single family Catabotryaceae. Catabotryales comprises saprobic species on dead leaves
and stems of tropical monocotyledons. This order is distinct from its sister orders in having
astromatic ascomata, broad cylindrical asci and ellipsoidal to cylindrical ascospores without a
mucilaginous sheath. The divergence time for Catabotryales has been estimated as 172 MYA (Fig.
2).
Cephalothecales Maharachch. & K.D. Hyde, ord. nov.
Index Fungorum number: IF557027; Facesoffungi number: FoF06900
An order within the class Sordariomycetes, subclass Sordariomycetidae occurring as saprobes
or epiphytes on rotting or dead plants or fungi, and causing systemic mycotic infection in humans
with burns. Sexual morph: Ascomata perithecial, solitary to gregarious, carbonaceous, covered by
sulphureous hyphae, without periphyses. Peridium cephalothecoid. Ascogenous hyphae septate.
Asci 8-spored, unitunicate, evanescent, apedicellate, without an apical ring. Ascospores irregularly
arranged, brown, variously shaped, without germ pores. Asexual morph: Hyphomycetous.
Mycelium branched septate. Conidiophores long, stiffly upright, cylindrical, septate. Conidiogenous
cells phialidic, cylindrical. Conidia in chains, cylindrical, ovate or obovate, with or without an
apiculate or truncate base, 1-celled.
Type family – Cephalothecaceae Höhn.
Notes – Cephalothecaceae was introduced by Höhnel (1917c) and is characterized by
ascomata with a cephalothecoid peridium. The peridial cells form plate-like complexes made up of
radiating groups of cells, where each plate is separated by well-defined lines of dehiscence
(Malloch & Cain 1970). Albertiniella, Cephalotheca, Cryptendoxyla, Phialemonium and the newly
introduced Victoriomyces (Davolos et al. 2019) are presently placed in the family. Although the
placement of Cephalothecaceae is not stable within subclass Sordariomycetidae, it has high support
in the MCC tree (Hyde et al. 2017a, Hongsanan et al. 2017). Cephalothecaceae has a divergence
time at 154 MYA (present study), thus, here we raise Cephalothecaceae to ordinal status.
Chaetosphaeriales Huhndorf, A.N. Mill. & F.A. Fernández, Mycologia 96(2): 378 (2004)
Huhndorf et al. (2004) introduced this order for Chaetosphaeriaceae. Linocarpaceae was
added to Chaetosphaeriales by Konta et al. (2017) to accommodate Neolinocarpon and Linocarpon,
which are previously accepted in Xylariales, genera incertae sedis (Maharachchikumbura et al.
2015, 2016b). Konta et al. (2017) also established Leptosporellaceae to accommodate
Leptosporella, which was maintained in Chaetosphaeriales genera incertae sedis by
Maharachchikumbura et al. (2016b). Wijayawardene et al. (2018a) accepted Leptosporellaceae and
Linocarpaceae in Chaetosphaeriales and this is confirmed in our phylogenetic analysis (Figs. 1, 8).
The polyphyletic genus Diplococcium was previously excluded from Sordariomycetes and accepted
in Helotiales by Maharachchikumbura et al. (2015). Wijayawardene et al. (2018a) listed
Diplococcium in Helminthosphaeriaceae. However, a phylogenetic analysis by Hernández-Restrepo
et al. (2017) confirmed that Diplococcium is not related to Helminthosphaeriaceae and was placed
in Helotiales. Lin et al. (2019) introduced Multiguttulispora and placed it within
Chaetosphaeriaceae. Neoleptosporella (incertae sedis) will be added to the order by Phukhamsakda
et al. (2020). Chaetosphaeriales grouped sister to Tracyllalales with a low statistical support in our
phylogenetic analysis (Figs 1, 8). The divergence time for Chaetosphaeriales has been estimated as
158 MYA (Fig. 2). Here we accept four families and 59 genera in the order.
Coniochaetales Huhndorf, A.N. Mill. & F.A. Fernández, Mycologia 96(2): 378 (2004)
This order consists of two families Coniochaetaceae representing the genera Barrina and
Coniochaeta and Cordanaceae which accommodates Cordana. Cordanaceae was previously
classified in Cordanales, however based on molecular clock evidence Hongsanan et al. (2017)
treated Cordanales as a synonym of Coniochaetales. The phylogenetic tree presented here supports
this synonymy (Fig. 9). The divergence time for Coniochaetales has been estimated as 131 MYA
(Fig. 2). Currently there are two families and five genera in this order (this paper).
375
Figure 7 – Phylogram generated from maximum likelihood analysis based on combined tub2, act,
ITS and LSU sequence data of Calosphaeriales, Togniniales, Jobellisiales and Tirisporallales. One
hundred and fifteen strains are included in the combined analyses which comprised 2296 characters
(572 characters for tub2, 269 characters for act, 586 characters for ITS, 869 characters for LSU)
after alignment. Three species of Ophiostomataceae are used as outgroup taxa. Single gene
analyses were also carried out and the phylogenies were similar in topology and clade stability. The
376
best RAxML tree with a final likelihood value of -29642.219830 is presented. Estimated base
frequencies were as follows: A = 0.222849, C = 0.289953, G = 0.265413, T = 0.221786;
substitution rates AC = 1.361161, AG = 3.471108, AT = 1.335451, CG = 1.139165, CT =
4.985065, GT = 1.000000; gamma distribution shape parameter a = 1.412987. Bootstrap support
values for ML greater than 60% are given near the nodes. Ex-type strains are in bold. The newly
generated sequences are indicated in blue.
Figure 8 – Phylogram generated from maximum likelihood analysis based on combined LSU and
ITS sequence data of Chaetosphaeriales and Tracyllalales taxa. Ninety-six strains are included in
the combined analyses which comprised 1695 characters (1081 characters for LSU, 614 characters
for ITS) after alignment. Neurospora crassa MUCL 19026 and Gelasinospora tetrasperma CBS
178.33 (Sordariaceae, Sordariales) are used as outgroup taxa. Single gene analyses were carried out
and the phylogenies were similar in topology and clade stability. Tree topology of the maximum
likelihood analysis is similar to the Bayesian analysis. The best RaxML tree with a final likelihood
value of - 23777.689886 is presented. Estimated base frequencies were as follows: A = 0.231060, C
= 0.264793, G = 0.308265, T = 0.195882; substitution rates AC = 1.388486, AG = 1.836207, AT =
1.649563, CG = 0.971659, CT = 6.316962, GT = 1.000000; gamma distribution shape parameter a
= 0.460297. Bootstrap support values for ML greater than 75% and Bayesian posterior probabilities
greater than 0.95 are given near the nodes. Ex-type strains are in bold. The newly generated
sequences are indicated in blue.
377
Figure 8 – Continued.
Conioscyphales Réblová & Seifert, Persoonia 37: 63 (2015)
Conioscyphales comprises the monotypic family Conioscyphaceae with species originating
from both terrestrial and freshwater habitats (Réblová et al. 2016c). Conioscyphales is
phylogenetically related to Pleurotheciales, Savoryellales and Fuscosporellales. The order was
initially introduced based on a multi-gene phylogeny using sequence data from five Conioscypha
species (Réblová et al. 2016c). Conioscyphales represents a distinct taxonomic entity which
accommodates species with astromatic ascomata, paraphyses, asci bearing J- apical ring and
transversely multiseptate ascospores. The conidia undergo schizolytic seccession (Réblová et al.
2016c). The divergence time for Conioscyphales has been estimated as 128.5 MYA (Fig. 2).
Currently there is one family and one genus in this order (this paper).
Coronophorales Nannf., Nova Acta R. Soc. Scient. upsal., Ser. 4 8(no. 2): 54 (1932)
= Melanosporales N. Zhang & M. Blackw., Mycol. Res. 111(5): 531 (2007)
Members of Coronophorales have a worldwide distribution as saprobes on wood (Mugambi
& Huhndorf 2010, Maharachchikumbura et al. 2016b, Nannfeldt 1932). Most species in this order
have superficial ascomata, sometimes with a subiculum (Mugambi & Huhndorf 2010). The
quellkörper is a conical structure sometimes present in the centrum of the cupulate or collapsed
ascomata and Munk pores are unique structures. These are rings of thickening between cells and
are important characters in peridium cells (Huang et al. 2019, Mugambi & Huhndorf 2010).
Coronophorales is a poorly studied group and was considered as synonym of Sordariales or
378
Nitschkiaceae (Barr 1990b, Müller & von Arx 1973, Nannfeldt 1932, 1975, Nannfeldt & Santesson
1975, Subramanian & Sekar 1990). Huhndorf et al. (2004a) used LSU sequence data to show that
Coronophorales is an independent member and lineage within Hypocreales, Hypocreomycetidae.
Multi-gene analyses were used to demonstrate the position of Coronophorales which
accommodates Bertiaceae, Ceratostomataceae, Chaetosphaerellaceae, Coronophoraceae,
Nitschkiaceae and Scortechiniaceae (Hongsanan et al. 2017, Huang et al. 2019, Hyde et al. 2017a,
Maharachchikumbura et al. 2015, Mugambi & Huhndorf 2010). However, Parasympodiellales
clusters as a sister group for Scortechiniaceae in our phylogenetic analysis (Fig. 11). The
divergence time for Coronophorales has been estimated as 192 MYA (Fig. 2). Currently there are
six families and 46 genera in this order (this paper).
Figure 9 – Phylogram generated from maximum likelihood analysis based on combined LSU and
ITS sequence data of Coniochaetales. Forty-two strains are included in the combined analyses
which comprised 2640 characters (521 characters for LSU, 567 characters for ITS) after alignment.
Chaetosphaeria garethjonesii (MFLUCC 15-1012) (Chaetosphaeriaceae, Chaetosphaeriales) is
used as outgroup taxon. Single gene analyses were carried out and the phylogenies were similar in
topology and clade stability. Tree topology of the maximum likelihood analysis is similar to the
Bayesian analysis. The best RaxML tree with a final likelihood value of - 6488.978861 is
presented. Estimated base frequencies were as follows: A = 0.249647, C = 0.238753, G =
0.279808, T = 0.231793; substitution rates AC = 1.587009, AG = 2.017143, AT = 1.817611, CG =
1.386630, CT = 7.265255, GT = 1.0000; gamma distribution shape parameter a = 0.484696.
Bootstrap support values for MP and ML greater than 50% and Bayesian posterior probabilities
greater than 0.9 are given near the nodes. Ex-type strains are in bold. The newly generated
sequences are indicated in blue.
379
Figure 10 – Phylogram generated from maximum likelihood analysis based on combined LSU,
SSU, ITS and rpb2 sequence data of Conioscyphales, Fuscosporellales, Parasympodiellales,
Pleurotheciales and Savoryellales. One hundred and twenty-nine strains are included in the
combined analyses which comprised 4253 characters (910 characters for LSU, 1499 characters for
SSU, 759 characters for ITS and 1085 characters for rpb2) after alignment. Lindra thalassiae
(AFTOL-ID 413) and Lulworthia uniseptata (AFTOL-ID 5014) are used as outgroup taxa. Single
gene analyses and the phylogenies were similar in topology and clade stability. Tree topology of
the maximum likelihood analysis is similar to the Bayesian analysis. The best RaxML tree with a
final likelihood value of -50774.805453 is presented. Estimated base frequencies were as follows:
A = 0.233897, C = 0.263834, G = 0.294235, T = 0.208033; substitution rates AC = 1.342378, AG
= 2.736022, AT = 1.339928, CG = 0.989078, CT = 6.301203, GT = 1.000000; gamma distribution
shape parameter a = 0.440908. Bootstrap support values for ML greater than 70% and Bayesian
posterior probabilities greater than 0.95 are given near the nodes. The tree is rooted with Lindra
thalassiae (AFTOL-ID 413) and Lulworthia uniseptata (AFTOL-ID 5014). Type strains are in bold
and black. The newly generated sequence is indicated in blue.
380
Figure 10 – Continued.
Delonicicolales R.H. Perera, Maharachch. & K.D. Hyde, Cryptog. Mycol. 38(3): 329 (2017)
Delonicicolales was introduced by Perera et al. (2017) to accommodate Delonicicolaceae
which comprises Delonicicola and Liberomyces. The divergence time estimates for the order are
181 (133–234) MYA and 95 (51–157) MYA (Perera et al. 2017). Voglmayr et al. (2019a)
introduced a new genus Furfurella to Delonicicolaceae and in the same study, Liberomyces was
synonymized under Leptosillia. Voglmayr et al. (2019a) also introduced Leptosilliaceae, which is
typified by Leptosillia, while rejecting Delonicicolales based on phylogenetic analyses with several
gene combinations (ITS-LSU and SSU-ITS-LSU-rpb1-rpb2-tef1-tub2). In our study, based on a
LSU-ITS-rpb2-tub2 matrix Delonicicolales forms a highly supported clade (100% ML) comprising
Delonicicolaceae and Leptosilliaceae, basal to Amphisphaeriales and Xylariales (Fig. 4). However,
381
in another combined gene analysis of LSU-SSU-tef1-rpb2, Delonicicolales forms an internal clade
in Xylariales (Fig. 1). The divergence time estimates in this study are a stem age of 165 Mya which
supports the order establishment by Perera et al. (2017). Currently there are two families and three
genera in this order (this paper).
Figure 11 – Phylogram generated from maximum likelihood analysis based on combined LSU, tef1
and ITS sequence data for Coronophorales. Related sequences are referred to Hongsanan et al.
(2017). Seventy-nine strains are included in the combined analyses which comprised 2211
characters (1044 characters for LSU, 627 characters for tef1, 540 characters for ITS) after
alignment. Members of Torpedosporales are used as outgroup taxa. Single gene analyses were
carried out and the phylogenies were similar in topology and clade stability. The best RaxML tree
with a final likelihood value of -23354.525618 is presented. Estimated base frequencies were as
follows: A = 0.228019, C = 0.288494, G = 0.281955, T = 0.201533; substitution rates AC =
1.194725, AG = 2.787601, AT = 1.703067, CG = 0.955075, CT = 3.781057, GT = 1.000000;
gamma distribution shape parameter a = 0.736138. Bootstrap support values for ML greater than
50% are given near the nodes. Ex-type strains are in bold. The newly generated sequences are
indicated in blue.
382
Figure 12 – Phylogram generated from maximum likelihood analysis based on combined LSU,
ITS, rpb2 and tef1 sequence data of Diaporthales. One hundred and fifteen strains are included in
the combined analyses which comprised 2890 characters (892 characters for LSU, 526 characters
for ITS, 1062 characters for rpb2, 407 characters for tef1) after alignment. Leotia lubrica (AFTOL
ID 1) is used as outgroup taxon. Single gene analyses were carried out and the phylogenies were
similar in topology and clade stability. Tree topology of the maximum likelihood analysis is similar
to the Bayesian analysis. The best RaxML tree with a final likelihood value of -41459.517722 is
presented. Estimated base frequencies were as follows: A = 0.250714, C = 0.246950, G =
0.282507, T = 0.219829; substitution rates AC = 1.688056, AG = 2.747632, AT = 1.764840, CG =
1.180905, CT = 6.854636, GT = 1.000000; gamma distribution shape parameter a = 0.339516.
Bootstrap support values for ML greater than 50% and Bayesian posterior probabilities greater than
0.90 are given near the nodes. Ex-type strains are in bold. The newly generated sequences are
indicated in blue.
383
Figure 12 – Continued.
Diaporthales Nannf., Nova Acta R. Soc. Scient. upsal., Ser. 4 8(no. 2): 53 (1932)
The order comprises terrestrial or aquatic taxa on plants, animals or in soil, which are
saprobes, endophytes, and pathogens (Senanayake et al. 2017a, 2018). This order has been
extensively revisited and new information on its taxonomy, morphology, ecology, mode of life, and
biotechnological potential have been reported (Alvarez et al. 2016, Senanayake et al. 2017a, 2018,
Braun et al. 2018, Carvalho et al. 2018, Fan et al. 2018, Pádua et al. 2019). This order is
characterized by sexual morph with solitary or aggregated perithecia sometimes with long papilla,
2–32-spored, unitunicate asci having a conspicuous refractive ring and the asexual morph is
generally coelomycetous, although rarely hyphomycetous (Rossman et al. 2007, Senanayake et al.
2018). Several research publications suggested that combined LSU, ITS, rpb2 and tef1 sequence
data provides the best resolution for the order. However, SSU, calM, tub2 and act have been used
in some cases to resolve the inter-generic taxonomic complications. The divergence time for
Diaporthales has been estimated as 180 MYA (Fig. 2). Currently, there are 30 families and 181
genera in this order (this paper).
384
Figure 13 – Phylogram generated from maximum likelihood analysis based on combined ITS,
LSU, rpb2 and tef1 sequence data of Distoseptisporales and Sporidesmiales. Forty-six strains are
included in the combined analyses which comprised 3153 characters (789 characters for LSU, 546
characters for ITS, 1038 characters for rpb2, 779 characters for tef1) after alignment. Single gene
analyses were carried out and the phylogenies were similar in topology and clade stability. Tree
topology of the maximum likelihood analysis is similar to the Bayesian analysis. The best RaxML
tree with a final likelihood value of -22582.382236 is presented. Estimated base frequencies were
as follows: A = 0.236961, C = 0.261709, G = 0.291323, T = 0.210007; substitution rates AC =
1.206796, AG = 2.260744, AT = 1.272906, CG = 1.049687, CT = 6.479176, GT = 1.000000;
gamma distribution shape parameter a = 0.651156. Bootstrap support values for ML greater than
75% and Bayesian posterior probabilities greater than 0.95 are given near the nodes. The tree is
rooted with Sordaria fimicola (FGSC 2918 and SMH 4106). Ex-type strains are in bold. The newly
generated sequences are indicated in blue.
Distoseptisporales Z.L. Luo, K.D. Hyde & H.Y. Su, Fungal Divers. 99: 482 (2019)
The monotypic order Distoseptisporales is placed in the class Sordariomycetes, subclass
Diaporthomycetidae and comprises a single family Distoseptisporaceae and Aquapteridospora
(Distoseptisporales genera incertae sedis). Distoseptisporaceae was established by Su et al. (2016b)
with a single genus Distoseptispora based on morphology and phylogeny. Yang et al. (2015)
385
introduced Aquapteridospora and was placed as Diaporthomycetidae genera incertae sedis. Luo et
al. (2019) raised Distoseptisporaceae to Distoseptisporales based on the morphology and
phylogenetic analysis of combined LSU, SSU, rpb2 and tef1 sequence data. The divergence time
for Distoseptisporales has been estimated as 102 MYA (Fig. 2), which falls in the range of family
status. The status may need revision following further study. Currently there is one family and one
genus in this order (this paper).
Falcocladiales R.H. Perera, Maharachch., Somrith., Suetrong & K.D. Hyde, Fungal Divers. 72:
218 (2015)
Falcocladiales was introduced to accommodate Falcocladiaceae by Maharachchikumbura et
al. (2015). Falcocladiaceae comprises a single genus Falcocladium (Jones et al. 2014).
Falcocladiales grouped sister to Coronophorales and Parasympodiellales with low statistical
support (63% ML) in our phylogenetic analysis (Fig. 1). The divergence time for Falcocladiales has
been estimated as 192 MYA. Currently there is one family with one genus in this order (this paper).
Fuscosporellales J. Yang, J. Bhat & K.D. Hyde, Cryptog. Mycol. 37(4): 457 (2016)
The order has been established in Hypocreomycetidae based on combined SSU, LSU and
rpb2 phylogenetic analyses and comprises one family, Fuscosporellaceae (Yang et al. 2016b). Six
genera, namely, Fuscosporella, Parafuscosporella, Mucispora, Bactrodesmiastrum, Plagiascoma
and Pseudoascotaiwania are accommodated within Fuscosporellaceae. Earlier, Bactrodesmiastrum
and Plagiascoma were placed in Sordariomycetes genera incertae sedis (Réblová et al. 2016c).
Yang et al. (2016b) showed that these two genera clustered with three newly introduced genera,
Fuscosporella, Parafuscosporella and Mucispora. The species Ascotaiwania persoonii also
grouped with the five genera, away from the type species of Ascotaiwania. This species was
therefore synonymized under Pseudoascotaiwania persoonia (Yang et al. 2016b). The six genera
form a monophyletic clade represented by Fuscosporellales, which is sister to Conioscyphales,
Pleurotheciales and Savoryellales (Yang et al. 2016b). The divergence time for Fuscosporellales
has been estimated as 190 MYA (Fig. 2). Currently there is one family and six genera in this order
(this paper).
Glomerellales Chadef., Stud. Mycol. 68: 170 (2011)
The ordinal name Glomerellales including Colletotrichum (= Glomerella) and three other
genera in a non-ranked group “Eu-Glomérellales” was introduced by Chadefaud (1960), but was
not validly published. Réblová et al. (2011) proposed Glomerellales to accommodate the families
Australiascaceae, Glomerellaceae and Reticulascaceae based on morphology, as well as analyses of
ITS, LSU, SSU and rpb2 sequence data. Maharachchikumbura et al. (2016b) established the
taxonomic position of Plectosphaerellaceae in Glomerellales. Tibpromma et al. (2018) introduced a
monotypic family, Malaysiascaceae to this order based on morphology and phylogeny. The
divergence time for Malaysiascaceae has been estimated as 44 MYA, which falls in the genus
status (Fig. 14). Therefore, the status of Malaysiascaceae may need revision following further
study. The divergence time for Glomerellales has been estimated as 216 MYA (Fig. 2). Currently
there are five families and 32 genera in this order (this paper).
Hypocreales Lindau, Nat. Pflanzenfam., Teil. I (Leipzig) 1(1): 343 (1897)
According to Wijayawardene et al. (2018a) Hypocreales comprised nine families.
Maharachchikumbura et al. (2016b) listed 223 genera belonging to families of Hypocreales based
on the literature and phylogenetic analysis. Later, two new genera were introduced to
Bionectriaceae: Bullanockia, and Paracylindrocarpon, based on DNA evidence and morphological
data (Crous et al. 2016a, b). Corinectria, Pleiocarpon, Varicosporella and Xenoleptographium
were also added in Nectriacae (González & Chaverri 2017, Aiello et al. 2017, Lechat & Fournier
2015, Crous et al. 2015d). Cryptoniesslia was introduced from previous research to Niessliaceae by
Scheuer (1993) and Crous et al. (2016b) added Mycophilomyces to Clavicipitaceae.
386
Blackwellomyces was added to Cordycipitaceae by Kepler et al. (2017). Maharachchikumbura et al.
(2016b) listed seven genera in Stachybotryaceae. Lombard et al. (2016) revisited the family and 36
genera were accepted in Stachybotryaceae, including 22 new genera which based on multigene
phylogeny and morphological data. Cylindriaceae, Sarocladiaceae and Xenodactylariaceae were
added to the order by Crous et al. (2018a, b). However, in our phylogentic anlysis, Cylindriaceae
groups within Amphisphaeriales (Xylariomycetidae), sister to Clypeophysalosporaceae and
Sporocadaceae (Figs 1, 2). Xenodactylariaceae does not appear to be related to Hypocreales as it
forms a separate lineage from the order, and basal to Torpedosporales (Figs 1, 2).
Flammocladiellaceae was introduced to accommodate the monotypic genus Flammocladiella by
Crous et al. (2015b). It grouped as sister clade to Clavicipitaceae with low statistical support (Crous
et al. 2015b), and similar results were obtained by Maharachchikumbura et al. (2016b).
Flammocladiellaceae grouped within Clavicipitaceae (Hongsanan et al. 2017, Sun et al. 2017),
hence it was not accepted as a family. Hongsanan et al. (2017) treated Flammocladiellaceae under
Clavicipitaceae based on phylogenetic data. However, Lechat & Fournier (2018a) observed that
Flammocladiellaceae is affiliated to Bionectriaceae. In our phylogenetic analysis,
Flammocladiellaceae forms a separate clade distant from Clavicipitaceae with low statistical
support (Figs 1, 15). Hence, we maintain Flammocladiellaceae as a separate family in Hypocreales
following Crous et al. (2015b). Here we accept 14 families in Hypocreales, namely: Bionectriaceae,
Calcarisporiaceae, Clavicipitaceae, Cocoonihabitaceae, Cordycipitaceae, Flammocladiellaceae,
Hypocreaceae, Myrotheciomycetaceae, Nectriaceae, Niessliaceae, Ophiocordycipitaceae,
Sarocladiaceae, Stachybotryaceae, and Tilachlidiaceae, based on molecular evidence. The
divergence time for Hypocreales has been estimated as 229 MYA (Fig. 2). Currently there are 303
genera in this order (this paper).
Jobellisiales M.J. D'souza & K.D. Hyde, Fungal Divers. 72: 219 (2015)
Jobellisiales was established by Maharachchikumbura et al. (2015) and comprises a single
family Jobellisiaceae based on perithecial, yellow, orange or brown ascomata and cylindrical asci
with brown ascospores (Ranghoo et al. 2001, Liu et al. 2012, Maharachchikumbura et al. 2016b).
Most members in Jobellisiales have been found in America in terrestrial and freshwater habitats
and no asexual morph is known (Ranghoo et al. 2001, Liu et al. 2012, Maharachchikumbura et al.
2016b). Jobellisiales species are similar to some members of Diaporthales in having brown, 1septate ascospores (Maharachchikumbura et al. 2015, Senanayake et al. 2017a). Phylogenetically,
Jobellisiales was a sister clade of Calosphaeriaceae (Maharachchikumbura et al. 2015, 2016b,
Hongsanan et al. 2017), and Hyde et al. (2017a) proposed that Jobellisiales is closely related to
Togniniaceae. Hongsanan et al. (2017) and Wijayawardene et al. (2018a) considered Jobellisiales
to be synonym of Calosphaeriales. However, Jobellisiales is an unstable clade, and we accept it as
an independent order and related to Calosphaeriales and Diaporthales (Fig. 12). The divergence
time for Jobellisiales is estimated as 138 MYA (Fig. 2). Currently there is one family with one
genus in this order (this paper).
Koralionastetales Kohlm., Volkm.-Kohlm., J. Campb. & Inderb., Mycol. Res. 113(3): 377 (2009)
Koralionastetales was introduced to accommodate the genera Pontogenia and Koralionastes
(Campbell et al. 2009). Pontogenia was initially classified as Sordariomycetes, genera incertae
sedis, while Koralionastes as Ascomycota, genera incertae sedis (Eriksson 2006). The order was
established based on the monophyletic clade formed by three Koralionastes species and one
Pontogeneia species in the LSU and SSU-based phylogeny (Campbell et al. 2009). Species of
Koralionastetales are distinct from taxa belonging to other orders in Sordariomycetes by the
formation of antheridia on their germinating ascospores (Zhang et al. 2006). Furthermore, all
Koralionastetales species are restricted to marine habitats, in association with corals, sponges and
algae (Campbell et al. 2009, Jones et al. 2019). The divergence time for Koralionastetales is
estimated as 210.5 MYA (Fig. 2). Currently there is one family and two genera in this order (this
paper).
387
Figure 14 – One of the 100 most phylogenetic tree generated by maximum parsimony analysis of
combined LSU, ITS and tef1 sequence data of species in Glomerellales. Thirty-eight strains are
included in the analyses, which comprise 2162 characters including gaps (800 characters for LSU,
558 characters for ITS, 793 characters for tef1) after alignment. Pleurothecium semifecundum (CBS
131271) and Pleurotheciella rivularia (CBS 125238) (Pleurotheciaceae, Pleurotheciales) are used
as outgroup taxa and the tree is rooted with. Single gene analyses were carried out and the
phylogenies were similar in topology and clade stability. Tree topology of the maximum parsimony
analysis is similar to the maximum likelihood and Bayesian analysis. The maximum parsimonious
dataset consisted of 1189 constant, 630 parsimony-informative and 343 parsimony-uninformative
characters. The parsimony analysis of the data matrix resulted in the maximum often equally most
parsimonious trees with a length of 2672 steps (CI=0.522, RI=0.666, RC=0.348, HI=0.478) in the
first tree. Bootstrap support values for MP and ML greater than 50% and Bayesian posterior
probabilities greater than 0.90 are given near the nodes. Ex-type strains are in bold. The newly
generated sequences are indicated in blue.
Lulworthiales Kohlm., Spatafora & Volkm.-Kohlm., Mycologia 92(3): 456 (2000)
Lulworthiales was introduced based on morphology and LSU and SSU phylogeny to
accommodate the genera Lulworthia and Lindra, which were initially accommodated in
Halosphaeriales (Kohlmeyer et al. 2000). All taxa referred to this order are marine aquatic fungi.
The order is accommodated in Lulworthiomycetidae, where it forms a strongly-supported clade
with Koralionastetales (Maharachchikumbura et al. 2016b) with a divergent age of 289 MYA
(Hongsanan et al. 2017). Lulworthia is polyphyletic, as observed from the combined LSU, SSU and
388
ITS phylogeny (Fig. 16). This is in accordance with other studies which also reported the polyphyly
of Lulworthia (Jones et al. 2008, 2009, 2019, Abdel-Wahab et al. 2010, Azevedo et al. 2017).
Spathulosporaceae, typified by Spathulospora is also reported to group in Lulworthiales (Jones et
al. 2019). Spathulosporaceae was initially placed in Spathulosporales based on morphology
(Kohlmeyer 1973), but molecular data of some Spathulosporaceae species have shown that the taxa
have a higher affinity to Lulworthiales, even though the type species of Spathulosporaceae was not
included, since it lacks sequence data (Inderbitzin et al. 2004, Campbell et al. 2005, Jones et al.
2009). Currently there is one family (the placement of Spathulosporaceae is not confirmed in
Lulworthiales) and 15 genera in this order with both asexual and sexual morphs (this paper).
Further studies are required to resolve the higher order rank of the marine algicolous parasites
Spathulospora.
Magnaporthales Thongk., Vijaykr. & K.D. Hyde, Fungal Divers. 34: 168 (2009)
Maharachchikumbura et al. (2016b) listed Magnaporthaceae, Ophioceraceae, and
Pyriculariaceae in Magnaporthales based on the literature and phylogenetic analysis.
Pseudohalonectriaceae was introduced as a novel family in Magnaporthales to accommodate
Pseudohalonectria based on phylogenetic and molecular dating evidence (Hongsanan et al. 2017).
The stem age of Pseudohalonectria falls within the range of family status (95 MYA) and has high
support in the phylogenetic and MCC trees. Silva et al. (2019) introduced a new genus
Bifusisporella, in Magnaporthaceae, to accommodate an endophytic fungus in Brazil. Hence,
Magnaporthales comprises five families, Ceratosphaeriaceae, Magnaporthaceae, Ophioceraceae,
Pseudohalonectriaceae, and Pyriculariaceae. The divergence time for Magnaporthales is estimated
as 190 MYA (Fig. 2). Currently there are 36 genera in this order (this paper).
Meliolales Gäum. ex D. Hawksw. & O.E. Erikss., Syst. Ascom. 5(1): 180 (1986)
Meliolales was introduced by Hawksworth & Eriksson (1986) accommodating a single
family Meliolaceae. Hosagoudar (2003) introduced Armatellaceae as a new family in this order
based on the morphology. Lumbsch & Huhndorf (2010) placed Meliolales in the class
Sordariomycetes. This was confirmed and followed by Hongsanan et al. (2015, 2017) and
Maharachchikumbura et al. (2016b). Hongsanan et al. (2015) provided a recent monograph that
reappraised the genera of Meliolales. Zeng et al. (2017) provided a checklist for identifying
Meliolales species, including current names of host plants with their corresponding Meliolales
species. Currently, Meliolales comprises two families, viz. Armatellaceae and Meliolaceae, but
sequence data is only available for Meliolaceae. The phylogenetic relationship of Meliolaceae in
the current study is different from Hongsanan et al. (2015) and Zeng et al. (2018) with regards to
the position of Irenopsis, which is sister to the main Meliola clade (Fig. 19). All recent studies
indicated that Meliola is polyphyletic and the clade including some Meliola species clusters in the
Appendiculella and Asteridiella lineages with low bootstrap support. The divergence time for
Meliolales is estimated as 219 MYA (Fig. 2). Currently there are two families and nine genera in
this order (this paper).
Microascales Luttr. ex Benny & R.K. Benj., Mycotaxon 12(1): 40 (1980)
Microascales comprises Ceratocystidaceae, Chadefaudiellaceae, Gondwanamycetaceae,
Halosphaeriaceae, Microascaceae and Graphiaceae (Réblová et al. 2011, Maharachchikumbura et
al. 2016b). In an outline of Ascomycota by Wijayawardene et al. (2018a), 99 genera have been
accepted under Microascales, distributed in six families, but only 89 have molecular data
(www.ncbi.nlm.nih.gov/Taxonomy/). A new genus Bretziella was introduced in Ceratocystidaceae
by de Beer et al. (2017) to accommodate B. fagacearum. Microascales may be paraphyletic, due to
the inclusion of Halosphaeriales (Zhang et al. 2006, Hibbett et al. 2007). However, in this study,
Microascales formed a monophyletic clade and this agrees with Maharachchikumbura et al. (2015,
2016b) and Hongsanan et al. (2017). The divergence time for Microascales is estimated as 216
MYA (Fig. 2). Currently there are seven families and 110 genera in this order (this paper).
389
Figure 15 – Phylogram generated from maximum likelihood analysis based on combined LSU,
ITS, cmdA, rpb2, tef1 and tub2 sequence data for Hypocreales. Two hundred sixty six strains are
included in the combined analysis which comprised 4731 characters (890 characters for LSU, 906
characters for ITS, 557 characters for cmdA, 862 characters for rpb2, 1163 characters for tef1, 348
characters for tub2) after alignment. Single gene analyses were carried out and the phylogenies
were similar in topology and clade stability. The best RaxML tree with a final likelihood value of 143259.087299 is presented. Estimated base frequencies were as follows: A = 0.238481, C =
0.265503, G = 0.275795, T = 0.220220; substitution rates AC = 1.361350, AG = 3.162200, AT =
1.584547, CG = 1.000927, CT = 7.415164, GT = 1.000000; gamma distribution shape parameter a
= 0.592521. Bootstrap support values for ML greater than 50% are given near the nodes. The tree is
rooted with Achaetomium macrosporum CBS 532.94, Chaetomium elatum CBS 374.66,
Gelasinospora tetrasperma AFTOL-ID 1287, and Neurospora crassa ICMP 6360. Ex-type strains
are in bold. The newly generated sequences are indicated in blue.
390
Figure 15 – Continued.
391
Figure 15 – Continued.
392
Figure 15 – Continued.
Myrmecridiales Crous, Persoonia 34: 219 (2015)
Myrmecridiales was established to accommodate Myrmecridium which was previously
placed in Hypocreomycetidae, genera incertae sedis (Crous et al. 2015, Maharachchikumbura et al.
2016b). Another genus Neomyrmecridium was later included in the order in a strongly supported
clade with Myrmecridium (Crous et al. 2018) and this is confirmed in this study (Fig. 3). The
divergence time for Myrmecridiales is estimated as 113 MYA (Fig. 2). Currently there are two
families and three genera in this order (this paper).
393
Figure 16 – Phylogram generated from maximum likelihood analysis based on combined LSU,
SSU and ITS sequence data of selected taxa from Koralionastetales, Lulworthiales and
Pisorisporiales. Sixty-nine strains are included in the combined gene analyses comprising 2780
characters after alignment (930 characters for LSU, 1059 characters for SSU and 784 characters for
ITS). Fuscosporella pyriformis (MFLUCC 16-0570) and Parafuscosporella mucosa (MFLUCC 160571) are used as outgroup taxa. Analyses of each single gene were performed and the topology of
each tree had clade stability. The tree topology of the maximum likelihood was similar to the
Bayesian and maximum parsimony analyses. Maximum likelihood analysis with 1000 bootstrap
replicates yielded a best tree with the likelihood value of -24852.531353. The matrix had 1582
distinct alignment patterns, with 38.88% of undetermined characters or gaps. Estimated base
frequencies were as follows; A = 0.249996, C = 0.239207, G = 0.291483, T = 0.219314;
substitution rates AC = 1.016404, AG = 2.006035, AT = 1.090073, CG = 1.243791, CT =
5.331398, GT = 1.000000; gamma distribution shape parameter α = 0.395174. Maximum
parsimony (black) and maximum likelihood (black) bootstrap values >65% and Bayesian posterior
probabilities (green) >0.90 (ML/BYPP) are given above the nodes. Ex-type strains are in bold and
new strains are in blue.
394
Figure 16 – Continued.
Ophiostomatales Benny & Kimbr., Mycotaxon 12(1): 48 (1980)
This order was introduced by Benny & Kimbrough (1980) for Ophiostomataceae, while
Kathistaceae was added by Malloch & Blackwell (1990). In our phylogenetic analyses generated
from LSU, ITS, tub2 and SSU sequence data Ophiostomataceae formed a monophyletic clade with
Kathistaceae with high statistical support (99% ML, 0.99 PP) (Fig. 20). Nevertheless, based on
phylogenetic analysis, Subbaromyces splendens (strain U63552), a species in Ophiostomataceae,
grouped with Kathistaceae. However, the type genus Subbaromyces lacks sequence data and
recollection and sequencing is needed to confirm its position. Ophiostomataceae comprises 13
genera of which sequence data is available for 11 genera (Wijayawardene et al. 2018a).
Kathistaceae comprises three genera and sequence data are available for only Kathistes
(Wijayawardene et al. 2018a). The divergence time for Ophiostomatales is estimated as 91 MYA
(Fig. 2). Currently there are two families and 16 genera in this order (this paper).
Pararamichloridiales Crous, Persoonia 39: 357 (2017)
Pararamichloridiales comprises the monotypic family Pararamichloridiaceae and includes a
single genus Pararamichloridium. In our phylogenetic analyses generated from LSU and ITS
sequence data, Pararamichloridiales formed a monophyletic clade with high statistical support
(100% MLBS/1.00PP), which is the same with Crous et al. (2017a) (Fig. 17). Pararamichloridiales
is characterised by branched, subhyaline to brown, septate conidiophores, with polyblastic, terminal
and intercalary conidiogenous cells that produce solitary, hyaline, aseptate, clavate to ellipsoid
conidia (Crous et al. 2017a). In this study, we accept only one family i.e. Pararamichloridiaceae
with one genus i.e. Pararamichloridium. The divergence time for Pararamichloridiales is estimated
as 101.5 MYA (Fig. 2), which falls in the range of family status. The status of this order may need
revision following further study.
Parasympodiellales Hern.-Restr., Gené, R.F. Castañeda & Crous, Stud. Mycol. 86: 87 (2017)
The monotypic order Parasympodiellales accommodates Parasympodiellaceae, typified by
Parasympodiella (Hernández-Restrepo et al. 2017). The order has been introduced based on a LSU
and ITS phylogeny for four Parasympodiella species, including the type species, P. laxa, all of
which grouped together to form a monophyletic clade (Hernández-Restrepo et al. 2017). In
addition, taxa of Parasympodiellales are characterised by stylaspergillus-like synasexual morphs,
supporting this order as a distinct taxonomic entity (Cheewangkoon et al. 2009, Hernández395
Restrepo et al. 2017). Parasympodiellales clusters as a sister group to Coronophorales in our
phylogenetic analysis (Fig. 2). Their status may need revision following further study. Currently
there is one family and one genus in this order (this paper).
Figure 17 – Phylogram generated from maximum likelihood analysis based on combined LSU and
ITS sequence data of Magnaporthales and Pararamichloridiales. Eighty-seven strains are included
in the combined analyses which comprised 1559 characters (875 characters for LSU, 684 characters
for ITS) after alignment. Diaporthe obtusifoliae (CPC 32336) and D. passiflorae (CBS 132527) are
used as outgroup taxa. Single gene analyses were carried out and the phylogenies were similar in
topology and clade stability. Tree topology of the maximum likelihood analysis is similar to the
Bayesian analysis. The best RaxML tree with a final likelihood value of – 12664.4150 is presented.
Estimated base frequencies were as follows: A = 0.2243, C = 0.2819, G = 0.2966, T = 0.1971;
substitution rates AC = 1.5257, AG = 2.2337, AT = 2.6637, CG = 1.0862, CT = 7.4361, GT =
1.0000; gamma distribution shape parameter a = 0.4747. Bootstrap support values for ML greater
than 50% and Bayesian posterior probabilities greater than 0.95 are given near the nodes. Ex-type
strains are in bold. The newly generated sequences are indicated in blue.
396
Figure 17 – Continued.
Phomatosporales Senan., Maharachch. & K.D. Hyde, Mycosphere 7(5): 631 (2016)
Phomatosporales comprises Phomatospora, Lanspora and Tenuimurus (Senanayake et al.
2016). Phomatospora and Lanspora cluster together with statistical support of 86% ML and 0.97
PP, while Tenuimurus forms a clade with the other two genera with good support (100% ML, 1.00
PP) (Fig. 20). Phomatosporales contains the single family Phomatosporaceae (Senanayake et al.
2016). The divergence time for Phomatosporales is estimated as 91 MYA (Fig. 2), which falls in
the range of family status. Its status may need revision following further study. Currently there is
one family and three genera in this order (this paper).
Phyllachorales M.E. Barr, Mycologia 75(1): 11 (1983)
Phyllachorales comprises three families viz. Phaeochoraceae, Phyllachoraceae and
Telimenaceae (Dayarathne et al. 2017, Mardones et al. 2017). In our phylogenetic analyses with
concatenated ITS, LSU and SSU sequence data, the Phaeochoraceae formed a monophyletic clade
with high statistical support (98% ML) (Fig. 21). Phaeochoraceae comprises Cocoicola,
Phaeochora, Phaeochoropsis and Serenomyces and sequence data are available only for two
Cocoicola strains and one Serenomyces species. However, the type genus Phaeochora lacks
sequence data. Phyllachoraceae formed a less stable clade within Phyllachorales. Out of 54 genera
in Phyllachoraceae, sequences data are available for only five genera. A few Polystigma strains
grouped within Phyllachorales in two different places which is similar to the study of Dayarathne et
al. (2017). However, Polystigma was excluded and placed within the reinstated Polystigmataceae
by Jones et al. (2019) and Bundhun et al. (2019). Telimenaceae species also formed a monophyletic
clade in our phylogram, but with low statistical support (Fig. 21). The divergence time for
Phyllachorales is estimated as 168 MYA (Fig. 2). Currently there are three families and 60 genera
in this order (this paper).
Pisorisporiales Réblová & J. Fourn., Persoonia 34: 43 (2014)
Phylogenetic analyses of strains from an aquatic environment, including LSU, SSU and rpb2
sequence data, grouped them in a monophyletic clade with Achroceratosphaeria species (Réblová
et al. 2015a). The new taxa were placed in the novel genus, Pisorisporium and a new family,
Pisorisporiaceae was erected to accommodate Pisorisporium and Achroceratosphaeria. The latter
genus was, upon its introduction, initially placed in Sordariomycetes incertae sedis (Réblová et al.
2010). This new family Pisorisporiaceae was, in turn, accommodated in Pisorisporiales based on its
distinct taxonomy and phylogeny at the ordinal level (Réblová et al. 2015a). Pisorisporiales forms a
397
moderately-supported
sister clade with Lulworthiales
and Koralionastetales
in
Lulworthiomycetidae (Hongsanan et al. 2017). The order has been reported to have a stem age
which falls between 250–300 MYA, thereby placing it at a subclass level (Hyde et al. 2017a,
Hongsanan et al. 2017). In this study, therefore Pisorisporiales is raised to the subclass
Pisorisporiomycetidae. Currently there is one family and two genera in this order (this paper).
Figure 18 – Phylogram generated from Bayesian inference based on combined LSU, SSU and ITS
sequence data of Meliolales. Twenty-one strains are included in the combined analyses which
comprised 2688 characters (869 characters for LSU, 1020 characters for SSU, 799 characters for
ITS) after alignment. Coniochaeta rosae (MFLUCC 17-0810) is used as outgroup taxa. Single gene
analyses were carried out and the phylogenies were similar in topology and clade stability. Tree
topology of the maximum likelihood analysis is similar to the Bayesian analysis. Estimated base
frequencies were as follows: A = 0.25335, C = 0.2223, G = 0.281553, T = 0.242798; substitution
rates AC = 1.021441, AG = 3.556619, AT = 2.123549, CG = 0.380727, CT = 6.542902, GT =
1.000000; gamma distribution shape parameter α = 0.549005. Bootstrap support values for ML
greater than 75% and Bayesian posterior probabilities greater than 0.95 are given near the nodes.
Ex-type strains are in bold. The newly generated sequences are indicated in blue.
Pleurotheciales Réblová & Seifert, Persoonia 37: 63 (2015)
Pleurotheciales was introduced based on a combined ITS, SSU, LSU, tub2, mcm7 and rpb2
dataset (Réblová et al. 2016c) and it presently accommodates Pleurotheciaceae which in turn
comprises ten genera. Taxa belonging to Pleurotheciales cannot be successfully differentiated based
on their overlapping sexual morphology. The type of conidiogenesis can, to a certain extent,
delineate species into groups within the order, since conidial secession is rhexolytic or schizolytic
with holoblastic, monoblastic or polyblastic (sympodially extending) conidiogenesis (Réblová et al.
2016c). Molecular data and/or cultures are essential to establish the placement of Pleurotheciales
taxa (Réblová et al. 2016c). In this study, Pleurothecium obovoideum does not cluster with other
Pleurothecium species, including the type species, P. recurvatum (Fig. 10). This result is in
agreement with Réblová et al. (2012). The divergence time for Pleurotheciales is estimated as 105
MYA (Fig. 2), which is evidence of family status. The status as an order may need revision
following further study. Currently there is one family and ten genera in this order (this paper).
398
Figure 19 – Phylogram generated from maximum likelihood analysis based on combined ITS,
LSU, SSU and tef1 sequence data of Microascales. One hundred and thirty-two strains are included
in the combined analyses which comprised 4538 characters (653 characters for ITS, 1215
characters for LSU, 1685 characters for SSU, 985 characters for tef1) after alignment.
Lecanicillium fungicola var. aleophilum (CBS 357.80) and L. fungicola var. fungicola (CBS
992.69) (Cordycipitaceae, Hypocreales) are used as outgroup taxa. Single gene analyses were
carried out and the phylogenies were similar in topology and clade stability. Tree topology of the
maximum likelihood analysis is similar to the Bayesian analysis. The best RaxML tree with a final
likelihood value of -46610.749560 is presented. Estimated base frequencies were as follows: A =
0.240650, C = 0.256729, G = 0.270623, T = 0.231998; substitution rates AC = 0.965367, AG =
2.062871, AT = 1.418856, CG = 0.960832, CT = 4.902766, GT = 1.000000; gamma distribution
shape parameter a = 0.478984. Bootstrap support values for ML greater than 65% and Bayesian
posterior probabilities greater than 0.95 are given near the nodes. Ex-type strains are in bold.
399
Figure 19 – Continued.
Pseudodactylariales Crous, Persoonia 39: 421 (2017)
Pseudodactylariales was introduced by Crous et al. (2017a) and Pseudodactylariaceae, one
genus, Pseudodactylaria, and three species were placed in this order. In our phylogenetic analyses
generated from LSU, ITS, SSU and rpb2 sequence data, Pseudodactylariales formed a
monophyletic clade with high statistical support (100% MPBS/1.00 PP) (Fig. 22). The divergence
time for Pseudodactylariales is estimated as 128 MYA (Fig. 2). Currently there is one family and
one genus in this order (this paper).
Savoryellales Boonyuen, Suetrong, Sivichai, K.L. Pang & E.B.G. Jones, Mycologia 103(6): 1368
(2011)
Savoryellales consists of Savoryellaceae with the genera Ascotaiwania, Canalisporium,
Dermatisporium and Savoryella (Réblová et al. 2016c, Luo et al. 2019). Species are found in
terrestrial, marine, brackish and freshwater habitats, as well as water-cooling towers (Jones &
Eaton 1969, Minoura & Muroi 1978, Hyde & Jones 1988, Chang et al. 1998, Ranghoo & Hyde
1998, Luo et al. 2019). The sexual morphs of Savoryellales species have perithecial ascomata with
elongate necks, while the asexual morphs are dematiaceous hyphomycetes with semimacronematous conidiophores and monoblastic conidiogenous cells (Réblová et al. 2016c,
Dayarathne et al. 2019a, Zhang et al. 2019). Savoryellales was initially placed in
Hypocreomycetidae based on multi-gene phylogeny (Boonyuen et al. 2011). However, later it was
raised to the subclass Savoryellomycetidae following molecular dating data with a stem age of 268
MYA, indicating a subclass status (Hongsanan et al. 2017). Currently there is one family and four
genera in this order (this paper).
400
Figure 20 – Phylogram generated from maximum likelihood analysis based on combined LSU,
SSU ITS and tub2 sequence data of Ophiostomatales and Phomatosporales. Sixty-six strains are
included in the combined analyses which comprised 2897 characters (584 characters for LSU, 985
characters for SSU, 737 characters for ITS, 591 characters for tub2) after alignment. Ophioceras
dolichostomum (CBS 114926) (Ophioceraceae, Magnaporthales) and Nakataea oryzae (ATCC
44754) (Magnaporthaceae, Magnaporthales) are used as outgroup taxa. Tree topology of the
maximum likelihood analysis is similar to the Bayesian analysis. The best RaxML tree with a final
likelihood value of - 21546.866789 is presented. Estimated base frequencies were as follows: A =
0.213168, C = 0.292921, G = 0.285293, T = 0.208619; substitution rates AC = 1.227359, AG =
2.074779, AT = 1.607139, CG = 1.020705, CT = 4.744232, GT = 1.000000; gamma distribution
shape parameter a = 0.294947. Bootstrap support values for ML greater than 70% and Bayesian
posterior probabilities greater than 0.90 are given near the nodes. Ex-type strains are in bold.
401
Figure 21 – Phylogram generated from maximum likelihood analysis based on combined LSU,
SSU and ITS sequence data of selected taxa from Phyllachorales. Related sequences are taken from
Dayarathne et al. (2017). One hundred and sixteen strains are included in the combined gene
analyses comprising 1326 characters after alignment (610 characters for LSU, 972 characters for
SSU and 657 characters for ITS). Gelasinospora tetrasperma (CBS 178.33), Neurospora crassa
(MUCL 19026) and Sordaria fimicola (CBS 508.50) are used as outgroup taxa. Analyses of each
single gene were performed and the topology of each tree had clade stability. Maximum likelihood
analysis with 1000 bootstrap replicates yielded a best tree with the likelihood value of 27773.000538. The matrix had 1583 distinct alignment patterns, with 57.76% of undetermined
characters or gaps. Estimated base frequencies were as follows; A = 0.250314, C = 0.239655, G =
0.269539, T = 0.240492; substitution rates AC = 1.057397, AG = 2.260386, AT = 1.375589, CG =
0.834187, CT = 4.235196, GT = 1.000000; gamma distribution shape parameter α = 0.534795.
Maximum likelihood bootstrap (ML, black) values > 60%) are given above the nodes. Ex-type
strains are in bold.
402
Figure 21 – Continued.
Sordariales Chadef. ex D. Hawksw. & O.E. Erikss., Syst. Ascom. 5(1): 182 (1986)
Sordariales was established by Hawksworth & Eriksson (1986) and clarified by Huhndorf et
al. (2004b) based on LSU sequences analysis. Chaetomiaceae, Sordariaceae and Lasiosphaeriaceae
are accommodated in this order and taxa are characterized by cleistothecial or perithecial ascomata
and hyaline or brown ascospores often with appendages or sheaths (Huhndorf et al. 2004b, Kruys et
al. 2015, Maharachchikumbura et al. 2016b, Huang et al. 2019). Sordariales members cluster in a
clade with Boliniales, Chaetosphaeriales and Phyllachorales (Hongsanan et al. 2017, Hyde et al.
2017a). The traditionally defined Chaetomiaceae and Lasiosphaeriaceae were found to be
polyphyletic (Huhndorf et al. 2004b, Kruys et al. 2015, Wang et al. 2016a). Some members of
Lasiosphaeriaceae, Triangularia, Cladorrhinum and Podospora, were placed in a new family
Podosporaceae (Wang et al. 2019a). In this study, Chaetomiaceae, Podosporaceae,
Lasiosphaeriaceae (I, II, III), and Sordariaceae form a distinct lineage (Fig. 23). However, more
data is needed to determine the phylogenetic affinities of this order. The divergence time for
Sordariales is estimated as 128 MYA (Fig. 2). Currently there are four families and 89 genera in
this order (this paper).
Spathulosporales Kohlm., Mycologia 65(3): 615 (1973)
Spathulosporales includes Hispidicarpomycetaceae and Spathulosporaceae, while,
Hispidicarpomycetaceae lacks sequence data in GenBank. Sequence data are available only for two
species of Spathulospora. Our phylogenetic analyses with combined LSU, SSU and ITS data
showed that Spathulospora and Rostrupiella species form a well-supported single clade (99% ML,
1.00 PP) within Lulworthiaceae (Fig. 16) and this is similar to Inderbitzin et al. (2004), Campbell et
403
al. (2005) and Jones et al. (2009, 2019). However, sequence data are not available for the type
species of Spathulospora hence, further collections are need from the type to resolve the
phylogenetic placement of this order. The divergence time for Sordariales is estimated as 121 MYA
(Fig. 2), which falls in the range of family status. The order status may need revision following
further study. Currently there are two families and three genera in this order (this paper).
Figure 22 – Phylogram generated from maximum likelihood analysis based on combined LSU,
SSU, ITS and rpb2 sequence data of Pseudodactylariales and Vermiculariopsiellales. Forty-seven
strains are included in the combined analyses which comprised 4320 characters (865 characters for
LSU, 1634 characters for SSU, 662 characters for ITS, 1159 characters for rpb2) after alignment.
Arthrinium arundinis (CBS 133509) and Arthrinium montagnei (AFTOL-ID 951) (Apiosporaceae,
Xylariales) are used as outgroup taxa. Single gene analyses were carried out and the phylogenies
were similar in topology and clade stability. Tree topology of the maximum likelihood analysis is
similar to the Bayesian analysis. The best RaxML tree with a final likelihood value of 29256.470401 is presented. Estimated base frequencies were as follows: A = 0.249068, C =
0.239305, G = 0.280423, T = 0.231203; substitution rates AC = 1.369489, AG = 2.705642, AT =
1.345736, CG = 1.341396, CT = 6.599852, GT = 1.000000; gamma distribution shape parameter a
= 0.220775. Bootstrap support values for ML greater than 75% and Bayesian posterior probabilities
greater than 0.95 are given near the nodes, respectively. Ex-type strains are in bold.
404
Figure 23 – Phylogram generated from maximum likelihood analysis based on combined LSU,
ITS, tub2 and rpb2 sequence data of Sordariales. Related sequences were taken from Wang et al.
(2016b). Two hundred and fifty-seven strains are included in the combined analyses which
comprised 2717 characters (855 characters for LSU, 480 characters for ITS, 860 characters for
tub2, 522 characters for rpb2) after alignment. Members of Amphisphaeriales are used as outgroup
taxa. Single gene analyses were carried out and the phylogenies were similar in topology and clade
stability. The best RAxML tree with a final likelihood value of -59082.079074 is presented.
Estimated base frequencies were as follows: A = 0.238795, C = 0.266829, G = 0.275292, T =
0.219085; substitution rates AC = 1.483472, AG = 3.436969, AT = 1.859704, CG = 1.064423, CT
= 6.934931, GT = 1.000000; gamma distribution shape parameter a = 0.757314. Bootstrap support
values for ML greater than 50% are given near the nodes. Ex-type strains are in bold. The newly
generated sequences are indicated in blue.
405
Figure 23 – Continued.
406
Figure 23 – Continued.
407
Sporidesmiales Crous, Persoonia 40: 377 (2018)
Sporidesmiales was introduced by Crous et al. (2018d) and presently only contains
Sporidesmiaceae with a single genus. Taxa are saprobic on woody debris in terrestrial and aquatic
habitats or mycoparasites of fungi. The divergence time for Sporidesmiales is estimated as 81
MYA (Fig. 2), which is evidence of family status but ranking it as an order may need revision
following further study. Currently there is one family and one genus in this order (this paper).
Tirisporellales Suetrong, E.B.G. Jones & K.L. Pang, Fungal Divers. 73(1): 42 (2015)
The monotypic order Tirisporellales is placed in the class Sordariomycetes, subclass
Diaporthomycetidae and comprises a single family Tirisporellaceae. Suetrong et al. (2015)
introduced Tirisporellaceae with two monotypic genera Tirisporella and Thailandiomyces. Jones et
al. (2015) introduced Tirisporellales to accommodate the genera Tirisporella, Thailandiomyces
with an additional genus Bacusphaeria. In our phylogram generated with concatenated LSU and
SSU sequence data, the genera Tirisporella (100% ML, 100% MP), Thailandiomyces (93% ML,
100% MP), and Bacusphaeria (100% ML, 100% MP) formed strongly supported clades within
Tirisporellales (Fig. 25). In Fig. 1 this order formed an internal clade of Diaporthales and this
observation is supported in Hongsanan et al. (2017). However, Tirisporellales was not formally
synonymized under Diaporthales by either Hyde et al. (2017a) or Hongsanan et al. (2017).
Tirisporellaceae is primarily aquatic with both freshwater and marine taxa with a hyphomycetous
asexual morphs which are not usually found in Diaporthales (Zhang et al. 2019). Therefore, we do
not place this order in Diaporthales. The divergence time for Tirisporellales is estimated as 142
MYA (Fig. 2). Currently there is one family and three genera in this order (this paper).
Togniniales Senan., Maharachch. & K.D. Hyde, Fungal Divers. 72: 220 (2015)
Togniniales comprises a single family Togniniaceae based on phylogenetic analyses and it is
similar to some members of Calosphaeriaceae in having perithecial ascomata and clavate, tiny asci
with hyaline ascogenous hyphae and cylindrical to allantoid ascospores (Maharachchikumbura et
al. 2015, Réblová et al. 2015b). Togniniaceae was placed in Diaporthales based on SSU and LSU
gene analyses (Mostert et al. 2006). However, multi-gene analyses showed that Togniniales as an
independent clade with Diaporthales and Calosphaeriales (Maharachchikumbura et al. 2015, 2016b,
Réblová et al. 2015b, Huang et al. 2019). Species of this order have a worldwide distribution and
are well-known pathogens of plants and animals (Rooney-Latham et al. 2005, Hoffmann & Voigt
2011, Kredics et al. 2011, Huang et al. 2018). The divergence time for Togniniales is estimated as
138 MYA (Fig. 2). Currently there is one family and two genera in this order (this paper).
Torpedosporales E.B.G. Jones, Abdel-Wahab & K.L. Pang, Fungal Divers. 73(1): 42 (2015)
The families Etheirophoraceae, Juncigenaceae and Torpedosporaceae form well-separated
(99% ML, 1.00 PP, 99% ML, 1.00 PP and 100% ML, 1.00 PP) clades within Torpedosporales in
the phylogram generated from combined LSU, SSU and rpb2 sequence data (Fig. 24). This
observation is supported by previous studies of Jones et al. (2015) and Maharachchkumbura et al.
(2016b). The divergence time for Torpedosporales is estimated as 185 MYA (Fig. 2). Currently
there are three families and ten genera in this order (this paper).
Tracyllalales Crous, Persoonia 40: 365 (2018)
Hernandez-Restrepo et al. (2016a) placed Tracylla aristata within Sordariomycetidae
incertae sedis based on LSU and ITS sequence data. With the description of a new species of
Tracylla from Eucalyptus urophylla in Colombia, Crous et al. (2018d) introduced Tracyllalales to
accommodate the monotypic family Tracyllaceae. Multigene phylogenetic analysis (ITS, LSU,
SSU) of T. aristata and T. eucalypti group these taxa in a well-supported clade distinct from known
orders within Sordariomycetidae. Tracyllalales is closely related to Chaetosphaeriales. The
divergence time for Tracyllalales is estimated as 154 MYA (Fig. 2). Currently there is one family
with one genus in this order (this paper).
408
Figure 24 – Phylogram generated from maximum likelihood analysis based on combined LSU,
SSU and rpb2 sequence data of selected taxa from Falcocladiales and Torpedosporales. Twenty-six
strains are included in the combined gene analyses comprising 3557 characters after alignment
(1142 characters for LSU, 1440 characters for SSU and 967 characters for rpb2). Nectria
haematococca (GJS89-70) is used as the outgroup taxon. Analyses of each single gene were
performed and the topology of each tree had clade stability. The tree topology in maximum
likelihood analysis was similar to the Bayesian and maximum parsimony analyses. Maximum
likelihood analysis with 1000 bootstrap replicates yielded a best tree with the likelihood value of 14936.746257. The matrix had 1096 distinct alignment patterns, with 25.55% of undetermined
characters or gaps. Estimated base frequencies were as follows; A = 0.247642, C = 0.231060, G =
0.304405, T = 0.216894; substitution rates AC = 1.746605, AG = 3.110551, AT = 1.540183, CG =
1.746965, CT = 7.476678, GT = 1.000000; gamma distribution shape parameter α = 0.291342.
Maximum parsimony (black) and maximum likelihood (black) bootstrap values >65% and
Bayesian posterior probabilities (blue) >0.90 (ML/BYPP) are given above the nodes. Ex-type
strains are in bold.
Vermiculariopsiellales Hern.-Restr., J. Mena, Gené & Crous, Stud. Mycol. 86: 91 (2017)
Hernández-Restrepo et al. (2017) introduced Vermiculariopsiellales and one family,
Vermiculariopsiellaceae, and one genus, Vermiculariopsiella, were accepted. ITS, LSU, SSU, actA
and tef1 sequence data are available for species of Vermiculariopsiella. In our phylogenetic
analyses generated from LSU, ITS, SSU and rpb2 sequence data, Vermiculariopsiellales formed a
monophyletic clade with high statistical support (100% MPBS/1.00 PP) (Fig. 22). The divergence
time for Vermiculariopsiellales is estimated as 131 MYA (Fig. 2). Currently there is one family
with one genus in this order (this paper).
409
Xenospadicoidales Hern.-Restr., J. Mena & Gené, Stud. Mycol. 86: 91 (2017)
Xenospadicoidales is shown as a strongly supported monophyletic group and the single
family Xenospadicoidaceae was proposed to replace Lentomitellaceae (Hernández-Restrepo et al.
2017, Réblová et al. 2018). The genera Xenospadicoides and Pseudodiplococcium were also
synonymized under Spadicoides (Réblová et al. 2018). Luo et al. (2019) introduced another
freshwater genus Neospadicoides. All genera formed a well-supported clade in this study (Fig. 3).
The divergence time for Xenospadicoidales is estimated as 106 MYA (Fig. 2). Currently there is
one family and five genera in this order (this paper).
Xylariales Nannf., Nova Acta R. Soc. Scient. upsal., Ser. 4 8(no. 2): 66 (1932)
Maharachchikumbura et al. (2016b) accepted Xylariales as the only order in the subclass
Xylariomycetidae, which comprised 22 families. However, Samarakoon et al. (2016) and
Hongsanan et al. (2017) provided phylogenetic and divergence time estimations for the subclass
Xylariomycetidae and accepted Amphispheriales and Xylariales. Wijayawardene et al. (2018a)
accepted 12 families in Xylariales, including Myelospermataceae, which lacks molecular data.
However, the phylogenetic placements of the families Oxydothidaceae and Polystigmataceae are
uncertain in two different analyses (Figs 1, 4). However, following the previous phylogenetic
treatments (Konta et al. 2016, Dayarathne et al. 2017) we accept Oxydothidaceae and
Polystigmataceae as families of Xylariales. We accept 15 families in Xylariales, including
Barrmaeliaceae,
Cainiaceae,
Clypeosphaeriaceae,
Coniocessiaceae,
Diatrypaceae,
Graphostromataceae, Hansfordiaceae, Hypoxylaceae, Induratiaceae, Lopadostomataceae,
Microdochiaceae, Polystigmataceae, Requienellaceae, Xylariaceae and Zygosporiaceae, while
Myelospermataceae is placed in the Xylariomycetidae families, incertae sedis. The divergence time
for Xylariales is estimated as 147 MYA (Fig. 2). Here we accept 160 genera in Xylariales.
The families and genera of Sordariomycetes
In this section we list the families of Sordariomycetes in alphabetical order, providing
descriptions, brief history and notes on the ecological and economic significance. The accepted
genera are listed, starting with the type genus and rest alphabetically, with brief notes.
Acrodictyaceae J.W. Xia & X.G. Zhang, Scientific Reports 7 (no. 7888): 2 (2017)
Index Fungorum number: IF818894; Facesoffungi number: FoF06600; 25 species.
Saprobic on terrestrial or freshwater habitats. Sexual morph: Undetermined. Asexual morph:
Hyphomycetous. Colonies on natural substrate effuse, hairy or velvety, yellowish brown to black.
Mycelia immersed, composed of pale brown, septate, branched hyphae. Conidiophores
macronematous, mononematous, erect, cylindrical, greenish brown to brown, septate, mostly
unbranched, straight or broadly curved, thick-walled. Conidiogenous cells monoblastic, integrated,
terminal, cylindrical, greenish brown to brown, thick-walled. Conidia acrogenous, solitary, dry,
greenish brown to dark brown, obovoid to pyriform, muriform, deeply constricted at septa; conidial
secession schizolytic (adapted from Xia et al. 2017).
Type genus – Acrodictys M.B. Ellis
Notes – Xia et al. (2017) studied acrodictys-like species and established Acrodictyaceae
based on molecular data. Acrodictyaceae is a monotypic family containing the type genus
Acrodictys. Acrodictyaceae species are characterized by macronematous, mostly unbranched
conidiophores and obovoid to pyriform, muriform conidia. Sexual morphs in this family have not
been reported.
Ecological and economic significance of Acrodictyaceae
Acrodictyaceae species often occur as saprobes in both freshwater and terrestrial habitats. As
decomposers, species are involved in nutrient cycling. Moreover, the ecological role of freshwater
fungi has increasingly been recognized (Hyde et al. 2016a). Freshwater fungi can cause soft rot
decay (Zare-Maivan & Shearer 1988b, Yuen et al. 2000, Bucher et al. 2004), therefore, lignicolous
410
freshwater species, such as A. aquatica, may play important functional roles in nutrient cycling of
woody material.
Genus included in Acrodictyaceae
Acrodictys M.B. Ellis, Mycological Papers 79: 5 (1961)
Index Fungorum number: IF7034; 25 morphological species (Species Fungorum 2020), 10
species with sequence data (Luo et al. 2019).
Type species – Acrodictys bambusicola M.B. Ellis
Notes – Ellis (1961) introduced Acrodictys, typified by A. bambusicola. Baker et al. (2002)
reviewed the status of Acrodictys and refined the generic concept in a stricter sense. Based on the
narrower concept, some species were excluded from Acrodictys and transferred to new genera, for
example, Junewangia, Rhexoacrodictys, Pseudoacrodictys and Ramoacrodictys (Baker et al. 2002,
Baker & Morgan-Jones 2003, Zhao et al. 2009). Seifert et al. (2011) provided a taxonomic key to
Acrodictys and similar genera. The sexual morph of A. aquatica is illustrated in this entry (Fig. 25).
Amphisphaeriaceae G. Winter, Rabenh. Krypt.-Fl., Edn 2 (Leipzig) 1.2: 259 (1885)
Index Fungorum number: IF80452; Facesoffungi number: FoF00673; 105 species.
Saprobic on leaves, twigs, and branches of deciduous shrubs and trees, conifers or
monocotyledons and occasionally hemibiotrophic or necrotrophic, appearing as slightly raised,
black dots on host surface, often surrounded by a darkened area. Sexual morph: Pseudostromata
when present made up of host cells and brown to black fungal hyphae, solitary, uni- to bi-loculate,
hemisphaerical, initially appearing as raised, pale brown areas, with small, black dots at the center,
becoming dark at maturity, glabrous. Ascomata perithecial, scattered to clustered, immersed in host
cortex, becoming raised, subglobose to lenticular, glabrous, dark brown to black, vegetative hyphae
surrounding the locules, ostioles individual, central. Papilla erumpent through host surface,
internally lined by hyaline, filamentous periphyses. Peridium of unequal thickness, thinner at the
base, thicker towards the mid-upper section of the ascomata, comprising several layers of dark
brown pseudoparenchymatous cells, the outer layer of thick-walled, brown cells of textura
prismatica, the inner layer of flattened, hyaline cells of textura prismatica. Paraphyses numerous,
septate, filamentous. Asci 8-spored, unitunicate, cylindrical, short pedicellate, apex rounded, with
J+ or J-, apical ring. Ascospores overlapping uniseriate, light to dark brown, ellipsoidal to fusiform,
1-septate. Asexual morph: Coelomycetous. Conidiomata solitary or aggregated, globose, dark
brown. Peridium comprising thick-walled, septate, brown mycelium. Conidiophores dichotomously
branched, septate, thick-walled, smooth, hyaline. Conidiogenous cells elongated, wide at the base
and narrow at the tip, thin-walled, hyaline. Conidia hyaline, 1-celled, smooth-walled, elongate to
fusiform, narrow at both ends (adapted from Maharachchimbura et al. 2016).
Type genus – Amphisphaeria Ces. & De Not.
Notes – Amphisphaeriaceae was introduced by Winter (1885a) as ‘Amphisphaerieae’ and
later established as Amphisphaeriaceae to accommodate the type genus Amphisphaeria and similar
genera, viz. Caryospora, Ohleria, Strickeria, Trematosphaeria and Winteria (Winter 1887). Kirk et
al. (2008) confirmed the family within the Xylariales comprising 32 genera with 499 species.
Senanayake et al. (2015) accepted only the type genus Amphisphaeria while excluding all other
genera based on their morphology and phylogeny. Some were transferred to Bartaliniaceae,
Discosiaceae, Iodosphaeriaceae, Sporocadaceae, and Phlogicylindriaceae. Jaklitsch et al. (2016b)
and Maharachchikumbura et al. (2016b) accepted only Amphisphaeria and Lepteutypa as belonging
to the family based on molecular phylogenetic studies.
Earlier, Amphisphaeriaceae was classified in Xylariales and thought to share a close
relationship with Cainiaceae, Clypeosphaeriaceae, and Hyponectriaceae (Jeewon 2002). However,
Senanayake et al. (2015) resurrected Amphisphaeriales, which was introduced by Eriksson &
Hawksworth (1986a) based on combined gene analyses. Amphisphaeriales was not followed in
Maharachchikumbura et al. (2016b) and Jaklitsch et al. (2016b). Samarakoon et al. (2016) and
Hongsanan et al. (2017) provided divergence estimates as additional information for
411
Amphisphaeriales. Wijayawardene et al. (2018a) accepted three genera Amphisphaeria,
Griphosphaerioma, and Lepteutypa in Amphisphaeriaceae (Amphisphaeriales).
Figure 25 – Acrodictys aquatica (Material examined – THAILAND, Chiang Rai Province, Muang,
Ban Nang Lae Nai, on decaying wood submerged in a freshwater stream, 31 December 2016, NG
Liu, CR004, MFLU 18-0040, holotype). a Specimen. b, c Colonies on substrate. d-f Conidiophores
and conidia. g, h Conidiogenous cells and conidia. i-m Conidia. Scale bars: b = 100 μm, c = 50 μm,
d-f = 15 μm, g, h = 10 μm, i-m = 5 μm.
412
Ecological and economic significance of Amphisphaeriaceae
Amphisphaeriaceous taxa play a major role as saprobes on dead plant material in terrestrial,
aquatic (submerged wood) and marine habitats (Wang et al. 2004, Liu et al. 2015, Senanayake et al.
2015, 2019, Jaklitsch et al. 2016b, Phookamsak et al. 2018, Samarakoon et al. 2018). Some species
(e.g. Lepteutypa hederae) are important decomposers of hardwood (Rappaz 1995).
Genera included in Amphisphaeriaceae
Amphisphaeria Ces. & De Not., Comm. Soc. crittog. Ital. 1(4): 223 (1863)
Index Fungorum number: IF173; 88 morphological species (Species Fungorum 2020), 18
morphological species (Senanayake et al. 2019), 6 species with sequence data.
Type species – Amphisphaeria umbrina (Fr.) De Not.
Notes – Amphisphaeria was introduced by Cesati & De Notaris (1863). Wang et al. (2004)
re-examined more than 170 type species described as Amphisphaeria and accepted 12 species.
Senanayake et al. (2015) reviewed the phylogenetic status of the genus, which is sister to
Lepteutypa (Jaklitsch et al. 2016b). Based on phylogenetic investigations, other six new species
have been added: A. doidgeae (Marincowitz et al. 2008), A. sorbi (Liu et al. 2015), A. mangrovei
(Phookamsak et al. 2019), A. flava, A. thailandica (Samarakoon et al. 2019a) and A. acericola
(Senanayake et al. 2019) respectively. The sexual morph of A. acericola and asexual morph of A.
sorbi are illustrated in this entry (Figs. 26, 27).
Figure 26 – Amphisphaeria sorbi (ex-type culture MFLUCC 13-0721, redrawn from Liu et al.
2015). a, b Conidiophores and conidiogenous cells with attached conidia. c Conidia. Scale bars: a-c
= 10 μm.
Griphosphaerioma Höhn., Ber. dt. bot. Ges. 36(7): 312 (1918)
Index Fungorum number: IF2140; 2 morphological species (Species Fungorum 2020).
Type species – Griphosphaerioma symphoricarpi (Ellis & Everh.) Höhn.
Notes – Wijayawardene et al. (2018a) accepted Griphosphaerioma in Amphisphaeriaceae
based on the known asexual morphs. Griphosphaerioma is characterised by unitunicate, cylindrical
asci with a J-, apical ring, and uniseriate, ellipsoidal, hyaline, 1-septate or rarely 2- to 3-septate
ascospores (Kang et al. 1999a). Ono & Kobayashi (2003) introduced G. zelkovicola from bark of
Zelkova serrata with a Sarcostroma asexual morph. However, the phylogenetic affinity of the
genus is still in doubt due to the lack of cultures and DNA sequences.
Lepteutypa Petr., Ann. Mycol. 21(3/4): 276. (1923)
Index Fungorum number: IF2758; 15 morphological species (Jaklitsch et al. 2016b, Species
Fungorum 2020), 5 species with sequence data.
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Figure 27 – Amphisphaeria acericola (Material examined – ITALY, Province of Forlì-Cesena,
Galeata, Strada San Zeno, on a branch of Acer campestre, 26 March 2014, E. Camporesi, IT 1779,
MFLU 16-2479, holotype). a, b Ascomata on the substrate. c, d Cross-sections of ascoma. e
Ostiole. f Peridium. g Paraphyses. h J+, apical ring. i-m Asci (m stained in Congo Red). n-r
Ascospores. Scale bars: a, b = 500 µm, c, d = 200 µm, e = 50 µm, f, i-m = 20 µm, g = 10 µm, h, n-r
= 5 µm.
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Type species – Lepteutypa fuckelii (G.H. Otth) Petr.
Notes – Lepteutypa species are saprobes with a worldwide distribution (Wijayawardene et al.
2017a). Jaklitsch et al. (2016b) revisited the genus, designated the neotype for L. fuckelii, and
introduced a new species, L. sambuci. Luo et al. (2019) introduced L. aquatica on decaying wood
submerged in freshwater habitats. Some of the sequences in GenBank are similar to species of
those members of Sporocadaceae and need further studies with molecular data.
Amplistromataceae Huhndorf, A.N. Mill., Greif & Samuels, Mycologia 101(6): 905 (2009)
Index Fungorum number: IF513238; Facesoffungi number: FoF00618; 31 species.
Saprobic on bark or wood or in highly acidic soil. Sexual morph: Stromata superficial,
turbinate, obovoid to irregularly pulvinate, texture soft or firm with polystichous or monostichous
ascomata, or stromata absent and ascomata single to clustered, with or without a hyphal subiculum.
Ascomata globose or subglobose, with a cylindrical or rostrate neck. Paraphyses abundant,
tapering, of broad cells. Asci 8-spored, unitunicate, cylindrical to clavate, pedicellate, apical ring J-.
Ascospores uniseriate, unicellular, tiny, globose, hyaline. Asexual morph: Hyphomycetous.
acrodontium-like. Conidiophores semi-macronematous or macronematous. Conidiogenous cells
lageniform to ampulliform, elongating sympodially forming a rachis; conidiogenesis holoblasticdenticulate. Conidia single, globose, or ellipsoidal to lacrimose, hyaline to pale brown (adapted
from Maharachchimbura et al. 2016b).
Type genus – Amplistroma Huhndorf, A.N. Mill., Greif & Samuels
Notes – Amplistromataceae was introduced by Huhndorf et al. (2009) to accommodate
Amplistroma and Wallrothiella and referred to Sordariomycetidae, family incertae sedis.
Wallrothiella differs from Amplistroma mainly by the lack of stromatal structures (Huhndorf et al.
2009). Maharachchikumbura et al. (2015) introduced Amplistromatales with two families
Amplistromataceae and Catabotryaceae. Phylogenetic analyses based on LSU, SSU, tef1 and rpb2
markers revealed that Amplistromatales is distinct from Meliolales and other members of
Sordariomycetidae and was referred to Sordariomycetes incertae sedis until further data is
available. Amplistromatales was accepted in Diaporthomycetidae, Sordariomycetes (Hongsanan et
al. 2017, Wijayawardene et al. 2017a). In this entry, the asexual morphs of Acidothrix acidophila
and sexual morph of Amplistroma erinaceum are illustrated (Figs. 28, 29).
Ecological and economic significance of Amplistromataceae
Amplistromataceae species play a major role as saprobes thriving on decaying wood or bark
(Amplistroma and Wallrothiella) and in extremely acidic soil (pH<3) (Acidothrix) (Huhndorf et al.
2009, Hujslová et al. 2014). However, there is potential to use the acidophilic fungi in
bioremediation processes (Hujslová et al. 2019). We could not find any published chemical
profiling associated with amplistromataceous taxa, and therefore the secondary metabolites
associated with these species adapted to extremely variable environments might be worth
investigating.
Genera included in Amplistromataceae
Acidothrix Hujslová & M. Kolařík, Mycol. Progr. 13(3): 824 (2014)
Index Fungorum number: IF805194; 1 species with sequence data.
Type species – Acidothrix acidophila Hujslová & M. Kolařík
Notes – Acidothrix was isolated from acidic soil from the Czech Republic (Hujslová et al.
2014). It is known only as the asexual morph (Fig. 28) and is morphologically similar to
acrodontium-like asexual morphs reported for Amplistroma and Wallrothiella.
Amplistroma Huhndorf, A.N. Mill., Greif & Samuels, Mycologia 101(6):907 (2009)
Index Fungorum number: IF513239; 9 morphological species (Species Fungorum 2020), 6
species with sequence data.
Type species – Amplistroma carolinianum Huhndorf, A.N. Mill., Greif & Samuels
415
Notes – Amplistroma was introduced by Huhndorf et al. (2009) as sister to Wallrothiella.
Members are saprobes with hyphomycetous, acrodontium-like asexual morphs.
Figure 28 – Acidothrix acidophila (ex-type culture CBS 136259, redrawn from Hujslová et al.
2014). a-c Conidiophores and conidia. d Conidia. Scale bars: 10 μm.
Wallrothiella Sacc., Syll. fung. (Abellini) 1: 455 (1882)
Index Fungorum number: IF5760; 21 morphological species (Species Fungorum 2020), 3
species with sequence data.
Type species – Wallrothiella congregata (Wallr.) Sacc.
Notes – Wallrothiella was introduced by Saccardo (1882) and re-described by Réblová &
Seifert (2004b) based on new collections; a neotype for W. congregata, the type species of the
genus, was proposed. Wallrothiella is characterized by the absence of stromata, globose to
subglobose ascomata with a rostrate neck, sometimes surrounded by a whitish subiculum, and tiny,
globose ascospores in cylindrical asci.
Annulatascaceae S.W. Wong, K.D. Hyde & E.B.G. Jones, Syst. Ascom. 16(1-2): 18 (1998)
Index Fungorum number: IF81939; Facesoffungi number: FoF01212; 38 species.
Saprobic on submerged wood; sometimes on bamboo and on other substrates in terrestrial
habitats. Sexual morph: Ascomata perithecial, scattered or densely aggregated in patches,
immersed, partially superficial, brown to black, ostiolate, unilocular, rarely clypeate, globose to
subglobose, ellipsoidal, smooth or rough-walled with short setose-like hyphae on the outside.
Ostiolar necks cylindrical to conical, black, black at the base and pale yellow or lighter at the apex
in some genera, glabrous, rarely with setae, hyphae or hairs. Peridium carbonaceous or coriaceous
composed of cells of textura angularis, textura intricata, textura epidermoidea or textura
prismatica with compressed cells. Paraphyses numerous, septate, branched or unbranched,
tapering. Asci 8-spored, unitunicate, cylindrical, pedicellate, usually with a massive J-, refractive,
cylindrical to flaring, doughnut-shaped, apical ring. Ascospores usually uniseriate, sometimes
overlapping, hyaline to pale yellow or brown, ellipsoidal to oval or fusiform or lunate or citriform,
unicellular or septate, septa mostly transverse, smooth or roughed, with or without appendages,
mucilaginous sheath and germ pores. Asexual morph: Hyphomycetous. taeniolella-like for
Chaetorostrum where Conidiophores are micronematous, mononematous. Conidia monoblastic,
elongate cylindrical, trans-septate, euseptate, brown, paler near the apex. Conidial secession
schizolytic (adapted from Maharachchimbura et al. 2016b).
Type genus – Annulatascus K.D. Hyde
Notes – Annulatascaceae was introduced by Wong et al. (1998d), with Annulatascus as the
type genus. Maharachchikumbura et al. (2015) formally placed it in Annulatascales based on
phylogenetic analyses. Many genera, having a relatively massive apical ring, were placed in
Annulatascaceae as listed in Maharachchikumbura et al. (2016b). Zhang et al. (2017a) excluded
some genera to other families and Diaporthomycetidae genera incertae sedis based on phylogenetic
416
analyses and divergence estimates. Currently, ten genera are accepted in this study (as listed
below). Annulatascaceae species clustered in a distinct clade, but with low bootstrap support as
shown in Luo et al. (2019) and this study.
Figure 29 – Amplistroma erinaceum (Material examined – PANAMA, La Chorrera, Llanito Verde,
on decaying wood of Anacardium excelsum (Anacardiaceae), E. Esquivel, 5 September 2012, AH
43901, paratype). a, b Herbarium details. c, d Stroma (long ostiolar necks are in black arrows). e
Longitudinal section of stroma. f Peridium. g Longitudinal section through the neck. h Asci. i
Ascospores. j Paraphyses and asci (h, j redrawn from Checa et al. 2013). Scale bars: c = 1000 µm,
d, e = 500 µm, g = 200 µm, f = 50 µm, h, j = 10 µm, i = 5 µm.
Ecological and economic significance of Annulatascaceae
Annulatascaceae species are generally saprobic on submerged wood in freshwater, sometimes
on bamboo and on other substrates in terrestrial habitats (Maharachchikumbura et al. 2016b).
417
Annulusmagnus triseptatus produces soft-rot cavities on balsa wood in culture (Campbell &
Shearer 2004). Species are involved in nutrient cycling (Tsui et al. 2000).
Genera included in Annulatascaceae
Annulatascus K.D. Hyde, Aust. Syst. Bot. 5(1): 118 (1992)
Index Fungorum number: IF25398; 17 morphological species (Species Fungorum 2020), 7
species with sequence data.
Type species – Annulatascus velatispora K.D. Hyde
Notes – Annulatascus is the type genus of Annulatascaceae and was introduced by Hyde
(1992b) based on two tropical freshwater taxa, A. velatispora as the type species and A. bipolaris.
However, the holotype of A. velatisporus was in poor condition as herbarium material had only a
few ascomata. Thus, Dayarathne et al. (2016a) designated an epitype, which was collected from
submerged wood in a river, at Millaa Millaa Falls, north Queensland, Australia, based on
morphological examination and DNA sequences. The genus presently contains 19 epithets (Index
Fungorum 2020) and molecular data has demonstrated that this genus is polyphyletic (AbdelWahab et al. 2011, Luo et al. 2015) and confirmed that placement of species in this genus based
solely on the relatively massive apical ascal ring is not very reliable (Abdel-Wahab et al. 2011, Luo
et al. 2015). In our phylogenetic analyses, Annulatascus nilensis does not cluster within
Annulatascus, but is basal to Ascitendus, Longicollum and Submersisphaeria with low bootstrap
support (Fig. 3). Annulatascus nilensis also formed a weakly-supported clade with members of
Annulatascus (Abdel-Wahab et al. 2011, Luo et al. 2019). Fresh collections and multigene
phylogenies are needed to resolve the taxonomy of this group of fungi. We introduce a new species,
Annulatascus thailandensis (Fig. 30) and redescribe the type species, A. velatisporus (Fig. 31).
Annulatascus thailandensis W. Dong, H. Zhang & K.D. Hyde, sp. nov.
Fig. 30
Index Fungorum number: IF556954; Facesoffungi number: FoF06860
Etymology – In reference to the host location in Thailand, where the holotype was collected.
Holotype – MFLU 18-1556.
Saprobic on decaying wood submerged in freshwater. Sexual morph: Ascomata 170–200 μm
high, 250–300 μm diam, solitary or gregarious, superficial, ellipsoidal or subglobose, black,
coriaceous, with a lateral neck, often lying horizontally or obliquely to the substrate surface. Neck
long or short, black, curving upwards or lying on the host surface, up to 600 μm long. Ostiole
eccentric, lateral, black, periphysate. Peridium 32–37 μm thick at upper part, comprising 5–8 layers
of brown to dark brown, thick-walled, compressed cells of textura angularis or round cells; 75–85
μm thick near the base where below the neck, comprising 12–15 layers of pale brown to brown,
thin-walled, compressed cells of textura angularis or irregular cells; 3–5 layers of hyaline,
compressed cells of textura angularis inwardly. Paraphyses ca 4–5.5 μm diam. near the base,
tapering distally, hypha-like, numerous, hyaline, unbranched, septate, constricted at the septa. Asci
275–375 × 13–15 μm (x̅ = 310 × 13.7 μm, n = 10), 8-spored, unitunicate, cylindrical, apically
rounded, with a long pedicel, up to 130 μm long, with a massive, wedge-shaped, refractive apical
ring, 3.5–4 μm high × 5.5–6 μm wide. Ascospores 28–33 × 10.5–11.5 μm (x̅ = 30.8 × 11.2 μm, n =
10), uniseriate, hyaline, ellipsoidal to fusiform, straight to curved, 1–2-septate, slightly constricted
at the septa, minutely guttulate, thin-walled, with a large, nearly ellipsoidal, mucilaginous sheath,
10–15 μm thick. Asexual morph: Undetermined.
Culture characteristics – On PDA, colony circular with filamentous margin, reaching 30 mm
in 35 days at 25°C, brown to white from above, white to pale brown from below, surface rough,
dry, raised, with dense mycelia, edge filiform.
Material examined – THAILAND, Nakhon Si Thammarat Province, on submerged wood in a
stream, 10 May 2018, W. Dong hat650 (MFLU 18-1556, holotype), ex-type living culture
MFLUCC 18-1248; on submerged wood in a stream, 10 May 2018, W. Dong hat650 (HKAS
105023, isotype), living culture KUMCC 19-0038.
GenBank numbers – ITS: MN733256, LSU: MN733254, SSU: MN733253
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Notes – Annulatascus thailandensis fits well within the generic concept of Annulatascus in
having a typical massive, apical ring and ellipsoidal to fusiform ascospores (Hyde 1992b,
Maharachchikumbura et al. 2016b). In our phylogenetic tree, A. thailandensis clustered with A.
hongkongensis with strong bootstrap support (Fig. 3). However, A. thailandensis has smaller
ascospores (28–33 × 10.5–11.5 μm vs. 35–37.5 × 12.5–15 μm) and longer and thinner asci (275–
375 × 13–15 μm vs. 250–275 × 25–30 μm). The ascospores of A. thailandensis are 1–2-septate,
whereas those of A. hongkongensis are 3-septate (Ho et al. 1999). Annulatascus thailandensis is
also similar to A. saprophyticus in asci and ascospore morphology and they have overlapping
ascospore size (Luo et al. 2015). However, they can be separated by characters of ascomata
(superficial, with long or short necks, lying horizontally or obliquely to the substrate surface vs.
partly immersed, with straight upright necks to the substrate surface), size of asci (275–375 × 13–
15 μm vs. 230.5–261 × 13.5–16.5 μm) and septa of ascospores (1–2-septate vs. 0–3-septate) (Luo
et al. 2015).
Annulusmagnus J. Campb. & Shearer, Mycologia 96(4): 826 (2004)
Index Fungorum number: IF28870; 1 species with sequence data.
Type species – Annulusmagnus triseptatus (S.W. Wong, K.D. Hyde & E.B.G. Jones) J.
Campb. & Shearer
Notes – The genus comprises a single species which occurs on submerged corticated or
decorticated wood in freshwater habitats and causes soft-rot cavities on balsa wood in culture
(Campbell & Shearer 2004)
Aqualignicola Ranghoo, C.K.M. Tsui & K.D. Hyde, Mycol. Res. 105(5): 628 (2001)
Index Fungorum number: IF28493; 2 morphological species (Species Fungorum 2020).
Type species – Aqualignicola hyalina Ranghoo, C.K.M. Tsui & K.D. Hyde
Notes – Aqualignicola was introduced by Ranghoo et al. (2001) to accommodate a single
species A. hyalina which was isolated from submerged wood in freshwater in Hong Kong, China.
Another species, A. vaginata, was introduced in the genus from submerged wood in Yunnan, China
(Hu et al. 2012). There is no sequence data for either species and the family placement requires
confirmation.
Ascitendus J. Campb. & Shearer, Mycologia 96(4): 829 (2004)
Index Fungorum number: IF28871; 2 species with sequence data.
Type species – Ascitendus austriacus (Réblová, Winka & Jaklitsch) J. Campb. & Shearer
Notes – Ascitendus was introduced by Campbell & Shearer (2004) to accommodate
Ascolacicola austriaca based on morphology and phylogenetic analyses of 28S rDNA sequence
data. Ascitendus aquaticus was introduced in the genus by Hyde et al. (2018b). Ascitendus
austriacus occurs on submerged corticated or decorticated wood in freshwater habitats and causes
soft-rot cavities on balsa wood in culture (Campbell & Shearer 2004).
Ayria Fryar & K.D. Hyde, Cryptog. Mycol. 25(3): 248 (2004)
Index Fungorum number: IF28835; 2 morphological species (Species Fungorum 2020).
Type species – Ayria appendiculata Fryar & K.D. Hyde
Notes – Ayria was collected from decaying wood submerged in brackish and seawater.
Another species, A. nubispora Raja, Ferrer & Shearer, was isolated from decorticated woody debris
submerged in a lake (Raja et al. 2009). There is no sequence data for either species and the family
placement needs confirmation.
Cataractispora K.D. Hyde, S.W. Wong & E.B.G. Jones, Mycol. Res. 103(8): 1019 (1999)
Index Fungorum number: IF28328; 5 morphological species (Species Fungorum 2020), 3
species with sequence data.
Type species – Cataractispora aquatica K.D. Hyde, S.W. Wong & E.B.G. Jones
419
Figure 30 – Annulatascus thailandensis (MFLU 18-1556, holotype). a-c Ascomata superficial on
host surface. d Vertical section of ascoma. e Structure of peridium near the base. f Structure of
peridium at upper part. g-j Unitunicate asci. k Paraphyses. l-o Ascospores. p Ascospore mounted in
India ink. q Colony on PDA (from front). r Colony on PDA (from reverse). Scale bars: d, e, g-i =
50 μm, f = 30 μm, j-p = 20 μm.
420
Figure 31 – Annulatascus velatisporus (Material examined – AUSTRALIA, North Queensland, on
submerged wood in a small river, 14 April 2015, S. Fryar & B. Cawson MR102A, MFLU 16-2204,
epitype) a Herbarium material. b, c Appearance of black ascomata on host. d Vertical section of
ascoma. e Structure of peridium. f Paraphyses. g-i Unitunicate asci. j-l Ascospores. Scale bars: d =
100 μm, e = 50 μm, f, i-l = 10 μm, g, h = 20 μm.
421
Notes – Cataractispora was introduced by Hyde et al. (1999a) to accommodate four
freshwater species, C. appendiculata (Brunei), C. aquatica (Brunei), C. bipolaris (Queensland) and
C. viscosa (Hong Kong), based on their unique appendage ontogeny. Ho et al. (2004) introduced a
fifth freshwater species, C. receptaculorum, from Hong Kong. The LSU sequence data of C.
receptaculorum is identical to an invalid species C. recepticuli, which was accommodated in
Pseudoproboscisporaceae by Zhang et al. (2017a) and they stated that it appears to be an invalidly
published synonym of Pseudoproboscispora based on morphology. The family placement of
Cataractispora needs confirmation with sequence data from the type species.
Chaetorostrum Zelski, Raja, A.N. Mill. & Shearer, Mycosphere 2(5): 594 (2011)
Index Fungorum number: IF563571; 1 morphological species.
Type species – Chaetorostrum quincemilense Zelski, Raja, A.N. Mill. & Shearer
Notes – The monotypic genus, Chaetorostrum was introduced by Zelski et al. (2011b) from
woody debris in a semi-aquatic, intermittent stream in Peru. It is the first member of the
Annulatascaceae that produces its asexual morph (taeniolella-like) in culture (Zelski et al. 2011b).
There is no sequence data for this species and the family placement needs confirmation.
Longicollum Zelski, F.R. Barbosa, Raja, A.N. Mill. & Shearer, Mycosphere 2(5): 540 (2011)
Index Fungorum number: IF563241; 1 species with sequence data.
Type species – Longicollum biappendiculatum Zelski, F.R. Barbosa, Raja, A.N. Mill. &
Shearer
Notes – The monotypic genus, Longicollum was introduced by Zelski et al. (2011a) from
submerged woody debris in Peru. It appears to be widely distributed in freshwater in the
Neotropics, i.e. Brazil, Costa Rica, Peru, and USA (Zelski et al. 2011a).
Submersisphaeria K.D. Hyde, Nova Hedwigia 62(1-2): 172 (1996)
Index Fungorum number: IF27640; 5 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Submersisphaeria aquatica K.D. Hyde
Notes – Sequences of a collection of S. aquatica (strain A95-1B) are available in GenBank
and placed the genus in Annulatascaceae (Zhang et al. 2017a; Fig. 3).
Vertexicola K.D. Hyde, Ranghoo & S.W. Wong, Mycologia 92(5): 1019 (2000)
Index Fungorum number: IF28465; 2 morphological species (Species Fungorum 2020)
Type species – Vertexicola caudata K.D. Hyde, S.W. Wong & Ranghoo
Notes – Vertexicola was introduced by Ranghoo et al. (2000) to accommodate V. caudata
which was isolated from submerged wood in China and Philippines. Vertexicola was listed in
Annulatascaceae by Maharachchikumbura et al. (2015) and Wijayawardene et al. (2018a).
However, it was placed in Diaporthomycetidae genera incertae sedis based on sequence data of one
strain of V. confuse, which was invalidly published (Nom. inval. in Index Fungorum 2020). Due to
absence of morphological data, the validity of sequence data used by Zhang et al. (2017a) is
unknown. We therefore, retain Vertexicola in Annulatascaceae until the sequence data of the type
species V. caudata are available.
Apiosporaceae K.D. Hyde, J. Fröhl., Joanne E. Taylor & M.E. Barr, Sydowia 50(1): 23 (1998)
Index Fungorum number: IF81935; Facesoffungi number: FoF00629; 114 species.
Saprobic or pathogenic on leaves, stems and roots of monocotyledons, mostly grasses, or
endophytic on plant tissues, lichens, and marine algae, occasionally infecting humans, or isolated
from soil. Sexual morph: Pseudostromata visible as raised, linear, blackened areas on the host
surface, with neck and upper surface visible through splits in the host tissue, mostly gregarious,
fusiform, ellipsoid to irregular, black, cells between ascomata and darkened layer above usually
thick, composed of brown cells of textura angularis. Ascomata solitary or usually gregarious in
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linear groups, immersed under pseudostromata, globose to subglobose, membranous, papillate,
ostiole with periphyses. Peridium composed of several layers, comprising smallish, brown, reddish
brown to hyaline cells of textura angularis. Paraphyses dense, hypha-like, long, broad, septate and
branched. Asci 8-spored, unitunicate, broad cylindrical to clavate or subglobose, without an apical
ring. Ascospores overlapping 1–3-seriate to irregularly arranged, hyaline, apiosporous, with a large,
straight or curved, upper cell and smaller lower cell, usually surrounded by a gelatinous sheath.
Asexual morph: Coelomycetous or hyphomycetous. Conidiomata sporodochial or acervuli, solitary
to gregarious, immersed, erumpent from host tissue when mature, irregular, black, carbonaceous,
coriaceous. Conidiomata wall composed of several layers of dark brown to hyaline cells of textura
angularis, thick at sides, thin at upper and lower walls. Setae absent, or occasionally present,
intermingled among conidiophores. Conidiophore mother cells ampulliform, verrucose wall,
producing a single conidiophore. Conidiophores hyphoid, cylindrical, 1–2-septate, verrucose,
flexuous. Conidiogenous cells basauxic, cylindrical, with or without verrucose wall. Conidia
globose to subglobose, dark brown, smooth-walled or with minute wall ornamentations, with a
truncate basal scar (adapted from Maharachchimbura et al. 2016b).
Type genus – Arthrinium Kunze
Notes – Apiosporaceae was introduced to accommodate taxa which produce apiospores and a
basauxic, arthrinium-like conidiogenesis (Samuels et al. 1981, Hyde et al. 1998a, Bahl 2006,
Senanayake et al. 2015, Dai et al. 2017, Wang et al. 2018, Pintos et al. 2019). Kirk et al. (2008),
Senanayake et al. (2015), Maharachchikumbura et al. (2016b) and Wijayawardene et al. (2018a)
accepted six genera viz. Appendicospora, Arthrinium (= Apiospora), Dictyoarthrinium, Endocalyx,
Scyphospora and Spegazzinia in this family.
Appendicospora was placed in this family based on its apiospores, and it differs from
Arthrinium in having ascospores with a bifurcate appendage at the lower cell (Hyde 1995b). This
genus includes two species, A. coryphae, and A. hongkongensis, however, only A. hongkongensis
and an unidentified species with sequence data. A blast search of SSU sequences in GenBank
shows Appendicospora is similar to Arthrinium, however, it more similar to Oxydothis based on the
LSU rDNA sequences. Phylogenetic analyses of Appendicospora are required to clarify its
relationship within Apiosporaceae. Dictyoarthrinium is a hyphomycetous genus with basauxic
conidiophores and conidiogenous cells producing 4-celled, cruciately septate conidia. Endocalyx
was established in Apiosporaceae on the basis of its basauxic conidiophores, however, species of
Endocalyx usually produce funnel-shaped to elongated cupulate sporodochia, which are not
observed in other genera of this family (Seifert et al. 2011). Spegazzinia produces basauxic
conidiophores and conidiogenous cells with a basal mother cell and stellate to disc-shaped conidia
with radiating spine-like appendages (Seifert et al. 2011). Tanaka et al. (2015) transferred
Spegazzinia to Didymosphaeriaceae based on the establishment of two species phylogenetically
assigned in this family. Two more species were included in Didymosphaeriaceae based on
morphology and phylogenetic analysis (Thambugala et al. 2017, Jayasiri et al. 2019), thus this
genus is no longer included in Apiosporaceae. Scyphospora was introduced by Kantschaveli (1928)
and typified by S. phyllostachydis. Index Fungorum (2020), however synonymized the type species
under Arthrinium hysterinum probably based on morphology. The hand drawings of both epithets
provided by Nag Raj (1974) and Kirk (1986) also show they are the same species. Therefore, we
accept the combination as in Index Fungorum (2020) and regard Scyphospora as a synonym of
Arthrinium. Nigrospora was introduced by Zimmermann (1902) and redrawn by Seifert et al.
(2011). Wang et al. (2017) transferred Nigrospora to Apiosporaceae based on morphology and
phylogeny. Thus, we conclude that Apiosporaceae accommodates five genera. We illustrate a new
species, Arthrinium neogarethjonesii with sexual and asexual morphs.
Ecological and economic significance of Apiosporaceae
Arthrinium has a widespread distribution and is an important genus comprising species
reported as endophytes or pathogens. Some species cause scabs or spots on leaves or culms of
crops, or even cause damping-off, such as Arthrinium sacchari (Speg.) M.B. Ellis (Mavagani et al.
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2007). Some species have been reported as human pathogens; for example, cutaneous mycosis is
caused by A. phaeospermum (Corda) M.B. Ellis (Rai 1989). Some species such as Arthrinium
arundinis (Corda) Dyko & B. Sutton, have been utilized in the pharmaceutical industry as
antifungal agents.
Genera included in Apiosporaceae
Appendicospora K.D. Hyde, Sydowia 47(1): 31 (1995)
Index Fungorum number: IF27559; 2 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Appendicospora coryphae (Rehm) K.D. Hyde
Notes – Appendicospora is similar to Arthrinium in having apiospores. However, it differs in
its ascospores having basal bifurcate appendages. Appendicospora coryphae was originally
collected from a palm in the Philippines. A second species, A. hongkongensis described by Yanna
et al. (1997) was collected from Hong Kong.
Arthrinium Kunze, Mykologische Hefte (Leipzig) 1: 9 (1817)
Index Fungorum number: IF7214; 73 morphological species (Kirk et al. 200, Index
Fungorum 2020), 57 species with sequence data.
Type species – Arthrinium caricicola Kunze & J.C. Schmidt
Notes – Crous & Groenewald (2013) showed that Apiospora is the sexual morph of
Arthrinium. Crous & Groenewald (2013) placed the younger sexual typified name, i.e. Apiospora
under the older asexual typified name. This adoption was followed in subsequent publications by
Dai et al. (2016), Maharachchikumbura et al. (2016b) and Wijayawardene et al. (2016b). Molecular
data has proved that Arthrinium is a speciose genus (Dai et al. 2017, Wang et al. 2018, Pintos et al.
2019). We introduce a new holomorph species, Arthrinium neogarethjonesii from bamboo.
Arthrinium neogarethjonesii D.Q. Dai & K.D. Hyde, sp. nov.
Fig. 32
Index Fungorum number: IF555481; Facesoffungi number: FoF05112
Etymology – Based on the similarities with species Arthrinium garethjonesii.
Holotype – HKAS 102408.
Saprobic on dead bamboo culms. Sexual morph: Stromata 1–2 mm long, 0.25–0.5 mm wide,
175–250 μm high, scattered to gregarious, immersed, becoming slightly erumpent through the host
tissue, black to dark brown, fusiform to elongated ellipsoid, with a central slit-like opening at the
top. Ascomata perithecial, 125–230 μm high, 120–230 μm diam., arranged in rows, clustered,
gregarious, with 2–7 perithecia forming groups immersed in stromata and surrounding by stromatic
tissue, ellipsoidal to subglobose, or obpyriform with a slightly flattened base, brown to reddishbrown, membranous. Ostiole raised from centre of perithecia, internally lined with periphyses.
Peridium 15–25 μm thick, composed of dark brown to reddish brown to hyaline cells of textura
angularis. Stromatic cells between perithecia 150–350 μm wide, comprising host and fungal
tissues, composed of elongated, brown cells of textura angularis. Paraphyses dense, long, 3.5–6
μm wide, septate, un-branched with the wider base and thinner apex. Asci 95–125 × 20–25 μm (x̅ =
97.6 × 21.3 μm, n = 20), 8-spored, unitunicate, clavate, apedicellate, apically rounded. Ascospores
25–30 × 9.5–11 μm (x̅ = 29.1 × 10.3 μm, n = 20), 2-seriate, overlapping, 1-septate, ellipsoidal, with
a small lower cell and a large upper cell, with many guttules, hyaline, smooth-walled, with a thick
gelatinous sheath, 3–10 µm wide. Asexual morph: Coelomycetous or Hyphomycetous on
substrates, Conidiomata 450–600 µm wide × 1–1.5 mm long, 150–250 µm high, solitary to
gregarious, immersed, then raised from host tissue when mature, ellipsoid to irregular, black,
coriaceous. Conidiomata wall composed of one layer of dark brown to brown to hyaline cells of
textura angularis. Conidiophores mother cells 4.5–6 × 3.5–4.5 µm ( x = 5.4 × 4.3 µm, n = 20),
cylindrical, aseptate. Conidiogenous cells 10–48 × 4–5.5 µm ( x = 35.4 × 4.3 µm, n = 20),
basauxic, cylindrical, discrete, smooth-walled. Conidia 20–35 × 15–30 µm ( x = 28.5 × 25.6 µm, n
= 20), globose to subglobose, dark brown, smooth-walled, with a truncate basal scar.
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Figure 32 – Arthrinium neogarethjonesii (HKAS 102408, holotype). a-c Appearance of stromata
on host. d Appearance of conidiomata on host. e Section of ascostroma. f Section of ascoma. g
Stromatic tissue. h-j Asci. k Paraphyses. l, m Ascospores surrounded by a gelatinous sheath. n
Conidia. o-r Conidia and conidiogenous cells. Scale bars: a = 1 mm, b-d = 5 µm, e = 100 µm, f-h =
50 µm, i-r = 10 µm.
Culture characters – Ascospores germinating on WA within 24 h and germ tubes developing
from the upper cell. Colonies on PDA reaching 50 mm in 1 week at 27°C, in dark, cottony, circular,
with irregular edge, white from above and below, becoming dark from centre after 2 weeks.
Mycelium superficial to immersed in media, with branched, septate, smooth hyphae.
Material examined – CHINA, Yunnan Province, Xishuangbanna Topical Botanical Garden,
Bamboo Collection, on bamboo culm, 2 April 2017, Dong-Qin Dai DDQ00403, HKAS 102408,
holotype; ex-type living culture KUMCC 18-0192.
GenBank numbers – ITS: MK070897, LSU: MK070898, SSU: MK070899.
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Notes – This species is similar to Arthrinium garethjonesii which was introduced by Dai et al.
(2016) based on morphological and phylogenetic analyses, and is only known from the southwest
of China. However, A. neogarethjonesii has smaller asci (95–125 × 20–25 μm vs. 125–154 × 35–42
μm) and ascospores (25–30 × 9.5–11 μm vs. 30–42 × 11–16 μm), and the conidia are larger when
compared with most Arthrinium species. Based on a Blast search using the ITS sequence in NCBI’s
GenBank, the closest hits with highest similarity is A. garethjonesii (GenBank KY356086;
similarity = 575/595 (97%), Gaps = 8/595 (1%)). A similar search using the LSU sequence showed
highest similarity to A. garethjonesii (GenBank KY356091, similarity = 851/854 (99%), no gaps),
and closest hits with A. ovatum (GenBank NG_042782; similarity = 849/875 (97%), no gaps). The
phylogenetic tree generated from maximum-parsimony analysis indicated that the new species is
close to A. garethjonesii and these two species cluster in two well-supported clades (MP/BYPP
100/1.00) (Fig. 4).
Dictyoarthrinium S. Hughes, Mycol. Pap. 48: 29 (1952)
Index Fungorum number: IF7993; 7 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Dictyoarthrinium quadratum S. Hughes
Notes – Dictyoarthrinium was erected by Hughes (1952) for two hyphomycetes, collected in
Ghana. The genus is characterized by terminal and lateral, septate conidiophores, basauxic
conidiogenous cells and muriform conidia.
Endocalyx Berk. & Broome, J. Linn. Soc., Bot. 15(1): 84 (1876)
Index Fungorum number: IF8158; 7 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Endocalyx thwaitesii Berk. & Broome
Notes – Petch (1908) emended the genus, and considered that E. thwaitesii and E. psilostoma
were identical. Okada & Tubaki (1984) described E. melanoxanthus var. grossus and E.
melanoxanthus var. melanoxanthus. The genus is characterized by funnel-shaped synnemata or
sporodochia, branched conidiophores and dark brown conidia with a germ slit (Seifert et al. 2011).
Nigrospora Zimm., Centbl. Bakt. ParasitKde, Abt. I 8: 220 (1902)
Index Fungorum number: IF9124; 25 species with morphological data (Species Fungorum
2020), 17 species with sequence data.
Type species – Nigrospora panici Zimm.
Notes – Seifert et al. (2011) revised the genus and mentioned Khuskia as its sexual morph.
Maharachchikumbura et al. (2016b) placed Nigrospora in Sordariomycetes, genera incertae sedis.
Two new collections of N. oryzae and N. musae were sequenced (Vu et al. 2019), while Wang et al.
(2017) published 12 new Nigrospora taxa and combined N. vietnamensis under Arthrinium and
placed the genus in Apiosporaceae. Raza et al. (2019) introduced five new species from sugarcane.
The genus is characterized by sphaerical to subsphaerical conidiogenous cells and black and
globose to subglobose conidia (Seifert et al. 2011, Wang et al. 2017).
Apiosporopsidaceae Senan., Maharachch. & K.D. Hyde, Stud. Mycol. 86: 234 (2017)
Index Fungorum number: IF821538; Facesoffungi number: FoF03455; 3 species.
Parasitic on living leaves and twigs. Sexual morph: Ascomata scattered, black, oval to almost
sphaerical, immersed in the leaf tissue beneath a thin, well-developed clypeus, without a neck or
only slightly papillate, periphysate. Peridium comprising 5–6 outer layers of dark, thick-walled
cells of textura angularis and inner, thin-walled, strongly flattened cells of textura angularis.
Paraphyses not seen. Asci 8-spored, unitunicate, short-pedicellate, blunt at the apex with a J-,
apical ring. Ascospores 1–2-seriate, elliptical to fusoid, often slightly flattened on one side,
unicellular, hyaline. Asexual morph: Coelomycetous. Stroma loculate, globose to irregular,
sometimes with beaks. Conidiogenous cells phialidic, short to elongate, simple or branched.
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Conidia oblong or cylindrical to allantoid, 1-celled, hyaline (adapted from Senanayake et al.
2017a).
Type genus – Apiosporopsis (Traverso) Mariani
Notes – Apiosporopsidaceae was introduced by Senanayake et al. (2017a) to accommodate
Apiosporopsis species which are morphologically distinct from other families in Diaporthales. The
asci initially form a hymenium-like fascicle within a papillate ascomata, asci have a bilobed apical
ring and residual stromatic tissue is present within the locules. This family is supported by
molecular data (Braun et al. 2018, Fan et al. 2018, Senanayake et al. 2017a, 2018, Tian et al. 2018).
Ecological and economic significance of Apiosporopsidaceae
Apiosporopsis species are generally associated with over-wintered plant parts as epifoliar
fungi and rarely on living leaves (Barr 1978, Reid & Dowsett 1990). Apiosporopsis species are
considered as saprobes or mild pathogens (Senanayake et al. 2018).
Genus included in Apiosporopsidaceae
Apiosporopsis (Traverso) Mariani, Atti Soc. ital. Sci. nat. 50: 165 (1911)
Index Fungorum number: IF268; 3 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Apiosporopsis carpinea (Fr.) Mariani
Notes – Traverso (1906) erected Apiosporopsis and Mariani (1911) designated A. carpinea as
the type species. Currently Apiosporopsis comprises three species: A. carpinea, A. coronillae and
A. saccardoana (Senanayake et al. 2018, Index Fungorum 2020). Senanayake et al. (2017b)
illustrated A. carpinea from Austria (IMI 11662), by the ascospore morphology of this collection
(IMI 57187) is quite different to A. carpinea, A. coronillae and A. saccardoana. The collection we
illustrated here has ascospores with a central septum and apical and basal ends with pores.
Apoharknessiaceae Senan., Maharachch. & K.D. Hyde, Stud. Mycol. 86: 234 (2017)
Index Fungorum number: IF821881; Facesoffungi number: FoF03457; 10 species.
Endophytic, saprobic or pathogenic on plants. Sexual morph: Undetermined. Asexual morph:
Coelomycetous. Conidiomata stromatic or eustromatic, subepidermal to immersed, solitary to
gregarious, subglobose to irregular, unilocular, pale brown. Conidiomata wall outer layer
composed of thin-walled, pale brown cells of textura angularis, with inner layer of pale yellow to
hyaline cells. Conidiophores hyaline, septate, cylindrical, and sparingly branched or reduced to
conidiogenous cells. Conidiogenous cells holoblastic, annellidic, cylindrical, lageniform to
ampulliform, hyaline, smooth, invested in mucus. Conidia obclavate, conical, aseptate, pale brown,
with a longitudinal band on the flat surface, thick- and smooth-walled, guttulate, with short hyaline
apiculus, with small globule of mucus on base or obtuse apex with a scar at the base (adapted from
Senanayake et al. 2017a).
Type genus – Apoharknessia Crous & S.J. Lee
Notes – Apoharknessiaceae was introduced by Senanayake et al. (2017a) to accommodate
Apoharknessia and Lasmenia. Lasmenia was previously assigned to Cryphonectriaceae
(Maharachchikumbura et al. 2015, 2016b). However, available sequence data in GenBank for
Lasmenia are not identified at the species level, thus, Wijayawardene et al. (2016b) placed
Lasmenia in Diaporthales genera incertae sedis. However, phylogenetic analysis by Senanayake et
al. (2017a) indicates that Apoharknessia and Lasmenia clearly belong in Diaporthales forming a
well-supported clade.
Ecological and economic significance of Apoharknessiaceae
Lasmenia causes rachis necrosis, flower abortion and necrotic spots on leaves of Nephelium
lappaceum and some Lasmenia species are associated with tropical fruits as pathogens (SerratoDiaz et al. 2011). Apoharknessia forms leaf spots on Eucalyptus species and some are reported
from leaf litter as saprobes (Lee et al. 2004).
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Figure 33 – Apiosporopsis carpinea (IMI 57187). a Herbarium packet. b Ascomata on substrate. c,
d Asci. e-g Ascospores. Scale bars: b = 500 µm, c-d = 10 µm, e-g = 5 µm.
Genera included in Apoharknessiaceae
Apoharknessia Crous & S.J. Lee, Stud. Mycol. 50(1): 239 (2004)
Index Fungorum number: IF500065; 3 species with sequence data.
Type species – Apoharknessia insueta (B. Sutton) Crous & S.J. Lee
Notes – Apoharknessia was established by Lee et al. (2004) to accommodate Harknessia
insueta, and the latter constitutes a distinct phylogenetic lineage from Harknessia sensu stricto.
Crous et al. (2017b) introduced A. eucalyptorum. as a second species. Marin-Felix et al. (2018)
introduced a new species as A. eucalypti. This is a foliar pathogenic genus known from Brazil,
Colombia, Cuba and Mauritius. Apoharknessia is similar to Harknessia, but differs in having a
hyaline, apical apiculus, and lacks prominent fluffy aerial mycelium in culture medium (Lee et al.
2004). In this entry we illustrate Apoharknessia insueta.
Lasmenia Speg., Anal. Soc. cient. argent. 22(4): 199 (1886)
Index Fungorum number: IF8714; 7 morphological species (Species Fungorum 2020),
molecular data available for an unnamed species.
428
Type species – Lasmenia balansae Speg.
Notes – Höhnel (1910) selected Lasmenia balansae as the lectotype and there are 12 species
epithets recorded under Lasmenia (Senanayake et al. 2018). Some Lasmenia species are associated
with tropical fruits as pathogens. Lasmenia was treated as a member in Apoharknessiaceae by
Senanayake et al. (2017a).
Figure 34 – Apoharknessia insueta reproduced from Lee et al. (2004). a Vertical section of
conidioma. b Peridium. c, d Conidiogenous cells and conidia. e Conidia. Scale bars: a = 50 µm, b-e
= 10 µm.
Armatellaceae Hosag., Sydowia 55(2): 165 (2003)
Index Fungorum number: IF82149; Facesoffungi number: FoF00723; 17 species.
Colonies epiphyllous or hypophyllous, mostly confluent. Hyphae superficial, brown, straight
to flexuous, septate, branched, reticulate, with hyphopodia. Hyphopodia alternate to unilateral,
mostly 2-celled, brown, with capitates head cells, with irregular lobes, stalk cells rarely septate.
Sexual morph: Ascomata superficial, globose to subglobose, verrucose. Peridium thick, comprising
black, verrucose outer stratum, and brown inner stratum of textura angularis. Asci 4–8-spored,
unitunicate, subcylindrical to ovoid, or clavate, with ascus wall attenuated or broken when mature.
Ascospores ellipsoidal to oblong, 1-septate, not constricted at the septum, hyaline when young,
becoming brown at maturity. Asexual morph: Undetermined (adapted from Maharachchimbura et
al. 2016b).
Type genus – Armatella Theiss. & Syd.
429
Notes – Armatellaceae was established by Hosagoudar (2003) with 16 species. It differs from
Meliolaceae by its 1-septate ascospores, verrucose ascomata and lacking phialides. There is no
sequence data for Armatellaceae. The taxonomic placement is supported by morphology. Armatella
litseae is illustrated for this entry.
Ecological and economic significance of Armatellaceae
There is no report of Armatellaceae species causing any economic problems. However,
similar to the Meliolaceae, the colonies of Armatellaceae cover the leaf surface, may reduce
photosynthesis and increase the temperature and respiration in those areas, while hyphopodia can
penetrate the leaf surface to gain nutrients from host plants.
Genus included in Armatellaceae
Armatella Theiss. & Syd., Annls mycol. 13(3–4): 235(1915)
Index Fungorum number: IF309; 17 morphological species (Species Fungorum 2020).
Type species – Armatella litseae (Henn.) Theiss. & Syd.
Notes – Armatella was placed in Polystigmellaceae (Dothideales) when introduced by
Theissen & Sydow (1915). Hansford (1946) transferred this genus to Meliolaceae. Hosagoudar
(2003) included this single genus in the newly introduced family Armatellaceae. This was followed
in Hongsanan et al. (2015) and Wijayawardene et al. (2017a, 2018a).
Asterosporiaceae Senan., Maharachch. & K.D. Hyde, Stud. Mycol. 86: 236 (2017)
Index Fungorum number: IF821539; Facesoffungi number: FoF03459; 4 species.
Endophytic or saprobic on Betulaceae, Fagaceae, Juglandaceae and Sapindaceae. Sexual
morph: Undetermined. Asexual morph: Coelomycetous. Conidiomata acervular, subepidermal,
erumpent at maturity, solitary, or occasionally confluent, unilocular, dark brown to black.
Conidiomata wall composed of thin-walled, brown cells of textura angularis. Conidiophores
cylindrical, branched at the base, septate, hyaline to pale brown. Conidiogenous cells holoblastic,
cylindrical, unbranched, integrated, determinate, hyaline to pale brown, smooth. Conidia terminal,
transversely distoseptate, consisting of four arms, with reduced lumina, brown, smooth-walled
(adapted from Senanayake et al. 2017a).
Type genus – Asterosporium Kunze
Notes – Asterosporiaceae is based on the single genus Asterosporium (Senanayake et al.
2017a) and is morphologically distinct from other families in Diaporthales having star-like arms,
bearing brown conidia and acervular conidiomata. Asterosporium species occur in temperate
regions. We illustrate Asterosporium asterospermum.
Ecological and economic significance of Asterosporiaceae
Cankers and branch dieback of beech may be caused by Asterosporium hoffmannii (Pirone
1978). Sieber (2007) reported that A. asterospermum may cause canker on Fagus crenata and F.
sylvatica as mild pathogens. However, most species are saprobes in decaying wood.
Genus included in Asterosporiaceae
Asterosporium Kunze, Flora, Regensburg 1: 225 (1819)
Index Fungorum number: IF7270; 4 morphological species (Species Fungorum 2020), 2
species with sequence data.
Type species – Asterosporium asterospermum (Pers) S. Hughes
Notes – Asterosporium was introduced and typified by A. hoffmannii. However, Hughes
(1958) regarded Stilbospora asterosperma as the older name for this taxon based on morphology.
Hence, a new combination, Asterosporium asterospermum was proposed. However according to
molecular analysis of this study, A. asterospermum and A. hoffmannii are phylogenetically
identical. Hence, we synonymise A. hoffmannii (1819) under A. asterospermum (1801) giving
priority to the older name. The sexual morph of Asterosporium has not been reported.
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Figure 35 – Armatella litseae (Material examined – JAPAN, Tokushima, Prov. Awa, Kigasumi, on
Litsea glauca (Thunb.) Siebold (Lauraceae), 25 December 1897, S. Kusano, S F70331, holotype). a
Label of specimen. b Host leaves. c, d Colonies with ascomata on leaf surface. e The ascoma. f
Hyphae with appressoria. g Ascus. h Ascospore. Scale bars: e = 100 μm, f-h = 20 μm.
Atractosporaceae H. Zhang, K.D. Hyde & Maharachch., Fungal Divers. 85: 88 (2017)
Index Fungorum number: IF553757; Facesoffungi number: FoF03334; 7 species.
Saprobic on submerged deciduous wood. Sexual morph: Ascomata solitary or aggregated in
small groups, immersed or semi-immersed to superficial, astromatic, subglobose to conical, dark
brown to black, with a lateral neck, often lying horizontally on the substrate surface. Ostiole
periphysate. Peridium leathery to fragile, two-layered. Paraphyses abundant, persistent, cylindrical.
Asci 8-spored, unitunicate, pedicellate, apically rounded or obtuse, with a J-, relatively small, apical
ring. Ascospores uniseriate, hyaline, fusiform, aseptate or transversely septate, smooth or
ornamented. Asexual morph: Undetermined (adapted from Zhang et al. 2017a).
Type genus – Atractospora Réblová & J. Fourn.
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Notes – Atractosporaceae was introduced in a new order Atractosporales by Zhang et al.
(2017a) with two genera i.e. Atractospora and Rubellisphaeria. Although Aquaticola
hongkongensis also nested in Atractosporaceae (Zhang et al. 2017a), it has never been validly
published (Ranghoo et al. 1999). This species needs to be recollected and characterized. This
family represents a distinct group of freshwater Sordariomycetes, which is strongly supported by its
stem age, ca 98 MYA in the MCC tree (Hyde et al. 2017a). Atractosporaceae is similar to
Pseudoproboscisporaceae in the ascomata often lying parallel to the host surface. However, the
ascospores of Atractosporaceae lack appendages, unlike in Conlariaceae and
Pseudoproboscisporaceae which have appendages at one or both ends. We illustrate Atractospora
thailandensis.
Figure 36 – Armatella litseae (redrawn from Hongsanan et al. 2015). a Colonies on the leaf. b The
peridium. c Hyphae with appressoria. d Ascospores. e Asci. Scale bars: b = 100 µm, d, e = 20 µm, c
= 10 µm.
Ecological and economic significance of Atractosporaceae
Atractospora and Rubellisphaeria have been reported as saprobic genera on submerged
deciduous wood (Réblová et al. 2016a).
Genera included in Atractosporaceae
Atractospora Réblová & J. Fourn., Mycol. Progr. 15(no. 21): 8 (2016)
Index Fungorum number: IF815026; 6 species with sequence data.
Type species – Atractospora reticulata Réblová & J. Fourn.
Notes – Atractospora was introduced by Réblová et al. (2016a) with three new species and a
new combination. Zhang et al. (2017a) extended the generic description to accommodate taxa with
both thick- and thin-walled ascospores as A. thailandensis was included based on phylogeny and
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otherwise similar morphology. Asexual morphs of this genus have not been reported from the
natural substrate, however, asexual structures were observed in cultures of A. decumbens and
characterized by ellipsoidal to subglobose conidia, brown, aseptate, and thick-walled often forming
in short chains of three to five cells (Réblová et al. 2016a).
Figure 37 – Asterosporium asterospermum (Material examined – ITALY, Forlı`-Cesena Province,
Santa Sofia, near Passo la Calla, on dead branch of Fagus sylvatica L. (Fagaceae), 29 September
2012, E. Camporesi, MFLU 15-3555). a Conidiomata on host substrate. b, c Vertical sections of
conidiomata. d-f Different stages of conidiogenesis. g-l Conidia. Scale bars: a = 1 mm, b = 400 μm,
c = 50 μm, d-f = 20 μm, g-l = 30 μm.
Rubellisphaeria Réblová & J. Fourn., Mycol. Progr. 15(no. 21): 13 (2016)
Index Fungorum number: IF815031; 1 species with sequence data.
Type species – Rubellisphaeria abscondita Réblová & J. Fourn.
Notes – The monotypic genus Rubellisphaeria was isolated from submerged coniferous wood
in a stream in France Réblová et al. (2016a). It formed a well-supported clade with Atractospora in
the phylogenetic analyses of, Luo et al. (2019) and this study (Fig. 3).
Auratiopycnidiellaceae Senan., Maharachch. & K.D. Hyde, Stud. Mycol. 86: 237 (2017)
Index Fungorum number: IF821540; Facesoffungi number: FoF03461; 1 species.
Foliicolous. Sexual morph: Undetermined. Asexual morph: Conidiomata amphigenous,
pycnidial, globose, orange-coloured with dark brown border on leaves. Peridium comprises pale
brown cells of textura angularis. Paraphyses hyaline, subcylindrical, septate, branched or
unbranched, with obtuse apex, constricted at septa. Conidiophores reduced to conidiogenous cells.
Conidiogenous cells hyaline, annellidic, smooth, ampulliform, with terminal truncate locus, thickwalled, sometimes proliferating percurrently. Conidia ellipsoidal, smooth, solitary, median 1septate, constricted at septum, obtuse at apex, truncate at base, thickened, at times with marginal
frill, becoming golden brown at germination and with solitary, brown, wavy, thick, bright banded,
germ-slit along the long-axis of the conidium (adapted from Senanayake et al. 2017a).
Type genus – Auratiopycnidiella Crous & Summerell
Notes – Following molecular analysis, Crous et al. (2012b) introduced Auratiopycnidiella
based on A. tristaniopsis and placed it in Diaporthales genera incertae sedis. Senanayake et al.
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(2017a) introduced Auratiopycnidiellaceae to accommodate Auratiopycnidiella. In this study we
illustrate A. tristaniopsis.
Figure 38 – Atractospora thailandensis (Material examined – THAILAND, Prachuap Khiri Khan,
on submerged wood in a small river, 30 July 2015, W. Dong 56B, HKAS 96226, holotype). a, b
Appearance of dark brown to black ascomata semi-immersed on host. c Vertical section of ascoma.
d Structure of peridium. e Paraphyses. f-h Unitunicate asci. i, j Ascospores. Scale bars: c = 50 μm,
d-f = 20 μm, g, h = 40 μm, i, j = 10 μm.
Ecological and economic significance of Auratiopycnidiellaceae
Auratiopycnidiella tristaniopsis is the only species in this family and is a leaf spotting
pathogen on Tristaniopsis laurina (Myrtaceae) in New South Wales, Australia (Crous et al. 2012b).
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Genus included in Auratiopycnidiellaceae
Auratiopycnidiella Crous & Summerell, Persoonia 28: 69 (2012)
Index Fungorum number: IF564733; 1 species with sequence data.
Type species – Auratiopycnidiella tristaniopsis Crous & Summerell
Notes – Auratiopycnidiella is monotypic and comprises only the type species A. tristaniopsis.
This taxon shows some similarities to taxa of Cryphonectriaceae in having orange conidiomata.
However, Auratiopycnidiella is distinct from other genera of Diaporthales in having subepidermal,
pycnidial, orange conidiomata with dark brown margins, holoblastic to annellidic conidiogenous
cells, ellipsoid and uni-septate conidia with a thickened hilum or minute marginal frill. Conidia
becoming brown at germination forming a thin, wavy germ slit, 90° to the long axis of the
conidium (Crous et al. 2012b).
Figure 39 – Auratiopycnidiella tristaniopsis (redrawn from Crous et al. 2012b). a Host. b Close-up
of orange pycnidia (top-view). c Conidiogenous cells giving rise to conidia. d Disarticulating
conidial cells. e Conidia with thick, bright banded germ slit. Scale bars: c-e = 10 μm.
Australiascaceae Réblová & W. Gams, Stud. Mycol. 68(1): 171 (2011)
Index Fungorum number: IF515430; Facesoffungi number: FoF01110; 8 species.
Pathogenic on tubers of sweet potato and saprobic on terrestrial plant leaves, branches, and
on spathes and stipes of ferns. Sexual morph: Ascomata solitary to gregarious, brown to black,
conical to obpyriform, glabrous or clothed with setae. Ostioles periphysate. Setae scant, acute,
thick-walled, septate, dark brown. Peridium 2-layered, fragile, textura epidermoidea to prismatica
in surface view. Paraphyses septate, persistent, branching. Asci 8-spored, unitunicate, cylindrical to
clavate, short-pedicellate, apex truncate, with a distinct, shallow, J-, apical ring. Ascospores
overlapping biseriate, hyaline, ellipsoidal to oblong, apiculate at both ends, septate, smooth-walled.
Asexual morph: Hyphomycetous. Conidiophores macronematous, pale to dark brown, septate.
Conidiogenous cells monophialidic, ampulliform to cylindrical, subhyaline, with minute collarette.
Conidia ellipsoid to cylindrical-ellipsoidal, smooth-walled, hyaline, septate, aggregated in slime or
in chains (adapted from Maharachchikumbura et al. 2016b).
Type genus – Monilochaetes Halst.
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Notes – The family was established in Glomerellales based on morphological and molecular
DNA data, to accommodate the holomorphic genus Australiasca with its asexual morph
Monilochaetes, with Dischloridium as its synonym (Réblová et al. 2011). The relationship between
Monilochaetes and Dischloridium was suggested by Rong & Gams (2000). The correct name of the
taxon is Monilochaetes (Maharachchikumbura et al. 2016b) as the name is older and more
commonly used, especially in the plant pathological community. Based on both phylogenetic and
MCC trees, the family is well-supported and related to the families Glomerellaceae,
Plectosphaerellaceae and Reticulascaceae in Glomerellales and with a stem age of 256 MYA
(Réblová et al. 2011, Maharachchikumbura et al. 2016b, Hongsanan et al. 2017).
Ecological and economic significance of Australiascaceae
Members of Australiascaceae are important inhabitants of commercial crops and medicinal
plants. Monilochaetes infuscans causes scurf disease or soil stain of sweet-potato (Ipomea batata)
in many countries (Harter 1916, Rong & Gams 2000). Monilochaetes nothapodytis is an endophyte
isolated from Nothapodytes pittosporoides in China, a plant used in Traditional Chinese Medicine
(Zhou et al. 2017).
Genus included in Australiascaceae
Monilochaetes Halst. J. Agric. Res., Washington 5: 791 (1916)
Index Fungorum number: IF8969; 8 morphological species (Species Fungorum 2020), 5
species with sequence data.
Type species – Monilochaetes infuscans Harter
Notes – Monilochaetes was experimentally linked with its sexual morph by Réblová et al.
(2011). Currently, eight taxa are included in Monilochaetes. Monilochaetes camelliae (=
Australiasca queenslandica) is illustrated in this entry.
Barbatosphaeriaceae H. Zhang, K.D. Hyde & Maharachch., Fungal Divers. 85: 94 (2017)
Index Fungorum number: IF553763; Facesoffungi number: FoF03341; 26 species.
Saprobic on decaying wood or other plant materials. Sexual morph: Ascomata astromatic,
leathery to fragile, dark brown to black, solitary or usually aggregated in circular to oval nests or in
short rows, globose to subglobose, glabrous or roughened, with elongate necks, venter or neck are
sparsely covered with a pubescence that disappears with age. Necks cylindrical, central or lateral,
straight to slightly flexuous, when in circular groups decumbent to perpendicular, covering,
piercing the periderm in a group. Ostiole periphysate. Peridium two-layered. Paraphyses abundant,
persistent, cylindrical, unbranched, septate, constricted at the septa, tapering to the apex. Asci 8spored, unitunicate, clavate or cylindrical, tapering towards the pedicel, with a distinct or indistinct,
refractive, J-, apical ring, floating freely in centrum at maturity. Ascospores 1- or 2-seriate, hyaline
to pale brown, oblong to ellipsoid, subcylindrical, reniform, straight, sometimes curved, allantoid,
U- to horseshoe-shaped or 3/4 circular, aseptate or septate, not- or slightly constricted at the
septum, smooth-walled. Asexual morph: Hyphomycetous, ramichloridium- and sporothrix-like.
Conidiophores semi-micronematous to macronematous, unbranched or branched, cylindrical to
flask- or irregularly-shaped, brown or hyaline. Conidiogenous cells polyblastic, integrated,
terminal, sympodially elongating with several denticles or forming rachis producing conidia
holoblastically. Conidia ellipsoidal to suballantoid, straight or curved, unicellular, hyaline (adapted
from Réblová 2007, Réblová et al. 2015c Zhang et al. 2017a).
Type genus – Barbatosphaeria Réblová
Notes – Barbatosphaeriaceae was introduced by Zhang et al. (2017a) with three genera,
Barbatosphaeria, Ceratostomella and Xylomelasma, in Diaporthomycetidae genera incertae sedis.
The family formed a strongly supported, distinct clade, with a stem age of ca 110 MYA in the
MCC tree (Hyde et al. 2017a, Zhang et al. 2017a). Barbatosphaeriaceae formed a sister clade to
Natantiella ligneola and Ophiostomatales in Zhang et al. (2017a), while it was a sister group to
Amplistromatales, Ophiostomatales and Phomatosporales in Senanayake et al. (2016). In this entry,
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the sexual morph of Barbatosphaeria varioseptata and the ramichloridium- and sporothrix-like
asexual morphs of B. barbirostris are illustrated.
Figure 40 – Monilochaetes queenslandica (Material examined – AUSTRALIA, Queensland,
Malanda, on branch of Camellia sinensis, 26 Mar 1988, J.L. Alcorn, BRIP 25190a, holotype, BRIP
24334c). a Perithecia on the host. b Peridium. c Ascospores. d-h Asci. i Conidiophores. j
Conidiogenous cells. k Conidia. Scale bars: a = 100 µm, b, d-i = 50 µm, c = 20 µm, j-k = 10 µm.
Ecological and economic significance of Barbatosphaeriaceae
Barbatosphaeriaceae comprises three genera which are saprobic on decaying wood and bark.
Genera included in Barbatosphaeriaceae
Barbatosphaeria Réblová, Mycologia 99(5): 727 (2008)
Index Fungorum number: IF505954; 9 morphological species (Species Fungorum 2020), 8
species with sequence data.
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Type species – Barbatosphaeria barbirostris (Dufour) Réblová
Notes – The genus was introduced by Réblová (2007) based on Barbatosphaeria barbirostris
and defined as a well-supported monophyletic clade in Sordariomycetidae, genera incertae sedis
(Réblová et al. 2015c). Zhang et al. (2017a) transferred the genus to the newly described
Barbatosphaeriaceae based on a stable clade in phylogenetic and MCC trees. A key to species
accepted in the genus was provided by Réblová et al. (2015c). Ramichloridium- and sporothrix-like
asexual morphs have been observed regularly in cultures of various species (Samuels &
Candoussau 1996, Réblová 2007, Réblová et al. 2015c).
Ceratostomella Sacc., Michelia 1(4): 370 (1878), Mycologia 98(1): 75 (2006)
Index Fungorum number: IF901; 14 morphological species (Species Fungorum 2020), 2
species with sequence data.
Type species – Ceratostomella rostrata (Tode: Fr.) Sacc.
Notes – Ceratostomella is a well-established genus that comprises 112 epithets in Index
Fungorum (2020). Ceratostomella was redescribed based on the lectotype species C. rostrata and
three other species were accepted, i.e. C. cuspidate, C. pyrenaica and C. rhynchophora. Other
species attributed to the genus that do not match the emended generic concept were revised.
Ceratostomella species with evanescent asci were transferred to Microascales and Ophiostomatales
(de Beer et al. 2013a, 2014). Ceratostomella species with persistent asci were revised by Réblová
(2011) and Réblová et al. (2018). Many species were excluded from Ceratostomella, some were
accommodated in other genera, and some remain as species of uncertain status and cannot presently
be reassigned (de Beer et al. 2013a, 2014, Réblová et al. 2011, 2018).
Xylomelasma Réblová, Mycologia 98(1): 87 (2006)
Index Fungorum number: IF501459; 3 morphological species (Species Fungorum 2020), 2
species with sequence data.
Type species – Xylomelasma sordida Réblová
Notes – Xylomelasma was introduced and delimited from the closely related genus
Ceratostomella by Réblová (2006), with two new species, X. novae-zelandiae and X. sordida, both
occurring on decaying wood. The third wood-inhabiting species, X. shoalensis, was described in
Hernández-Restrepo et al. (2016c). Xylomelasma moderata was transferred to Calyptosphaeria
subdenudata in the Xenospadicoidaceae (Réblová et al. 2018).
Barrmaeliaceae Voglmayr & Jaklitsch, Mycol. Progr. 17(1–2): 162 (2018)
Index Fungorum number: IF822042; Facesoffungi number: FoF06772; 12 species.
Saprobic on wood or bark. Sexual morph: Stroma if present mostly in wood and blackening
the surface in wide areas or in elongate bands, sometimes darker around the ostioles; entostroma
prosenchymatous, poorly developed, without KOH extractable pigments. Ascomata perithecial,
globose, sometimes raising the substrate, singly, in small groups or gregarious. Peridium
melanised, pseudoparenchymatous to prosenchymatous. Paraphyses numerous persistent, septate.
Asci 8-spored, cylindrical, persistent, with J-, or infrequently J+, apical ring. Ascospores yellow to
dark brown, unicellular with or without germ slit (Barrmaelia), or 2-celled with septum near one
end, small cell hyaline, large cell dark brown and with an apical germ apparatus consisting of radial
slits (Entosordaria); allantoid or ellipsoid, inequilateral, slightly inequilateral or nearly equilateral,
with narrowly or broadly rounded ends. Asexual morph: libertella-like where known (Barrmaelia;
Rappaz 1995, adapted from Voglmayr et al. 2018).
Type genus – Barrmaelia Rappaz
Notes – Barrmaeliaceae was introduced by Voglmayr et al. (2018) to accommodate the type
genus Barrmaelia and is a sister genus to Entosordaria (Rappaz 1995). Barrmaelia macrospora, B.
moravica, B. rhamnicola and Entosordaria perfidiosa were epitypified by Voglmayr et al. (2018).
In addition, a new species B. rappazii and Entosordaria quercina were described in the family
(Voglmayr et al. 2018).
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Figure 41 – Barbatosphaeria varioseptata (Material examined – CZECH REPUBLIC, Southern
Moravia, distr. Břeclav, Milovice, Milovický les Nature Reserve, north slopes of Mt Špičák 293 m
asl, on decaying wood and bark of a branch of Quercus sp., 18 November 2010, M. Réblová, PRM
924376, holotype). a Herbarium material with label. b Appearance of superficial black ascomata on
host. c Structure of peridium. d Vertical section of ascoma. e Ostiole. f Paraphyses. g-j Unitunicate
asci. k-o Ascospores. Scale bars: c, d = 100 μm, e = 50 μm, f-j = 20 μm, k-o = 5 μm.
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Figure 42 – Asexual morphs of Barbatosphaeria barbirostris, ramichloridium-like (a), sporothrixlike (b, c) (redrawn from Réblová 2007) a-c Conidiophores and conidia. Scale bars: a-c = 10 µm.
Ecological and economic significance of Barrmaeliaceae
Barrmaeliaceae comprises saprobes, especially wood and bark inhabiting species, which are
important as decomposers (Rappaz 1995, Voglmayr et al. 2018). Among the Entosordaria species,
E. jacobi-felicis was reported on Coffea liberica leaves from Ivory Coast (Moreau & Moreau
1951).
Genera included in Barrmaeliaceae
Barrmaelia Rappaz, Mycol. Helv. 7(1): 130 (1995)
Index Fungorum number: IF27434; 8 morphological species (Voglmayr et al. 2018), 5
species with sequence data.
Type species – Barrmaelia rhamnicola Rappaz
Notes – Rappaz (1995) introduced Barrmaelia which resembles Diatrypaceae due to its
allantoid ascospores, but in contrast, has short-pedicellate asci and J-, ascal rings. Rappaz (1995)
combined six species in Barrmaelia (B. macrospora, B. moravica, B. oxyacanthae, B. picacea, B.
pseudobombarda, and B. sustenta) and described one new species, which he also selected as the
generic type, B. rhamnicola based on morphology. Maharachchikumbura et al. (2016b) regarded
Barrmaelia as a xylariaceous genus with a conidial morph that did not belong to nodulisporiumlike or geniculosporium-like taxa. Wendt et al. (2018) placed Barrmaelia in Xylariales genera
incertae sedis due to its unconfirmed affinities to the other families. A study on natural
classification of Graphostromataceae, Hypoxylaceae, Lopadostomataceae and Xylariaceae has
placed the genus in the Diatrypaceae based on their morphology and tentative phylogeny
(Daranagama et al. 2018). However, the clade comprising Barrmaelia and Entosordaria within
Xylariaceae sensu lato has been introduced as Barrmaeliaceae (Voglmayr et al. 2018).
Barrmaelia rhamnicola, the type species of the genus was introduced by Rappaz (1995) and
epitypified by Voglmayr et al. (2018). It is characterized by often slightly curved, relatively large
ascospores, which are filled with conspicuous oil drops and lack a germ slit. The known libertellalike asexual morph has been shown for B. rhamnicola. Barrmaelia rhamnicola is illustrated in this
entry (Fig. 43).
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Figure 43 – Barrmaelia rhamnicola (redrawn from Voglmayr et al. 2018). a Ostioles protruding
through the blackened wood surface. b Cross section of ascoma. c, d Asci e-h Ascospores. Scale
bars: b = 150 µm, c, d = 10 µm, e-h = 5 µm.
Entosordaria (Sacc.) Höhn., Sber. Akad. Wiss. Wien, Math.-naturw. Kl., Abt. 1129: 167 (1920),
emend.
Index Fungorum number: IF1839; 4 morphological species (Species Fungorum 2020), 2
species with sequence data.
Type species – Entosordaria perfidiosa (De Not.) Höhn.
Notes – Voglmayr et al. (2018) re-described and validated Entosordaria in Barrmaeliaceae.
Species are saprobes. The combined ITS, LSU, rpb2 and tub2 phylogenetic analyses revealed that
the genus is sister to Barrmaelia.
Batistiaceae Samuels & K.F. Rodrigues, Mycologia 81(1): 54 (1989)
Index Fungorum number: IF81989; Facesoffungi number: FoF01321; 1 species.
Saprobic on wood. Sexual morph: Ascomata cleistothecial, stipitate-capitate, solitary to
gregarious, superficial, black, globose, long stipitate, carbonaceous, without an ostiole. Peridium
cephalothecoid. Asci 8-spored, unitunicate, globose, lacking a visible discharge ring and
deliquescent. Ascospores irregularly arranged, brown, ellipsoidal to oblong, 1-celled, without germ
slits or germ pores, smooth-walled. Asexual morph: Hyphomycetous. Synnemata determinate, dark
brown to black. Hyphae of stipe parallel, dark brown, septa simple. Conidiophores forming a
compact, grey palisade. Capitulum terminal, globose, in vertical section composed of a darkly
pigmented dome of angular cells giving rise to loose textura intricata of hyaline, more or less
swollen hyphae, a compact subhymenium of similar hyphae and then a conidiogenous hymenium.
Conidiogenous cells phialidic, single or in terminal pairs, light brown, cylindrical, with an abruptly
terminated apex and cylindrical collarette, smooth-walled. Conidia in dry chains without
connectives, pale olivaceous, light brown to grey in mass, ellipsoidal, globose, subglobose or
oblong, 1-celled, with thin, smooth-walls (adapted from Maharachchimbura et al. 2016b).
Type genus – Batistia Cif.
Notes – Batistiaceae, introduced in 1989 and classified in Sordariales, is typified by Batistia
(Samuels & Rodrigues 1989, Eriksson 2001). Kirk et al. (2001) excluded the family from
Sordariales and placed in Sordariomycetidae families incertae sedis. However, based on molecular
441
analyses, Huhndorf et al. (2004b) placed Batistiaceae in Pezizomycotina family incertae sedis. This
classification was followed by Lumbsch & Huhndorf (2010) and Wijayawardene et al. (2012).
However, Maharachchikumbura et al. (2015) determined that the sequence data published by
Huhndorf et al. (2004b) was contaminated. The classification of Batistiaceae was verified in
Sordariomycetidae family incertae sedis (Maharachchikumbura et al. 2015, 2016b, Hongsanan et
al. 2017, Wijayawardene et al. 2018a). A lack of ex-type cultures and sequence data in GenBank
makes it difficult to place Batistiaceae in a phylogenetic scheme (Kirk et al. 2008,
Maharachchikumbura et al. 2015, 2016b, Hongsanan et al. 2017). We follow its classification in
Sordariomycetidae family incertae sedis until other evidence emerges (Maharachchikumbura et al.
2015, 2016b). Fresh collections, especially of neotype or epitype strains, together with molecular
data are needed to clarify the species and family. In this entry Batistia annulipes is illustrated.
Ecological and economic significance of Batistiaceae
Batistiaceae are likely to be saprobes involved in nutrient cycling.
Genus included in Batistiaceae
Batistia Cif., Atti Ist. bot. Univ. Lab. crittog. Pavia, Ser. 5 15: 166 (1958)
Index Fungorum number: IF522; 1 species with sequence data (likely contaminant).
Type species – Batistia annulipes (Mont.) Cif.
Notes – Batistia is typified by Batistia annulipes. Maharachchikumbura et al. (2016b)
synonymized Acrostroma under Batistia based on the report by Samuels & Rodrigues (1989) and a
study of herbarium type materials of both Acrostroma and Batistia.
Beltraniaceae Nann., Repert. mic. uomo: 498 (1934)
Index Fungorum number: IF80516; Facesoffungi number: FoF05312; 80 species.
Saprobic on plant tissues. Sexual morph: Ascomata pale yellow, solitary to aggregated on OA
and PDA, globose to somewhat papillate, with central ostiole; wall of 3–4 layers of subhyaline cells
of textura angularis to intricata. Paraphyses septate, cellular, anastomosing, distributed among
asci. Asci 8-spored, unitunicate, subcylindrical, sessile. Ascospores tri- to multi-seriate, hyaline,
obovoid, granular, smooth, aseptate with dissolving mucoid sheath. Asexual morph:
Hyphomycetous. Mycelium immersed to superficial, composed of subhyaline to brown, thin-walled
hyphae. Stromata usually 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 or
sometimes branched, erect, septate, pale brown, arising from the base of setae or separate, with or
without radially lobed basal cell. Conidiogenous cells holoblastic, polyblastic, pale brown,
integrated, denticulate. Conidia biconic, lageniform to navicular, subhyaline to red-brown, with
transverse band of pale pigment at widest part of the conidium, base rounded, 1-denticulate or
rostrate, apex spicate, apiculate or truncate (adapted from Crous et al. (2015a) and
Maharachchikumbura et al. (2016b)).
Type genus – Beltrania Penz.
Notes – Beltrania, Beltraniella, Beltraniopsis, Hemibeltrania, Parapleurotheciopsis,
Porobeltraniella, Pseudobeltrania, Pseudosubramaniomyces and Subsessila, are accepted in the
family (Maharachchikumbura et al. 2015, 2016b, Hongsanan et al. 2017, Lin et al. 2017a). We
introduce and illustrate a new species, Beltraniella ramosiphora based on phylogenetic data (Figs.
4, 45).
Ecological and economic significance of Beltraniaceae
Most Beltraniaceae species are saprobes. However, Beltrania rhombica was reported to be a
pathogen which can cause a leaf spot on Tibouchina semidecandra in China (Shi et al. 2012).
Milagres et al. (2018) reported Pseudobeltrania cedrelae, as the causal agent of pseudobeltrania
spot on Cedrela fissilis in Brazil.
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Figure 44 – Batistia annulipes a-g MNHN-PC-PC0167686 MC10053b (Material examined –
BRAZIL, Rio de Janeiro, Corcovado, at cortices, Montagne, 1831-1833, MNHN-PC-PC0167685
(MC10053a), MNHN-PC-PC0167686 (MC10053b) MNHN-PC-PC0167687 (MC10053c),
holotype, MNHN-PC-PC0167688 (MC10054), MNHN-PC-PC0167689 (MC10055)) h-m NY
00936852 (Dumont-VE 6722) (Material examined – VENEZUELA, EDO. SURE., Trail from Los
Pocitos through la Roma to town of Rio Grande Arriba, on unidentified wood, 12 July 1972, K. P.
Dumont, G. J. Samuels and G. Morillo, and J. Farfan, Dumont-VE 5017, NY 00936853, paratype).
a Ascomata on the host. b Ascoma. c Vertical section of ascoma. d, e Peridium. f Asci. g
Ascospores. h Synnemata on the host. i Synnema longitudinal section. j Conidiogenous cells. k, l
Conidia. m Detail of capitulum tissues in vertical section. Scale bars: a = 1 mm, b, c = 200 µm, d, i
= 50 µm, e= 10 µm, f, g, j-l = 3µm, h = 500 µm, m = 20 µm.
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Genera included in Beltraniaceae
Beltrania Penz., Nuovo G. bot. ital. 14: 72 (1882)
Index Fungorum number: IF7355; 15 morphological species (Species Fungorum 2020), 5
species with sequence data.
Type species – Beltrania rhombica Penz.
Notes – Beltrania is characterised by dark and mostly unbranched setae with radially lobed
basal cells, unbranched conidiophores arising from basal cells of setae or from separate radially
lobed basal cells; polyblastic, sympodial and denticulate conidiogenous cells; swollen separating
cells and biconic, spicate or apiculate conidia with a hyaline transverse band (Seifert et al. 2011,
Lin et al. 2017b).
Beltraniella Subram., Proc. Indian Acad. Sci., Sect. B 36: 227 (1952)
Index Fungorum number: IF7356; 27 morphological species (Species Fungorum 2020), 11
species with sequence data.
Type species – Beltraniella odinae Subram.
Notes – Beltraniella species are characterised by setae that are sterile extensions of
conidiophores or occur among conidiophores or are lacking, unbranched and arising from radially
lobed basal cells. Conidiophores are branched, often with setiform apex, arising from radially lobed
basal cells. Conidiogenous cells are polyblastic and sympodial and produce acropleurogenous,
turbinate or biconic conidia with a distinct hyaline transverse band.
Crous et al. (2016a) described a new Beltraniella species, B. acaciae based on morphology
and phylogenetic analysis of ITS sequence data, revealed that B. acaciae was intermediate between
Beltraniella and Beltrania. However, Beltraniella acaciae grouped together with other Beltraniella
species within the Beltraniella clade in our phylogenetic analysis of combined ITS and LSU
sequence data.
Beltraniella ramosiphora C.G. Lin & K.D. Hyde, sp. nov.
Figs 45, 46
Index Fungorum number: IF555603; Facesoffungi number: FoF03885
Etymology – Name reflects the branched conidiophores.
Holotype – MFLU 17-2649.
Saprobic on decaying leaves. Sexual morph: Undetermined. Asexual morph: Colonies on
plant substrate effuse, thin, pale brown. Mycelium mostly immersed in the substratum. Setae
numerous, erect, arising from radially lobed basal cells, straight or flexuous, unbranched, single,
thick-walled, smooth, dark brown, 155–280 μm long, 4.5–7 μm wide at the base, tapering to a
pointed apex, arising from a dark brown, swollen, radially lobed cell, 13–23 μm diam.
Conidiophores macronematous, long setiform and short non-setiform; long conidiophores single,
straight, septate, smooth, thick-walled, unbranched, dark brown at the base and paler at the apex,
260–365 μm long, swollen at the base and 18–22.5 μm wide, 7.5–9 μm wide just above the swollen
base, slightly tapering to a pointed apex; short conidiophores, non-setiform, branched, septate,
smooth-walled, subhyaline to pale brown, thin-walled, 20–55 μm long, swollen at the base and 4.5–
12 μm wide. Conidiogenous cells polyblastic, integrated, determinate, terminal, cylindrical, oblong,
hyaline, smooth, 5–22 µm long, 3–7.5 µm wide at the base. Separating cells ovoid or obovoid,
thin-walled, smooth, hyaline, 7.5–11.5 µm long, 3.5–4.6 µm wide in the broadest part. Conidia
arise directly from conidiogenous cells or from separating cells, aggregated, acrogenous, simple,
dry, straight, smooth, thin-walled, biconic, turbinate to pyriform, rostrate to pointed at proximal
end, truncate at distal end, hyaline with a hyaline supraequatorial transverse band, 15.5–21 µm
long, 4.5–6.5 µm wide in the broadest part.
Culture characteristics – Conidia germinating on PDA within 12 h. Colonies on PDA effuse,
pale white from above, light yellow to dark brown from below, reaching a diam. of 4–5 cm in 3
days at 25°C.
Material examined – THAILAND, Chang Wat Lampang, Amphoe Mueang Pan, Tambon
Chae Son, on decaying leaf, 24 Sep 2016, CG Lin LCG10-2 (MFLU 17-2649, holotype), ex-type
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living culture, MFLUCC 17-2582.
GenBank numbers – ITS: MG717500, LSU: MG717502.
Notes – Three Beltraniella species have two types of conidiophores (long setiform and short
non-setiform), viz., B. botryospora, B. clara and B. fertilis. Beltraniella ramosiphora differs from
B. clara by smooth setae and biconic, turbinate to pyriform conidia, whereas setae are absent and
conidia are lageniform in B. clara (Onofri 1983). The long setiform conidiophores of B.
ramosiphora are unbranched, while they are branched at the apex in B. fertilis (Heredia et al. 2002).
The short non-setiform conidiophores of B. ramosiphora are longer than B. botryospora, and they
are branched in B. ramosiphora (Shirouzu et al. 2010).
Beltraniopsis Bat. & J.L. Bezerra, Publicações Inst. Micol. Recife 296: 4 (1960)
Index Fungorum number: IF7357; 11 morphological species (Species Fungorum 2020), 2
species with sequence data.
Type species – Beltraniopsis esenbeckiae Bat. & J.L. Bezerra
Notes – Beltraniopsis was introduced by Batista & Bezerra (1960). It is characterised by
unbranched, setiform conidiophores arising from radially lobed basal cells, sympodial and
denticulate conidiogenous cells, swollen separating cells and biconic, rostrate conidia with a
hyaline transverse band (Seifert et al. 2011). Most Beltraniopsis species were collected from
terrestrial litter, however, B. tanzaniensis was reported from decaying submerged leaves in India
(Patil & Borse 2015).
Hemibeltrania Piroz., Mycol. Pap. 90: 30 (1963)
Index Fungorum number: IF8498; 8 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Hemibeltrania cinnamomi (Deighton) Piroz.
Notes – Hemibeltrania was established by Pirozynski (1963) to accommodate H. cinnamomi
and H. nectandrae. The genus is characterised by unbranched or sparingly branched conidiophores
arising from radially lobed basal cells, sympodial and denticulate conidiogenous cells, and broadly
ellipsoidal, limoniform, ovoid, obovoid, cymbiform, navicular, biconic or fusiform, pale
olivaceous, aseptate conidia without a hyaline transverse band (Seifert et al. 2011). In addition,
Hemibeltrania lacks a separating cell. A synopsis of accepted Hemibeltrania species was provided
by Rajeshkumar et al. (2016b). Lin et al. (2017b) provided sequence data for H. cinnamomi, which
was collected from Thailand.
Parapleurotheciopsis P.M. Kirk, Trans. Br. mycol. Soc. 78(1): 63 (1982)
Index Fungorum number: IF9219; 4 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Parapleurotheciopsis inaequiseptata (Matsush.) P.M. Kirk
Notes – The genus was introduced by Kirk (1982) for P. ilicina and P. inaequiseptata. Crous
et al. (2015d) placed this genus within Beltraniaceae. Parapleurotheciopsis is characterised by
unbranched, percurrent conidiophores arising from radially lobed basal cells, sympodial and
denticulate conidiogenous cells and catenate conidia without a transverse band, lacking setae and
swollen separating cells (Seifert et al. 2011).
Porobeltraniella Gusmão, Mycologia 96(1): 151 (2004)
Index Fungorum number: IF28821; 2 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Porobeltraniella porosa (Piroz. & S.D. Patil) Gusmão
Notes – Porobeltraniella was introduced by Fernando & Gusmão (2004) to accommodate P.
patilii and P. porosa. Rajeshkumar et al. (2016a) provided sequence data for P. porosa and
confirmed the placement of the genus within Beltraniaceae. Porobeltraniella is characterised by
unbranched or verticillate, setiform conidiophores arising from radially lobed basal cells,
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polyblastic, discrete conidiogenous cells, solitary or catenate, turbinate conidia with circular pores
near the broadest part (Seifert et al. 2011).
Figure 45 – Phylogram generated from maximum likelihood analysis based on combined LSU and
ITS sequence data of Beltraniaceae. Fifty strains are included in the combined analyses which
comprised 1546 characters (884 characters for LSU, 662 characters for ITS) after alignment. Tree
topology of the maximum likelihood analysis is similar to the Bayesian and maximum parsimony
analyses. The best RaxML tree with a final likelihood value of -7145.370801 is presented.
Estimated base frequencies were as follows: A = 0.252192, C = 0.214861, G = 0.266786, T =
0.266162; substitution rates AC = 1.363640, AG = 2.557848, AT = 1.973142, CG = 1.035845, CT =
5.942470, GT = 1.000000; gamma distribution shape parameter α = 0.171878. Bootstrap support
values for maximum likelihood and maximum parsimony greater than 50% and Bayesian posterior
probabilities greater than 0.8 are indicated above or below the nodes as MLBS/MPBS/PP. The tree
is rooted with Anthostomella leucospermi (CBS 110126). Ex-type strains are in bold and black. The
newly generated sequences are indicated in blue.
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Figure 46 – Beltraniella ramosiphora (MFLU 17-2649, holotype). a Host material. b, c
Conidiophores on host surface. d Setae. e Long conidiophore. f, g Short conidiophores with
conidiogenous cells. h, i Separating cell and conidia. Scale bars: d, e = 50 μm, f-i = 10 μm.
Pseudobeltrania Henn., Hedwigia 41: 310 (1902)
Index Fungorum number: IF9555; 9 morphological species (Species Fungorum 2020), 2
species with sequence data.
Type species – Pseudobeltrania cedrelae Henn.
Notes – Pseudobeltrania is characterised by unbranched conidiophores arising from radially
lobed basal cells, polyblastic and denticulate conidiogenous cells and biconic, apiculate conidia
with a transverse band, lacking of setae and swollen separating cells (Seifert et al. 2011). A key to
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species was provided by Heredia et al. (2002). Pseudobeltrania cedrelae was reported to be a
pathogen which can cause leaf spot on Cedrela fissilis (Milagres et al. 2018).
Pseudosubramaniomyces Crous, Persoonia 39: 375 (2017)
Index Fungorum number: IF823465; 1 species with sequence data.
Type species – Pseudosubramaniomyces fusisaprophyticus (Matsush.) Crous
Notes – The genus was introduced by Varghese & Rao (1979b) to accommodate S. indicus
and S. navicularis. It is characterised by unbranched or sparingly branched, pale brown
conidiophores, sympodial and denticulate conidiogenous cells and catenate conidia without a
transverse band, lacking setae and swollen separating cells (Seifert et al. 2011). Rajeshkumar et al.
(2016a) placed Subramaniomyces within Beltraniaceae based on phylogenetic analysis of S.
fusisaprophyticus. However, Crous et al. (2017a) introduced a new genus Pseudosubramaniomyces
to accommodate S. fusisaprophyticus and it differs from by lacking lateral conidiogenous cells
along the length of the conidiophore stipe, and tends to have pale brown conidiophores.
Subsessila C.G. Lin & K.D. Hyde, Mycol. Progr. 16: 396 (2017)
Index Fungorum number: IF552504; 1 species with sequence data.
Type species – Subsessila turbinata C.G. Lin & K.D. Hyde
Notes – Subsessila is separated from Beltrania and other similar genera by the absence of
distinct conidiophores and conidia without a hyaline transverse band (Lin et al. 2017a).
Bertiaceae Smyk, Ukr. bot. Zh. 38(6): 47 (1981)
Index Fungorum number: IF82053; Facesoffungi number: FoF01111; 38 species.
Saprobic on wood in freshwater and terrestrial habitats. Sexual morph: Ascomata perithecial,
dark brown to black, solitary or gregarious, superficial to erumpent, cupuliform, globose to
subglobose, carbonaceous to membranaceous, turbinate or tuberculate or smooth, collabent or
collapsing laterally or not collapsing, ostiolate. Peridium thick, Munk pores present or absent, outer
layer (pseudoparenchymatous cells) hard, composed of dark tissues, basal part mixed with host
cells; inner layer composed of brown to dark brown to hyaline cells of textura angularis.
Paraphyses few, inflated, branched. Asci 8-spored, unitunicate, clavate, long pedicellate, apical ring
indistinct or absent. Ascospores 2–3-seriate or irregularly arranged, hyaline to brown, cylindrical to
fusiform, aseptate to multi-septate, smooth-walled, mostly with guttules. Asexual morph:
Undetermined (adapted from Maharachchimbura et al. 2016b).
Type genus – Bertia De Not.
Notes – Bertiaceae was introduced by Smyk (1981) and is typified by Bertia (De Notaris
1844). It is characterized by superficial, black, turbinate ascomata and clavate asci with hyaline,
fusiform ascospores, irregularly arranged in the upper region of the ascus (Maharachchikumbura et
al. 2016b). Nannfeldt & Santesson (1975) reported that Nitschkiaceae consists of
Acanthonitschkea, Bertia, Coronophora, Gaillardiella and Nitschkia based on similarity of
ascomata. Nannfeldt & Santesson (1975) also postulated that Bertia should be given generic status
in the family based on different morphs of the ascoma without Quellkörper and 8-spored asci.
Smyk (1981) was of the same view and introduced Bertiaceae. Mugambi & Huhndorf (2010) reevaluated Coronophorales based on multi-gene analysis and proposed Bertia and Gaillardiella in
Bertiaceae. Gaillardiella has superficial, brown ascomata with a roughened papulose peridium
(Miller & Huhndorf 2009). Maharachchikumbura et al. (2015) analysed a combined DNA sequence
dataset to confirm the placement of Bertia and Gaillardiella in Bertiaceae, in a highly supported
clade in the Coronophorales. In this study, Bertiaceae is closely related to Nitschkiaceae based on
combined LSU-tef1-ITS sequence data analysis (Fig. 11). Bertia moriformis is illustrated in this
entry.
Ecological and economic significance of Bertiaceae
Most Bertiaceae species are saprobic on wood, widespread with most known from America
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and Europe (De Notaris 1844, Patouillard & Lagerheim 1895, Viégas 1947, Nannfeldt & Santesson
1975, Mugambi & Huhndorf 2010, Vasilyeva et al. 2015).
Genera included in Bertiaceae
Bertia De Not., Gior. Bot. Ital. 1(1): 334 (1844)
Index Fungorum number: IF551; 32 morphological species (Species Fungorum 2020), 9
species with sequence data.
Type species – Bertia moriformis (Tode) De Not.
Notes – The type species, Bertia moriformis, is characterized by tuberculate ascomata and
clavate asci and brown ascospores (Mugambi & Huhndorf 2010, Maharachchikumbura et al.
2016b). We provide a multi-gene phylogenetic tree (Fig. 11), which reveals that Bertia is closely
related to members of Nitschkiaceae (Nitschkia, Acanthonitschkea and Fracchiaea) with good
support (Fig. 11).
Figure 47 – Bertia moriformis (Material examined – ITALY, Riva, Valdobbia; on Rhododendron
sp., Carestia, 30 April 1859, S-F298). a Ascomata on host. b Tuberculate ascomata. c Ascoma cross
section. d Peridium. e Munk pores (arrows). f, g Asci. h-j Ascospores (f, i-j are stained in Melzer’s
reagent). Scale bars: a = 1 mm, b = 500 µm, c = 200 µm, d = 100 µm, f, g = 20 µm, h-j = 10 µm, e
= 5 µm.
Gaillardiella Pat., Bull. Soc. mycol. Fr. 11(4): 226 (1895)
Index Fungorum number: IF2027; 6 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Gaillardiella pezizoides Pat.
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Notes – Gaillardiella is characterized by turbinate ascomata, clavate to cylindrical asci with
short stalks and ring at the apex and hyaline ascospores.
Bionectriaceae Samuels & Rossman, Stud. Mycol. 42: 15 (1999)
Index Fungorum number: IF82088; Facesoffungi number: FoF01367; 579 species.
Herbicolous, corticolous, lichenicolous or fungicolous, some are coprophilous, most occur in
terrestrial or freshwater habitats, less common in marine habitats. Sexual morph: Ascomata
superficial on the substratum or embedded in a weakly or well-developed erumpent stroma, solitary
or densely aggregated, crowded, perithecial, rarely cleistothecial, if perithecial, globose, subglobose
to pyriform, if cleistothecial, globose, white, yellow, orange, reddish-brown, greenish or purple to
violet, without changing colour in KOH or lactic acid, smooth to rough, or warted, ostiolate or
lacking ostioles. Periphyses present or absent. Peridium composed of 1–3 layers. Asci (2–)8spored, unitunicate, clavate, saccate, cylindrical, sessile or short pedicellate, rarely evanescent, apex
simple or with an inconspicuous or distinct, J-, apical ring. Ascospores uniseriate, biseriate, multiseriate or irregular, hyaline, aseptate to multi-septate, sometimes muriform, globose, fusiform,
ellipsoid or broadly ellipsoid, smooth-walled, spinulose to tuberculate or striate. Asexual morph:
Hyphomycetous or less commonly ceolomycetous, acremonium-, gliocladium-like, gyrostroma-like
or penicillium-like, sometimes conidia produced on hyphae. Conidiophores dimorphic or
monomorphic, mostly sporodochial or synnematous, hyaline, subhyaline to brown or blackish
brown, smooth to finely echinulate-walled. Conidiogenous cells phialidic. Phialides cylindrical to
flask-shaped. Conidia unicellular to multi-septate, ellipsoidal, fusiform to subfusiform, sometimes
with papillate or truncate ends, hyaline to greenish hyaline or olivaceous grey, smooth or striatewalled. Chlamydospores present or absent (adapted from Rossman et al. 1999).
Type genus – Clonostachys Corda
Notes – Rossman et al. (1999) introduced Bionectriaceae to accommodate 26 genera
including five cleistothecial genera. Four cleistothecial genera were accepted in the family based on
phylogenetic analysis of the LSU gene by Rossman et al. (2001). Rossman et al. (2001) reported
that Bionectriaceae is monophyletic within Hypocreales, by including those genera and related
asexual morphs. Maharachchikumbura et al. (2015, 2016b) listed 39 genera in the family.
Wijayawardene et al. (2018a) accepted Didymostilbe and Virgatospora in Stachybotryaceae.
Stromatocrea and Vesicladiella, previously included in Bionectriaceae by Maharachchikumbura et
al. (2015, 2016b), were not considered by Wijayawardene et al. (2018a). Wijayawardene et al.
(2018) accepted 39 genera in the family including Bullanockia, Fusariella, Paracylindrocarpon
and Synnemellisia. Spicellum was listed under Bionectriaceae in Wijayawardene et al. (2018).
However, Spicellum was synonymized under Trichothecium (Myrotheciomycetaceae, Hypocreales)
based on morphology and DNA sequence data by Summerbell et al. (2011). LSU is commonly
used and the available gene for phylogenetic analysis of Bionectriaceae. However, few additional
sequence data are available for the family except for well-studied genera such as Geosmithia and
Clonostachys (Voglmayr & Jaklitsch 2019). Many genera need to be recollected and sequenced and
their placement in, or exclusion from Bionectriaceae or synonymies confirmed.
Ecological and economic significance of Bionectriaceae
Only few plant-pathogenic species are recognized in Bionectriaceae e.g. Nectriella pironi
causing galls on stems and leaves of woody plants (Alfieri et al. 1980). Acremonium species are
causal agents of rose dieback in Iran (e.g. A. sclerotigenum) (Domsch et al. 2007, Mirtalebi et al.
2017). Some species of Acremonium and Clonostachys are effective biological control agents (Auer
& Ludwig-Müller 2014, Sutton & Mason 2017, Bobeck & Pearce 2017). Clonostachys rosea
(strain BVT Cr-7) is a beneficial biological control agent for fungal plant pathogens such as
Botrytis cinerea, Sclerotinia sclerotiorum, Pythium, Alternaria, Monilinia, Colletotrichum,
Cladosporium, Rhizoctonia., Streptomyces, Didymella and Fusarium (Sutton & Mason 2017).
Furthermore, strain 88-710 helps to protect plants against environmental stresses, hence promoting
plant growth and productivity (Sutton & Mason 2017, Bobeck & Pearce 2017). Acremonium
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alternatum strains are used as biological control agents in several countries, to control clubroot
pathogen, Plasmodiophora brassicae, on Chinese cabbage (Brassica rapa) and oilseed rape
(Brassica napus) (Auer & Ludwig-Müller 2014).
Genera included in Bionectriaceae
We only list the genera in Bionectriaceae. Notes will be provided with species illustrations.
Acremonium Link, Mag. Gesell. naturf. Freunde, Berlin 3(1–2): 15 (1809)
Index Fungorum number: IF7028; 113 morphological species (Species Fungorum 2020), 72
species with sequence data.
Type species – Acremonium alternatum Link
Anthonectria Döbbeler, Mycologia 102(2): 405 (2010)
Index Fungorum number: IF514117; 1 morphological species.
Type species – Anthonectria mammispora Döbbeler
Aphanotria Döbbeler, Mycol. Res. 111(12): 1408 (2007)
Index Fungorum number: IF510591; 1 morphological species.
Type species – Aphanotria paradoxa Döbbeler
Battarrina (Sacc.) Clem. & Shear, Gen. fung., Edn 2 (Minneapolis): 279 (1931)
Index Fungorum number: IF526; 1 morphological species.
Type species – Battarrina inclusa (Berk. & Broome) Clem. & Shear
Bryocentria Döbbeler, Mycol. Progr. 3(3): 247 (2004)
Index Fungorum number: IF28841; 15 morphological species (Species Fungorum 2020), 2
species with sequence data.
Type species – Bryocentria brongniartii (P. Crouan & H. Crouan) Döbbeler
Bryotria Döbbeler & P.G. Davison, Nova Hedwigia 106(1-2): 232 (2017)
Index Fungorum number: IF821323; 5 morphological species (Species Fungorum 2020).
Type species – Bryotria urophora Döbbeler & P.G. Davison
Bullanockia Crous, Persoonia 37: 327 (2016)
Index Fungorum number: IF819077; 1 species with sequence data.
Type species – Bullanockia australis Crous
Clibanites (P. Karst.) P. Karst., Bidr. Känn. Finl. Nat. Folk 19: 14, 167 (1871)
Index Fungorum number: IF1107; 1 morphological species.
Type species – Clibanites paradoxa (P. Karst.) P. Karst.
Clonostachys Corda, Pracht-Fl. Eur. Schimmelbild.: 31 (1839)
Index Fungorum number: IF7701; 41 morphological species (Species Fungorum 2020), 20
species based on molecular data.
Type species – Clonostachys araucaria Corda
Notes – Bionectria was linked to Clonostachys by Rossman et al. (2013) and priority given to
the older and commonly used name Clonostachys. We illustrate the sexual morph of Clonostachys
rosea and asexual morph of C. byssicola (Figs. 48, 49).
Dimerosporiella Speg., Revta Mus. La Plata 15: 10 (1908)
Index Fungorum number: IF1585; 7 morphological species (Species Fungorum 2020).
Type species – Dimerosporiella paulistana Speg.
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Figure 48 – Clonostachys rosea (Material examined – THAILAND, Phayao Province, Chiang
Kham District, on twigs of a dicotyledonous plant, R.H. Perera, 11 September 2017, Bion 17
(MFLU 19-0960), living culture MFLUCC 17-2632). a Herbarium material. b, c Ascomata on host
(a dry, b after wet with water). d Section of ascoma. e Section of ostiole. f Section of peridium. g, h
Asci. i-n Ascospores. o Germinating ascospore. p, q Colony on PDA. Scale bars: b, c = 200 µm, d
= 100 µm, e, f = 50 µm, g, h = 20 µm, i-n = 10 µm, o = 20 µm.
452
Figure 49 – Clonostachys byssicola (Material examined – THAILAND, Chiang Mai Province, on a
dead stem of Chromolaena odorata, 2 February 2017, R.H. Perera BN4, MFLU 18-2729; living
culture MFLUCC 17-2033). a Herbarium material. b Sporodochia on the host. c Secondary
conidiophores, penicillate. d Secondary conidiophores and verticillium-like primary conidiophores.
e Phialides with conidia. f Conidia. g Germinating conidium. h, i Colony on PDA. Scale bars: c, d =
50 μm, e = 20 μm, f, g = 10 μm.
Fusariella Sacc., Atti Inst. Veneto Sci. lett., ed Arti, Sér. 6 2: 463 (1884)
Index Fungorum number: IF8282; 17 morphological species (Species Fungorum 2020), 8
species with sequence data.
Type species – Fusariella atrovirens (Berk.) Sacc.
Geonectria Lechat & J. Fourn., Ascomycete.org 10(2): 81 (2018)
Index Fungorum number: IF824881; 1 morphological species.
Type species – Geonectria subalpina Lechat, J. Fourn., M. Vega & Priou
453
Geosmithia Pitt, Can. J. Bot. 57(19): 2021 (1979)
Index Fungorum number: IF8324; 23 morphological species (Species Fungorum 2020), 16
species with sequence data.
Type species – Geosmithia lavendula (Raper & Fennell) Pitt
Gliomastix Guég., Bull. Soc. mycol. Fr. 21: 240 (1905)
Index Fungorum number: IF8345; 20 morphological species (Species Fungorum 2020), 5
species with sequence data.
Type species – Gliomastix chartarum (Cooke) Guég.
Globonectria Etayo, Biblthca Lichenol. 84: 47 (2002)
Index Fungorum number: IF28686; 1 morphological species.
Type species – Globonectria cochensis Etayo
Gracilistilbella Seifert, Stud. Mycol. 45: 18 (2000)
Index Fungorum number: IF28455; 3 morphological species (Species Fungorum 2020).
Type species – Gracilistilbella clavulata (Mont.) Seifert
Halonectria E.B.G. Jones, Trans. Br. Mycol. Soc. 48(2): 287 (1965)
Index Fungorum number: IF2206; 1 morphological species.
Type species – Halonectria milfordensis E.B.G. Jones
Heleococcum C.A. Jørg., Bot. Tidsskr. 37: 417 (1922)
Index Fungorum number: IF2253; 4 morphological species (Species Fungorum 2020), 2
species with sequence data.
Type species – Heleococcum aurantiacum C.A. Jørg.
Hydropisphaera Dumort., Comment. bot. (Tournay): 89 (1822)
Index Fungorum number: IF2407; 27 morphological species (Species Fungorum 2020), 11
species with sequence data.
Type species – Hydropisphaera peziza (Tode) Dumort.
Ijuhya Starbäck, Bih. K. svenska VetenskAkad. Handl., Afd. 3 25(no. 1): 30 (1899)
Index Fungorum number: IF2482; 22 morphological species (Species Fungorum 2020), 14
species with sequence data.
Type species – Ijuhya vitrea Starbäck
Kallichroma Kohlm. & Volkm.-Kohlm., Mycol. Res. 97(6): 759 (1993)
Index Fungorum number: IF22429; 4 species with sequence data.
Type species – Kallichroma tethys (Kohlm. & E. Kohlm.) Kohlm. & Volkm.-Kohlm.
Laniatria Döbbeler & P.G. Davison, Nova Hedwigia 106(1-2): 239 (2017)
Index Fungorum number: IF821328; 1 morphological species
Type species – Laniatria myxostoma Döbbeler & P.G. Davison
Lasionectria (Sacc.) Cooke, Grevillea 12(no. 64): 111 (1884)
Index Fungorum number: IF2652; 13 morphological species (Species Fungorum 2020), 10
species with sequence data.
Type species – Lasionectria mantuana (Sacc.) Cooke
Lasionectriella Lechat & J. Fourn., Ascomycete.org 8(2): 59 (2016)
Index Fungorum number: IF815673; 2 species with sequence data
454
Type species – Lasionectriella rubioi Lechat & J. Fourn.
Mycoarachis Malloch & Cain, Can. J. Bot. 48(10): 1820 (1970)
Index Fungorum number: IF3304; 2 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Mycoarachis inversa Malloch & Cain
Mycocitrus Möller, Bot. Mitt. Trop. 9: 297 (1901)
Index Fungorum number: IF3313; 2 species with sequence data.
Type species – Mycocitrus aurantium Möller
Nectriella Nitschke, Jb. nassau. Ver. Naturk. 23–24: 175 (1870)
Index Fungorum number: IF3432; 52 morphological species (Species Fungorum 2020), 3
species with sequence data.
Type species – Nectriella fuckelii Nitschke
Nectriopsis Maire, Annls mycol. 9(4): 323 (1911)
Index Fungorum number: IF3434; 72 morphological species (Species Fungorum 2020), 11
species with sequence data.
Type species – Nectriopsis violacea (J.C. Schmidt) Maire
Nigrosabulum Malloch & Cain, Can. J. Bot. 48(10): 1822 (1970)
Index Fungorum number: IF3507; 1 species with sequence data.
Type species – Nigrosabulum globosum Malloch & Cain
Ochronectria Rossman & Samuels, Stud. Mycol. 42: 53 (1999)
Index Fungorum number: IF28315; 2 species with sequence data.
Type species – Ochronectria calami (Henn. & E. Nyman) Rossman & Samuels
Ovicuculispora Etayo, Bull. Soc. linn. Provence 61: 110 (2010)
Index Fungorum number: IF565893; 2 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Ovicuculispora parmeliae (Berk. & M.A. Curtis) Etayo
Paracylindrocarpon Crous, Roets & L. Lombard, Persoonia 36: 367 (2016)
Index Fungorum number: IF817039; 4 species with sequence data.
Type species – Paracylindrocarpon aloicola Crous et al.
Paranectria Sacc., Michelia 1(no. 3): 317 (1878)
Index Fungorum number: IF3707; 8 morphological species (Species Fungorum 2020).
Type species – Paranectria affinis (Grev.) Sacc.
Periantria Döbbeler & P.G. Davison, Nova Hedwigia 106(1-2): 242 (2017)
Index Fungorum number: IF821330; 2 morphological species (Species Fungorum 2020).
Type species – Periantria frullaniae (Racov.) Döbbeler & P.G. Davison
Peristomialis (W. Phillips) Boud., Hist. Class. Discom. Eur. (Paris): 116 (1907)
Index Fungorum number: IF3830; 6 morphological species (Species Fungorum 2020),
molecular data available for an unnamed species.
Type species – Peristomialis berkeleyi Boud.
Pronectria Clem., Gen. fung., Edn 2 (Minneapolis): 78, 282 (1931)
455
Index Fungorum number: IF4370; 45 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Pronectria lichenicola (Cooke) Clem.
Protocreopsis Yoshim. Doi, Kew Bull. 31(3): 551 (1977)
Index Fungorum number: IF4384; 12 morphological species (Species Fungorum 2020), 5
species with sequence data.
Type species – Protocreopsis zingibericola Yoshim.
Rhopalocladium Schroers, Samuels & W. Gams, Mycologia 91(2): 375 (1999)
Index Fungorum number: IF28338; 1 morphological species.
Type species – Rhopalocladium myxophilum Schroers, Samuels & W. Gams
Roumegueriella Speg., Revue mycol., Toulouse 2 (no. 1): 18 (1880)
Index Fungorum number: IF4798; 3 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Roumegueriella muricospora Speg.
Selinia P. Karst., Meddn Soc. Fauna Flora fenn. 1: 57 (1876)
Index Fungorum number: IF4996; 5 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Selinia pulchra (G. Winter) Sacc.
Stephanonectria Schroers & Samuels, Sydowia 51(1): 116 (1999)
Index Fungorum number: IF28373; 1 species with sequence data.
Type species – Stephanonectria keithii (Berk. & Broome) Schroers & Samuels
Stilbocrea Pat., Bull. Soc. mycol. Fr. 16: 188, 186 (1900)
Index Fungorum number: IF5263; 7 morphological species (Species Fungorum 2020), 3
species with sequence data.
Type species – Stilbocrea dussii Pat.
Stromatonectria Jaklitsch & Voglmayr, Mycologia 103(2): 435 (2011)
Index Fungorum number: IF518755; 1 species with sequence data.
Type species – Stromatonectria caraganae (Höhn.) Jaklitsch & Voglmayr
Synnemellisia N.K. Rao, Manohar. & Goos, Mycologia 80(6): 896 (1989)
Index Fungorum number: IF11201; 2 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Synnemellisia hyalospora N.K. Rao, Manohar. & Goos.
Trichonectria Kirschst., Verh. bot. Ver. Prov. Brandenb. 48: 60 (1907)
Index Fungorum number: IF5563; 21 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Trichonectria aculeata Kirschst.
Verrucostoma Hirooka, Tak. Kobay. & P. Chaverri, Mycologia 102(2): 422 (2010)
Index Fungorum number: IF513353; 2 species with sequence data.
Type species – Verrucostoma freycinetiae Hirooka, Tak. Kobay. & P. Chaverri
Xanthonectria J. Fourn. & P.-A. Moreau, Ascomycete.org 8(4): 173 (2016)
Index Fungorum number: IF816960; 1 species with sequence data
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Type species – Xanthonectria pseudopeziza (Desm.) Lechat, J. Fourn. & P.-A. Moreau
Boliniaceae Rick, Brotéria, sér. bot. 25: 65 (1931)
Index Fungorum number: IF80526; Facesoffungi number: FoF01129; 46 species.
Saprobic on decorticated decaying wood, occasionally hypersaprobic on polypores, in
terrestrial habitats. Sexual morph: Stroma absent or present, immersed to superficial, often
erumpent, clypeate to irregular in shape, light brown to black, coriaceous, membranaceous or
powdered or furfuraceous, with or without brown, septate interwoven hyphae, if present
surrounding the apex of the ascomata. Ascomata perithecial, solitary to gregarious, immersed
within the stromata, monostichous or polystichous or erumpent to superficial, brown to black,
globose, cylindrical to obpyriform, coriaceous or membranaceous, smooth, with or without papilla,
occasionally of stellate appearance. Ostiole periphysate. Peridium two-layered, outer layer
composed of brown, thick-walled cells of textura angularis or textura intricata; inner layer
composed of hyaline, thin-walled cells of textura prismatica. Paraphyses abundant, persistent,
filamentous, tapering, septate, branched. Asci 8-spored, unitunicate, cylindrical to clavate, long
pedicellate, some with an apical ring. Ascospores 2–3-seriate, hyaline or brown, with an
inconspicuous apical germ pore at one or both ends that may be covered by an indistinct hyaline
cap, ellipsoid to cylindrical, straight to slightly curved to suballantoid, sometimes laterally
flattened, aseptate or septate, smooth-walled, mostly with guttules. Asexual morph: Undetermined
(adapted from Maharachchimbura et al. 2016b).
Type genus – Camarops P. Karst.
Notes – Boliniaceae was introduced by Rick (1931) based on black stroma and cylindrical
asci with smooth-walled ascospores, and is typified by Camarops (Karsten 1873). Earlier, this
family was placed in Xylariales based on its morphological similarities (Barr 1990b, Romero &
Samuels 1991). Andersson et al. (1995) concluded that Boliniaceae was more closely related to
Sordariales based on SSU sequence data. This family was subsequently placed in Boliniales by
Kirk et al. (2001). Based on phylogenetic and morphological data, six genera were accepted in
Boliniaceae (Lumbsch & Huhndorf 2010). Untereiner et al. (2013) added Apiorhynchostoma and
Pseudovalsaria based on LSU rDNA sequence data. A key to Boliniaceae genera was given in
Untereiner et al. (2013).
Ecological and economic significance of Boliniaceae
Most Boliniaceae species are saprobic on wood and widespread in temperate regions
(Maharachchikumbura et al. 2016b).
Genera included in Boliniaceae
Apiocamarops Samuels & J.D. Rogers, Mycotaxon 28(1): 54 (1987)
Index Fungorum number: IF25015; 3 morphological species (Species Fungorum 2020).
Type species – Apiocamarops alba Samuels & J.D. Rogers
Notes – Apiocamarops is characterized by ascomata surrounded by furfuraceous outer
coating, cylindrical asci and brown ascospores (Samuels & Rogers 1987). Four species were
accommodated in Apiocamarops (A. alba, A. cryptocellula, A. luquilloensis, and A. pulvinata) and
they are related to Camarops with turbinate stromata and ascospores with a germ pore at one end
(Samuels & Rogers 1987, Rogers & Samuels 1988, Reagan & Waide 1996, Rogers & Ju 2003).
Apiocamarops has furfuraceous, hyaline ascomata and 1-septate ascospores and differs from
Camarops which has black ascomata and aseptate ascospores (Samuels & Rogers 1987).
Apiorhynchostoma Petr., Annls mycol. 21(3/4): 185 (1923)
Index Fungorum number: IF261; 3 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Apiorhynchostoma apiculatum (Sacc.) Petr.
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Notes – Untereiner et al. (2013) revalidated Apiorhynchostoma based on herbarium material
and Apiorhynchostoma curreyi.
Figure 50 – Apiocamarops alba (Material examined – BRAZIL, Amazonas, Plateau of Serra
Araca. Alt. 1250 m. (4101 ft.) 00° 54' 06"N, 63° 22' 29"W (0.902, -63.375), Cloud forest, on
decorticated wood, 19 February 1984, collected by G. J. Samuels 486 with G. T. Francis, J. Pipoly,
NY no. 00986029, holotype). a Material label. b Material. c Ascomata. d Ascoma cross section. e
Peridium. f Paraphyses. g-i Asci. j-m Ascospores. Scale bars: b = 2000 µm, c-d = 200 µm, e = 50
µm, g-i = 20 µm, f = 10 µm, j-m = 5 µm.
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Camaropella Lar.N. Vassiljeva, Mikol. Fitopatol. 31(1): 6 (1997)
Index Fungorum number: IF27727; 2 species with sequence data.
Type species – Camaropella pugillus (Schwein.) Lar.N. Vassiljeva
Notes – Camaropella was introduced to accommodate two species, C. lutea and C. pugillus
occurring on dead wood (Vassiljeva 1997). They could not be clearly distinguished from other
Camarops species in the family based on LSU-ITS-tub2 sequence analysis (Fig. 6).
Camarops P. Karst., Bidr. Känn. Finl. Nat. Folk 23: 6, 53 (1873)
Index Fungorum number: IF775; 23 morphological species (Species Fungorum 2020), 9
species with sequence data.
Type species – Camarops hypoxyloides P. Karst.
Notes – Karsten (1873) introduced Camarops based on C. hypoxyloides. The name Bolinia
was first introduced as a subgenus by Nitschke (1867) and raised to generic rank by Saccardo
(1882). Shear (1938) considered Bolinia tubulina to be a homonym of Camarops tubulina.
Nannfeldt (1972) reviewed the taxonomy of these taxa and concluded that Camarops and Bolinia
are two different genera, but having similar characters. Lumbsch & Huhndorf (2010) proposed
Bolinia to be a synonym of Camarops.
Cornipulvina Huhndorf, A.N. Mill., F.A. Fernández & Lodge, Fungal Divers. 20: 61 (2005)
Index Fungorum number: IF501318; 1 species with sequence data.
Type species – Cornipulvina ellipsoides Huhndorf, A.N. Mill., F.A. Fernández & Lodge
Notes – The monotypic Cornipulvina was included as a member of Boliniaceae based on
LSU sequences analysis and the unique hyaline ascospores without germ pores in this family
(Huhndorf et al. 2005).
Endoxyla Fuckel, Jb. nassau. Ver. Naturk. 25-26: 321 (1871)
Index Fungorum number: IF1813; 8 morphological species (Species Fungorum 2020), 3
species with sequence data.
Type species – Endoxyla macrostoma Fuckel
Notes – Endoxyla was established by Fuckel (1872) based on Endoxyla macrostoma.
Untereiner et al. (2013) provided the type strains of Endoxyla macrostoma, E. mallochii and E.
operculata.
Mollicamarops Lar.N. Vassiljeva, Mycotaxon 99: 160 (2007)
Index Fungorum number: IF510602; 1 species with sequence data.
Type species – Mollicamarops stellata Lar.N. Vassiljeva
Notes – Vasilyeva (2007) introduced the monotypic genus Mollicamarops from dead wood in
Russia.
Neohypodiscus J.D. Rogers, Y.M. Ju & Læssøe, Mycologia 86(5): 684 (1994)
Index Fungorum number: IF27387; 3 morphological species (Species Fungorum 2020).
Type species – Neohypodiscus rickii (Lloyd) J.D. Rogers, Y.M. Ju & Læssøe
Notes – Neohypodiscus was introduced by Rogers et al. (1994) in Amphisphaeriaceae and
Andersson et al. (1995) transferred it to Boliniaceae based on resemblance of characters to
Camarops.
Pseudovalsaria Spooner, Trans. Br. mycol. Soc. 86(3): 405 (1986)
Index Fungorum number: IF25775; 2 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Pseudovalsaria foedans (P. Karst.) Spooner
Notes – Spooner (1986) introduced Pseudovalsaria as a member of Trichosphaeriaceae, Barr
(1994) transferred it to Clypeosphaeriaceae and Rappaz (1995) referred it to the Boliniaceae.
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Untereiner et al. (2013) provided sequence data for Pseudovalsaria ferruginea which was closely
related to Apiorhynchostoma (Boliniaceae) (Fig. 6).
Cainiaceae J.C. Krug, Sydowia 30 (1–6):123 (1978)
Index Fungorum number: IF80542; Facesoffungi number: FoF00687; 50 species.
Saprobic on dead grasses, bamboo and other monocotyledons and fabaceous dicotyledons,
appearing as shiny black dots, slightly effuse from the substrate, or pathogenic. Sexual morph:
Pseudostromata poorly developed or lacking, or sometimes clypeate, scattered, superficial, domeshaped or slightly effuse, dark brown to black. Ascomata immersed, solitary or aggregated, globose
to subglobose, coriaceous, brown, ostiolate. Ostiolar papilla short, internally lined with hyaline
periphyses. Peridium one or two-layered, outer layer comprising thick-walled, brown cells of
textura angularis and inner layer comprising hyaline thin-walled cells of textura angularis.
Paraphyses abundant, filamentous, slightly constricted at the septa. Asci 8-spored, unitunicate,
cylindrical to broadly cylindrical, short pedicellate, with a complex, J+, cylindrical apical ring or
series of rings. Ascospores uniseriate, overlapping uniseriate to biseriate, hyaline when young and
dark brown at maturity, sphaerical to ellipsoidal, unicellular to 1-septate, slightly constricted at the
septum, wall ornamented with longitudinal germ slits or germ pores, surrounded by a gelatinous
sheath. Asexual morph: Coelomycetous. Conidiomata pycnidial, scattered, immersed, globose to
subglobose, black. Conidiophores hyaline, denticulate, sympodially proliferating. Conidiogenous
cells with 1–3 phialides, filiform, branched or simple, septate, hyaline. Conidia elongate fusiform,
falcate to lunate, unicellular or septate, hyaline, with pointed ends (adapted from
Maharachchimbura et al. 2016b).
Type genus – Cainia Arx & E. Müll.
Notes – Cainiaceae was introduced by Krug (1978) to accommodate species of Cainia with
unique apical rings in the asci, consisting of a series of rings, and ascospores with longitudinal
germ slits. Kang et al. (1999b) revived Cainiaceae and included the genera Arecophila,
Atrotorquata, Cainia, Ceriophora, Reticulosphaeria and Ommatomyces. The phylogenetic study
conducted by Jeewon et al. (2003b) provides molecular evidence to support the generic status of
Arecophila in Cainiaceae. Seynesia was included in the family based on phylogenetic analyses
(Maharachchikumbura et al. 2015). Several phylogenetic and morphological studies accepted
Amphibambusa, Arecophila, Atrotorquata, Cainia and Seynesia in this family (Smith et al. 2003,
Maharachchikumbura et al. 2015, 2016b, Liu et al. 2015, Senanayake et al. 2015, Wijayawardene
et al. 2017a). Maharachchikumbura et al. (2015, 2016b), Senanayake et al. (2015) and Jaklitsch et
al. (2016b) placed the family within the Xylariales Hongsanan et al. (2017) provided divergence
estimates and referred the family to Xylariomycetidae family incertae sedis. We introduce a new
monotypic genus Alishanica with the type A. miscanthii collected from Taiwan.
Ecological and economic significance of Cainiaceae
Cainiaceae species are commonly recorded on dead culms of monocotyledons, and leaves and
dead branches of dicotyledons as saprobes and important as decomposers in tropical and temperate
regions. There are a few records of some pathogenic species in the group such as Cainia desmazieri
(Krug 1978, Kang et al. 1999a).
Genera included in Cainiaceae
Alishanica Karun., C.H. Kuo & K.D. Hyde, gen. nov.
Index Fungorum number: IF556756; Facesoffungi number: FoF06335; 1 species with
sequence data.
Etymology – The specific epithet reflects the name of Ali Mountain from where the species
was collected.
Saprobic on dead sheaths of Miscanthus sinansis (Poaceae). Sexual morph: Ascomata
immersed beneath blackened aggregated clypeus on the surface of dead sheath, loosely aggregated
or rarely solitary; dark brown to black, globose to subglobose, slightly depressed, uniloculate with
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distinct centrally erumpent ostiole with periphyses, surrounded by distinct, shiny, black flanges the
tissue spreading down along the papilla. Peridium comprising 4–5 cell layers of thin-walled, brown
cells of textura angularis, inwardly lighter. Paraphyses filamentous, distinctly septate, embedded
in a gelatinous matrix. Asci 8-spored, unitunicate, cylindrical, short pedicellate, slightly truncate at
the apex with a wedge-shaped, J+, subapical ring. Ascospores overlapping uniseriate, brown,
ellipsoidal, slightly tapering at the ends, equally 2-celled and guttulate at both cells, constricted at
the septum, with striations, surrounded by a thick, hyaline mucilaginous sheath, parallel to the
margin of the spore. Asexual morph: Undetermined.
Type species – Alishanica miscanthii A. Karun., C.H. Kuo & K.D. Hyde
Notes – Alishanica miscanthi resembles Cainiaceae in having clypeate, ostiolate ascomata,
unitunicate, cylindrical asci, and apical ring and ascospores with a number of longitudinal germslits resembling ridges. The genus is phylogenetically well-supported with bayesian posterior
probability (0.94 PP) and moderate maximum likelihood (53%) and maximum parsimony (53%)
support (Fig. 4).
Alishanica miscanthii Karun., C.H. Kuo & K.D. Hyde, sp. nov.
Fig. 51
Index Fungorum number: IF556757; Facesoffungi number: FoF06776
Etymology – The specific epithet reflects the host genus Miscanthus.
Holotype – MFLU 19-2333.
Saprobic on dead sheaths of Miscanthus sinansis (Poaceae). Sexual morph: Ascomata 277–
272 µm high × 288–285 µm diam. (x̅ = 275 × 287 µm, n = 8), immersed beneath blackened
aggregated clypeus of the surface of dead sheath, loosely aggregated or rarely solitary; dark brown
to black, globose to subglobose, slightly depressed, uniloculate. Ostiole 110–108 μm long, 52–51
μm diameter (x̅ = 108 × 52 μm, n = 5), centrally erumpent, with periphyses, surrounded by distinct
shiny black flanges the tissue spreads down along the papilla. Peridium 52–60 μm wide,
comprising 4–5 cell layers of thin-walled, brown cells of textura angularis, inwardly lighter.
Paraphyses filamentous, distinctly septate, embedded in a hyaline gelatinous matrix. Asci 173–179
× 23–31 μm (x̅ = 176 × 28 μm, n = 30), 8-spored, unitunicate, cylindrical, short pedicellate, slightly
truncate at the apex, with a wedge-shaped, J+, subapical ring. Ascospores 59–62 × 19–21 μm (x̅ =
61 × 20 μm, n = 40), overlapping, uniseriate, ellipsoidal, slightly tapering at the ends, equally 2celled and guttulate at both cells, constricted at the septum, brown with striations, surrounded by a
thick, hyaline mucilaginous sheath, subglobose, parallel to the margin of the spore. Asexual morph:
Undetermined.
Cultural characteristics – Ascospores germinating on PDA within 12 h and germ tubes
produced from both ends. Colonies growing on PDA, cottony, white to pale brown, mycelium
superficial, irregular edge; asexual spores or sexual spores were not formed within 90 days.
Material examined – Taiwan, Chiayi Province, Ali Mountain, Kwang Hwa, on dead sheaths
of Miscanthus sinansis (Poaceae), 5 May 2018, A. Karunarathna AKTW 44, MFLU 19-2333,
holotype, FU31025, isotype; extype living culture FU31025.
GenBank numbers – ITS: MK503821, LSU: MK503827, SSU: MK503833.
Notes – Alishanica and Cainia (C. desmazieresii and C. anthoxanthis) share similar
ascospore characters by ellipsoid with slightly tapering ends, 2 equal cells with guttules at both
ends, being constricted at the septum, brown with striations and surrounded by a thick, hyaline
mucilaginous sheath. However, Alishanica miscanthii is distinct from Cainia desmazieresii and C.
anthoxanthis in having an aggregated clypeus, distinct long ostioles with periphyses, distinct shiny
black flanges around the ostioles, cylindrical asci and with uniseriate, slightly curved and distinctly
closely striated ascospores (Krug 1978, Senanayake et al. 2015). Alishanica miscanthii can be
distinguished from Seynesia by the absence of apical mucilaginous appendages and spore
dimensions (Hyde 1995c). Atrotorquata shares more similarities with Cainia and Alishanica and
has distinct in spore characters (Krug 1978, Kohlmeyer & Volkmann-Kohlmeyer 1993,
Senanayake et al. 2015). Furthermore, the separation of A. miscanthii is supported by sequence
data.
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Amphibambusa D.Q. Dai & K.D. Hyde, Fungal Divers. 72: 7 (2015)
Index Fungorum number: IF550940; 1 species with sequence data.
Type species – Amphibambusa bambusicola D.Q. Dai & K.D. Hyde
Notes – Liu et al. (2015) introduced Amphibambusa, a bambusicolous saprobic genus from
Thailand. The monotypic Amphibambusa is characterized by immersed ascomata surrounded by a
small blackened clypeus and ostiolar opening surrounded by a white margin, and cylindrical asci
with fusiform ascospores surrounded by a wide gelatinous sheath.
Figure 51 – Alishanica miscanthii (MFLU 19-2333 holotype). a Appearance of ascomata on the
host (Miscanthus gigantius). b Clypeus with distinct flanges. c Section through ascoma. d Section
through peridium. e Section through ostiole. f Section through flange. g Pseudoparaphyses. h-k
Different developing stages of the asci. l Apical ring. m-p Ascospores. q Ascospore surrounded by
mucilaginous sheath. r Germinating ascospore. s, t Culture characteristics on PDA (s = from below,
t = from above). Scale bars: c-g = 50 μm, h-r = 10 μm.
Arecophila K.D. Hyde, Nova Hedwigia 63(1–2):82 (1996)
Index Fungorum number: IF27653; 14 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Arecophila gulubiicola K.D. Hyde
Notes – Arecophila was introduced to accommodate species from monocotyledons that have
striate or verrucose ascospores (Hyde 1996b). Lee et al. (2013) investigated the potential antiinflammatory effect of A. saccharicola.
Atrotorquata Kohlm. & Volkm.-Kohlm., Syst. Ascom. 12(1–2):8 (1993)
Index Fungorum number: IF26457; 2 species with sequence data.
Type species – Atrotorquata lineata Kohlm. & Volkm.-Kohlm.
Notes – Atrotorquata is characterized in unitunicate asci with a J+, subapical ring and brown
2-celled ascospores similar to Amphisphaeria (Kohlmeyer & Volkmann-Kohlmeyer 1993). There
are two accepted species with known sexual and undetermined asexual morphs (Kohlmeyer &
Volkmann-Kohlmeyer 1993).
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Cainia Arx & E. Müll., Acta bot. neerl. 4(1): 111 (1955)
Index Fungorum number: IF719; 5 morphological species (Species Fungorum 2020), 4
species with sequence data.
Type species – Cainia graminis (Niessl) Arx & E. Müll.
Notes – The genus was placed in a distinct family because of its longitudinal germ slits in the
ascospores and the complex ascus ring comprising a series of rings (Krug 1978). Senanayake et al.
(2015) designated a reference specimen for the type Cainia graminis and provided phylogenetic
placement to accommodate genera in Cainiaceae.
Seynesia Sacc., Syll. fung. (Abellini) 2:668 (1883)
Index Fungorum number: IF5017; 27 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Seynesia nobilis (Welw. & Curr.) Sacc.
Notes – Seynesia species are mostly confined to palms and characterized by the welldeveloped clypeus, the ascospores having cap-like appendages and germ slits in each cell, and the
asci having a complex wedge-shaped subapical ring (Hyde 1995c). There are no recent studies on
this genus and cultures and sequences are unavailable for the type.
Calcarisporiaceae Jing Z. Sun, X.Z. Liu & K.D. Hyde, Mycol Prog 16 (4): 435 (2017)
Index Fungorum number: IF817660; Facesoffungi number: FoF02511; 8 species.
Fungicolous, parasitic, endophytic or saprobic on higher ascomycetes and basidiomycetes,
and leaves and branches of trees. Sexual morph: Undetermined. Asexual morph: Conidiophores
macronematous, mononematous, erect, unbranched or verticillate, branched hyaline, straight or
flexuous. Conidiogenous cells holoblastic, polyblastic, discrete, subulate, hyaline. Conidia ovoid
to ellipsoid, or acerose to narrowly obclavate, aseptate, hyaline, straight to curved (adapted from
Sun et al. 2017).
Type genus – Calcarisporium Preuss
Notes – Multi-gene analysis revealed that Calcarisporium species form a monophyletic
lineage which is distinct from other families in Hypocreales. This family is dist inguished from
other families by its verticillate conidiophores with a narrow apical part and monoblastic to
polyblastic, sympodial and denticulate conidiogenous cells at the apex (Sun et al. 2017).
Calcarisporium arbuscula is illustrated in the entry.
Ecological and economic significance of Calcarisporiaceae
Calcarisporium species are fungicolous, caulicolous and foliicolous. Some Calcarisporium
species are saprobes of decaying leaves (C. phaeopodium) and dead bark (C. acerosum).
Calcarisporium arbuscula is a common parasite on higher ascomycetes and basidiomycetes,
occasionally occurring on wood (Sutton 1973, de Hoog 1974). In addition, C. arbuscula has been
reported as an endophyte in sporophores of Lactarius and Russula (Watson 1955).
Genus included in Calcarisporiaceae
Calcarisporium Preuss, Linnaea 24: 124 (1851)
Index Fungorum number: IF7465; 8 morphological species (Species Fungorum 2020), 4
species with sequence data.
Type species – Calcarisporium arbuscula Preuss
Notes – Calcarisporium occurs on Pezizaceae (Preuss 1851), Russulaceae (Anke & Sterner
1988) and Cordyceps (Sun et al. 2016, 2019), and litter leaf (Somrithipol & Jones 2006). Species
are distinguished by hyaline, erect, verticillate conidiophores with sympodial and denticulate,
polyblastic conidiation.
Calosphaeriaceae Munk, Dansk bot. Ark. 17(no. 1): 278 (1957)
Index Fungorum number: IF80548; Facesoffungi number: FoF01133; 67 species.
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Saprobic on wood or decorticated wood in terrestrial habitats, sometimes hypersaprobic on
old stromata or ascomata of other fungi, and several species have been isolated from wood of fruit
trees showing canker symptoms. Sexual morph: Stroma absent. Ascomata perithecial, solitary to
gregarious, dark brown to black, superficial or typically erumpent or immersed, globose to
subglobose, coriaceous or membranous, tuberculate or smooth or with brown, septate, hyphal
coating, papillate or with a cylindrical neck. Ostioles periphysate. Peridium two-layered, outer
layer coriaceous or membranous, composed of dark brown, reddish-brown to brown cells of textura
angularis or prismatica or porrecta; inner layer membranous, composed of hyaline cells of textura
prismatica. Paraphyses numerous, broad, septate, unbranched, tapering. Ascogenous hyphae
discrete, hyaline, smooth, branched, producing a sympodial sequence of hyaline, ovoid to
ellipsoidal cells, often with mucronate apex, in dense clusters, each giving rise to an ascus. Asci 8spored, unitunicate, clavate, long or short pedicellate, mostly in fascicles, with a conspicuously
thickened apex lacking a visible discharge mechanism. Ascospores 2-seriate or in a fascicle, hyaline
to light brown, allantoid, suballantoid, oblong to subcylindrical, aseptate or transversely septate,
smooth-walled. Asexual morph: Hyphomycetous, Acremonium- and phialophora-like.
Conidiophores micronematous or semi-macronematous, brown or hyaline, straight or flexuous,
septate, branched or unbranched. Conidiogenous cells phialides or adelophialdes, terminal,
intercalary or lateral, hyaline to subhyaline, ampulliform to subcylindrical. Conidia aseptate,
allantoid, oblong or subcylindrical, hyaline, smooth (adapted from Réblová et al. 2015b).
Type genus – Calosphaeria Tul. & C. Tul.
Notes – Calosphaeriaceae was introduced by Munk (1957) for perithecial taxa with a unique
ascoma centrum in Sordariomycetes. Lumbsch & Huhndorf (2010) included eight genera
(Calosphaeria, Conidiotheca, Jattaea, Kacosphaeria, Phragmocalosphaeria, Sulcatistroma,
Togniniella, Wegelina) in this family. Réblová (2011) proposed Phragmocalosphaeria and
Wegelina as synonyms of Jattaea based on similar septate ascospores. Maharachchikumbura et al.
(2016b) accepted Calosphaeria, Jattaea, Kacosphaeria, Sulcatistroma, Togniniella and
Tulipispora. A survey of Calosphaeriaceae was undertaken by Réblová et al. (2015b) and the
genera Calosphaeria, Flabellascus, Jattaea and Togniniella were accepted. Their asexual morphs
are dematiaceous phialidic hyphomycetes and numerous sexual-asexual relationships have been
experimentally proven (Damm et al. 2008, Réblová 2011, Réblová et al. 2004, 2015b).
Calosphaeriophora and Phaeocrella were originally described as asexual morphs for Calosphaeria
and Togniniella (Réblová et al. 2004), however, in accordance with the abolishment of dual
nomenclature for pleomorphic fungi, the names of their sexual morphs were selected following the
principle of priority (Réblová et al. 2015b). In the same publication, three other genera listed in the
Outline of Ascomycota (Lumbsch & Huhndorf 2010), i.e. Conidiotheca, Sulcatistroma and
Kacosphaeria, were excluded from the broadly perceived Calosphaeriaceae, however, without
DNA sequence data they could not be reassigned to other families. In addition, the asexual genus
Tulipispora was considered a member of Calosphaeriaceae (Maharachchikumbura et al. 2016b,
Wijayawardene et al. 2018a). Unfortunately, sequence data for Tulipispora is not available. We
therefore exclude it from Calosphaeriaceae.
Ecological and economic significance of Calosphaeriaceae
Most Calosphaeriaceae species are saprobic on wood and widespread in temperate regions,
especially in Europe and western Asia (Réblová et al. 2004, Damm et al. 2008, Trouillas et al.
2010a, 2012, Réblová 2011, Chakusary et al. 2015). Some species of Calosphaeria and Jattaea,
such as Calosphaeria pulchella, Jattaea algeriensis and J. prunicola have been reported as plant
pathogens associated with disease of woody plants (Damm et al. 2008, Chakusary et al. 2015).
Genera included in Calosphaeriaceae
Calosphaeria Tul. & C. Tul., Select. fung. carpol. (Paris) 2: 108 (1863)
Index Fungorum number: IF753; 39 morphological species (Species Fungorum 2020), 2
species with sequence data.
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Type species – Calosphaeria princeps Tul. & C. Tul.
Notes – Calosphaeria was introduced by Tulasne & Tulasne (1863) and typified by C.
princeps. The acremonium-like asexual morphs, formerly referred to Calosphaeriophora, have
been experimentally linked to Calosphaeria africana (Damm et al. 2008) and C. pulchella
(Réblová et al. 2004). Jattaea discreta is illustrated in this entry for the sexual morph of the family.
Figure 52 – Calcarisporium arbuscula (redrawn from de Hoog (1974) and Sun et al. (2017)). a
Conidiophore at low magnification. b Tip of conidiophore. c Conidiophores with conidia. d
Conidiophores without conidia. e Conidiogenous scars and conidia. f Conidia. Scale bars: a = 100
µm, b-f = 10 µm.
Flabellascus Réblová, PLoS ONE 10(12): e0144616, 15 (2015)
Index Fungorum number: IF814416; 1 species with sequence data.
Type species – Flabellascus tenuirostris Réblová
Notes – The monotypic genus Flabellascus is characterized by tiny asci with ascogenous
hyphae and ampulliform phialides with aseptate, allantoid conidia (Réblová et al. 2015b). The
sexual morph of Flabellascus is similar to Jattaea and they are distinguished by the different
asexual morphs. However, Flabellascus constitutes an independent clade within Calosphaeriaceae
based on a multi-gene analysis (Réblová et al. 2015b; Fig. 7).
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Figure 53 – Jattaea discreta (Material examined – ITALY, Belluno, Veneta, on the bark of
branches, Autumn 1879, NY no.00986029, isotype). a Material label. b Ascomata. c Ascoma in
cross section. d Peridium. e Asci and paraphyses attached to ascogenous hyphae with ellipsoid to
obpyriform cells (arrow head indicates ascogenous hyphae). f, g Asci. h-l Ascospores. Scale bars: b
= 500 µm, c = 200 µm, d = 50 µm, e = 20 µm, f, g = 10 µm, h-l = 5 µm.
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Jattaea Berl., Icon. fung. (Abellini) 3(1-2): 6 (1900)
Index Fungorum number: IF2528; 26 morphological species (Species Fungorum 2020), 11
species with sequence data.
Type species – Jattaea algeriensis Berl.
Notes – Jattaea has long-necked ascomata containing hyaline, cylindrical paraphyses,
branched ascogenous hyphae, clavate asci and oblong to subglobose ascospores. The asexual
morph has always been referred to phialophora-like (Damm et al. 2008). The monophyletic Jattaea
is related to Calosphaeria and Togniniella within Calosphaeriaceae based on multi-gene analysis
(Réblová et al. 2015b; Fig. 7).
Figure 54 – Phialophora-like asexual morph of Jattaea aurea (redrawn from Réblová et al. 2015b)
Conidiophores with conidia of Phialophora asexual morph. Scale bar = 10 µm.
Togniniella Réblová, L. Mostert, W. Gams & Crous, Stud. Mycol. 50(2): 543 (2004)
Index Fungorum number: IF500157; 1 species with sequence data.
Type species – Togniniella microspora (Ellis & Everh.) Réblová
Notes – Togniniella has glabrous ascomata with branched ascogenous hyphae, paraphyses
longer than asci and suballantoid ascospores. After re-examining the type material of T. acerosa,
Réblová (2011) concluded that Togniniella microspora (synonym of Ceratostomella microspora) is
identical to T. acerosa and accepted T. microspora as the only species in Togniniella. Réblová et al.
(2004) and Réblová (2011) found that Togniniella constitutes an independent clade and is related to
Calosphaeria. In this study, Flabellascus is closely related with this genus based on tub2-ITS-actLSU analysis (Fig. 7).
Castanediellaceae Hern.-Restr., Guarro & Crous, Stud. Mycol. 86: 93 (2017)
Index Fungorum number: IF820354; Facesoffungi number: FoF06777; 16 species.
Foliicolous, saprobic or associated to leaf spots. Sexual morph: Undetermined. Asexual
morph: Conidiophores macronematous or semi-macronematous, mononematous or aggregated in
sporodochia, branched or unbranched, brown to pale brown. Conidiogenous cells mono- to
polyblastic, sympodial, integrated, solitary, or in whorls, cylindrical to lageniform, hyaline to
subhyaline, smooth. Conidial secession schizolytic. Conidia unicellular or transversely septate,
fusiform or lunate, hyaline (adapted from Hernández-Restrepo et al. 2017).
Type genus – Castanediella Hern.-Restr.
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Notes – LSU based phylogenetic analyses revealed that Castanediellaceae is monophyletic
and represents a distinct taxonomic group at the family level closely related to Beltraniaceae in
Xylariales (Hernández-Restrepo et al. 2017). Lin et al. (2019a) provided a synopsis of hitherto
described Castanediella species and accepted 16 species in the genus.
Ecological and economic significance of Castanediellaceae
Castanediellaceae species have been collected mainly on dead wood, leaves, leaf spots and
rotten leaves, including leaf litter of Eucalyptus species, as saprobes. Since some species have been
isolated from leaf spots, their parasitic nature might be hidden. However, they have a significance
role as saprobes (Crous et al. 2015c, 2016a, b, Hernández-Restrepo et al. 2016b, 2017, Wanasinghe
et al. 2018, Hyde et al. 2019a, Lin et al. 2019a).
Genus included in Castanediellaceae
Castanediella Hern.-Restr., Crous & M.J. Wingf., Persoonia 34: 187 (2015)
Index Fungorum number: IF811878; 16 morphological species (Species Fungorum 2020),
11 with sequence data.
Type species – Castanediella acaciae Crous, Hern.-Restr. & M.J. Wingf.
Notes – Castanediella was introduced by Crous et al. (2015c) and placed in Xylariales genera
incertae sedis. The genus is characterized by a hyphomycetous asexual morph with branched or
unbranched conidiophores, mono- to polyblastic conidiogenous cells and straight to slightly curved,
0–1-septate conidia. The species are commonly isolated from leaf litter, leaf spots and wood.
Currently, the genus comprises 16 species (Crous et al. 2015c, 2016a, b, 2018c, HernándezRestrepo et al. 2016b, 2017, Wanasinghe et al. 2018, Hyde et al. 2019a, Lin et al. 2019a). In this
entry we illustrate Castanediella brevis (Fig. 55).
Figure 55 – Castanediella brevis (Material examined – THAILAND, Lampang, Amphoe Mueang
Pan, Tambon Chae Son, on decaying leaves, 24 September 2016, Chuangen Lin, LCG 10-1, MFLU
18-1695, holotype). a Host material. b-e Conidiophores on the host surface, conidiophores,
conidiogenous cells with conidia. f-i Conidia. Scale bars: b-d = 5 μm, e-i = 10 μm.
Catabotryaceae Petr., Mycotaxon 39: 83 (1990)
Index Fungorum number: IF90924; Facesoffungi number: FoF01372; 2 species.
Saprobic on tropical monocotyledons. Sexual morph: Stromata solitary or irregularly
scattered, conspicuous, multi-loculate, superficial, with base slightly penetrating the epidermis at
regular intervals, discoid to pulvinate, reddish brown to black, surface scurfy, flat or slightly
convex. Ascomata deeply imbedded in stromatic columns, globose, with a long, periphysate,
ostiolar neck. Peridium thick, composed of several reddish brown outer layer cells of textura
globosa and textura epidermoidea, with pale brown to hyaline internal cells of textura angularis.
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Paraphyses hypha-like, numerous, tapering towards the apex, not embedded in a gelatinous matrix.
Asci 8-spored, unitunicate, broad cylindrical, short pedicellate, apically rounded or truncate, with a
J-, discoid, refractive, apical ring. Ascospores bi-seriate, hyaline, 1-celled, ellipsoidal to cylindrical,
smooth-walled. Asexual morph: Undetermined.
Type genus – Catabotrys Theiss. & Syd.
Notes – Catabotryaceae was validated by Barr (1990b) following Petrak’s (1954) invalid
publication, which is monotypic and accommodates a single species Catabotrys deciduum. Barr
(1990b) considered the centrum of Catabotrys as similar to that of taxa in Sordariales, and therefore
proposed a monotypic family Catabotryaceae to accommodate and further placed it in Sordariales.
Hyde & Cannon (1999) suggested Catabotryaceae could be included in Xylariales, as the centrum
and stromatic features are more similar to Xylariaceae and Diatrypaceae. With phylogenetic
analysis of the taxa in Sordariales, Catabotryaceae has been found transitory and placed in
Sordariomycetidae family incertae sedis (Huhndorf et al. 2004b, Miller & Huhndorf 2005) and
assigned in Amplistromatales (Maharachchikumbura et al. 2015). In a study of ranking fungi by
using divergence estimates (Hyde et al. 2017a), Catabotryaceae has been shown to have a stem age
of 165 MYA and therefore can be considered as an order. In this study the stem age was 171.92
MYA and therefore we place Catabotryaceae in a new order Catabotryales (Fig. 2).
Ecological and economic significance of Catabotryaceae
Catabotrys species have a pantropical distribution, and mainly appear on dead plant material
with striking superficial stromata. Wu et al. (2014b) has reported endophytic isolates of Catabotrys
from Fortunearia sinensis authorities in China. However, the frequent appearance on dead plant
tissue indicates the saprobic nutrition may dominate and occupy a large part of its life cycle, while
it may also at first be an endophyte within living plant tissue.
Genus included in Catabotryaceae
Catabotrys Theiss. & Syd., Annls mycol. 13(3/4): 297 (1915)
Index Fungorum number: IF834; 2 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Catabotrys palmarum (Pat.) Theiss. & Syd.
Current name – Catabotrys deciduum (Berk. & Broome) Seaver & Waterston
Notes – Catabotrys was erected by Theissen & Sydow (1915) based on Bagnisiella
palmarum Pat. and has been placed in Dothideaceae (Dothideae). Petrak (1934) transferred
Catabotrys to Hypocreales because its thin-walled asci and bright coloured stroma are similar to the
latter. Seaver & Waterston (1946) synonymized an earlier name Hypoxylon deciduum as
Catabotrys deciduum. Morphologically, Catabotrys is characterized by superficial, erumpent,
multi-loculate stromata, and simple hyaline, 1-celled spores. Catabotrys deciduum is illustrated in
this entry (Fig. 56).
Cephalothecaceae Höhn., Annls mycol. 15(5): 362 (1917)
Index Fungorum number: IF80571; Facesoffungi number: FoF01330; 27 species.
Saprobic on decaying wood or bark and soil or hypersaprobic on other fungi, particularly
polypores, some species are opportunistic pathogens causing systemic mycotic infection in animals
and humans. Sexual morph: Ascomata cleistothecial, solitary to gregarious, superficial, subglobose
to globose, dark brown to black, glabrous, sometimes covered or surrounded at base by subiculum
consisting of sulphureous hyphae, lacking ostioles. Peridium cephalothecoid, comprising 2–3
layers, outer layer composed of dark brown cells of textura angularis to prismatica, inner layer
composed of elongated, hyaline cells of textura prismatica. Paraphyses comprising branched, and
septate ascogenous hyphae. Asci forming upon the ascogenous hyphae, 8-spored, unitunicate,
pyriform to subglobose to globose, evanescent, apedicellate, without an apical ring. Ascospores
irregularly arranged, brown, variously-shaped, unicellular, without germ pores, smooth-walled.
Asexual morph: Hyphomycetous. Mycelium pale-yellowish to brownish-yellow, branched septate.
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Conidiophores macronematous, semi-macronemous or micronematous, cylindrical, stiffly upright,
septate, unbranched or verticillate. Conidiogenous cells phialidic, cylindrical, strongly tapering and
thick-walled, hyaline, smooth-walled. Conidia in chains, hyaline to brown, cylindrical, ovate or
obovate, with or without an apiculate or truncate base, 1-celled, smooth-walled (adapted from
Maharachchikumbura et al. 2016b, Davolos et al. 2019).
Type genus – Cephalotheca Fuckel
Notes – Cephalothecaceae was previously accommodated in the Sordariales by Suh &
Blackwell (1999), but this study was based only on SSU, LSU sequence analysis. This has been
revised with time and based on its uncertain phylogenetic placement and different morphology,
Maharachchikumbura et al. (2015) tentatively referred the Cephalothecaceae to family incertae
sedis in Sordariomycetes. Similar results were obtained by Hongsanan et al. (2017) who reported
that the divergence of Cephalothecaceae was at 175 MYA. Five genera, Albertiniella,
Cephalotheca, Cryptendoxyla, Phialemonium and Victoriomyces are currently included in
Cephalothecaceae (Fuckel 1872, Kirschstein 1936, Malloch & Cain 1970, Gams & McGinnis 1983,
Maharachchikumbura et al. 2016b, Davolos et al. 2019). The genus is placed in Cephalothecales in
this study.
Ecological and economic significance of Cephalothecaceae
Cephalothecaceae species are emerging as fungal opportunistic pathogens of humans and
other animals and they can affect both immunocompromised and immunocompetent hosts (Gavin
et al. 2002, Proia et al. 2004, Suh et al. 2006, Perdomo et al. 2011, Řehulka et al. 2016, Sun et al.
2019). The infections caused by members of Cephalothecaceae were reported as peritonitis,
endocarditis, osteomyelitis, and cutaneous infections of wounds following burns. In addition, C.
sulfurea can produce Gibberellins, which are well known for plant growth promotion (Hamayun et
al. 2012).
Genera included in Cephalothecaceae
Albertiniella Kirschst., Annls mycol. 34(3): 183 (1936)
Index Fungorum number: IF109; 2 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Albertiniella reticulata Kirschst.
Notes – Albertiniella was introduced by Kirschstein (1936) to accommodate a fungus
developing ascomata on rotting Polyporus applanatus. It is characterized by globose, black,
glabrous, carbonaceous cleistothecium, subglobose to globose asci and ovate to globose, aseptate,
hyaline to yellowish ascospores.
Cephalotheca Fuckel, Jb. nassau. Ver. Naturk. 25-26: 297 (1871)
Index Fungorum number: IF877; 15 morphological species (Species Fungorum 2020), 3
species with sequence data.
Type species – Cephalotheca sulfurea Fuckel.
Notes – The asexual morph of Cephalotheca was reported as tritirachium-like and
phialemonium-like (Limber 1940, Suh et al. 2006). In this entry we illustrate Cephalotheca
sulfurea from the isotype (Figs. 57, 58).
Cryptendoxyla Malloch & Cain, Can. J. Bot. 48(10): 1816 (1970)
Index Fungorum number: IF1309; 2 species with sequence data.
Type species – Cryptendoxyla hypophloia Malloch & Cain
Notes – Cryptendoxyla was introduced by Malloch & Cain (1970) in family of
Pseudeurotiaceae, which was typified by Cryptendoxyla hypophloia. Based on the results of
phylogenetic investigations, Cryptendoxyla was placed in Cephalothecaceae (Wijayawardene et al.
2012, Maharachchikumbura et al. 2016b). It is characterized by ascospores which are cylindrical to
oblong and brown to dark brown with small guttules.
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Figure 56 – Catabotrys deciduum (Material examined – THAILAND, Chumphon, Amphoe Pathio,
on rachis of Salacca sp. (Arecaceae), 1 December, 2016, S.N. Zhang SNT33B (MFLU 18-1072);
THAILAND, Ranong, Amphoe Mueang Ranong, on decaying petiole of oil palm (Arecaceae), 29
August 2017, S.N. Zhang SNT214, MFLU 18-1073, HKAS 97484; living culture MFLUCC 180463). a, b Appearance of stromata on host surface. c, d Vertical section through the stromata with
ascomata. e Ostiole with periphyses. f Structure of peridium. g-k Asci. k Asci in Cotton blue
reagent. l Paraphyses. m Ascus apex in Melzer’s reagent, with J-, subapical ring. n-p Ascospores.
Scale bars: a = 1000 μm, b-d = 500 μm, e = 50 μm, f-l = 20 μm, m = 10 μm, n-p = 5 μm.
Phialemonium W. Gams & McGinnis, Mycologia 75(6): 978 (1983)
Index Fungorum number: IF11160; 7 species with sequence data.
Type species – Phialemonium obovatum W. Gams & McGinnis
Notes – Phialemonium was introduced by Gams & McGinnis (1983) to accommodate three
species. The genus is characterized by phialides that are often inflated at the base, with conspicuous
collarettes and conidia arranged in slimy heads or in long chains, and may have connectives at both
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ends. Phialemonium was revised by Perdomo et al. (2013b), its species are pathogens and can
infect both immunocompromised and immunocompetent hosts (Perdomo et al. 2011).
Figure 57 – Cephalotheca sulfurea (Material examined – GERMANY, on rotting planks of Oak,
lying on damp ammoniacal ground, S-F90775, isotype). a Herbarium material. b, c Ascomata on
the host. d Squash mount of ascoma. e Peridium. f Immature asci on the ascogenous hyphae. g
Ascogenous hyphae. h-j Asci. k-l Ascospores. Scale bars: b = 1000 µm, c = 100 µm, d = 20 µm, e,
g = 10 µm, f, h-l = 5 µm.
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Victoriomyces D Davolos, B Pietrangeli, AM Persiani & O Maggi, Int. J. Syst. Evol. Microbiol.
69(4):1099-1110 (2019)
Index Fungorum number: IF823713; 1 species with sequence data.
Type species – Victoriomyces antarcticus Maggi, Davolos & Persiani
Notes – The type species Victoriomyces antarcticus was isolated from soil and is
characterized by metarhizium-like morphology, dark red-coloured disk-like structures, immature
bodies and the production of an intense red pigment in the growth media. Its placement in
Cephalothecaceae was confirmed by phylogenetic analysis of a combined LSU, SSU, rpb2 dataset.
Figure 58 – Different asexual morphs of Cephalothecaceae, Cephalotheca purpurea, C. sulfurea
(redrawn from Chesters 1935). a-c Fertile hyphae and radulo-conidia. d Conidiophores with
conidiogenous cells and conidia. e Conidia. Scale bars: a, d = 10 µm, b, c, e = 5 µm.
Ceratocystidaceae Locq., Stud. Mycol. 68(1): 188 (2011)
Index Fungorum number: IF515438; Facesoffungi number: FoF01248; 161 species.
Saprobic or pathogenic on plant materials and plants, parasitic in beetles, flies, or mites, and
isolated from soil. Sexual morph: Ascomata perithecial, initially immersed to semi-immersed,
becoming superficial, solitary or clustered; ascomatal bases globose, subglobose to obpyriform or
ovoid, light brown or dark brown to black, clothed with spines or digitate or stellate appendages,
ornamented or unornamented or with undifferentiated ornamental hyphae. Necks usually filiform,
tapering and paler upwards. Ostiolar hyphae straight or divergent to convergent, aseptate, light
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brown to hyaline. Paraphyses lacking. Asci unitunicate, evanescent. Ascospores aseptate, hyaline,
hat-shaped or varied in shape, ellipsoidal or elongate to slightly curved, with rounded ends, or
oblong, cylindrical or narrowly fusiform to spindle-shaped, with eccentric wall thickening or
surrounded by a sheath, accumulating in masses at tips of ostiole. Asexual morph: Conidiophores
mononematous, single or aggregated in sporodochia or synnematous, septate, tapering towards
apex, hyaline to pale brown or dark brown, unbranched or branched; in some genera such as
Ceratocystis and Huntiella two types of conidiophores (primary and secondary) occur.
Conidiogenous cells phialidic, borne terminally or laterally on vegetative hyphae, lageniform,
tubular, rectangular, oblong cylindrical to flask-shaped, sometimes with a slightly flared collarette,
subhyaline or pale brown, tapering towards the apex. Conidia unicellular, varied in shape,
cylindrical to oblong, globose to subglobose, rectangular, single or formed in chains, with rounded
or truncate ends, hyaline to pale brown, or becoming grey at maturity; in some genera with two
types of conidia: (i) primary or bacilliform conidia hyaline, aseptate, cylindrical, and (ii) secondary
or barrel-shaped conidia cylindrical to oblong, hyaline or becoming grey, aseptate, mostly in
chains; aleurioconidia (some genera) globose to subglobose, ovoid to pyriform, single or in chains,
hyaline or pale brown to brown (adapted from Réblová et al. 2011, Maharachchikumbura et al.
2016b).
Type genus – Ceratocystis Ellis & Halst.
Notes – Ceratocystidaceae (as “Ceratocystaceae”) was introduced by Locquin (1972), but
was not validly published but validated by Réblová et al. (2011). It is currently placed in
Microascales in the subclass Hypocreomycetidae (Réblová et al. 2011, de Beer et al. 2013b,
Maharachchikumbura et al. 2016b). Within the order, it forms a monophyletic group, distinct from
Gondwanamycetaceae based on strong bootstrap support (Réblová et al. 2011). Historically,
Ceratocystidaceae and especially Ceratocystis included a highly heterogeneous group of species
based on similar morphology. With the aid of multigene phylogenetic analyses, morphological
characteristics and ecological preferences, several genera were delimited and placed in
Ceratocystidaceae, which include Ambrosiella, Berkeleyomyces, Bretziella, Ceratocystis,
Chalaropsis, Davidsoniella, Endoconidiophora, Huntiella, Meredithiella, Phialophoropsis and
Thielaviopsis (Peyronel 1916, de Beer et al. 2014, 2017, Mayers et al. 2015, Nel et al. 2018).
Some asexual genera defined by similar characters are not readily identifiable which in the
absence of molecular data, lead to difficulties in generic identification. DNA sequence data, the
60S ribosomal protein RPL10 (60S), nuclear ribosomal DNA large subunit (LSU) and minichromosome maintenance complex component 7 (MCM7) are suggested to delimit genera (de Beer
et al. 2014, Marin-Felix et al. 2017). In addition, the internal transcribed spacer (ITS) regions and
the 5.8S gene, partial β-tubulin (tub2), translation elongation factor 1α (tef1), second largest
subunits of RNA polymerase II (rpb2), and/or the guanine nucleotide-binding protein subunit betalike protein (MS204) gene regions are used to resolve taxa at the species level (de Beer et al. 2014,
Marin-Felix et al. 2017, Liu et al. 2018). Huntiella chinaeucensis is illustrated in this study.
Ecological and economic significance of Ceratocystidaceae
Members of Ceratocystidaceae, particularly species of Ceratocystis, are destructive plant
pathogens. They cause serious diseases in several economic crops, trees and ornamental plants such
as coffee, cacao, eucalyptus, giant taro, and occur worldwide (Roux et al. 2004, Van Wyk et al.
2010, Li et al. 2016, Mbenoun et al. 2016). For example, Ceratocystis fimbriata causes a serious
wilt and die-back disease on Eucalyptus clones in the Republic of Congo in Central Africa (Roux et
al. 2000). Ceratocystis fimbriata attacks a wide variety of plants; for example, it causes a severe
leaf blight disease on Alocasia macrorrhiza in Yunnan, China (Li et al. 2016). The disease
outbreaks result with extensive crop losses and severely affects the economy.
Genera included in Ceratocystidaceae
Ambrosiella Brader, Mycopath. Mycol. appl. 25: 314 (1965)
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Index Fungorum number: IF7119; 12 morphological species (Species Fungorum 2020), 10
species with sequence data.
Type species – Ambrosiella xylebori Brader.
Notes – Ambrosiella was invalidly described by Brader (1964) as A. xylebori from a gallery
of Xylosandrus compactus in Coffea canephora from the Ivory Coast; no type was designated. The
generic type was subsequently illustrated and designated based on Brader’s isolate (CBS 110.61)
by von Arx & Hennebert (1965). Ambrosiella was previously a polyphyletic group. However, the
genus was delimited based on DNA sequence data and new combinations and new species were
proposed (Six et al. 2009, Harrington et al. 2010, 2014, Mayers et al. 2015, 2017, 2019).
Ambrosiella species are obligate symbionts of the ambrosia beetle tribe Xyleborini (Mayers et al.
2015).
Berkeleyomyces W.J. Nel, Z.W. de Beer, T.A. Duong & M.J. Wingf., Pl. Path. 67(4): 876 (2018)
Index Fungorum number: IF822838; 2 species with sequence data.
Type species – Berkeleyomyces basicola (Berk. & Broome) W.J. Nel, Z.W. de Beer, T.A.
Duong & M.J. Wingf.
Notes – Berkeleyomyces was introduced with two species, B. basicola and B. rouxiae (Nel et
al. 2018). The type species which was previously identified as Thielaviopsis basicola has a chaotic
taxonomic history and was classified in several genera (Ferraris 1912, Carmichael et al. 1980,
Crane & Miller 2016, Nel et al. 2018). It was treated in Berkeleyomyces by Nel et al. (2018) based
on phylogenetic analyses and morphology. The fungus is an important plant pathogen which causes
serious black root rot of numerous plants such as tobacco and groundnuts (Geldenhuis et al. 2006,
Coumans et al. 2011).
Bretziella Z.W. de Beer, Marinc., T.A. Duong & M.J. Wingf., MycoKeys 27: 10 (2017)
Index Fungorum number: IF822520; 1 species with sequence data.
Type species – Bretziella fagacearum (Bretz) Z.W. de Beer, Marinc., T.A. Duong & M.J.
Wingf.
Notes – de Beer et al. (2017) reclassified the Ceratocystidaceae (Microascales) based on
multi-gene phylogenetic inference and found that the oak wilt fungus Ceratocystis fagacearum
(current name Bretziella fagacearum) formed a well-supported monophyletic clade distinct from all
other genera in Ceratocystidaceae. They also confirmed that Chalara quercina (the first name
applied to the fungus) and Endoconidiophora fagacearum (the name applied when the sexual
morph was discovered) are conspecific based on their axenic cultures from the same host trees and
geographical area. Thus, de Beer et al. (2017) introduced Bretziella to accommodate the oak wilt
fungus with the single species B. fagacearum found on Quercus sp.
Ceratocystis Ellis & Halst., New Jersey Agric. Coll. Exp. Sta. Bull. 76: 14 (1890)
Index Fungorum number: IF888; 101 morphological species (Species Fungorum 2020), 53
species with sequence data.
Type species – Ceratocystis fimbriata Ellis & Halst.
Notes – Ceratocystis is characterized by black, globose ascomatal bases with filiform,
elongated necks terminating in an ostiole and sticky, hat-shaped ascospores (Upadhyay 1981,
Seifert et al. 1993, de Beer et al. 2014). The asexual morph of most Ceratocystis species is chalaraor thielaviopsis-like and characterized by phialidic conidiogenous cells producing chains of
hyaline, single-celled, cylindrical conidia called endoconidia, and in some cases secondary dark,
barrel-shaped, thick-walled aleurioconidia which are commonly produced that facilitate survival in
wood or soil during dry seasons (Hedgcock 1906, Harrington 2013, de Beer et al. 2014,
Maharachchikumbura et al. 2016b).
The history of Ceratocystis was discussed in Réblová et al. (2011), de Beer et al. (2014),
Maharachchikumbura et al. (2016b) and Holland et al. (2019). Ceratocystis species are important
plant pathogens and act as causal agents of sap stain in timber and symbiotic associates of insects.
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For example, C. platani is an invasive alien pathogen of Platanus trees in Europe (Ocasio-Morales
et al. 2007), whereas C. albifundus is a virulent pathogen of Acacia mearnsii in Africa (Roux &
Wingfield 2013). The ITS, tub1, tef1, rpb2, ms204 gene regions are used to identify species
boundaries in Ceratocystis (Marin-Felix et al. 2017, Barnes et al. 2018). The loci rpb2 and ms204
provide stronger resolution among species than tef1 and tub1, but also need to be used in
combination with ITS (Fourie et al. 2015).
Chalaropsis Peyronel, Staz. Sper. Argar. Ital. 49: 595 (1916)
Index Fungorum number: IF7602; 3 species with sequence data.
Type species – Chalaropsis thielavioides Peyronel.
Notes – Chalaropsis has been reported as a hyphomycetous genus and the sexual morph is so
far undetermined. Although C. ovoidea, C. populi and C. thielavioides are confirmed species with
sequence data, none of the sequence data were derived from type specimens (Paulin-Mahady et al.
2002, Wingfield et al. 2013, de Beer et al. 2014).
Davidsoniella Z.W. de Beer, T.A. Duong & M.J. Wingf., Stud. Mycol. 79: 210 (2014)
Index Fungorum number: IF810235; 4 species with sequence data.
Type species – Davidsoniella virescens (R.W. Davidson) Z.W. de Beer, T.A. Duong & M.J.
Wingf.
Notes – de Beer et al. (2014) emended four species, viz. Davidsoniella australis, D. eucalypti,
D. neocaledoniae and D. virescens based on combined sequence data of 60S, LSU and MCM7, and
introduced a new holomorphic genus with D. virescens as the type species. Only Davidsoniella
virescens and D. eucalypti have known sexual morphs, while D. australis and D. neocaledoniae are
known only as asexually fungi (Kiffer & Delon 1983, Samuels 1993, de Beer et al. 2014).
Endoconidiophora Münch, Naturwiss. Z. Forst-Landw. 5: 564 (1907)
Index Fungorum number: IF1794; 9 species with sequence data.
Type species – Endoconidiophora coerulescens Münch
Notes – The generic description of Endoconidiophora was emended and described from both
sexual and asexual morphs lacking aleurioconidia (de Beer et al. 2014). Eight species were
confirmed with molecular data mostly from ex-types, and E. coerulescens was confirmed from a
new collection (de Beer et al. 2014). Epitypification of E. coerulescens which is the type species of
Endoconidiophora, is required.
Huntiella Z.W. de Beer et al. Stud. Mycol. 79: 211 (2014)
Index Fungorum number: IF810236; 18 morphological species (Species Fungorum 2020), 17
species with sequence data.
Type species – Huntiella moniliformis (Hedgc.) Z.W. de Beer, T.A. Duong & M.J. Wingf.
Notes – Huntiella species occur on a wide range of hosts and are distributed worldwide on
Acacia mangium in Indonesia, Eucalyptus obliqua in Australia, E. saligna in South Africa,
Mangifera indica in Oman, Picea spinulosa in Bhutan (de Beer et al. 2014), Eucalyptus exserta,
Acacia confusa in China (Liu et al. 2018), and Tectona grandis in Thailand (this study). The
species are saprobes, weak pathogens and are also commonly found on tree wounds (Mbenoun et
al. 2016, Liu et al. 2018). Huntiella chinaeucensis, is illustrated in this entry (Fig. 59).
Meredithiella McNew, C. Mayers & T.C. Harr., Fungal Biology 119(11): 1086 (2015)
Index Fungorum number: IF812574; 3 species with sequence data.
Type species – Meredithiella norrisii McNew, C. Mayers & T.C. Harr.
Notes – Meredithiella norrisii was collected from insect galleries and from male Corthylini
punctatissimus beetles from Michigan and Iowa (Mayers et al. 2015). Meredithiella fracta and M.
guianensis were subsequently introduced and their type cultures isolated from mycangium of
Corthylus papulans and gallery of Corthylus crassus, respectively (Mayers et al. 2018). The genus
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is a symbiont with ambrosia beetle of the tribe Corthylini (Mayers et al. 2018). So far, members of
the genus are only asexual morphs.
Figure 59 – Huntiella chinaeucensis (Material examined – THAILAND, Chiang Rai Province,
Mae Suai District, Mae Lao garden, on stumps of Tectona grandis (Lamiaceae), 24 December
2012, M. Doilom, MFLU 15-3204). a, b Globose to subglobose ascoma bases with elongated
ostiolar necks on Tectona grandis (teak) wood. c Cream-coloured ascospore mass at the tip of the
ascoma neck on teak wood. d Immature globose ascoma. e Close up of ascoma base with conical
spines. f Ascoma neck with longitudinal striations. g, h Mature ascoma with conical spines on the
surface of bases. i Hat-shaped ascospores released through ostiolar hyphae. j, k Hat-shaped
ascospores. l Flask-shaped conidiogenous cells. m Bacilliform conidia. n Barrel-shaped conidia in
chains. o Colony on PDA. Scale bars: a = 200 µm, b = 300 µm, c = 500 µm, d, g = 50 µm, e, f, l =
20 µm, h, i = 100 µm, j, k, m, n = 5 µm.
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Phialophoropsis L.R. Batra, Mycologia 59(6): 1008 (1968)
Index Fungorum number: IF9344; 1 species with sequence data.
Type species – Phialophoropsis trypodendri L.R. Batra
Notes – Phialophoropsis was resurrected by Mayers et al. (2015) to accommodate P.
trypodendri (type species) from Trypodendron scabricollis and P. ferruginea from Trypodendron
lineatum. Both species are associated with ambrosia beetles of the tribe Xyloterini (Mayers et al.
2015). Other two species, P. cambrensis and P. nipponica were treated as uncertain or excluded
species which would not be consistent with the current concept of Phialophoropsis (Mayers et al.
2015). The general concept of Phialophoropsis is accommodated ambrosia beetle symbionts with
deep-seated phialides and the absence of aleurioconidia (Mayers et al. 2015).
Thielaviopsis Went, Meded. Proefstn Suik Riet W. Java 5: 4 (1893)
Index Fungorum number: IF10210; 7 species with sequence data.
Type species – Thielaviopsis ethacetica Went
Notes – Went (1893) introduced Thielaviopsis and the generic description was emended to
include both sexual and asexual morphs with aleurioconidia (de Beer et al. 2014). The species are
plant pathogens. For example, T. paradoxa has been reported to cause trunk rot and black scorch
diseases of palms (Elliott 2006, Al-Naemi et al. 2014).
Ceratosphaeriaceae Z.L. Luo, H.Y. Su & K.D. Hyde, Fungal Divers 99: 490 (2019)
Index Fungorum number: IF555643; Facesoffungi number: FoF05415; 26 species.
Saprobic on wood. Sexual morph: Stromata absent. Ascomata globose to pyriform, deeply
immersed to almost superficial, dark brown to black, carbonaceous, with a long cylindrical, black
or yellow crystals neck. Periphyses well-developed. Peridium composed of a large number of
layers of very thick-walled rather small cells in the neck region. Interascal tissue of paraphyses
thin-walled, probably evanescent at maturity. Asci 8-spored, unitunicate, cylindrical. Ascospores
biseriate, narrowly cylindric-fusiform, or filiform, ends acute, septate, smooth-walled. Asexual
morph: Hyphomycetous. harpophora-like. Phialides or short conidiophores arising on aerial
hyphae, with conidial heads slimy, inconspicuous, and transparent. Conidiogenous cells phialidic,
ampulliform to lageniform, terminal or intercalary, cylindrical. Conidia cylindrical, hyaline,
aseptate, smooth (adapted from Luo et al. 2019).
Type genus – Ceratosphaeria Niessl
Notes – Réblová (2006) accommodated Ceratosphaeria in Magnaporthaceae based on
phylogenetic analyses of combined LSU and SSU sequence data. In the phylogenetic study of Luo
et al. (2019), Ceratosphaeria species formed a distinct clade in Magnaporthales and they
introduced Ceratosphaeriaceae to accommodate Ceratosphaeria. Presently, there is only one genus
(Ceratosphaeria) accepted in this family.
Ecological and economic significance of Ceratosphaeriaceae
Ceratosphaeriaceae are saprobes which have the ability to decompose lignocellulose matter in
woody litter, resulting in softening of the wood and releasing nutrients in the form of simple
molecules that go back into the soil and can be reused by plants and all other organisms (Yuen et al.
1998, Bucher et al. 2004). Thus, they play an important role in nutrient and carbon cycling,
biological diversity and ecosystem functioning (Palmer et al. 1997, Wong et al. 1998a).
Genus included in Ceratosphaeriaceae
Ceratosphaeria Niessl, Verh. nat. Ver. Brünn 14: 203 (1876)
Index Fungorum number: IF897; 26 morphological species (Species Fungorum 2020), 3
species with sequence data.
Type species – Ceratosphaeria lampadophora (Berk. & Broome) Niessl
Notes – The genera was established to accommodate recombined species, Ceratosphaeria
lampadophora (= Sphaeria lampadophora), species found from wood in Great Britain (Niessl
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1876). Ceratosphaeria is characterized by globose to pyriform, immersed to almost superficial,
dark coloured stromatic ascomata, leathery to fragile perithecial walls, cylindric-clavate, shortstipitate asci, truncate to broadly rounded at the apex, with an apical ring and allantoid to
suballantoid, pale brown, aseptate ascospores and a harpophora-like asexual morph with phialidic
conidiogenesis (Hyde et al. 1997b, Niessl 1876, Réblová 2006). Five species in this genus have
been recorded from freshwater habitats (Luo et al. 2019).
Ceratostomataceae G. Winter, Rabenh. Krypt.-Fl., Edn 2 (Leipzig) 1.2: 247 (1885)
Index Fungorum number: IF80575; Facesoffungi number: FoF01803; 134 species.
Saprobic or weakly parasitic, often growing on other fungi and commonly isolated from soil.
Sexual morph: Ascomata perithecial or cleistothecial, yellow to pale brown, transparent, with or
without ostiole: often with long-necks, smooth ostiolar setae. Interascal tissue absent. Peridium
membranaceous, pale yellow to pale yellowish-brown, cells of a textura angularis or textura
globulosa. Asci 8-spored, unitunicate, thin-walled, clavate, without an apical ring, deliquescing.
Ascospores biseriate, brown to dark brown, ellipsoidal to citriform, occasionally discoid or
fusiform, 1-celled, usually with 2 germ pores at each end, smooth to strongly ornamented, without
sheath. Asexual morph: Hyphomycetous. Conidiophores simple, semi-macronematous,
mononematous, some genera branched towards the apex, in some genera conidiophore with several
roughened swellings along the entire length, reddish-brown to dark. Conidiogenous cells integrated,
terminal, monoblastic, some genera with phialides, singly on aerial hyphae or rarely on
conidiophores, lageniform, hyaline. Conidia globose, spindle-shaped, ovate to pyriform, some
genera aggregated in small globose heads at the apices of phialides, 1-celled, some genera with 2–
3-transverse septa, hyaline, brown and verrucose (adapted from Maharachchimbura et al. 2016b).
Type genus – Melanospora Corda
Notes – Ceratostomataceae (= Melanosporaceae) was introduced by Winter (1885b).
Hawksworth et al. (1995) placed Ceratostomataceae in Sordariales based on characters. Jones &
Blackwell (1998) placed Ceratostomataceae in Hypocreales based on molecular phylogenetic
studies. Zhang & Blackwell (2002) considered Ceratostomataceae was derived from the
Hypocreales clade and they included it in Hypocreomycetidae. Hibbett et al. (2007) introduced
Melanosporales to accommodate Ceratostomataceae which clustered with Coronophorales.
Hongsanan et al. (2017) treated Melanosporales as a synonym of Coronophorales based on
phylogenetic analysis, which maintained monophyly with Melanosporales. Wijayawardene et al.
(2018a) accepted nine genera in Ceratostomataceae (Arxiomyces, Erythrocarpon, Gonatobotrys,
Melanospora, Pteridiosperma, Pustulipora, Rhytidospora, Setiferotheca, Vittatispora), when
merging both asexual and sexual genera into one outline. Marin-Felix et al. (2018) re-examined the
most relevant genera of Ceratostomataceae based on phylogenetic relationships, when redefining
Melanospora, and re-established Microthecium and introduced three new genera to
Ceratostomataceae and accepted 12 genera in this family. So far, several species and genera in
Ceratostomataceae were emended and introduced, and some genera were recombined (Réblová et
al. 2016a, Marin-Felix et al. 2018). Consequently, 14 genera are accepted in Ceratostomataceae
based on morphological characteristic and DNA-sequence data, these include Arxiomyces,
Dactylidispora, Echinusitheca, Erythrocarpon, Harzia, Melanospora (= Gonatobotrys),
Microthecium (= Pteridiosperma), Pseudomicrothecium, Pustulipora, Rhytidospora, Scopinella,
Setiferotheca, Syspastospora and Vittatispora (Réblová et al. 2016a, Marin-Felix et al. 2018,
Wijayawardene et al. 2018a). Many of the species in this family were isolated from dung and five
genera lack sequence data. Fresh collections with sequence data are therefore needed.
Ecological and economic significance of Ceratostomataceae
Most of the species in Ceratostomataceae are saprobic on plant material. Certain species have
a widespread host range, and are closely associated with other fungi, on soil, rotting vegetation and
some species are considered as potential biocontrol agents as they can grow on other plant
pathogens (Cannon & Kirk 2007).
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Figure 60 – Ceratosphaeria aquatica (Material examined – CHINA, Yunnan Province, saprobic on
decaying wood submerged in a freshwater river, April 2015, Z.L. Luo, S-639, MFLU 18-2323). a
Appearance of necks on substrate. b, c Section through ascomata. d Paraphyses. e Structure of
peridium. f, g Asci. h-l Ascospores. Scale bars: b, c = 200 μm, d, e, h-l = 20 μm, f, g = 50 μm.
480
Genera included in Ceratostomataceae
Arxiomyces P.F. Cannon & D. Hawksw., Trans. Br. mycol. Soc. 81(3): 644 (1983)
Index Fungorum number: IF25489; 3 morphological species (Species Fungorum 2020).
Type species – Arxiomyces vitis (Fuckel) P.F. Cannon & D. Hawksw.
Notes – Arxiomyces was introduced by Cannon & Hawksworth (1982) to replace Phaeostoma
with the type species Arxiomyces vitis, on bark in Europe (Lumbsch & Huhndorf 2010).
Arxiomyces campanulatus (on Stachybotrys chartarum, fungi from cultivated soil from Honshu and
A. zuberiensis, fungi on dead stems of Phragmites australis in Iraq were also introduced (Index
Fungorum 2020). The genus is characterised by ovoid to ellipsoidal ascospores rounded at the apex
and with a truncate base with a broad germ pore that bears a mucilaginous and collapsing
appendage (Cannon & Hawksworth 1982, 1983). The asexual morph of the genus is undetermined.
Dactylidispora Y. Marín, Stchigel, Guarro & Cano, MycoKeys 44: 89 (2018)
Index Fungorum number: IF812079; 3 morphological species (Species Fungorum 2020), 2
species with sequence data.
Type species – Dactylidispora ellipsospora (Takada) Y. Marín, Stchigel, Guarro & Cano
Notes – The genus was introduced by Marin-Felix et al. (2018) to establish three new species
combinations from Microthecium. They are Dactylidispora ellipsospora (= Microthecium
ellipsosporum), a species isolated from forest soil in North Solomons, Dactylidispora collipora (=
Microthecium collipora), a species isolated from soil in Rajasthan, and Dactylidispora
singaporensis (= Microthecium singaporensis), a species isolated from soil in Singapore (MarinFelix et al. 2018). The most distinctive characteristic of the genus is short, conical ascomata, with a
crown of setae surrounding the ostiole, smooth-walled and fusiform ascospores with a germ pore at
each end surrounded by a raised rim, and the asexual morph with phialidic, flask-shaped
conidiogenous cells and subglobose to ovoid conidia (Marin-Felix et al. 2018).
Echinusitheca Y. Marín, Stchigel, Dania García, Guarro, A.N. Mill. & Cano, MycoKeys 44: 91
(2018)
Index Fungorum number: IF812084; 1 species with sequence data.
Type species – Echinusitheca citrispora Y. Marín, Stchigel, Dania García, Guarro, A.N. Mill.
& Cano
Notes – Echinusitheca was introduced to accommodate a single species E. citrispora, isolated
from forest soil in North Carolina, USA. The genus is characterised by strongly setose, dark
ascomata lacking ostioles with and unicellular, ellipsoidal ascospores with a depressed germ pore at
each end. The asexual morph of the genus is undetermined (Marin-Felix et al. 2018).
Erythrocarpon Zukal, Verh. zool.-bot. Ges. Wien 35: 337 (1886)
Index Fungorum number: IF1901; 1 morphological species.
Type species – Erythrocarpon microstomum Zukal
Notes – Erythrocarpon was introduced by Zukal (1886) as monotypic genus to accommodate
a species, Erythrocarpon microstomum, found on rotten branches in Hütteldorf, Austria. The genus
is characterised by cylindrical, 8-spored asci and the asexual morph is characterised by spindleshaped conidia, with 3 transverse septa.
Harzia Costantin, Mucéd. Simpl. (Paris): 42 (1888)
Index Fungorum number: IF8458; 12 morphological species (Species Fungorum 2020), 10
species with sequence data.
Type species – Harzia acremonioides (Harz) Costantin
Notes – Harzia was introduced by Costantin et al. (1888) with the type species H.
acremonioides (≡ Monosporium acremonioides), a seed-borne species from soil. The genus is
characterised by hyaline mycelium and conidiophores and 1-celled, brown to golden brown, ovoid
to subglobose, usually smooth-walled conidia, but sometimes with a slight wrinkling/exposure, and
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each species tending to vary in conidia size (Horne & Williamson 1923, Domsch et al. 1980). In the
phylogenetic analysis, the genus formed a monophyletic clade in the family phylogenetic tree, and
is sister to genus Melanospora with high bootstrap support (99%ML; Fig. 11)
Melanospora Corda, Icon. fung. (Prague) 1: 24 (1837)
Index Fungorum number: IF3085; 69 morphological species (Species Fungorum 2020), 9
species with sequence data.
Type species – Melanospora zamiae Corda
Notes – Melanospora was established by Corda (1837) to accommodate Ceratostoma
chioneum (= Melanospora chionea) and two new species, Melanospora zamiae and M. leucotricha,
with the former chosen as the type species (Kowalski 1965). Melanospora is the largest genus of
this family. Most Melanospora species are parasitic and associated with a wide host range, such as
basidiomycetes, sexual and asexual ascomycetes, as well as with other fungi and some species
obtain nutrients by fusing with the host protoplasts, an interaction called fusion biotrophism
(Jeffries & Young 1994, Sun et al. 2019). Harveson (1999) reported Melanospora species as
potential biocontrol agents, which can grow on pathogenic fungi. This genus is distinguished by
translucent ascomata with a neck composed of intermixed hyphae and with an apical crown of
setae, and smooth or ornamented ascospores with an apiculate germ pore at each end, and a
phialidic asexual morph (Marin-Felix et al. 2018). The morphologically similar genera to
Melanospora and other genera in Ceratostomataceae was re-examined and discussed in MarinFelix et al. (2018).
Microthecium Corda, Icon. fung. (Prague) 5: 30, 74 (1842)
Index Fungorum number: IF3196; 23 morphological species (Species Fungorum 2020), 13
species with sequence data.
Type species – Microthecium zobelii Corda
Notes – Microthecium was introduced by Corda (1842) with M. zobelii as the type species,
found on Rhizopogon albus in Germany. The genus is characterised by yellowish to reddish,
globose ascomata and short necks or necks lacking, with setae around the ostiole, clavate and
evanescent asci, fusiform, navicular, citriform, plataniform or spindle-shaped ascospores, which are
hyaline and become brown to dark brown when mature, with a terminal apiculate or depressed
germ pore at each end. The asexual morph comprises hyaline phialides with pale orange to reddishorange conidia.
Pseudomicrothecium Marin-Felix, Stchigel, Guarro & Cano, MycoKeys 44: 114 (2018)
Index Fungorum number: IF812108; 1 species with sequence data.
Type species – Pseudomicrothecium subterraneum (L. Fan, C.L. Hou, P.F. Cannon & Y. Li)
Y. Marín, Stchigel, Guarro & Cano
Notes – Pseudomicrothecium was introduced by Marín-Felix et al. (2018) as a monotypic
genus to accommodate a single species, P. subterranean (= Melanospora subterranean) found on
litter in Japan. The genus is characterised by globose, pale brown to dark brown ascomata, lacking
ostioles when mature and shrouded with setae, 2-spored, clavate, evanescent asci with a short
pedicel and lacking apical structures and ellipsoidal to citriform, 1-celled, dark brown to blackish
ascospores, umbonate and truncate at both ends, with a terminal indistinct germ pore at each end
(Marín-Felix et al. 2018). The asexual morph of the genus is undetermined.
Pustulipora P.F. Cannon, Mycotaxon 15: 526 (1982)
Index Fungorum number: IF4561; 1 morphological species.
Type species – Pustulipora corticola P.F. Cannon
Notes – The monotypic genus Pustulipora was introduced by Cannon (1982) with P.
corticola occurring on bark. Pustulipora can be distinguished from Dactylidispora by its blistered,
482
rarely cushion-like structures surrounding the germ pore (Cannon 1982). The asexual morph of the
genus is undetermined.
Figure 61 – Pustulipora corticola (Material examined – UK, Nottinghamshire, Clumber Park, from
unidentified bark, 7 September 1956, W.D. Graddon IMI 284968, isotype). a, b Herbarium packet.
c-g Ascospores. h Ascoma. i Ascus. j Ascospores (h-j: redraw from Cannon 1982). Scale bars: c =
100 µm, d = 50 µm, e-g = 20 µm.
Rhytidospora Jeng & Cain, Mycotaxon 5(1): 278 (1977)
Index Fungorum number: IF4744; 5 morphological species (Species Fungorum 2020).
Type species – Rhytidospora tetraspora Jeng & Cain
Notes – Rhytidospora was introduced by Jeng & Cain (1977) based on a single species R.
tetraspora, isolated from burro dung in Venezuela. Rhytidospora bispora, R. cainii, R. citriformis,
and R. inordinata, were added to this genus by (Doveri 2014). The genus is characterized by
cleistothecial ascomata with a cephalothecoid peridium, globose to subglobose, irregularlyevanescent asci, and thick-walled, ellipsoidal ascospores with a wrinkled a germ pore (Krug & Jeng
1979). The asexual morph of the genus is undetermined.
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Figure 62 – Rhytidospora tetraspora (Material examined – VENEZUELA, Edo. Sucre, trail
between Manacal and Los Pocitos, NW of Irapa, on burro dung, 13 July 1972, K.P. Dumont, R.F.
Cain, G.J. Samuels, G. Morillo, J. Farfan, TRTC052086, holotype). a-e Herbarium package details.
f-i Ascomata. j, k Ascospores (f-k, observed from slides of herbarium package). Scale bars: d =
2000 µm, e = 1000 µm, f-i = 100 µm, j, k = 20 µm.
Scopinella Lév., Dict. Univ. Hist. Nat. 8: 493 (1846)
Index Fungorum number: IF4962; 9 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Scopinella pleiospora (J. Schröt.) Sacc.
Notes – The type species was described growing on rabbit dung in Norway. The genus is
characterised by long necked ascomata, evanescent asci and cuboid-ellipsoidal ascospores with two
prominent germ slits. Thee asexual morph of the genus is undetermined.
Setiferotheca Matsush., Matsush. Mycol. Mem. 8: 34 (1995)
Index Fungorum number: IF27340; 1 morphological species.
Type species – Setiferotheca nipponica Matsush.
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Notes – Setiferotheca nipponica is characterised by ovoid to ellipsoidal ascospores rounded at
the apex and a truncate base with a broad germ pore that bears a small basal appendage and
ascomata with a crown of dark brown setae surrounding the ostiole (Matsushima 1995).
Syspastospora P.F. Cannon & D. Hawksw., J. Linn. Soc., Bot. 84(2): 152 (1982)
Index Fungorum number: IF5345; 4 morphological species (Species Fungorum 2020), 2
species with sequence data.
Type species – Syspastospora parasitica (Tul.) P.F. Cannon & D. Hawksw.
Notes – Syspastospora was introduced by Cannon & Hawksworth (1982) to accommodate
Melanospora parasitica, with three additional species, S. boninensis, S. cladoniae, and S.
tropicalis. The genus has ascomata with long necks composed of parallel arranged hyphae and
cylindrical ascospores with a large terminal, slightly sunken germ pore at each end (Cannon &
Hawksworth 1982). The asexual morph of the genus is undetermined.
Vittatispora P. Chaudhary, J. Campb., D. Hawksw. & K.N. Sastry, Mycologia 98(3): 461 (2006)
Index Fungorum number: IF500705; 1 species with sequence data.
Type species – Vittatispora coorgii P. Chaudhary, J. Campb., D. Hawksw. & K.N. Sastry
Notes – Vittatispora was introduced to accommodate a single species, V. coorgii, isolated
from soil in India. The taxon was observed from culture and has light brow, perithecia, with a long
cylindrical neck, 8-spored, clavate asci and ellipsoidal to citriform, brown to dark brown ascospores
with two apical germ pores; germ pores surrounded by a raised rim; with a thick hyaline ridge
running vertical length of the ascospore between the germ pores (Chaudhary et al. 2006). The
asexual morph of the genus is undetermined.
Chadefaudiellaceae Faurel & Schotter, Mycotaxon 12(1): 46 (1980)
Index Fungorum number: IF80580; Facesoffungi number: FoF01666; 6 species.
Saprobic on mammal dung. Sexual morph: Stromata absent. Ascomata elongate or
perithecial, hemisphaerical, embedded. Interascal tissue of undifferentiated hyphae, basal
perithecial envelope black, globular, carbonaceous, foot cylindrical, light yellow to brown,
translucent, striated, brown apex, the upper part formed from anastomosing setae, lacking ostioles.
Peridium pseudoparenchymatous, composed of textura angularis embedded in the substrate and
with an aerial “capillitium”. Asci 8-spored, catenulate, globose or clavate, evanescent. Ascospores
overlapping, pale yellow to brown, aseptate, hyaline to pale brown, ellipsoidal or fusiform, lacking
germ pores, smooth-walled or with striations, dextrinoid or not dextrinoid, turning to reddish brown
in Melzer’s reagent (adapted from Benny & Kimbrough 1980, Maharachchikumbura et al. 2016).
Asexual morph: Hyphomycetous, forming arthrospores (Wijayawardene et al. 2017b).
Type genus – Chadefaudiella Faurel & Schotter
Notes– Chadefaudiellaceae was validated by Benny & Kimbrough (1980) and typified by
Chadefaudiella. The family consists of species with immersed ascomata lacking ostioles, with a
pseudoparenchymatous peridium and aerial thick-walled ‘capillitium’, enclosing catenulate,
evanescent asci with non-dextrinoid ascospores devoid of germ pores (Cannon & Kirk 2007). The
species was originally found on gazelle dung at Koudou, Chad (Africa) (Benny & Kimbrough
1980). The classification of Faurelina has been problematic and despite the similarities with
Chadefaudiella noted by Locquin-Linard (1975), it has undergone many changes in terms of its
higher order placement. The placement of Faurelina was debatable and has been speculated to have
affinities with many different groups such as Sordariaceae, Testudinaceae, Pithoascaceae and even
in Dothideomycetes (Locquin-Linard 1975, Parguey-Leduc & Locquin-Linard 1976, von Arx
1978). Finally, the genus was placed in Chadefaudiellaceae along with Chadefaudiella (Cannon &
Kirk 2007, Tang et al. 2007, Maharachchikumbura et al. 2015).
Réblová et al. (2011) excluded Faurelina from Chadefaudiellaceae. However, they suggested
that further molecular analysis was required and revision of the family. Wijayawardene et al.
(2017a), in their checklist included an arthrographis-like asexual morph for Faurelina (Locquin485
Linard 1975) and confirmed the placement of Faurelina in Chadefaudiellaceae. In Chadefaudiella
ascomata have apical anastomosing setae and ascospores are not dextrinoid.
Ecological and economic significance of Chadefaudiellaceae
Ecologically the species of Chadefaudiellaceae are coprophilous, in desert environments,
where they degrade dung.
Genera included in Chadefaudiellaceae
Chadefaudiella Faurel & Schotter, Revue Mycol., Paris 30: 339 (1959)
Index Fungorum number: IF928; 2 morphological species (Species Fungorum 2020).
Type species – Chadefaudiella quezelii Faurel & Schotter
Notes – Chadefaudiella is placed in the Chadefaudiellaceae, Microascales because it
produces asci in chains and has other characteristics of Microascaceae taxa. The ascospore wall of
C. quezelii is striate and non-dextrinoid (Locquin-Linard 1973, 1975). Usually Chadefaudiella
species are saprobic on mammalian dung and was found on Gazelle dung from Koudou, Chad
(Africa) (Benny & Kimbrough 1980). The genus is characterized by large elongate or
hemisphaerical, pale yellow to brown, carbonaceous ascomata, which are immersed at the base, and
lack ostioles. The peridium is composed of cells of textura angularis embedded in the substrate and
an aerial “capillitium”. Asci are 8-spored, globose and evanescent. Ascospores are aseptate,
fusiform, thick, striate and do not turn reddish-brown in Melzer’s reagent (non dextrinoid). The
asexual morph is undetermined (Benny & Kimbrough 1980, Maharachchikumbura et al. 2016b).
Faurelina Locq.-Lin., Revue Mycologique, Paris 39(2): 127 (1975)
Index Fungorum number: IF1981; 4 morphological species (Species Fungorum 2020), 3
species with sequence data.
Type species – Faurelina fimigena Locq.-Lin.
Notes – The asexual morph is arthrographis-like, and coprophilous. The sexual morph is
undetermined. Taxonomic revisions were undertaken by Kirk et al. (2013). The genus is accepted
in Chadefaudiellaceae by Cannon & Kirk (2007), Tang et al. (2007), Rossman et al. (2012), Kirk et
al. (2013), Maharachchikumbura et al. (2015, 2016b), and Wijayawardene et al. (2017a, 2018a).
Faurelina differs as ascomata lack apical anastomosing setae and ascospores are dextrinoid.
Chaetomiaceae G. Winter, Rabenh. Krypt.-Fl., Edn 2 (Leipzig) 1.2: 153 (1885)
Index Fungorum number: IF80582; Facesoffungi number: FoF01842; 506 species.
Saprobic or parasitic on plant debris, straw, seeds, dung, feathers of birds, soil, paper,
textiles, in air, on mushrooms, rabbits, and humans. Sexual morph: Ascomata perithecial or
cleistothecial, colourless to light brown or carbonaceous, egg-yellow, grey olivaceous to (greenish)
grey, golden brown, or brown to black, solitary to scattered to gregarious, superficial or immersed
to semi-immersed, subglobose to obpyriform, globose, ellipsoidal, ovoid to pyriform, cylindrical,
covered with hairs, setae or glabrous, hairs may be simple or branched and of one or two types,
ostiolate or lacking ostioles. Peridium with thick walls, comprising several layers of hyaline or
brown cells of textura epidermoidea to textura intricata. Paraphyses absent or greatly reduced, if
present septate, hyaline. Asci 4- or 8-spored, unitunicate, clavate, cylindrical, obovate or ellipsoidal,
pedicellate, without apical structures or with an indistinct thickened ring, evanescent, with wall
dissolving at maturity. Ascospores irregularly arranged, at first colourless and dextrinoid
(translucent, greenish, dark), brown to black, opaque when mature, ellipsoidal, globose,
subglobose, oval, fusiform or triangular, with single or sometimes two germ pores, aseptate, with
thick, smooth walls. Asexual morph: Hyphomycetous. Conidiophores absent or when present
simple, hyaline, branched, septate, smooth-walled or slightly rough. Conidiogenous cells phialidic
or blastic, verticillate or solitary, hyaline, consisting of a lageniform or ellipsoid swollen basal
portion, cylindrical. Conidia dimorphic, first type holoblastic, hyaline to brown, smooth, globose-
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Figure 63 – Faurelina fimigena (redrawn from Melo et al. 2016). a Fruiting bodies on habit. b
Closed cleistothecium. c External superficial layer of peridium. d Cells of the inner layer of the
peridium. e Mature ascospores. Scale bars: a = 500 µm, b = 30 µm, c = 10 µm, d = 5 µm, e = 2 µm.
-to obclavate, 1-celled, formed on hyphae or cylindrical conidiogenous cells, single or racemose,
clusters; second type produced from phialides, hyaline to brown, lutescent, or dark brown,
subglobose to globose or ellipsoidal, occasionally cylindrical, pyriform or fusiform, formed singly
or in chains, 1-celled, with thick, smooth wall (adapted from Maharachchimbura et al. 2016b).
Type genus – Chaetomium Kunze
Notes – Chaetomiaceae was introduced by Winter (1885a) as “Chaetomieae”. It belongs to
Sordariales (Hawksworth & Wells 1973, Mehrotra & Aneja 1990, Huhndorf et al. 2004b, Kirk et
al. 2008, Lumbsch & Huhndorf 2010, Maharachchikumbura et al. 2015, 2016b, Hongsanan et al.
2017). A key for 20 genera of Chaetomiaceae was provided by Maharachchikumbura et al.
(2016b). Based on the phylogenetic analyses of rpb2, tub2, ITS and LSU gene regions, Wang et al.
(2016a) introduced five new genera: Amesia, Arcopilus, Collariella, Dichotomopilus and
Ovatospora in Chaetomiaceae. In a recent paper, Wijayawardene et al. (2018a) listed 26 genera as
belonging to this family. A new genus, Arxotrichum, was introduced by Crous et al. (2018d) for a
strain of Chaetomium succineum and a new isolate of Arxotrichum yomingense. Based on the
results of rpb2 and a combined rpb2, tub2, ITS and LSU dataset, Mycothermus was validated as a
new genus and the taxonomic status of Remersonia was determined in Chaetomiaceae by Wang et
al. (2019b).
Ecological and economic significance of Chaetomiaceae
Species of Chaetomiaceae are important pathogens of humans and animals (Ahmed et al.
2016, Plumlee et al. 2017, Jeragh et al. 2018, Pote et al. 2018, Wang et al. 2019b). They cause
serious opportunistic infections in immunocompromised patients, and cause equine mycotic
encephalitis (Ahmed et al. 2016, Plumlee et al. 2017). Species of Chaetomiaceae have been shown
to have high potential for use in biological control and production of bioactive secondary
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metabolites (Soytong et al. 2001, Khumkomkhet et al. 2009, Sibounnavong et al. 2012, Wang et al.
2012, Li et al. 2014).
Genera included in Chaetomiaceae
Achaetomium J.N. Rai, J.P. Tewari & Mukerji, Can. J. Bot. 42(6): 693 (1964)
Index Fungorum number: IF38; 14 morphological species (Species Fungorum 2020), 5
species with sequence data.
Type species – Achaetomium globosum J.N. Rai & J.P. Tewari
Notes – Achaetomium was established by Rai et al. (1964) to accommodate A. globosum, A.
luteum and A. strumarium and placed in Achaetomiaceae (Achaetomiales), however, the family
was invalidly published. Achaetomiaceae was validated by Mukerji (1978). Cannon (1986)
accepted only A. globosum in Achaetomium, and suggested that Achaetomium might have been
derived from Sordariaceae. von Arx et al. (1988) disagreed with and accepted the species in
Achaetomiaceae, while Lee et al. (1999) recommended that Achaetomium should be placed in
Chaetomiaceae based on molecular data. Based on the analyses of the LSU gene, Rodríguez et al.
(2004) showed Achaetomium sharing a common ancestor with Chaetomium within Chaetomiaceae.
Later studies (Maharachchikumbura et al. 2015, 2016b, Wang et al. 2016a, 2019b, Wijayawardene
et al. 2018a) showed that Achaetomium is an independent genus in Chaetomiaceae. Achaetomium is
characterized by globose to pyriform, ostiolate ascomata covered with yellowish hypha-like hairs,
with a relatively thick peridium consisting of cells of textura intricata, cylindrical asci and dark
brown ascospores that are opaque with an apical germ pore and colonies usually showing a reddishbrown exudate (Rodriguez et al. 2004).
Acrophialophora Edward, Mycologia 51(6): 784 (1961)
Index Fungorum number: IF7037; 18 morphological species (Species Fungorum 2020), 7
species with sequence data.
Type species – Acrophialophora nainiana Edward
Notes – Acrophialophora was reintroduced by Samson & Mahmood (1970) as a
thermotolerant genus with A. fusispora, A. levis and A. nainiana. Based on combined analysis of
ITS, SSU and tub2 sequence data, Zhang et al. (2015a) showed that Acrophialophora is a
monophyletic genus in Chaetomiaceae and they synonymized Taifanglania under
Acrophialophora. The genus is characterised by conidiophores that are absent or unbranched or
sparingly branched conidiogenous cells that are monophialidic or polyphialidic with two necks,
flask-shaped or in whorls in verticillate species. Conidia are unicellular, subglobose, elliptical to
fusiform, echinulate, with spiral bands or smooth, in basipetal chains or slimy heads and with
schizolytic secession (Zhang et al. 2015a). The sexual morphs are undetermined.
Allobotryotrichum Raza M., Z.F. Zhang, K.D. Hyde, Y.Z. Diao & L. Cai, Fungal Divers. 99: 74
(2019)
Index Fungorum number: IF556672; 1 species with sequence data.
Type species – Allobotryotrichum blastospora M. Raza & L. Cai
Notes – Allobotryotrichum was introduced by Raza et al. (2019) based on phylogenetic
analysis of Botryotrichum and its sterile setae and conidia being different from that of
Myceliophthora and Thielavia. This genus is characterized by hyaline to pale brown or brown, erect
or flexuous, sterile setae, holoblastic, cylindrical or broadly irregular conidiogenous cells and
globose to subglobose conidia produced on aerial hyphae.
Amesia X. Wei Wang, Samson & Crous, Stud. Mycol. 84: 156 (2016)
Index Fungorum number: IF818829; 4 species with sequence data.
Type species – Amesia atrobrunnea (L.M. Ames) X. Wei Wang & Samson
Notes – Amesia was introduced by Wang et al. (2016a) to accommodate A. atrobrunnea, A.
cymbiformis, A. gelasinospora and A. nigricolor, which were transferred from Chaetomium. The
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genus is characterised by superficial, sphaerical, ellipsoid or ovate, ostiolate ascomata, with
apically flexuous, spirally coiled or undulate hairs. Asci are broadly clavate and evanescent and
ascospores are brown, mostly fusiform, often ovate to elongate ovate, with an apical or subapical
germ pore (Wang et al. 2016a).
Arcopilus X. Wei Wang, Samson & Crous, Stud. Mycol. 84: 159 (2016)
Index Fungorum number: IF818835; 7 species with sequence data.
Type species – Arcopilus aureus (Chivers) X. Wei Wang & Samson
Notes – Arcopilus was established by Wang et al. (2016a) to accommodate A. aureus, A.
cupreus, A. flavigenus, A. fusiformis and A. turgidopilosus, which were transferred from
Chaetomium. Based on the unique phylogenetic and morphologic analyses, Raza et al. (2019)
introduced A. globulus and A. tangerinicapillus, which were isolated from Saccharum officinarum.
The genus is characterised by superficial, subglobose to ovate, ostiolate ascomata, with hairs at
apical region which are arcuate, with apices incurved, circinate to coiled, at lateral sides flexuous or
apically incurved. Asci are clavate and evanescent and ascospores brown, inequilateral, fusiform,
navicular, reniform, lunate or limoniform with one or two apical germ pores.
Arxotrichum A. Nováková & M. Kolařík, Persoonia 40: 259 (2018)
Index Fungorum number: IF824080; 2 species with sequence data.
Type species – Arxotrichum wyomingense A. Nováková & M. Kolařík
Notes – Arxotrichum was established by Crous et al. (2018d) for A. wyomingense which was
isolated from soil, based on the analysis of LSU sequence data. Ascomata if present are pale
ochraceous to olivaceous grey, superficial, sphaerical to ovate and ostiolate with numerous
flexuous, undulate or spirally coiled, verrucose to echinulate, septate, pale ochraceous or brown
hairs. Asci are obovate-clavate and evanescent with short pedicels and ascospores are ellipsoidalfusoid to brown, with distinct apical germ pores. The asexual morph has septate conidiophores with
basal yellowish brown and apically colourless, ramified, racemose branched stipes and solitary,
aseptate, subglobose conidia that are rough-walled to rugose (Crous et al. 2018d).
Botryotrichum Sacc. & Marchal, Bull. Soc. R. Bot. Belg. 24(1): 66 (1885)
Index Fungorum number: IF7431; 11 morphological species, 4 species with sequence data.
Type species – Botryotrichum piluliferum Sacc. & Marchal
Notes – Botryotrichum has rarely been reported (Kirk et al. 2008, Wijayawardene et al.
2018a). Based on the phylogenetic analyses of ITS, LSU, rpb2 and tub2 gene regions, Wang et al.
(2016a) re-organized the genus and included some species of the genera Emilmuelleria and
Chaetomium. The genus is characterised by superficial, subglobose to ovate, olivaceous, ostiolate
or ascomata lacking ostioles, with flexuous or undulate, often circinate, unbranched, lateral setalike hairs, fusiform, long pedicellate, evanescent asci and fasciculate, olivaceous brown, ellipsoidalfusiform ascospores, with an apical germ pore. Conidiophores are setose, terminating in setae,
branched, macronematous with clusters of thick-walled conidia that have a conspicuous germ pore
(Wang et al. 2016a, 2019b).
Chaetomium Kunze, Mykologische Hefte (Leipzig) 1: 15 (1817)
Index Fungorum number: IF953; 211 morphological species (Species Fungorum 2020), 106
species with sequence data.
Type species – Chaetomium globosum Kunze.
Notes – Chaetomium is the largest genus in Chaetomiaceae. The genus was reviewed by
Maharachchikumbura et al. (2016b) and Wang et al. (2016a, b, 2019b). The genera Bommerella,
Chaetomidium and Chaetomiopsis were proposed as synonyms of Chaetomium (Moustafa &
Abdul-Wahid 1990, Doveri 2008a, Wang et al. 2016a, b, 2019b). Chaetomium contains more than
150 species (Wijayawardene et al. 2017a); they grow on soil and decaying twigs (Wang et al.
2016a, b, Zhang et al. 2017b). Wang et al. (2016a, b, 2019b) described novel Chaetomiaceae taxa
489
collected from indoor environments, which were described as new species. Many new species,
collected from soil, were published by (Zhang et al. 2017b). The genus is characterised by globose,
ellipsoid to ovate or obovate, ostiolate ascomata with or lacking ostioles, hypha-like, flexuous,
undulate, coiled to simply or dichotomously branched ascomatal hairs, clavate or fusiform,
evanescent asci, and limoniform to globose, bilaterally flattened ascospores. Asexual morphs, if
produced, are acremonium-like (Wang et al. 2016a).
Figure 64 – Chaetomium globosum (a, d, f-h redrawn from Wang 2016b and
(https://www.canada.ca/content/dam/eccc/documents/pdf/pded/chaetomium-globosum/2018-02-20C.%20Globosum-EN.pdf), b, c, e redrawn from Whiteside 1961). a Ascoma. b, c Sections of
ascomata. d Asci. e Paraphyses. f Ascomal hair. g Ascospores. h Peridium. Scale bars: a = 100 µm,
d, h = 10 µm, f, g = 5 µm, b = × 240, c = × 300, e = × 1000.
490
Figure 65 – Asexual morph of Chaetomium spp. (redrawn from Wang et al. 2016b). a, b
Chaetomium subaffine. c, d C. rectangulare. e, f C. elatum. a, c, e Conidiophores with conidia. b, d,
f Conidia. Scale bars: a, c, e = 10 µm, b, d, f = 5 µm.
Collariella X. Wei Wang, Samson & Crous, Stud. Mycol. 84: 177 (2016)
Index Fungorum number: IF818839; 9 species with sequence data.
Type species – Collariella bostrychodes (Zopf) X. Wei Wang & Samson
Notes – Based on both phylogenetic and morphological analyses, Collariella was established
by Wang et al. (2016b) to accommodate one new species and six species transferred from
Chaetomium. The genus is characterised by superficial, ovate, obovate, subglobose, ampulliform or
cylindrical, ostiolate ascomata, fusiform or clavate, pedicellate, evanescent asci and fasciculate
olivaceous brown, broadly limoniform to quadrangular, bilaterally flattened ascospores, with one or
two apical or subapical germ pores and no known asexual morphs (Wang et al. 2016a).
Corynascella Arx & Hodges, Stud. Mycol. 8: 23 (1975)
Index Fungorum number: IF1256; 3 morphological species (Species Fungorum 2020), 2
species with sequence data.
Type species – Corynascella humicola Arx & Hodges.
Notes – Corynascella was introduced by von Arx (1975) to accommodate the single species
C. humicola, in which the ascospores have strikingly thickened walls around the germ pores. This
genus was shown to cluster in a separate branch in phylogenetic analysis (van den Brink et al.
2012, Maharachchikumbura et al. 2015, Wang et al. 2016a, 2019b). The genus is characterised by
superficial, sphaerical, cleistothecial ascomata, covered with dark, septate, appendage-like hairs,
broadly clavate or obovate, thin-walled, evanescent asci and ellipsoidal, sphaerical or reniform, 1celled ascospores, rounded at the ends and with a large de Bary bubble, and 2(-4) apical germ pores
(von Arx 1975).
Crassicarpon Y. Marín, Stchigel, Guarro & Cano, Mycologia 107(3): 629 (2015)
Index Fungorum number: IF809487; 2 species with sequence data.
Type species – Crassicarpon thermophilum (Fergus & Sinden) Y. Marín, Stchigel, Guarro &
Cano
Notes – Crassicarpon is characterized by a thermophilic habit, blackish ascomata with a thick
wall of textura angularis, broadly clavate asci with 4–6 ascospores, broadly ellipsoidal ascospores
with a germ pore at each end, and hyaline, smooth-walled conidia, yellow in mass.
Dichotomopilus X. Wei Wang, Samson & Crous, Stud. Mycol. 84: 185 (2016)
Index Fungorum number: IF818840; 12 species with sequence data.
Type species – Dichotomopilus indicus (Corda) X. Wei Wang & Samson.
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Figure 66 – Dichotomopilus ramosissimus (Material examined – INDIA, Andaman and Nicobar
Islands, North Andaman, Diglipur, Near Sita Nagar (13˚11’14.2” N 92˚53’11” E), isolated on
decaying twig of Dolichandrone spathacea, 17 May 2018, M. Niranjan & V.V. Sarma, PUFNI
18725, (AMH-10059). a-c Ascomata on host. d, e Mature ascomata in lactophenol mount. f, g
Terminal ascomatal hair. h Ascospores. i-j Colonies on malt extract agar. Scale bars: d = 200 μm, e
= 100 μm, f = 20 μm, g, h = 10 μm.
Notes – The genus Dichotomopilus was established by Wang et al. (2016a) to include two
new species, D. pseudofunicla and D. pseudoerectus, and another ten species transferred from
Chaetomium. A new collection of D. ramosissimum was recorded on Clematis vitalba in the UK by
492
Phukhamsakda et al. (2020). The genus is characterised by superficial, sphaerical, ellipsoid or
ovate, ostiolate ascomata, with seta-like, branched, often ornamented terminal hairs, unbranched,
seta-like lateral hairs tapering towards the apex, clavate to ovate, pedicellate, evanescent asci and
fasciculate brown, ovate, bilaterally flattened ascospores attenuated at one or both ends and with
apical or slightly subapical germ pores. Asexual morphs are undetermined. Dichotomopilus
ramosissimus is illustrated in this entry (Fig. 66).
Guanomyces M.C. González, Hanlin & Ulloa, Mycologia 92(6): 1139 (2000)
Index Fungorum number: IF28466; 1 species with sequence data.
Type species – Guanomyces polythrix M.C. González, Hanlin & Ulloa
Notes – Based on morphology and phylogenetic analysis of LSU sequences, Guanomyces
was described in Sordariales by González et al. (2000) to accommodate species which were
isolated from bat dung. Stchigel et al. (2006) reported that Guanomyces was in Chaetomiaceae
(Sordariales). The following studies were concerned with secondary metabolites of Guanomyces
polythrix, with the taxonomic status rarely mentioned (Macías et al. 2000, 2001, Mata et al. 2003).
The genus is characterised by sexual morph with ascomata that are superficial, ovoidal, ostiolate,
with long neck, composed of glandular hairs, asci that are clavate, deliquescent, J- and ascospores
that are ellipsoidal, unicellular, multinucleate and hyaline, not appendaged and without germ pores
or slits. Asexual morphs have chlamydospores that are sphaerical or obovate and dark brown.
Humicola Traaen, Nytt Mag. Natur. 52: 31 (1914)
Index Fungorum number: IF8566; 72 morphological species (Species Fungorum 2020), 24
species with sequence data.
Type species – Humicola fuscoatra Traaen.
Notes – Humicola was introduced by Traaen (1914) to accommodate two species, H.
fuscoatra and H. grisea. A comprehensive study by Wang et al. (2019b) revised the phylogenetic
status of the genus. The genus is characterised sexual morphs with superficial, ostiolate ascomata
covered by aerial hyphae. Terminal hairs are seta-like, flexuous, undulate, coiled or arcuate with
apices incurved to coiled. Asci are clavate, early evanescent and ascospores are limoniform to
quadrangular, bilaterally flattened, with an apical germ pore. Asexual morphs producing conidia
arise laterally, intercalary or terminally from hyphae without differentiated conidiophores,
sometimes together with acremonium-like conidiophores, occasionally only acremonium-like
conidiophores are produced.
Madurella Brumpt, Compt.-Rend. Séances Mém. Soc. Biol. 58: 999 (1905)
Index Fungorum number: IF8824; 15 morphological species (Species Fungorum 2020), 3
species with sequence data.
Type species – Madurella mycetomatis (Laveran) Brumpt
Notes – Madurella was introduced by Brumpt (1905) to accommodate the species
Streptothrix mycetomatis. Madurella species are the causative agents of mycetoma of human’s
worldwide (Findlay et al. 1979, Mukerji & Manoharachary 2010, de Hoog et al. 2012, Bolis et al.
2018, Norina et al. 2018, Mekoguem et al. 2019). The genus is characterised by asexual morphs
with slow growing, white to brown colonies, hyphae producing chlamydospores, and conidiophores
and conidia have not been observed (Seifert et al. 2011).
Melanocarpus Arx, Stud. Mycol. 8: 17 (1975)
Index Fungorum number: IF3063; 5 morphological species (Species Fungorum 2020), 4
species with sequence data.
Type species – Melanocarpus albomyces (Cooney & R. Emers.) von Arx
Notes – Based on previous studies and phylogenetic analyses of combined rpb2, tub2, ITS
and LSU sequence datasets, Wang et al. (2016a, 2019b) consolidated its placement in
Chaetomiaceae (Maharachchikumbura et al. 2015, 2016b, Wijayawardene et al. 2018a). The genus
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is characterised by ascomata that are superficial, or enclosed within white aerial mycelium, often
aggregated, globose, black, glabrous or with a few hypha-like hyaline hairs, lacking ostioles; asci
that are fasciculate, ovate to broadly ovate, pedicellate, early evanescent; ascospores that are brown
at maturity, ovate to broadly ovate, bilaterally flattened, with an apical pore; and asexual morphs
undetermined.
Myceliophthora Costantin, C. r. hebd. Séanc. Acad. Sci., Paris 114: 849 (1892)
Index Fungorum number: IF9013; 5 morphological species (Species Fungorum 2020), 3
species with sequence data.
Type species – Myceliophthora lutea Costantin
Notes – Based on the phylogenetic analyses of sequences from ITS, tef1 and rpb2, MarinFelix et al. (2015) split Myceliophthora into four genera: Corynascus (a revalidated genus),
Crassicarpon, Myceliophthora and Thermothelomyces. However, according to the molecular
analyses, Corynascus was linked with Myceliophthora and all Corynascus species are proposed to
be placed under Myceliophthora (van den Brink et al. 2012, Maharachchikumbura et al. 2015,
2016b, Wijayawardene et al. 2018a). Wang et al. (2016a, 2019b) hinted that all three genera be
included under Myceliophthora. The genus is characterised by hyaline and smooth-walled conidia
and sexual morph is undetermined.
Mycothermus D.O. Natvig, J.W. Taylor, A. Tsang, M.I. Hutch. & A.J. Powell, Stud. Mycol. 93:
107 (2018)
Index Fungorum number: IF807381; 2 species with sequence data.
Type species – Mycothermus thermophilus (Cooney & R. Emers.) X. Wei Wang, Houbraken
& D.O. Natvig
Notes – Mycothermus was published by Natvig et al. (2015) but as an invalid genus. The
genus was validated by Wang et al. (2019b) based on multi-gene phylogenetic analyses with rpb2,
tub2, ITS and LSU. The genus is characterised by its thermophilic habit, hyaline to subhyaline
hyphae, holothallic chlamydospores that are thick-walled, brown, smooth to verrucose and
doliiform, globose, subglobose, oblong or obovoid. The sexual morphs are undetermined.
Ovatospora X. Wei Wang, Samson & Crous, Stud. Mycol. 84: 185 (2016)
Index Fungorum number: IF818850; 6 species with sequence data.
Type species – Ovatospora brasiliensis X. Wei Wang & Samson
Notes – Ovatospora was introduced by Wang et al. (2016a) to accommodate O. brasiliensis,
O. medusarum, O. mollicella, O. pseudomollicella, O. senegalensis, and O. unipora. The genus is
characterised by superficial, subglobose or ovate ascomata, ostiolate, with brown walls of cells of
textura angularis. Terminal hairs are usually coiled and often branched and lateral hairs are
flexuous. Asci are fasciculate, cylindrical or clavate and evanescent and ascospores are brown at
maturity, broadly ovate, bilaterally flattened, rounded at one end, attenuate or apiculate at other end
and have an apical germ pore. No asexual morphs are known (Wang et al. 2016a).
Remersonia Samson & Seifert, Can. J. Bot. 75(7): 1160 (1997)
Index Fungorum number: IF27809; 2 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Remersonia thermophila (Fergus) Seifert & Samson
Notes – Remersonia was introduced by Seifert et al. (1997) for the thermophilic synnematous
hyphomycete Stilbella thermophila. Based on previous studies and phylogenetic analyses, Wang et
al. (2019b) confirmed the placement of Remersonia as a monotypic genus in Chaetomiaceae. The
genus is characterised by synnemata that are cylindrical to clavate, capitate, unbranched and
hyaline. Conidiophores are unbranched or simply branched, have percurrently proliferating
conidiogenous cells and conidia are hyaline, oblong, clavate or ellipsoidal with truncate or rounded
bases (Seifert et al. 1997). The sexual morph is undetermined.
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Staphylotrichum J. Mey. & Nicot, Bull. trimest. Soc. mycol. Fr. 72: 322 (1957)
Index Fungorum number: IF10065; 9 morphological species (Species Fungorum 2020), 7
species with sequence data.
Type species – Staphylotrichum coccosporum J.A. Mey. & Nicot
Notes – Staphylotrichum was introduced by Nicot & Meyer (1956) to accommodate S.
coccosporum. Phylogenetic analyses of ITS and LSU sequence data in Nonaka et al. (2012)
reported the placement of Staphylotrichum in Chaetomiaceae. Wang et al. (2019b) confirmed this
placement in this family based on analysis of combined rpb2, tub2, ITS and LSU datasets for eight
isolates. The genus is characterised by ascomata that are superficial with long necks comprised of a
fused basal part of terminal hairs. Asci are 8-spored, clavate to fusiform and ascospores are broad
limoniform to globose, with an apical germ pore. The asexual morphs have two types of
conidiophores, micronematous that arise from hyphae and macronematous from apically branched,
seta-like conidiophores. Conidia are thick-walled, holoblastic, hyaline to pale brown and globose to
subglobose or obovoid (Wang et al. 2019b).
Subramaniula Arx, Proc. Indian Acad. Sci., Pl. Sci. 94(2-3): 344 (1985)
Index Fungorum number: IF25699; 9 morphological species (Species Fungorum 2020), 8
species with sequence data.
Type species – Subramaniula thielavioides (Arx, Mukerji & N. Singh) Arx
Notes – Subramaniula was introduced by von Arx (1985) to accommodate Achaetomium
thielavioides. The study of Ahmed et al. (2016) indicated a relationship of Subramaniula within
Chaetomiaceae. Phylogenetic analyses of Wang et al. (2016a) confirmed the placement and
expanded the number of species. The genus is characterised by ascomata that are globose and
glabrous without necks, with translucent walls and a wide ostiole surrounded by a hyaline collar.
Asci are clavate, short pedicellate, very thin-walled, evanescent and ascospores are widely
fusiform, brownish, thick-walled and 1-celled, with a conspicuous subapical germ pore (von Arx et
al. 1978, Wang et al. 2016a). The asexual morph in undetermined.
Thermothelomyces Y. Marín, Stchigel, Guarro & Cano, Mycologia 107(3): 630 (2015)
Index Fungorum number: IF809489; 4 species with sequence data.
Type species – Thermothelomyces thermophilus (Apinis) Y. Marín, Stchigel, Guarro & Cano
Notes – Thermothelomyces was introduced by Marin-Felix et al. (2015) and includes four
species. Phylogenetic analyses of combined ITS, tef1 and rpb2 datasets showed that
Thermothelomyces belongs in Chaetomiaceae (Marin-Felix et al. 2015). The genus is characterised
by a thermophilic habit, ascomata that are immersed or semi-immersed, globose, cleistothecial,
black, glabrous, with a wall of textura epidermoidea. Asci are ellipsoidal, thin-walled, pedicellate,
and evanescent and ascospores are ellipsoidal, dark brown to black at maturity, thick- and smoothwalled, with a single germ pore. In the asexual morph, conidiophores are micronematous or
semimacronematous and conidia are holoblastic, brown, subglobose or obovoid to ellipsoidal,
ornamented or rarely smooth.
Thielavia Zopf, Verh. bot. Ver. Prov. Brandenb. 18: 101 (1876)
Index Fungorum number: IF5450; 44 morphological species (Species Fungorum 2020), 23
species with sequence data.
Type species – Thielavia basicola Zopf
Notes – Thielavia was introduced by Zopf (1876) to accommodate a species isolated from the
roots of Senecio elegans and lupins. Revisions of the genus have been proposed by numerous
investigators (Booth 1961, Malloch & Cain 1973, Mouchacca 1973, von Arx 1975, Figueras &
Guarro 1988, Stchigel et al. 2002). Wang et al. (2016a, 2019b) confirmed Thielavia as a monotypic
genus in Chaetomiaceae. The genus is characterised by ascomata lacking ostioles and ascospores
that are unicellular, darkly pigmented with one or two germ pores. The asexual morph has conidia
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are hyaline or brightly coloured and produced as simple phialoconidia, aleurioconidia, or
arthroconidia.
Trichocladium Harz, Bull. Soc. Imp. nat. Moscou 44(1): 125 (1871)
Index Fungorum number: IF10278; 38 morphological species (Species Fungorum 2020), 10
species with sequence data.
Type species – Trichocladium asperum Harz
Notes – Trichocladium was introduced by Harz (1871). A review of Trichocladium was
undertaken by Goh & Hyde (1999) who accepted 18 species. Trichocladium species are
polyphyletic within Chaetomiaceae, and other families (Shearer & Crane 1971, Kirk et al. 2008,
Seifert et al. 2011, Jones et al. 2015). Based on the analyses of rpb2 and a combined rpb2, tub2,
ITS and LSU datasets, 22 strains of Trichocladium formed a lineage, which includes four closely
related subclades in Chaetomiaceae and they show morphologically diverse characters, and are
phylogenetically distinct. The asexual morphs have conidiophores that are hyaline, developed
laterally or terminally from the hyphae, and cylindrical, unbranched or branched, sometimes
verticillate. Conidia are 1- to 2-celled, rarely 3-celled, obovate, pyriform, ellipsoid, constricted at
the septa, conspicuously warted and olivaceous brown to dark brown, with paler basal cells. The
sexual morph of Trichocladium is characterized by superficial or immersed ascomata in a thick
mycelium, with or lacking ostioles. Asci are typically cylindrical, evanescent, with 8 (4) uniseriate
ascospores which are typically broadly ovate, bilaterally flattened, sometimes ellipsoidal and nonflattened, with an apical germ pore (Wang et al. 2019b).
Chaetosphaerellaceae Huhndorf, A.N. Mill. & F.A. Fernández, Mycol. Res. 108(12): 1387 (2004)
Index Fungorum number: IF82145; Facesoffungi number: FoF01114; 6 species.
Saprobic on woody substrates in terrestrial habitats. Sexual morph: Ascomata perithecial,
dark brown to black, scattered to densely gregarious, superficial, sitting in a subiculum or
subiculum absent, pyriform, obpyriform or ovoid, turbinate, coriaceous, tuberculate or smooth,
with or without, brown, branched or unbranched setae, papillate or papilla lacking, collabent or not
collapsing, ostiolate. Subiculum abundant or sparse, sometimes lacking, brown to dark brown,
consisting of septate, branched or unbranched with spiny hyphae. Peridium with Munk pores, outer
layer composed of dark brown to brown cells of textura angularis; inner layer composed of hyaline
cells of textura prismatica. Paraphyses inflated, often present only in young ascomata. Asci 8spored, unitunicate, clavate, long or short pedicellate, apical ring distinct, indistinct or absent,
evanescent. Ascospores 1–3-seriate or overlapping, with brown or brown median cells and hyaline
end cells, oblong cylindrical, ellipsoid or fusiform, 1–3-septate. Asexual morph Conidiophores
macronematous, mononematous, erect, straight, solitary or divaricate with 2–4 metulae, brown,
septate, branched, percurrent, with a terminal, ampulla. Conidiogenous cells holoblastic or
enteroblastic, integrated, terminal. Conidia hyaline to dark brown, oval or elliptical, solitary or
catenate (adapted from Maharachchikumbura et al. 2016b).
Type genus – Chaetosphaerella E. Müll. & C. Booth.
Notes – Chaetosphaerellaceae, typified by Chaetosphaerella (Müller & Booth 1972), was
introduced by Huhndorf et al. (2004a) for fungi with superficial, ostiolate ascomata siting on a
subiculum, clavate or cylindrical asci, with pigmented ellipsoid ascospores, and enteroblastic or
holoblastic conidiogenesis. In a taxonomic re-evaluation of Chaetosphaeria species, Réblová
(1999a, b, c, d) accepted Chaetosphaerella phaeostroma and C. fusca in the genus and placed them
in the Helminthosphaeriaceae based on peridium anatomy, setae, ascal anatomy, absence of
Quellkörper and conidiogenesis. Chaetosphaerella fusispora Sivanesan was excluded from the
genus and transferred to Crassochaeta based on difference in ascus, hamathecium and ascospore
anatomy by Réblová (1999b). Chaetosphaerella is associated with asexual morphs formerly
referred to Oedemium and Veramycina. Réblová (1999a) discussed relationships of
Chaetosphaerella with Nitschkiaceae based on the peridium. A phylogeny based on LSU rDNA
sequence data showed that Chaetosphaerella and Crassochaeta clustered in the same clade, for
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which Chaetosphaerellaceae was described (Huhndorf et al. 2004a). Mugambi & Huhndorf (2010)
referred Spinulosphaeria to Sordariomycetes genera incertae sedis, based on morphology and
analysis of LSU sequence data of S. nuda. In a combined LSU, SSU, tef1 and rpb2 dataset, S. nuda
formed a sister group to Chaetosphaerella species with high support in Chaetosphaerellaceae
(Mugambi & Huhndorf 2010, Maharachchikumbura et al. 2015, 2016b). Réblová et al. (2016b)
proposed Chaetosphaerella as a correct name for the genus and recommended Chaetosphaerella
for protection over Oedemium and Veramycina.
Ecological and economic significance of Chaetosphaerellaceae
Most Chaetosphaerellaceae species are saprobic on wood, widespread around the world,
although most have been found in Europe (Sivanesan 1974, Réblová 1999a, b, d, Huhndorf et al.
2004a, Mugambi & Huhndorf 2010).
Genera included in Chaetosphaerellaceae
Chaetosphaerella E. Müll. & C. Booth, Trans. Br. mycol. Soc. 58(1): 76 (1972)
Index Fungorum number: IF969; 2 species with sequence data.
Type species – Chaetosphaerella phaeostroma (Durieu & Mont.) E. Müll. & C. Booth
Notes – Chaetosphaerella was introduced by Müller & Booth (1972) and included two
species, C. phaeostroma and C. fusca. The type species C. phaeostroma, is characterized by black
ascomata surrounded by a subiculum, with tuberculate or roughened perithecia, clavate asci and
pigmented ascospores. Sivanesan (1976) and Varghese & Rao (1979a) included C. fusispora and C.
indica in Chaetosphaerella based on the characters, however, Réblová (1999a, b) excluded them
based on different morphology of the peridium, setae, asci and conidiogenesis. Chaetosphaerella is
closely related to Crassochaeta in Fig. 11.
Crassochaeta Réblová, Mycotaxon 71: 46 (1999)
Index Fungorum number: IF28310; 2 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Crassochaeta nigrita (Sacc.) Réblová
Notes – Crassochaeta nigrita is characterized by tuberculate ascomata surrounded by needleshaped setae, cylindrical asci and subglobose to ellipsoid, versicolored ascospores. The asexual
morph was reported as arthrinium-like Réblová (1999d). The phylogeny based on LSU sequence
data of C. nigrita was provided by Huhndorf et al. (2004a) and reveals a close relationship of
Crassochaeta to Chaetosphaerella within Chaetosphaerellaceae (same results in Fig. 11).
Spinulosphaeria Sivan., Trans. Br. mycol. Soc. 62(1): 5 (1974)
Index Fungorum number: IF5149; 2 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Spinulosphaeria thaxteri (Pat.) Sivan.
Notes– Spinulosphaeria, typified by Spinulosphaeria thaxteri, was established by Sivanesan
(1974) and has tuberculate ascomata, clavate asci and subglobose to ellipsoid ascospores. The
second species, Spinulosphaeria nuda is characterized by tuberculate ascomata and verrucose
ascospores with mucilaginous sheath (Mugambi & Huhndorf 2010). Phylogenetic analysis of LSU
sequence data of S. nuda reveals that Spinulosphaeria is related to Chaetosphaerellaceae and
Nitschkiaceae (Mugambi & Huhndorf 2010). In this study, Spinulosphaeria is closely related to
Crassochaeta and Chaetosphaerella within Chaetosphaerellaceae (Fig. 11).
Chaetosphaeriaceae Réblová, M.E. Barr & Samuels, Sydowia 51(1): 56 (1999)
Index Fungorum number: IF82100; Facesoffungi number: FoF01139; 658 species.
Saprobic on wood or decaying plant parts including leaves, fruits in terrestrial and aquatic
habitats, sometimes isolated from soil and fresh plant parts such as leaves, less commonly -
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Figure 67 – Spinulosphaeria thaxteri (Material examined – USA, Florida, Dayton; on rotten wood,
1898, collected by Patouillard, FH 16-10965, holotype). a Material label. b Material and line
drawing. c Ascomata on wood. d Ascoma. e Ascoma vertical section. f Peridium. g-i Asci with
paraphyses. j-l Ascospores. Scale bars: d-f = 200 µm, g-i = 20 µm, j = 10 µm, k, l = 5 µm.
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-pathogenic. Sexual morph: Ascomata perithecial, gregarious or scattered, solitary, superficial or
basally immersed on a thin stroma or on a thin subiculum, ovoid, globose to subglobose,
carbonaceous, coriaceous or membranaceous, dark brown to black, rough, smooth or with short
setae, papillate, the apex collapsing when dry. Ostiole lined with hyaline periphyses or periphyses
absent. Subiculum thin, scanty, brown to dark brown, septate, of unbranched hyphae. Peridium
composed of two layers, outer layer comprising brown cells of textura epidermoidea or angularis,
carbonaceous; inner layer comprising hyaline cells of textura prismatica, thin, membranaceus.
Paraphyses numerous, septate, unbranched, tapering, filiform or cylindrical. Asci 8-spored,
unitunicate, thin-walled, clavate to cylindrical, long or short pedicellate, with J-, apical ring.
Ascospores 2–3-seriate, fusiform, cylindrical to ellipsoid, straight or sometimes curved, 0–3septate, with or without fragmenting, hyaline or brown or becoming dark coloured in part, smooth
or striate, with guttules, sheath or appendages. Asexual morph: Hyphomycetous or coelomycetous.
When hyphomycetous, Conidiophores macronematous, mononematous, scattered or gregarious,
dark brown or hyaline, straight or flexuous, septate, branched or unbranched, with short encircling
collar hyphae. Conidiogenous cells monophialidic or polyphialidic, holoblastic or enteroblastic,
proliferating percurrently or sympodial, hyaline, with a distinct funnel-shaped collarette, smoothwalled. Conidia aggregated, continuous or mucilaginous, aseptate to multi-septate, flexuous,
fusiform, allantoid, cylindrical or doliiform, curved or straight, hyaline to dark brown. When
coelomycetous, Conidiomata stromatic, scattered or aggregated, superficial, cupuliform or globose,
unilocular, setose, comprising black to dark brown cells of textura angularis or intricata. Setae
numerous, black to brown, septate, ovoid to cylindrical or arising from the outer elements of
excipulum, smooth, thick-walled, multi-septate. Conidiophores lining the basal stroma in a dense
layer or arising from conidiomatal cavity, brown, 4–6-septate, unbranched, cylindrical, thin-walled,
smooth. Conidiogenous cells integrated, determinate, holoblastic or enteroblastic, phialidic with
conspicuous periclinal thickening at an attenuated apex, brown, smooth, subcylindrical to
lageniform. Conidia aseptate, globose to subglobose or ellipsoid, fusiform to allantoid, curved or
straight, obtuse to sub-obtusely rounded at apex, truncate at base, eguttulate or guttulate, hyaline to
brown, thin, smooth-walled, with a single, cellular, unbranched, flexuous, with tubular appendage
at each end, separated by a septum, with basal asymmetrically located appendage.
Type genus – Chaetosphaeria Tul. & C. Tul.
Notes – Chaetosphaeriaceae was introduced without any description by Locquin (1984) to
accommodate five genera including Chaetosphaeria, Loramyces, Niesslia, Rhagadostoma, and
Zignöella. This was not considered as a validly published family (Hawksworth & David 1989 –
Art. 36.1, Grueter et al. 1994). Hence, Réblová et al. (1999) re-introduced the family based on
Chaetosphaeria, to accommodate another six genera: Ascocodinaea, Melanochaeta,
Melanopsammella, Porosphaerella, Porosphaerellopsis, and Striatosphaeria. Réblová et al. (1999)
maintained Chaetosphaeriaceae in Sordariales based on morphology. Based on LSU sequence data,
Huhndorf (2004b) placed the family in Chaetosphaeriales. Subsequently more genera were added
to the family and, Maharachchikumbura et al. (2016b) accepted 37 genera. Maharachchikumbura et
al. (2016b) aknowledged the taxonomic confusion of genera in the family and the need for a
monograph with molecular support for accepted genera. Crous et al. (2016a) introduced
Adautomilanezia and Eucalyptostroma to Chaetosphaeriaceae. Ma et al. (2016b) introduced the
hyphomycetes genus Anacacumisporium based on phylogenetic analyses. Five genera,
Codinaeopsis, Dictyochaetopsis, Phaeostalagmus, Phialogeniculata and Zignoëlla, previously
listed by Maharachchikumbura et al. (2016b) in Chaetosphaeriaceae, were not considered by
Wijayawardene et al. (2018a). Wijayawardene et al. (2018a) listed 38 genera in Chaetosphaeriaceae
including Adautomilanezia, Anacacumisporium, Conicomyces, Eucalyptostroma, Menisporopsis
and Pseudolachnella. Yang et al. (2018a) placed Nawawia and Phialosporostilbe in
Chaetosphaeriaceae based on LSU and ITS sequence data. A new genus Multiguttulispora was
added to the family by Lin et al. (2019b). Crous et al. (2018d) provided first DNA sequence data
for Polynema and placed it in Chaetosphaeriaceae. In this entry the sexual morph of
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Chaetosphaeria jonesii and the asexual morph of a new species, Chloridium submersum, is
illustrated.
Ecological and economic significance of Chaetosphaeriaceae
Chaetosphaeriaceae members are saprobes in terrestrial and aquatic habitats (Réblová et al.
1999, Maharachchikumbura et al. 2016b). As examples Dictyochaeta, Tainosphaeria and
Menisporopsis are aquatic, while Adautomilanezia, Chaetosphaeria and Thozetella are terrestrial
saprobes (Crous et al. 2016a, Ho et al. 2001, Liu et al. 2016, Perera et al. 2016b, Réblová et al.
1999). Chaetosphaeriaceae species have the ability to decompose lignocellulose substrates in
woody litter and release nutrients (Yuen et al. 1998, Hyde et al. 2016a, Liu et al. 2016). Hence,
they play an important role in nutrient and carbon cycling and ecosystem functioning (Hyde et al.
2016a, Palmer et al. 1997, Wong et al. 1998a). Two Eucalyptostroma species (E. eucalypti and E.
eucalyptorum) were isolated from leaf spots of Eucalyptus (Crous et al. 2016a, 2018d). However,
there is no study confirming any pathogenicity in Chaetosphaeriaceae species. Some
Dinemasporium species have been reported to produce secondary metabolites (Yamaguchi et al.
2005, Krohn et al. 2008, Hashimoto et al. 2015a).
Genera included in Chaetosphaeriaceae
Adautomilanezia Gusmão, S.S. Silva, Fiuza, L.A. Costa & T.A.B. Santos, Persoonia 37: 229
(2016)
Index Fungorum number: IF815142; 1 species with sequence data.
Type species – Adautomilanezia caesalpiniae Gusmão, S.S. Silva, Fiuza, L.A. Costa &
T.A.B. Santos
Notes – The monotypic, hyphomycetous genus Adautomilanezia was introduced by Crous et
al. (2016a) based on LSU sequence data. The type species, A. caesalpinia, was isolated from
decaying twigs of Caesalpina echina in Brazil (Crous et al. 2016a). The genus is characterised by
sporodochial conidiomata with setae, enteroblastic conidiogenous cells and oblong to clavate,
multi-septate conidia (Crous et al. 2016a).
Anacacumisporium Y.R. Ma & X.G. Zhang, Cryptog. Mycol. 14: 8 (2016)
Index Fungorum number: IF811418; 1 species with sequence data.
Type species – Anacacumisporium appendiculatum Y.R. Ma & X.G. Zhang
Notes – The monotypic, hyphomycetous genus Anacacumisporium was established by Ma et
al. (2016b) with the single species A. appendiculatum, which was isolated from decaying wood in
China (Ma et al. 2016b). The genus is characterized by mononematous, macronematous
conidiophores, inconspicuous phialidic conidiogenous cells and euseptate conidia with an
appendage at the tip and base, aggregated in slimy masses (Ma et al. 2016b).
Ascochalara Réblová, Sydowia 51(2): 212 (1999)
Index Fungorum number: IF28371; 1 morphological species.
Type species – Ascochalara gabretae Réblová
Notes – Ascochalara was introduced by Réblová (1999e) to accommodate the lignicolous
species, A. gabretae, characterised by black, papillate ascomata, covered with greyish to whitish
powder, cylindrical-clavate asci with a J-, refractive apical ring and fusiform ascospores
disarticulating into part ascospores within the asci. The asexual morph of the fungus is charala-like,
with conidiophores mononematous, macronematous, phialidic conidiogenous cells and subcylindrical to wedge-shaped, aseptate conidia.
Brunneodinemasporium Crous & R.F. Castañeda, Persoonia, Mol. Phyl. Evol. Fungi 28: 128
(2012)
Index Fungorum number: IF800158; 2 species with sequence data.
Type species – Brunneodinemasporium brasiliense Crous & R.F. Castañeda
500
Notes – The coelomycetous genus Brunneodinemasporium was introduced by Crous et al.
(2012e) based on a dinemasporium-like species. Lu et al. (2016) added B. jonesii based on LSU
and ITS sequence data. Brunneodinemasporium species are characterised by cupulate, setose
conidiomata, with tightly aggregated brown conidiogenous cells and pale brown aseptate conidia
(Crous et al. 2012e, Lu et al. 2016). Species of this genus occur on decaying wood and leaves in
terrestrial and freshwater habitats (Crous et al. 2012e, Lu et al. 2016).
Catenularia Grove, Syll. fung. (Abellini) 4:303 (1886)
Index Fungorum number: IF7506; 13 morphological species (Species Fungorum 2020).
Type species – Catenularia simplex Grove
Notes – Saccardo (1886) established the hyphomycetous genus Catenularia to accommodate
the species C. simplex and C. atra. Species are lignicolous and found on dead wood and decaying
leaves and herbaceous substrates (Hughes & Kendrick 1965, Subramanian & Bhat 1987,
Matsushima 1971, Li et al. 2017a). Catenularia species are characterised by macronematous,
mononematous, simple or flexuous conidiophores, monophialidic, collarette conidiogenous cells
and aseptate phragmoconidia formed solitary or in chains (Saccardo 1886, Subramanian & Bhat
1987, Li et al. 2017a, Holubová-Jechová 1982).
Chaetosphaeria Tul. & C. Tul., Select. fung. carpol. (Paris) 2: 252 (1863)
Index Fungorum number: IF970; 107 morphological species (Species Fungorum 2020), 50
species with sequence data.
Type species – Chaetosphaeria innumera Berk. & Broome
Notes – Species of the genus are characterized by black ascomata with surface smooth or
covered by setae, 8-spored asci, with a small, distinct, refractive, J-, apical ring and septate, hyaline
ascospores (Maharachchikumbura et al. 2016b, Fernández et al. 2006). The asexual morph of
Chaetosphaeria is hyphomycetous with macronematous or mononematous conidiophores,
monophialidic or polyphialidic, conidiogenous cells, aseptate to multi-septate, guttulate or
eguttulate conidia, with or without appendages (Maharachchikumbura et al. 2016b, Fernández et al.
2006). Chaetosphaeria species are saprobes on decaying plant material in terrestrial and freshwater
habitats (Fernández & Huhndorf 2005, Perera et al. 2016b, Atkinson et al. 2007). The sexual morph
of Chaetosphaeria jonesii is illustrated (Fig. 68).
Chloridium Link, Mag. Gesell. naturf. Freunde, Berlin 3(1–2): 13 (1809)
Index Fungorum number: IF7624; 28 morphological species (Species Fungorum 2020), 5
species with sequence data.
Type species – Chloridium viride Link
Notes – Chloridium is characterised by mononematous or macronematous conidiophores,
monophialidic or polyphialidic conidiogenous cells, with a funnel-shaped collarette and hyaline to
brown, eguttulate or guttulate, appendaged or non-appendaged conidia (Link 1809, Seifert et al.
2011, Wei et al. 2018). Réblová et al. (2016b) pointed out that the generic type of Chloridium, C.
viride is congeneric with Melanopsammella inaequalis, the type of Melanopsammella (Réblová &
Winka 2000, Fernández et al. 2006, Crous et al. 2012e). Hughes (1951c) revealed that the generic
type of Gonytrichum, G. caesium, is the asexual morph of Melanopsammella inaequalis. By
considering the above facts, Réblová et al. (2016b) proposed that Gonytrichum and
Melanopsammella are synonyms of Chloridium. Hence, the widely used generic name Chloridium
was conserved over Gonytrichum and Melanopsammella (Réblová et al. 2016b, Wei et al. 2018).
Wijayawardene et al. (2018a) incorrectly treated Chloridium, Gonytrichum and Melanopsammella
as three distinct genera. Here we follow Réblová et al. (2016b) which is also confirmed in our
phylogenetic analysis of combined ITS and LSU sequence data (Fig. 8). Furthermore, Chloridium
species seems to be polyphyletic within Chaetosphaeriaceae (Fig. 8). We introduce a new species
of Chloridium in this entry.
501
Figure 68 – Chaetosphaeria jonesii (Material examined – THAILAND, Chiang Mai Province, on
decorticated wood, 5 August 2015, S. Boonmee RHP 121, MFLU 16-1020, holotype). a Herbarium
material. b Appearance of ascomata on host substrate. c Section of ascoma. d Setae. e Paraphyses.
f-h Asci. i Close up of apical ascus in Melzer’s reagent. j-l Ascospores. Scale bars: b = 200 µm, c =
100 µm, d-h = 50 µm, i-l = 10 µm.
Chloridium submersum Z.L. Luo, K.D. Hyde & H.Y. Su, sp. nov.
Fig. 69
Index Fungorum number: IF556752; Facesoffungi number: FoF06866
Etymology – Referring to the submerged habitat of this fungus.
Holotype – MFLU 18-1609.
Saprobic on submerged decaying wood. Sexual morph: Undetermined. Asexual morph:
Colonies effuse, brown, with long hairy mycelium, with white glistening conidial mass. Mycelium
partly immersed, partly superficial, consisting of branched, septate, brown hyphae. Conidiophores
502
116–264(–350) µm ( x = 190 µm, SD = 74, n = 10) long, 4–6 μm ( x = 5 µm, SD = 1, n = 10) wide,
macronematous, mononematous, erect, straight or slightly flexuous, septate, brown at the base,
gradually becoming paler towards apex, smooth. Conidiogenous cells integrated, terminal, polyenteroblastic, with a conspicuous outer collarette, hyaline. Conidia 3.5–4.5 µm ( x = 4 µm, SD =
0.5, n = 30) long, 2–3 μm ( x = 2.5 µm, SD = 0.5, n = 30) wide, acrogenous, aggregated in slimy
mass at the apex of the conidiophore, ellipsoid, hyaline, aseptate, guttulate, smooth.
Material examined – CHINA, Yunnan Province, saprobic on decaying submerged wood in
Dulong river, May 2015, H.Y. Su, H D5-13-1, S-510, MFLU 18-1609, holotype; ex-type living
culture, MFLUCC 16-1344.
GenBank numbers – ITS MN860551, LSU MN860556.
Notes – Chloridium submersum resembles C. phaeosporum in having macronematous,
mononematous, erect, septate conidiophores which are brown at the base, gradually becoming paler
towards apex, terminal, hyaline conidiogenous cells and ellipsoid, hyaline, aseptate conidia (Wu &
Zhang 2013). However, C. submersum differs from C. phaeosporum by its longer conidiophores
(116–264(–350) vs 70–120 µm), polyblastic, denticulate conidiogenous cells and guttulate, hyaline
conidia, while the latter species have monophialidic conidiogenous cells constricting abruptly and
expanding in a flaring collarette and pale brown conidia without guttules. Chloridium submersum is
phylogenetically related to C. aquaticum, C. gonythichii and C. aseptatum (Fig. 8).
Codinaea Maire, Publ. Inst. Bot. 3(4): 15 (1937)
Index Fungorum number: IF7720; 16 morphological species (Species Fungorum 2020), 4
species with sequence data.
Type species – Codinaea aristata Maire
Notes – Codinaea was introduced based on C. aristata (Maire 1937). Gamundi et al. (1977)
examined several specimens of Dictyochaeta fuegiana and proposed Codinaea as a synonym of
Dictyochaeta. Their treatment was accepted by most authors and many Codinaea taxa have been
moved to Dictyochaeta (Kirschner & Chen 2002, Cruz et al. 2008, Liu et al. 2016, Whitton et al.
2012). However, re-examination of the two collections, on which Gamundí et al. (1977) based their
description of D. fuegiana (LPS 38629, LPS 38630), revealed a fungus was misidentified as the
type of Dictyochaeta fuegiana (Réblová 2004). Réblová (2004) re-examined the type specimen of
Dictyochaeta fuegiana and gave detailed descriptions. Molecular studies by Réblová & Winka
(2000) revealed that Dictyochaeta species with or without setulae clustered into distinct subgroups
in the phylogenetic tree. Réblová (2000) proposed to retain Codinaea for species with setulae and
placed species without setulae in Dictyochaeta. Seifert et al. (2011) and Li et al. (2012) accepted
Réblová (2000) suggestion on segregation of Codinaea and Dictyochaeta as separate genera
delineated by the presence or absence of conidial setulae and accepted both Codinaea and
Dictyochaeta as valid genera.
Wijayawardene et al. (2018a) and Hernández-Restrepo (2017) also treated them as two
distinct genera. Currently, 104 species of Dictyochaeta and 53 species of Codinaea have been
described (Index Fungorum 2020). However, only few have DNA sequence data. There are no extype sequence data are available for the generic types of Codinaea or Dictyochaeta. Our
phylogenetic analysis based on the available sequences of Dictyochaeta and Codinaea species,
revealed that species of both genera are polyphyletic within Chaetosphaeriaceae (Fig. 8). Therefore,
a detailed morpho-molecular analysis with more taxonomic sampling would be required to finally
confirm the phylogenetic status of Codinaea and Dictyochaeta. Here we keep the Codinaea and
Dictyochaeta as separate genera until the generic types of Codinaea and Dictyochaeta are
recollected, sequenced and confirmed the link by molecular data.
Conicomyces R.C. Sinclair, Eicker & Morgan-Jones, Mycologia 75(6): 1100 (1983)
Index Fungorum number: IF11019; 4 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Conicomyces transvaalensis R.C. Sinclair, Eicker & Morgan-Jones
503
Figure 69 – Chloridium submersum (MFLU 18-1609, holotype). a Colonies on wood. b-d
Conidiophores with conidia. e, f Conidiogenous cells. g-k Conidia. l, m Germinating conidia. n, o
Culture on PDA from surface and reverse. Scale bars: b-d = 100 μm, e, f = 30 μm, g = 10 μm, h-m
= 5 μm.
504
Notes – The hyphomycetous genus Conicomyces was introduced by Sinclair & Eicker (1983).
Species of Conicomyces are lignicolous being saprobic on decaying woody substrates (HolubováJechová 1973a, Sinclair & Eicker 1983, Illman & White 1985, Liu et al. 2015). Conicomyces
species are characterised by cone-shaped, setose synnemata, macronematous, enteroblastic
conidiophores and aseptate, fusiform conidia bearing an apical appendage (Sinclair & Eicker 1983).
Craspedodidymum Hol.-Jech., Česká Mykol. 26(2): 70 (1972)
Index Fungorum number: IF7804; 14 morphological species (Species Fungorum 2020).
Type species – Craspedodidymum elatum Hol.-Jech.
Notes – The hyphomycetous genus Craspedodidymum was introduced by Holubová-Jechová
(1972) based on C. elatum. Craspedodidymum species are saprobic fungi occur on decaying plant
material (Pinruan et al. 2004, Ma et al. 2011, Yanna et al. 2000). Species of this genus are
characterised by macronematous conidiophores and apically swollen conidiogenous cells with a
large funnel-shaped terminal collarette (Pinruan et al. 2004).
Cryptophiale Piroz., Can. J. Bot. 46: 1123 (1968)
Index Fungorum number: IF7833; 21 morphological species (Species Fungorum 2020), 2
species with sequence data.
Type species – Cryptophiale kakombensis Piroz.
Notes – The hyphomycetous genus Cryptophiale was introduced by Pirozynski (1968) based
on C. kakombensis. Cryptophiale species are saprobes occur on decaying leaves, wood, roots and
bark (Pirozynski 1968, Goh & Hyde 1996). Species of the genus are characterised by unbranched
or apically dichotomous or verticillate, setiform, conidiophores, monophialidic, obscured
conidiogenous cells in two rows and unicellular to multiseptate, hyaline conidia produced in slimy
masses on one side of the conidiophore (Pirozynski 1968, Seifert et al. 2011, Yang et al. 2018a).
Cryptophialoidea Kuthub. & Nawawi, Trans. Br. mycol. Soc. 89(4): 581 (1987)
Index Fungorum number: IF11061; 5 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Cryptophialoidea secunda (Kuthub. & B. Sutton) Kuthub. & Nawawi
Notes – Kuthubutheen & Nawawi (1987) introduced asexual morph genus Cryptophialoidea
based on C. secunda. The genus is characterised by setiform, brown conidiophores, monophialidic
or polyphialidic phialidic, sessile conidiogenous cells, arranged on one side of the conidiophore
and, hyaline, falcate conidia (Kuthubutheen & Nawawi 1987, Yang et al. 2018a).
Dendrophoma Sacc., Michelia 2(no. 6): 4 (1880)
Index Fungorum number: IF7944; 59 morphological species (Species Fungorum 2020), 6
species with sequence data.
Type species – Dendrophoma cytisporoides Sacc.
Notes – Dendrophoma was proposed to be synonymous with Dinemasporium as
Dendrophoma cytisporoides was related to Dinemasporium graminum (Sutton 1965). However,
Crous et al. (2012e) epitypified Dendrophoma cytosporoides and confirmed that it is different from
Dinemasporium based on morphological and DNA sequence data. Species of Dendrophoma occur
on woody substrates and leaves, while some of them reported as leaf pathogens (Howard &
Albregts 1973, Moricca et al. 2016, Spegazzini 1880, Vu et al. 2019). The genus is characterised by
stromatic, stipitate, setose conidiomata, discrete, or integrated, lageniform to subcylindrical
conidiogenous cells and naviculate to botuliform conidia bearing an unbranched cellular appendage
at each end (Crous et al. 2012e).
Dictyochaeta Speg., Physis, Rev. Soc. Arg. Cienc. Nat. 7: 18 (1923)
Index Fungorum number: IF7996; 104 morphological species (Species Fungorum 2020), 13
species with sequence data.
505
Type species – Dictyochaeta fuegiana Speg.
Notes – Dictyochaeta was erected by Spegazzini (1923) based on Dictyochaeta fuegiana.
Species of this genus are saprobes on decaying plant parts (Whitton et al. 2000, Réblová & Winka
2000, Kuthubutheen & Nawawi 1991). For more details, see note under Codinaea. Dictyochaeta is
characterised by macronematous, mononematous, pale brown to dark brown conidiophores, monoor polyphialidic, sympodially proliferating conidiogenous cells with flared collarettes and hyaline,
clavate conidia without setulae (Réblová & Winka 2000).
Dictyochaetopsis Aramb. & Cabello, Mycotaxon 38: 12 (1990)
Index Fungorum number: IF11255; 14 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Dictyochaetopsis apicalis (Berk. & M.A. Curtis) Aramb. & Cabello
Notes – Arambarri & Cabello (1990) established Dictyochaetopsis based on D. apicalis to
accommodate species producing lateral phialides. Species of the genus are characterised by
macronematous, setiform conidiophores with fertile or sterile apices; discrete (rarely integrated),
monophialidic or rarely polyphialidic conidiogenous cells occurring on the conidiophore or on
lateral branches and, fusoid to cylindrical, 0-multiseptate, hyaline conidia (Arambarri & Cabello
1990, Whitton et al. 2000). Whitton et al. (2000) synonymized Codinaeopsis under
Dictyochaetopsis. Seifert et al. (2011) did not accept Codinaeopsis and Dictyochaetopsis as valid
genera. However, Lin et al. (2019a) treated Dictyochaetopsis as a distinct genus in
Chaetosphaeriaceae and is followed here.
Dinemasporium Lév., Annls Sci. Nat., Bot., sér. 3 5: 274 (1846)
Index Fungorum number: IF8040; 77 morphological species (Species Fungorum 2020), 20
species with sequence data.
Type species – Dinemasporium graminum (Lib.) Lév. (Pers.) Sacc.
Notes – Dinemasporium species are coelomycetes, mainly occurring on various plant and
woody substrates, including decaying wood (Nag Raj 1993, Hashimoto et al. 2015a, Crous et al.
2012e, Duan et at. 2007). Few were isolated from soil and human sputum (Nag Raj 1993,
Hashimoto et al. 2015a, Crous et al. 2012e). The genus is characterised by superficial, setose,
cupulate conidiomata, discrete or integrated conidiogenous cells and fusiform, naviculate or
allantoid, aseptate conidia, with one setulae at each end, with or without lateral appendages (Crous
et al. 2012e).
Eucalyptostroma Crous & M.J. Wingf., Persoonia 37: 311 (2016)
Index Fungorum number: IF819066; 2 species with sequence data.
Type species – Eucalyptostroma eucalypti Crous & M.J. Wingf.
Notes – The hypomycetes genus Eucalyptostroma was established by Crous et al. (2016a).
This genus comprises two species and both were isolated from symptomatic leaves of Eucalyptus
pellita (Crous et al. 2016a, 2018d). Eucalyptostroma species are characterised by sporodochial
conidiomata lacking setae, penicillate conidiogenous apparatus, and aseptate hyaline conidia,
without appendages (Crous et al. 2016a, 2018d).
Exserticlava S. Hughes, N.Z. J Bot. 16(3): 332 (1978)
Index Fungorum number: IF8242; 7 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Exserticlava vasiformis (Matsush.) S. Hughes
Notes – The hyphomycetous genus Exserticlava has been reported from decaying woody
substrates in terrestrial and freshwater habitats (Cai & Hyde 2007a, Ren et al. 2011, Hughes 1978,
Tsui et al. 2001a). The genus is characterised by simple conidiophores bearing single polyblastic,
funnel-shaped, conidiogenous cells and disto-septate conidia (Tsui et al. 2001a, Hughes 1978, Kirk
1985).
506
Hemicorynespora M.B. Ellis, Mycol. Pap. 131: 19 (1972)
Index Fungorum number: IF8499; 13 morphological species (Species Fungorum 2020).
Type species – Hemicorynespora deightonii M.B. Ellis
Notes – Most Hemicorynespora species are saprobes which have been collected from dead
leaves, branches, rachides or petioles, from tropical localities (Ellis 1972, Mercado et al. 1997,
Delgado et al. 2007, Ma et al. 2012, 2016b). The genus is characterized by macronematous,
mononematous conidiophores, determinate or percurrently elongated conidiogenous cells and,
acrogenous, 0–1-septate, conidia, seceding schizolytically (Ellis 1972, Seifert et al. 2011, Ma et al.
2016b).
Infundibulomyces Plaingam, Somrith. & E.B.G. Jones, Can. J. Bot. 81(7): 732 (2003)
Index Fungorum number: IF28753; 2 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Infundibulomyces cupulata Plaingam, Somrith. & E.B.G. Jones
Notes – The coelomycete genus Infundibulomyces is known only from the tropics and are
considered as saprobes (Plaingam et al. 2003, Somrithipol et al. 2008). Species of this genus are
characterised by stromatic, cupulate to infundibuliform conidiomata, irregularly branched
conidiophores, cylindrical conidiogenous cells and holoblastic cylindrical conidia, bearing two
filiform appendages at each end (Plaingam et al. 2003, Somrithipol et al. 2008).
Kionochaeta P.M. Kirk & B. Sutton, Trans. Br. mycol. Soc. 85(4): 712 (1986)
Index Fungorum number: IF11111; 12 morphological species (Species Fungorum 2020), 5
species with sequence data.
Type species – Kionochaeta ramifera (Matsush.) P.M. Kirk & B. Sutton
Notes – Species of Kionochaeta occur on decaying leaves and twigs in freshwater and
terrestrial habitats (Crous et al. 1994b, Kuthubutheen et al. 1988, Goh & Hyde 1997, Yanna &
Hyde 2002). Kionochaeta species are characterised by macronematous, mononematous, solitary
conidiophores, densely clustered, monophialidic, ampulliform, and hyaline to pale yellow conidia
aggregated into a slimy drop (Kirk & Sutton 1985, Goh & Hyde 1997, Okada et al. 1997).
Lecythothecium Réblová & Winka, Mycologia 93(3): 481 (2001)
Index Fungorum number: IF28485; 1 species with sequence data.
Type species – Lecythothecium duriligni Réblová & Winka
Notes – Lecythothecium is a monotypic genus typified by L. duriligni, which was collected
from decaying wood of Quercus sp. (Réblová & Winka 2001). Ths genus is characterised by
immersed, flask-shaped ascomata, 8-spored, cylindrical asci with a refractive, J-, apical ring and
ellipsoidal to fusiform transversely septate, versicolorous ascospores (Réblová & Winka 2001). The
asexual morph is hyphomycetous with macronematous, mononematous, unbranched conidiophores,
proliferating percurrently and holoblastic, cylindrical to clavate or obclavate, 5–11-pseudoseptate,
dark brown to reddish brown conidia (Réblová & Winka 2001).
Menispora Pers., Mycol. eur. (Erlanga) 1: 32 (1822)
Index Fungorum number: IF8902; 19 morphological species (Species Fungorum 2020), 6
species with sequence data.
Type specie – Menispora glauca (Link) Pers.
Notes – Menispora is a hyphomycete genus established by Persoon (1822) based on M.
glauca. Menispora species occur on decayed wood, the inner side of bark or on decayed leaves
(Hughes & Kendrick 1968, Holubová-Jechová 1973b, Réblová & Seifert 2008). Menispora is
characterised by macronematous, brown conidiophores, phialides in a lateral or terminal position
with a tapering, strongly recurved apex and a incospicous collarette and hyaline conidia without or
with polar setulae (Réblová & Seifert 2008).
507
Menisporopsis S. Hughes, Mycol. Pap. 48: 59 (1952)
Index Fungorum number: IF8904; 11 morphological species (Species Fungorum 2020), 3
species with sequence data.
Type species – Menisporopsis theobromae S. Hughes
Notes– Species of this hyphomycetous genus occur on decaying wood and leaf litter (Hughes
& Kendrick 1968, Ruiz et al. 2001). The genus is characterised by synnematous conidiophores with
a central, simple, subulate, tall seta, phialidic conidiogenous cells and appendaged conidia
aggregated into a slimy mass (Ellis 1971, Tsui et al. 1999).
Miyoshiella Kawam., Jap. J. Bot. 4: 295 (1929)
Index Fungorum number: IF3228; 3 morphological species (Species Fungorum 2020).
Type species – Miyoshiella fusispora (Kawam.) Kawam.
Notes – Miyoshiella species are saprobes on decayed wood (Réblová 1999c; Kawamura et al.
1929). They are wide spread in temperate Europe and Asia (Réblová 1999c). Species of this genus
are characterised by astromatic, papillate ascomata, asci with refractive, J-, apical ring and 3-septate
hyaline ascospores (Réblová 1999c). The asexual morph is hyphomycetous with semimacronematous conidiophores, monoblastic, ampulliform conidiogenous cells and holoblastic,
septate, cylindrical to clavate, medium to dark brown conidia (Réblová 1999c).
Morrisiella Saikia & A.K. Sarbhoy, Mycologia 77(2): 318 (1985)
Index Fungorum number: IF11132; 1 species with sequence data.
Type species – Morrisiella indica Saikia & A.K. Sarbhoy
Notes – Saikia & Sarbhoy (1985) introduced the monotypic genus Morrisiella based on M.
indica. Species of this genus are characterised by dark brown conidiophores, monoblastic, discrete,
ampulliform conidiogenous cells, lateral or terminal on the conidiophore, and obclavate, golden
brown to olivaceous brown, multipseudoseptate conidia (Saikia & Sarbhoy 1985). Shenoy et al.
(2010) confirmed phylogenetic placement of Paliphora in Chaetosphaeriaceae based on LSU
sequence data and this was accepted by Seifert et al. (2011). Similar results were obtained by Lin et
al. (2019) and is followed in this paper.
Nawawia Marvanová, Trans. Br. mycol. Soc. 75(2): 227 (1980)
Index Fungorum number: IF9081; 5 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Nawawia filiformis (Nawawi) Marvanová
Notes – Nawawia species occur on dead plant parts including stems, wood and leaves in
freshwater habitats (Hyde et al. 1996, Goh et al. 2014, Peng et al. 2016, Yang et al. 2018a). Yang et
al. (2018a) has recollected and provided sequence data for the generic type species, Nawawia
filiformis. Their phylogenetic analysis confirmed the placement of Nawawia within
Chaetosphaeriaceae. Nawawia malaysiana was the only Nawawia species with sequence data
(Crous et al. 2009). However, Yang et al. (2018a) observed that N. malaysiana is unrelated with
Nawawia, and introduced new genus Neonawawia, to accommodate N. malaysiana,
Chaetosphaeriales genera incertae sedis. Nawawia is characterised by brown, macronematous
conidiophores, monophialidic or sometimes with annellidic conidiogenous cells and turbinatetetrahedral to obpyramidal conidia with the blunt corners at the distal end with a distinct hair-like
appendage at each end (Hyde et al. 1996, Peng et al. 2016, Yang et al. 2018a).
Neopseudolachnella A. Hashim. & Kaz. Tanaka, Mycologia 107(2): 385 (2015)
Index Fungorum number: IF808687; 3 species with sequence data.
Type species – Neopseudolachnella acutispora A. Hashim. & Kaz. Tanaka
Notes – The coelomycetous genus Neopseudolachnella was established to accommodate
species with acervuloid, setose conidiomata lacking a peridium, when compared to Pseudolachnea
and Pseudolachnella species (Hashimoto et al. 2015b). Neopseudolachnella species have septate
508
conidia bearing a single to multiple appendages at each end (Hashimoto et al. 2015b).
Neopseudolachnella species are associated with Poaceae hosts in Japan (Hashimoto et al. 2015b).
Paliphora Sivan. & B. Sutton, Trans. Br. mycol. Soc. 85(2): 249 (1985)
Index Fungorum number: IF11148; 7 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Paliphora aurea Sivan. & B. Sutton
Notes – Sivanesan & Sutton (1985) introduced this genus based on P. aurea. Paliphora is
characterised by setiform conidiophores, integrated, determinate, polytretic, intercalary or terminal
conidiogenous cells and, unicellular or euseptate, cylindrical to subfusiform or subacerose, hyaline,
conidia, accumulated in slimy masses (Sivanesan & Sutton 1985, Malosso et al. 2018).
Phialosporostilbe Mercado & J. Mena, Revta Jardín bot. Nac., Univ. Habana 6(3): 57 (1985)
Index Fungorum number: IF11162; 5 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Phialosporostilbe turbinata Mercado & J. Mena
Notes – Mercado et al. (1985) introduced the hyphomycetous genus Phialosporostilbe based
on P. turbinata, which was isolated from dead stems of Arthrostylidium in Cuba. The placement of
Phialosporostilbe within Chaetosphaeriaceae was confirmed by DNA molecular data by Yang et al.
(2018a). The genus is characterized by synnematous conidiomata, monophialidic conidiogenous
cells and subhyaline, turbinate or cordiform conidia with three apical setulae (Sierra & Portales
1985, Yang et al. 2018a).
Polynema Lév., Annls Sci. Nat., Bot., sér. 3 5: 274 (1846)
Index Fungorum number: IF25008; 9 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Polynema ornata (De Not.) Lév.
Notes – The coelomycetes genus Polynema was introduced by Léveillé (1846). The genus
was redescribed by Sutton (1968) and accepted Neobarclaya as a synonym of Polynema. Nag Raj
(1978) identified Neobarclaya is distinct from Polynema and segregated it from Polynema. Crous
et al. (2018d) confirmed the placement of Polynema in Chaetosphaeriaceae based on phylogenetic
analysis of ITS sequence data. Members of Polynema are mainly saprobes associated with plant
leaves, culms and dead wood (Nag Raj 1993, Crous et al. 2018d). The genus is characterized by
acervuloid, cupulate, setose conidiomata, hyaline conidiophores, discrete or integrated, cylindrical
to clavate, hyaline conidiogenous cells and fusoid to subcylindrical, aseptate or 3 septate, mostly
hyaline conidia with attenuated or filiform appendages at each end (Nag Raj 1993, Crous et al.
2018d).
Pseudodinemasporium A. Hashim. & Kaz. Tanaka, Mycologia 107(2): 390 (2015)
Index Fungorum number: IF808707; 1 species with sequence data.
Type species – Pseudodinemasporium fabiforme A. Hashim., G. Sato & Kaz. Tanaka
Notes – The monotypic, coelomycetous genus Pseudodinemasporium was introduced to
include P. fabiform, which occurs on dead twigs of Betula platyphylla (Hashimoto et al. 2015b).
The genus is characterised by stromatic, acervular, setose conidiomata, phialidic conidiogenous
cells and aseptate conidia, bearing an appendage at each end (Hashimoto et al. 2015b).
Pseudolachnea Ranoj., Annls mycol. 8(3): 393 (1910)
Index Fungorum number: IF9587; 7 morphological species (Species Fungorum 2020), 2
species with sequence data.
Type species – Pseudolachnea insignis Velen.
Notes – The coelomycetous genus Pseudolachnea occur on twigs or wood of various
angiosperms (Ranojević 1910, Sutton 1980, Nag Raj 1993, Hashimoto et al. 2015b). The genus is
509
characterised by stromatic, acervular, setose conidiomata, phialidic conidiogenous cells and oneseptate conidia, bearing a filiform appendage at each end.
Pseudolachnella Teng, Sinensia, Shanghai 7: 775 (1936)
Index Fungorum number: IF9588; 17 morphological species (Species Fungorum 2020), 12
species with sequence data.
Type species – Pseudolachnella scolecospora (Teng & C.I. Chen) Teng
Notes – Pseudolachnella species mostly occur on bamboo, while some species occur on
twigs or wood of various angiosperms (Zhao et al. 2004, Sato et al. 2008, Hashimoto et al. 2015b).
The genus is characterized by stromatic, acervular conidiomata with marginal setae, phialidic
conidogenous cells and multiseptate conidia with single to multiple appendages at each end
(Hashimoto et al. 2015b).
Pyrigemmula D. Magyar & Shoemaker, Mycol. Progr. 10(3): 310 (2011)
Index Fungorum number: IF517148; 1 species with sequence data.
Type species – Pyrigemmula aurantiaca D. Magyar & Shoemaker
Notes – Pyrigemmula is a monotypic genus based on hyphomycete P. aurantiaca (Magyar et
al. 2011) and was isolated from inner bark or bark fissures of grapevines and also from other
woody hosts (Magyar et al. 2011). Even though P. aurantiaca occurs on fresh plant material, the
ecological role of the species is not understood (Magyar et al. 2011). Pyrigemmula aurantiaca is
characterised by pear-shaped, brown conidiogenous cells and distoseptate, reddish brown conidia
without appendages (Magyar et al. 2011).
Rattania Prabhug. & Bhat, Mycotaxon 108: 218 (2009)
Index Fungorum number: IF512876; 1 species with sequence data.
Type species – Rattania setulifera Prabhug. & Bhat
Notes – The monotypic genus Rattania was introduced based on an endophytic hyphomycete
R. setulifera isolated from fresh leaves of Calamus thwaitesii (Prabhugaonkar & Bhat 2009). The
genus is characterised by sporodochial, setose conidiomata, monoblastic conidiogenous cells and
0–multiseptate, setulate conidia (Prabhugaonkar & Bhat 2009).
Sporoschisma Berk. & Broome, Gard. Chron., London: 540 (1847)
Index Fungorum number: IF10042; 21 morphological species (Species Fungorum 2020), 8
species with sequence data.
Type species – Sporoschisma mirabile Berk. & Broome
Notes– The hyphomycetous genus Sporoschisma has cosmopolitan distribution (Goh et al.
1997, Ho et al. 2001, 2002, Zelski et al. 2014, Luo et al. 2016, Yang et al. 2016a). Species of the
genus mostly occur on submerged wood in freshwater habitats, while some species have been
collected from rotting leaves (Hughes 1966, Goh et al. 1997, Ho et al. 2001, 2002, Zelski et al.
2014, Luo et al. 2016, Yang et al. 2016a). Sporoschisma species are characterized by cylindrical,
stipitate conidiophores, with a swollen venter and a long, cylindrical neck, monophialidic
conidiogenous cells and cylindrical conidia occurring in endogenously chains with basipetal
succession (Zelski et al. 2014, Luo et al. 2016).
Striatosphaeria Samuels & E. Müll., Sydowia 31(1–6): 131 (1979)
Index Fungorum number: IF5282; 1 species with sequence data.
Type species – Striatosphaeria codinaeaphora Samuels & E. Müll.
Notes – The monotypic genus Striatosphaeria was based on lignicolous fungus S.
codinaeaphora which was isolated from bark of unidentified dead tree, in Brazil (Samuels &
Müller (1978). The genus is characterised by ostiolate ascomata seated on a small, basal stroma, 4–
8-spored, cylindrical asci and 1-septate, brown, ascospores without appendages.
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Tainosphaeria F.A. Fernández & Huhndorf, Fungal Divers. 18: 44 (2005)
Index Fungorum number: IF28947; 5 species with sequence data.
Type species – Tainosphaeria crassiparies F.A. Fernández & Huhndorf
Notes – This genus is typified by T. crassiparies, which was isolated from a Hymenaea seed
pod (Fernández & Huhndorf 2005). Species of this genus are saprobes and recorded from both
terrestrial and freshwater habitats (Fernández & Huhndorf 2005, Liu et al. 2016, Lu et al. 2016).
Species of Tainosphaeria are characterized by subglobose to ovoid ascomata, cylindrical,
pedicellate asci, with a J-, apical ring, and narrow-fusiform, septate ascospores (Fernández &
Huhndorf 2005). The asexual morph is hyphomycetous, with mononematous, unbranched
conidiophores, phyalidic conidiogenous cells with a collarette and ellipsoidal to clavate, or falcate,
hyaline conidia (Fernández & Huhndorf 2005, Liu et al. 2016, Lu et al. 2016).
Thozetella Kuntze, Revis. gen. pl. (Leipzig) 2: 873 (1891)
Index Fungorum number: IF10214; 26 morphological species (Species Fungorum 2020), 17
species with sequence data.
Type species – Thozetella nivea (Berk.) Kuntze
Notes– Most Thozetella species have been reported from decaying plant material and soil in
tropical habitats, while some species were reported in temperate habitats (Silva & Grandi 2013).
Species of this genus are characterised by sporodochial or synnematous conidiomata, phialidic
conidiogenous cells, and aseptate conidia, with unbranched setula at each end and sterile
microawns (Sutton & Cole 1983, Paulus et al. 2004, Jeewon et al. 2009).
Umbrinosphaeria Réblová, Mycotaxon 71: 17 (1999)
Index Fungorum number: IF28309; 1 species with sequence data.
Type species – Umbrinosphaeria caesariata (Clinton & Peck) Réblová
Notes – The monotypic, sexual morph genus Umbrinosphaeria was introduced to
accommodate the saprobic taxon U. caesariata, which was collected on decaying wood of Fagus
sylvatica (Réblová 1999c, Réblová & Winka 2001). Species of this genus are characterised by
papillate, setose ascomata, 8-spored, cylindrical asci with refractive, J-, apical ring and multiseptate, fusiform, versicoloured ascospores (Réblová 1999c, Réblová & Winka 2001).
Verhulstia Hern.-Restr., Persoonia 39: 449 (2017)
Index Fungorum number: IF823032; 1 species with sequence data.
Type species – Verhulstia trisororum Hern.-Restr.
Notes – Crous et al. (2017a) introduced monotypic genus Verhulstia based on V. trisororum.
This genus is charactrised by sporodochial, setose conidiomata, pale brown conidiophores,
phialidic, integrated, terminal, lageniform to subcylindrical, conidiogenous cells and, aseptate,
cylindrical to ellipsoidal, or obovoid, hyaline conidia Crous et al. (2017a).
Zanclospora S. Hughes &W.B. Kendr., N.Z. J Bot. 3: 151 (1965)
Index Fungorum number: IF10468; 9 morphological species (Species Fungorum 2020), 3
species with sequence data.
Type species – Zanclospora novae-zelandiae S. Hughes & W.B. Kendr.
Notes – Species of this genus are saprobes on plant parts such as dead wood and leaves
(Almeida et al. 2013, Hughes & Kendrick 1965). Zanclospora species are characterized by sterile
setae, setiform conidiophores, with a verruculose, dark brown apex, setiform conidiophores and
bacilliform, aseptate conidia (Almeida et al. 2013).
Clavicipitaceae Earle, Contr. U.S. natnl. Herb. 6: 170 (1901)
Index Fungorum number: IF82061; Facesoffungi number: FoF01313; 536 species.
Obligate saprotrophs, parasites and symbionts of insects, fungi, grasses, nematodes, rushes
or sedges. Sexual morph: Stromata or subiculum darkly or brightly coloured, fleshy or tough.
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Ascomata perithecial, superficial to completely immersed, ordinal or oblique in arrangement. Asci
cylindrical with thickened apex. Ascospores usually cylindrical and multiseptate, disarticulating
into part-spores or non-disarticulating. Asexual morph: Aschersonia, Ephelis, Metarhiziopsis,
Metarhizium, Neotyphodium, Nomuraea, Pochonia, paecilomyces-like, Rotiferophthora, Sphacelia,
and verticillium-like.
Type genus – Claviceps Tul.
Notes – As the former Hypocreaceae subfamily, the name Clavicipitaceae was recognized by
Earle (1901) without any description. Nannfeldt (1932) and Eriksson (1982) accepted the name, but
no one validated it (Eriksson 1982). Rogerson (1970) affirmed the name and then Eriksson &
Hawksworth (1985) validated it. Rogers (1979) removed it and classified it as two distinct orders,
Clavicipitales and Hyporeales. Clavicipitaceae was clearly placed under Hypocreales by Spatafora
& Blackwell (1993), based on SSU and LSU analyses and this was confirmed by Rehner &
Samuels (1995). Diehl (1950) proposed three subfamilies (Oomycetoideae, Clavicipitoideae, and
Cordycipitoideae) under Clavicipitaceae, but Sung et al. (2007) did not accept this. Sung et al.
(2007) divided Clavicipitaceae into three monophyletic groups (Clavicipitaceae, Cordycipitaceae
and Ophiocordycipitaceae) based on phylogenetic analyses.
Clavicipitaceae has been treated differently by many authors. Rogerson (1970) listed 13
genera, while White et al. (2000) made a clear definition for six genera based on morphology and
phylogenetic analyses. Nigrocornus was introduced to accommodate a species of Balansia which
was significantly different from that of the type and all other Balansia species (Ryley 2003). Sung
et al. (2007) listed 22 genera in Clavicipitaceae and also listed Berkelella, Cavimalum, Dussiella,
Epicrea, Helminthascus, Konradia, Moelleriella, Mycomalus, Neobarya, Neocordyceps,
Podocrella, Romanoa, Sphaerocordyceps, and Stereocrea as genera incertae sedis. Kirk et al.
(2008) recorded 43 genera in this family within Hypocreales. Chaverri et al. (2008) introduced
Samuelsia in Clavicipitaceae based on morphology and phylogenetic analyses. Ustilaginoidea was
introduced by Brefeld (1985) and revised by Tanaka et al. (2008) as a genus in Clavicipitaceae.
Conoideocrella and Orbiocrella were described by Johnson et al. (2009). Chamaeleomyces isolated
from the liver of Chamaeleo calyptratus was introduced as a new monotypic genus based on
morphology and phylogenetic analyses (Sigler et al. 2010). Periglandula was introduced with P.
ipomoeae as the type species isolated from Ipomoea asarifolia in Ecuador (Steiner et al. 2011).
Lumbsch & Huhndorf (2010) listed 32 genera under Clavicipitaceae. According to phylogenetic
analyses, Kepler et al. (2012) added Tyrannicordyceps to Clavicipitaceae which lives on sclerotia
of Claviceps (Kepler et al. 2012). Collarina was introduced by Crous et al. (2014b) based on
morphology and phylogenetic analyses. Maharachchikumbura et al. (2015) listed 46 genera in this
family: Aciculosporium, Albomyces, Aschersonia, Atkinsonella, Balansia, Cavimalum,
Chamaeleomyces, Claviceps, Conoideocrella, Corallocytostroma, Dussiella, Ephelis, Epichloë,
Epicrea, Helminthascus, Heteroepichloë, Hypocrella, Konradia, Loculistroma, Metacordyceps,
Metarhiziopsis, Metarhizium, Metapochonia, Moelleriella, Mycomalus, Myriogenospora,
Neobarya, Neoclaviceps, Neocordyceps, Neotyphodium, Nigrocornus, Nomuraea, Orbiocrella,
Parepichloë, Periglandula, Pochonia, Pseudomeria, Regiocrella, Romanoa, Samuelsia,
Shimizuomyces, Sphacelia, Sphaerocordyceps, Stereocrea, Tyrannicordyceps and Ustilaginoidea.
Chamaeleomyces was synonymized under Metarhizium by Kepler et al. (2014) and Neotyphodium
was also synonymised under Epichloe (Leuchtmann et al. 2014). Sphacelia was combined into
Claviceps (Rossman et al. 2016). Two new genera Helicocollum and Nigelia were introduced under
this family (Luangsa-ard et al. 2017a, b). Neoclaviceps and Cepsiclava were combined into
Aciculosporium (Píchová et al. 2018). Although no formal transfer had been made, molecular
analyses by Zare & Gams (2001) and Kepler et al. (2014) showed that Rotiferophthora belongs in
Clavicipitaceae and not Cordycipitaceae. Linearistroma, Metacordyceps, Nomuraea and Stereocrea
were transferred to Metarhizium (White and Glenn (1994, Kepler et al. 2014). The family is
therefore in need of revision and the asexual and sexual morph names need resolving. In this entry,
Aschersonia insperata (asexual morph) and Atkinsonella hypoxylon (sexual morh) are illustrated.
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Ecological and economic significance of Clavicipitaceae
The genera Aschersonia, Balansia, Claviceps, Dothichloe and Hypocrella were earlier
considered economically important as most of their species infect grasses, and are harmful to cattle
and other animals (e.g. insects) and cause ergot toxicity (Tulasne 1853, Atkinson 1905, Seaver
1920, Springer & Clardy 1980). Most species of Claviceps produce ergots and often produce
beneficial drugs (Molitoris 1994, Kren & Cvak 2003). Indole-diterpenes and ergot alkaloids
isolated from Claviceps cynodontis cause tremors in cattle (Uhlig et al. 2009). The secondary
metabolites of Claviceps, the ergot alkaloids, have been widely researched in the chemical industry
because of their toxicity (Hulvová et al. 2013). Some species are pathogenic on economic plants
(Hulvová et al. 2013).
Genera included in Clavicipitaceae
Aciculosporium I. Miyake, Bot. Mag., Tokyo 22: (307) (1908)
Index Fungorum number: IF41; 4 morphological species (Species Fungorum 2020), 3 species
with sequence data.
Type species – Aciculosporium take I. Miyake
Notes – This genus contains bamboo specific endophytic pathogens (Oguchi 2001, Tanaka et
al. 2003). The species produce multi-septate ascospores with no obvious part spores (Oguchi 2001).
Neoclaviceps and Cepsiclava were combined under Aciculosporium based on phylogenetic and
evolutionary analyses (Píchová et al. 2018).
Aschersonia Mont., Annls Sci. Nat., Bot., sér. 3 10: 121 (1848)
Index Fungorum number: IF7236; 175 morphological species (Species Fungorum 2020), 12
species with sequence data.
Type species – Aschersonia tahitensis Mont.
Notes – The genus comprises insect parasitic fungi. The sexual morphs are hypocrella-like
(Mongkolsamrit et al. 2009). Some species may control insect pests (Meekes et al. 2002, Wei et al.
2016). Rossman et al. (2016) recommended protecting Hypocrella over Aschersonia although
Aschersonia is an earlier name. However, the name Aschersonia was wildly used and many
molecular data were provided to the species of this genus. To respect the earlier name Aschersonia,
we suggest preserving Aschersonia over Hypocrella. Species infect scale insect and whiteflies and
produce bright, pulvinate or discoid stromata or conidiomata, superficial or embedded perithecia,
cylindrical asci, non-disarticulating ascospores, and fusiform conidia with acute ends (Chaverri et
al. 2008). In this entry we illustrate Aschersonia insperata.
Atkinsonella Diehl, Agriculture Monogr., US Dept Agric. 4: 48 (1950)
Index Fungorum number: IF449; 2 species with sequence data.
Type species – Atkinsonella hypoxylon (Peck) Diehl
Notes – This grass pathogen was separated from Epichloe based on morphology, producing
both micro- and macroconidia (Diehl 1950) and is characterised by stromata immersed and
cylindrical spores (Ambrose et al. 2014). The asexual morph is acremonium-like (Ginns 2011). In
this entry, Atkinsonella hypoxylon is illustrated. Atkinsonella hypoxylon was described as Epichloe
hypoxylon from infected grass (Peck 1875). Diehl (1950) transferred it to Atkinsonella hypoxylon
based on morphology. Ginns (2011) accepted Balansia hypoxylon as the correct name but this was
not confirmed by Ambrose et al. (2014). The specimen (DAOM 211269) was identified as Balansia
hypoxylon because of its perithecia and mature ascospores (Fig. 71, Ginns 2011).
Balansia Speg., Anal. Soc. cient.argent. 19(1): 45 (1885)
Index Fungorum number: IF497; 35 morphological species (Species Fungorum 2020), 14
species with sequence data.
Type species – Balansia claviceps Speg.
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Figure 70 – Aschersonia insperata (Material examined – Philippines, on larva of unidentified
homopteran on leaf of Maoutia setosa Wedd. (Urticaceae), 6 October 1998, M.C. Rombach, CUP
67438, paratype). a Herbarium material. b Overview of the stroma and host. c Vertical section of
the stroma. d Stroma e Conidiogenous cells, phialides. f Hyphal bodies. g, h Conidia. Scale bars: d
= 50 µm, e, f = 20 µm, g, h = 5 µm.
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Figure 71 – Atkinsonella hypoxylon (Material examined – Canada, on living and dead stems of
current year’s growth of Danthonia spicata, 9 October 1983, J. Ginns, DAOM 211269). a
Herbarium material. b-d Overview of the stromata and host. e Vertical section of the ascomata. f
Stromata. g-i Immature to mature asci. j, k Secondly ascospores. Scale bars: c, d = 1 mm, e, f = 500
µm, g-i = 50 µm, j, k = 20 µm.
Notes – Most species are grass-associated (Sung et al. 2007). The asexual morph is linked to
Ephelis (Glenn et al. 1996). Species are characterised by brown to black, superficial or immersed
stromata, with or without stipes (James & White 1994).
Cavimalum Y oshim. Doi, Dargan & K.S. Thind, Bull. natn. Sci. Mus., Tokyo, B 3(1): 23 (1977)
Index Fungorum number: IF861; 2 morphological species (Species Fungorum 2020).
Type species – Cavimalum indicum Yoshim. Doi, Dargan & K.S. Thind
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Notes – The genus comprises two poorly known species which infect bamboo and produce
immersed stromata, and irregularly septate ascospores (Doi et al. 1977).
Claviceps Tul., Annls Sci. Nat., Bot., sér. 3 20: 43 (1853)
Index Fungorum number: IF1092; 63 morphological species (Species Fungorum 2020), 40
species with sequence data.
Type species – Claviceps purpurea (Fr.) Tul.
Notes – Doubts concerning the taxonomic status of this genus were resolved in recent years
with molecular phylogenies and Claviceps was confirmed as the type genus of Clavicipitaceae
(Píchová & Parbery 1999, Yokoyama et al. 2006, Sung et al. 2007, Tanaka et al. 2008). Claviceps
species can infect about 600 species of monocotyledonous plants (Hulvová et al. 2013) and the
asexual morph, growing on honey dew, is reported in Sphacelia. The type of Sphacelia, S. segetum,
has long been regarded as the asexual morph of C. purpurea (Tulasne 1853). The genus infects
grasses and cereal crops and produces multi-colored sclerotia with 1-celled, hyaline, ephelidial
conidia and stipitate and capitulum stromata with partially embedded perithecia (White et al. 2003).
Collarina Giraldo, Gené & Guarro, Persoonia, Mol. Phyl. Evol. Fungi 33: 271 (2014)
Index Fungorum number: IF809407; 1 species with sequence data.
Type species – Collarina aurantiaca Giraldo, Gené & Guarro
Notes – This genus was isolated from soil and is characterised by conidiophores with simple,
shorter phialides and funnel-shaped collarettes, setae and large and brownish conidia (Crous et al.
2014b).
Conoideocrella D. Johnson, G.H. Sung, Hywel-Jones & Spatafora, Mycol. Res. 113(3): 286 (2009)
Index Fungorum number: IF512028; 3 morphological species (Species Fungorum 2020),
2species with sequence data.
Type species – Conoideocrella luteorostrata (Zimm.) D. Johnson, G.H. Sung, Hywel-Jones
& Spatafora
Notes – The genus was introduced for two entomophagous species from Torrubiella based on
phylogenetic analyses (Johnson et al. 2009). Mongkolsamrit et al. (2016) introduced a third species,
Conoideocrella krungchingensis. The genus is characterised by pulvinate to planar stromata and
elongated, conical-shaped perithecia, while the hosts are whiteflies and scale insects (Johnson et al.
2009).
Corallocytostroma Y.N. Yu & Z.Y. Zhang, Acta microbiol sin. 20(3): 232 (1980)
Index Fungorum number: IF7773; 2 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Corallocytostroma oryzae Y.N. Yu & Z.Y. Zhang
Notes – The genus was introduced with one species from Yunnan Province, China. Shivas et
al. (1997) described C. ornithocopreoides from Australia. Both species are plant pathogens. The
genus is characterised by coralloid conidiomata with convoluted locules and holoblastic conidia
(Shivas et al. 1997).
Dussiella Pat., Bull. Soc. mycol. Fr. 6: 107 (1890)
Index Fungorum number: IF1724; 3 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Dussiella tuberiformis (Berk. & Ravenel) Pat.
Notes – Two species were added to this monotypic genus by Rick (1906) and Höhnel (1907a)
respectively. The type species was considered as entomopathogenic, occurring on scale insects, and
host jumping onto close plant fungi species (Kepler et al. 2012). The genus is characterised by
superficial, subcylindrical perithecia, cylindrical asci with refractive apices, guttulate until the
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ascospores develop, multi-septate ascospores, breaking into part-spores and 1-septate, lunate
conidia, swelling in water (Patouillard 1890, Höhnel 1907a, Kepler et al. 2012).
Ephelis Fr., Summa veg. Scand., Section Post. (Stockholm): 370 (1849)
Index Fungorum number: IF8185; 4 morphological species (Species Fungorum 2020), 2
species with sequence data.
Type species – Ephelis mexicana Fr.
Notes – The sexual morph has been linked to Atkinsonella, Balansia, Epichloe and
Myriogenospora (White et al. 2003). The genus infects grasses, and produces holoblastic conidia in
a slender, filiform palisade layer of conidiogenous cells (White et al. 2003, Hernández-Restrepo et
al. 2016b).
Epichloe (Fr.) Tul. & C. Tul., Select. fung. carpol. (Paris) 3: 24 (1865)
Index Fungorum number: IF622348; 45 morphological species (Species Fungorum 2020), 32
species with sequence data.
Type species – Epichloe typhina (Pers.) Brockmใ
Notes – Epichloe species are endophytes which form symbiotic associations with cool-season
grasses (Poaceae subfamily Pooideae) and produce yellow to orange ascomata (White 1994, Scott
2001, White & Reddy 1998). The asexual morph has been linked to Ephelis (White et al. 2003).
The genus produces bright stromata without stipes, with numerous perithecia and yellow to orange
conidiomata and conidia on the surface of leaves (Leuchtmann et al. 2014).
Epicrea Petr., Sydowia 4(1-6): 325 (1950)
Index Fungorum number: IF1857; 1 morphological species.
Type species – Epicrea insignis Petr.
Notes – This monotypic genus is parasitic on Hypocrella chusqueae in Ecuador and is
characterised by dense, ovoid to ellipsoid perithecia and cylindrical asci (Petrak 1950).
Helicocollum Luangsa-ard, Mongkols., Noisrip. & Thanakitp., Mycol Prog 16(4): 424 (2017)
Index Fungorum number: IF817483; 3 species with sequence data.
Type species – Helicocollum surathaniense Luangsa-ard, Mongkols., Noisrip. & Thanakitp.
Notes – This genus is closely related to Aschersonia, Moelleriella, Regiocrella, and
Samuelsia, all pathogens of scale insects and whiteflies in Clavicipitaceae (Luangsa-ard et al.
2017a). Helicocollum differs from other scale insect pathogens in having phialides with helical
necks and synnematous or sporodochial pycnidial conidiomata. The morphological features of its
phialides resemble Hirsutella in the Ophiocordycipitaceae (Luangsa-ard et al. 2017a).
Helminthascus Tranzschel, Trudy S. Petersb. Obschch. Est. Otd. Bot. 28: 331 (1898)
Index Fungorum number: IF2258; 1 morphological species.
Type species – Helminthascus arachnophthorus Tranzschel
Notes – This genus comprises a single entomogenous species (Saccardo 1902, Popov et al.
2012) and is characterised by discoid stroma, immersed perithecia, cylindrical asci and septate
ascospores (Saccardo 1902).
Heteroepichloe E. Tanaka, C. Tanaka, Gafur & Tsuda, Mycoscience 43(2): 92 (2002)
Index Fungorum number: IF622360; 2 species with sequence data
Type species – Heteroepichloe bambusae (Pat.) E. Tanaka, C. Tanaka, Gafur & Tsuda
Notes – Based on phylogenetic analyses, the genus was established to accommodate two
species transferred from Epichloe. The asexual morph is ephelis-like and infects bamboo and
produces black stromata, embedded perithecia and cylindrical asci with septate ascospores (Tanaka
et al. 2002).
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Konradia Racib., Parasit. Alg. Pilze Java's (Jakarta) 2: 15 (1900)
Index Fungorum number: IF2581; 2 morphological species (Species Fungorum 2020).
Type species – Konradia bambusina Racib
Notes – The genus comprises two plant pathogenic species. No asexual morph has been
reported. The genus infects bamboo and producing black stromata, immersed, globose perithecia,
clavate asci and filiform and yellowish ascospores, becoming brown to dark brown after breaking
into part-spores (Saccardo 1902).
Loculistroma F. Patt. & Charles, Bull. Bureau Plant Industry U.S. Dep. Agric. 171: 11 (1910)
Index Fungorum number: IF2913; 1 morphological species.
Type species – Loculistroma bambusae F. Patt., Charles & Veihmeyer
Notes – The genus was introduced for a single plant pathogenic species with a cladosporiumlike asexual morph (White et al. 2003). It infects bamboo and produces green to black, cylindrical,
subsessile stromata, subglobose subimmersed perithecia, clavate to cylindrical asci and fusiform,
olivaceous, 3-septate ascospores (Patterson et al. 1910).
Metapochonia Kepler, Rehner & Humber, Mycologia 106(4): 820 (2014)
Index Fungorum number: IF806070; 9 morphological species (Species Fungorum 2020), 7
species with sequence data.
Type species – Metapochonia suchlasporia (W. Gams & Dackman) Kepler, Rehner &
Humber
Notes – This genus was introduced to accommodate five species. Zhang et al. (2017c)
described a new cave-inhabiting species. No sexual morph has been identified (Kepler et al. 2014).
The genus infects nematodes and rotifers and has been isolated from soil and plant roots and
produces conidia on slender, awl-shaped phialides (Kepler et al. 2014).
Metarhiziopsis D.W. Li, R.S. Cowles & C.R. Vossbrinck, Mycologia 100(3): 462 (2008)
Index Fungorum number: IF511393; 1 species with sequence data.
Type species – Metarhiziopsis microspora D.W. Li, R.S. Cowles & C.R. Vossbrinck
Notes – Metarhiziopsis was introduced as a monotypic entomopathogenic genus based on
morphology and phylogenetic analyses (Li et al. 2008). No sexual morph has been reported. It
infects insects producing white to buff, cupulate sporodochia, surrounded by differentiated septate
setae, cylindrical, unicellular phialides and unicellular, catenulate conidia (Li et al. 2008).
Metarhizium Sorokīn, Veg. Parasitenk. Mensch Tieren 2: 268 (1879)
Index Fungorum number: IF8912; 47 morphological species (Species Fungorum 2020), 36
species with sequence data.
Type species – Metarhizium anisopliae (Metschn.) Sorokīn
Notes – This genus was emended by Rombach et al. (1987) and Kepler et al. (2014). It
includes green-spored, Nomuraea (N. rileyi), several species formerly included in Paecilomyces
and Chameoleomyces. The sexual morphs were described as metacordyceps-like and most species
in Metacordyceps have been transferred to Metarhizium (Kepler et al. 2014), including the type
species. The genus infects arthropods and nematodes, can be found in the soil, colonizes plant roots
and act as endophytes, producing variously branched conidiophores without synnemata and hyaline
to brown or green conidia (Kepler et al. 2014). The sexual morph is characterised by tough, whitish
greenish yellow to greenish, cylindrical to enlarging stromata, partially or completely immersed
perithecia and ascospores which may or may not break into part-spores (Sung et al. 2007).
Moelleriella Bres., Boll. Soc. bot. ital. 44: 292 (1897)
Index Fungorum number: IF3231; 39 morphological species (Species Fungorum 2020), 17
species with sequence data.
Type species – Moelleriella sulphurea (Bres.) Bres.
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Notes – This genus is entomopathogenic (Chaverri et al. 2008). The asexual morphs are
aschersonia-like (Chaverri et al. 2008). It infects scale insects and whiteflies and produces bright
coloured, pulvinate or discoid stromata or conidiomata, superficial or embedded perithecia,
cylindrical asci, filiform multiseptate ascospores that disarticulate at maturity inside the ascus and
fusiform conidia (Chaverri et al. 2008).
Mycomalus Möller, Bot. Mitt. Trop. 9: 300 (1901)
Index Fungorum number: IF3324; 1 morphological species.
Type species – Mycomalus bambusinus Möller
Notes – This monotypic genus was introduced as a plant pathogen . It infects bamboo and
produces white, subglobose stromata, immersed, obpyriform perithecia and ascospores which
disarticulate into fusiform part-spores (Möller 1901).
Mycophilomyces Crous & M.J. Wingf., Persoonia 37: 325 (2016)
Index Fungorum number: IF819075; 1 species with sequence data.
Type species – Mycophilomyces periconiae Crous & M.J. Wingf.
Notes – Mycophilomyces was introduced with a single species, hyperparasite on Periconia on
leaves of Albizia falcataria. It has multiseptate conidiophores and obclavate solitary conidia (Crous
et al. 2016a). No sexual morph is known.
Myriogenospora G.F. Atk., Bull. Torrey bot. Club 21(5): 225 (1894)
Index Fungorum number: IF3371; 4 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Myriogenospora paspali G.F. Atk.
Notes – Myriogenospora atramentosa causes “tangle top disease” of mang grasses (White et
al. 2003). The asexual morph is ephelis-like (White & Glenn 1994). The genus infects grasses
producing linear, hard, black stromata, immersed, subglobose perithecia and ascospores which
disarticulate into cylindrical part-spores (White and Glenn 1994).
Neobarya Lowen, Syst. Ascom. 5(1): 121 (1986)
Index Fungorum number: IF25587; 11 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Neobarya parasitica (Fuckel) Lowen
Notes – The asexual morphs are acremonium-like, lecanicillium-like, paecilomyces-like,
calcarisporium-like or torrubiella-like (Candoussau et al. 2007). This genus was reported as hostspecific parasites of lichens and non lichenized fungi (Lawrey et al. 2015). It produces soft, bright
coloured stromata, sessile, superficial perithecia and filiform, aseptate or septate, entire ascospores
(Candoussau et al. 2007).
Neocordyceps Kobayasi, J. Jap. Bot. 59(6): 187 (1984)
Index Fungorum number: IF25653; 1 morphological species.
Type species – Neocordyceps kohyasanensis Kobayasi
Notes – This genus was introduced as a subgenus of Cordyceps with a single plant pathogenic
species. It has a hymenostilbe-like asexual morph (White et al. 2003). The genus is characterised by
clavate, stipitate stromata with an elongate fertile part, pyriform to ovoid, purple to black,
immersed perithecia and fusiform, septate, entire ascospores (Kobayasi 1981, White et al. 2003).
Nigelia Luangsa-ard, Tasan. & Thanakitp., i Mycol. Progr. 16(4): 378 (2017)
Index Fungorum number: IF823565; 2 species with sequence data
Type species – Nigelia aurantiaca Luangsa-ard, Thanakitp. & Tasan.
Notes – Nigelia was introduced as an invertebrate-pathogenic genus based on morphology
and phylogenetic analyses. The genus is characterised by yellow to reddish brown, cylindrical,
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stipitate, stromata with a cylindrical, clavate to irregularly shapedfertile part, ovoid with curved,
immersed perithecia and filiform, septate, whole ascospores or fragmenting into part spores
(Luangsa-ard et al. 2017b). The asexual morph is characterised by verticillately or irregularly
branched conidiophores and solitary conidiogenous cells or phialides with or without lateral necks
(Luangsa-ard et al. 2017b).
Nigrocornus Ryley & Langdon, Mycology Series (New York) 19: 266 (2003)
Index Fungorum number: IF28732; 1 morphological species.
Type species – Nigrocornus scleroticus (Pat.) Ryley
Notes – The genus was established to accommodate a single plant pathogenic species
transferred from Balansia. The genus is characterised by black, corniform, sessile stromata,
lageniform to obpyriform, immersed perithecia and filiform, 7-septate ascospores, ultimately
becoming 3-septate (White et al. 2003). The asexual morph bears simple, indeterminate
conidiophores and holoblastic, obclavate, multiguttulate, aseptate conidia (White et al. 2003).
Orbiocrella D. Johnson, G.H. Sung, Hywel-Jones & Spatafora, Mycol. Res. 113(3): 286 (2009)
Index Fungorum number: IF512031; 1 species with sequence data
Type species – Orbiocrella petchii (Hywel-Jones) D. Johnson, G.H. Sung, Hywel-Jones &
Spatafora
Notes – This entomophagous genus contains a single species transferred from Torrubiella
(Johnson et al. 2009). The asexual morph is simplicillium-like (Johnson et al. 2009). The genus is
characterised by ring-like, sessile stroma, superficial, flask-shaped perithecia and filiform,
multiseptate ascospores, not disarticulating into part-spores (Johnson et al. 2009).
Parepichloe J.F. White & P.V. Reddy, Mycologia 90(2): 231 (1998)
Index Fungorum number: IF622359; 4 morphological species (Species Fungorum 2020), 2
species with sequence data.
Type species – Parepichloe cinerea (Berk. & Broome) J.F. White & P.V. Reddy
Notes – The genus was established to accommodate epibiotic species transferred from
Epichloe. No asexual morph is known (White et al. 2003). The species infect grasses and produce
cylindrical to fusiform, gray to black, sessile stromata, immersed, ovoid perithecia and filiform,
multiseptate ascospores, not disarticulating into part-spores (White & Reddy 1998).
Periglandula U. Steiner, E. Leistner & Leuchtm., Mycologia 103(5): 1137 (2011)
Index Fungorum number: IF561039; 2 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Periglandula ipomoeae U. Steiner, E. Leistner & Schardl
Notes – The genus forms a symbiosis with plants (Steiner et al. 2011). It produces epibiotic
mycelium, chlamydospore-like and synnema-like structures, but conidia have not been observed
(Steiner et al. 2011).
Pochonia Bat. & O.M. Fonseca, Publicações Inst. Micol. Recife 462: 4 (1965)
Index Fungorum number: IF9479; 3 morphological species (Species Fungorum 2020), 2
species with sequence data.
Type species – Pochonia humicola Bat. & O.M. Fonseca
Notes – The sexual morph of this genus is metacordyceps-like (Zare & Gams 2007). The
genus infects nematodes and produces mostly prostrate, rarely erect conidiophores, verticillate or
solitary, aculeate phialides, subglobose, ellipsoidal to rod-shaped conidia and
dictyochlamydospores (Batista & Fonsêca 1965).
Pseudomeria G.L. Barron, Can. J. Bot. 58(4): 443 (1980)
Index Fungorum number: IF9589; 1 morphological species.
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Type species – Pseudomeria mucosa G.L. Barron
Notes – This monotypic genus is a parasite of rotifers (Barron 1980). The sexual morph is
undetermined. The genus is characterised by unbranched, septate conidiophores and globose
conidia with a conspicuous basal slime pad (Barron 1980).
Regiocrella P. Chaverri & K.T. Hodge, Mycologia 97(6): 1232 (2006)
Index Fungorum number: IF500786; 2 species with sequence data.
Type species – Regiocrella camerunensis P. Chaverri & H.C. Evans
Notes – This entomophagous genus has a sphacelia-like asexual morph (Chaverri et al. 2005).
It infects scale insects and produces pale orange, sessile stromata, semi-immersed, obpyriform
perithecia, cylindrical, not capitate asci and unicellular, fusiform ascospores (Chaverri et al. 2005).
Romanoa Thirum., R.C. Ist. Sup. Sanitä, (Rome) 17: 1326 (1954)
Index Fungorum number: IF4779; 1 species with sequence data.
Type species – Romanoa terricola Thirum.
Notes – The genus was isolated from soil.
Rotiferophthora G.L. Barron, Can. J. Bot. 69(3): 495 (1991)
Index Fungorum number: IF25566; 26 morphological species (Species Fungorum 2020), 2
species with sequence data.
Type species – Rotiferophthora globospora G.L. Barron
Notes – The genus was established by to accommodate diheterospora-like endoparasites of
rotifers (Barron 1991). There is no sexual morph linked to this genus. The genus infects rotifers and
produces phialides with short conidiiferous necks and dictyochlamydospores (Barron 1991).
Samuelsia P. Chaverri & K.T. Hodge, Stud. Mycol. 60: 59 (2008)
Index Fungorum number: IF511385; 6 morphological species (Species Fungorum 2020), 3
species with sequence data.
Type species – Samuelsia rufobrunnea P. Chaverri & K.T. Hodge
Notes – This entomophagous genus contains five species based on morphology and
phylogenetic analyses. The asexual morph is aschersonia-like (Chaverri et al. 2008).
Mongkolsamrit et al. (2011) described a new species, Samuelsia mundiveteris. The genus infects
scale insects and whiteflies and produces bright coloured, pulvinate or discoid stromata or
conidiomata, superficial or embedded perithecia, cylindrical asci, long-fusiform, non-disarticulating
ascospores and small allantoid conidia (Chaverri et al. 2008).
Shimizuomyces Kobayasi, Bull. natn. Sci. Mus., Tokyo, B 7(1): 1 (1981)
Index Fungorum number: IF5024; 2 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Shimizuomyces paradoxus Kobayasi
Notes – This is a plant pathogenic genus. Sung et al. (2007, 2010) provided molecular data
and described the culture characteristics of Shimizuomyces paradoxus. The genus has no asexual
morph. It is characterised by fleshy, stipitate stromata, clavate to cylindrical stroma, immersed
perithecia and fusiform, septate ascospores (Kobayasi 1981).
Sphaerocordyceps Kobayasi, Bull. natn. Sci. Mus., Tokyo, B 7(1): 2 (1981)
Index Fungorum number: IF5093; 2 morphological species (Species Fungorum 2020).
Type species – Sphaerocordyceps palustris (Berk. & Broome) Kobayasi
Notes – This genus was segregated from Cordyceps. There is no known asexual morph. The
genus infects insects and is characterised by brownish-purple or flesh-colour, stipitate stromata, and
globose perithecia (Kobayashi 1981).
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Tyrannicordyceps Kepler & Spatafora, Index Fungorum 12: 1 (2012)
Index Fungorum number: IF550079; 5 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Tyrannicordyceps fratricida (Tanda & Kobayasi) Kepler & Spatafora
Notes – This fungicolous genus was introduced for five species transferred from Cordyceps
based on morphology and phylogenetic analyses. The asexual morph is verticillium-like (Kepler et
al. 2012). It infects both plants and insects and produces pallid, yellow or sometimes vermilion
stromata and filiform, septate ascospores which may break into part-spores (Kepler et al. 2012).
Ustilaginoidea Bref., Unters. Gesammtgeb. Mykol. (Liepzig) 12: 194 (1895)
Index Fungorum number: IF10364; 15 morphological species (Tanaka et al. 2008), 3 species
with sequence data.
Type species – Ustilaginoidea oryzae (Pat.) Bref.
Notes – This is a plant pathogenic genus. Tanaka et al. (2008) reported the holomorph and
provided clear illustrations of the common rice disease caused by Ustilaginoidea virens . The genus
infects grasses and produces dark, hard sclerotia and dark green or brown, subglobose conidia
(White et al. 2003).
Clypeophysalosporaceae A. Giraldo & Crous, Mycol. Progr. 16(4): 340 (2017)
Index Fungorum number: IF818514; Facesoffungi number: FoF06867; 33 species.
Endophytic, saprobic or pathogenic mostly on leaves of Eucalyptus spp. Sexual morph:
Ascomata perithecial, brown or black, immersed, solitary, rarely in pairs, globose, sometimes with
clypeus or pseudoclypeus. Peridium of several layers of brown and flattened cells. Paraphyses
septate, numerous, hyaline, hypha-like, flexuose. Asci 8-spored, unitunicate, cylindrical or
obclavate, hyaline, with apical ring staining in Melzer’s reagent. Ascospores uniseriate or biseriate,
hyaline, fusoid-ellipsoidal, unicellular, commonly surrounded by a mucilaginous sheath. Asexual
morph: Coelomycetous or hyphomycetous. Conidiomata pycnidial, globose, solitary to aggregated,
brown. Conidiophores differentiated or reduced to conidiogenous cells (pycnidial form), forming
fascicules or rosettes on leaves, or solitary on the hyphae (hyphomycetous form), cylindrical to
subcylindrical, brown to pale grey-brown. Conidiogenous cells phialidic, ampulliform, lageniform
or subcylindrical, terminal and intercalary, pale to medium brown, apex with flared collarette.
Conidia solitary or in slimy heads, subcylindrical, curved, hyaline, obtuse apex and truncate to
subtruncate base (adapted from Giraldo & Crous 2017a).
Type genus – Clypeophysalospora H.J. Swart
Notes – Clypeophysalosporaceae was introduced by Giraldo & Crous (2017a) to
accommodate Bagadiella, Clypeophysalospora, Neophysalospora and Plectosphaerella. These
genera share characteristics in terms of both sexual and asexual morphs, host specificity (mainly
Eucalyptus spp.) and distribution (Australia, South Africa, Giraldo & Crous 2017a).
Neophysalospora eucalypti has also been isolated from Corymbia henryi (Myrtaceae) (Swart 1981,
Cheewangkoon et al. 2009, Crous et al. 2011, 2014b).
Ecological and economic significance of Clypeophysalosporaceae
Members in this family usually grow on living or dead leaves of various plants, and are
mostly endophytes, saprobes or pathogens (Swart et al. 1981, Cheewangkoon 2009).
Genera included in Clypeophysalosporaceae
Bagadiella Cheew. & Crous, Persoonia 23: 59 (2009)
Index Fungorum number: IF513840; 4 species with sequence data.
Type species – Bagadiella lunata Cheewangkoon & Crous
Notes – Cheewangkoon et al. (2009) introduced this genus with Bagadiella lunata as the type
species and there are four epithets recorded under Bagadiella (Cheewangkoon et al. 2009, Crous et
al. 2011). All species have been reported with asexual morphs and B. koalae is coelomycetous.
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Bagadiella koalae was reported from leaves of Eucalyptus globulus eaten by koala and B. victoriae
was reported from leaves of Eucalyptus sp. in Australia (Crous et al. 2011). Bagadiella is
characterised by caespituli pale, suprastomatal, pseudoparenchyma with a substomatal cavity
froming a rosette of conidiophores. Conidiophores are micronematous, cylindrical and
dichotomously branched. Conidiogenous cells are terminal, monophialidic, branched,
subcylindrical to lageniform and conidia are hyaline, lunate, curved, with a round apices, tapering
towards a subtruncate base and borne in slimy heads.
Clypeophysalospora H.J. Swart, Trans. Br. mycol. Soc. 76(1): 93 (1981)
Index Fungorum number: IF1121; 1 species with sequence data.
Type species – Clypeophysalospora latitans (Sacc.) H.J. Swart
Notes – Kang et al. (1999c) considered Clypeosphaeria to be similar with
Clypeophysalospora, and C. latitans was accommodated in Clypeosphaeriaceae. Senanayake et al.
(2015) used LSU and ITS sequence data of C. uniseptata to consider the affinity of this family with
Amphisphaeriales. The concept of Clypeosphaeriaceae (based on C. uniseptata; Senanayake et al.
2015) was not followed in Jaklitsch et al. (2016a). Giraldo et al. (2017a) included the sequence of
C. uniseptata in a LSU analysis and this species is not related at the family level with
Clypeophysalospora latitans, and therefore they introduced a new family to accommodate the latter
taxon.
Neophysalospora Crous & M.J. Wingf., Persoonia 33: 247 (2014)
Index Fungorum number: IF810608; 1 species with sequence data.
Type species – Neophysalospora eucalypti Crous & M.J. Wingf.
Notes – Crous et al. (2014b) introduced Neophysalospora eucalypti from leaves of Corymbia
henryi (Myrtaceae) in Mozambique. This species has endophytic and plant pathogenic lifestyles.
Neophysalospora is a holomorphic genus. Asci are unitunicate with apical J+ rings and asexual
morphs comprise conidiophores lining the inner conidiomatal wall, which are straight to curved,
hyaline to pale brown.
Plectosphaera Theiss., Annls mycol. 14(6): 413 (1917)
Index Fungorum number: IF4196; 27 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Plectosphaera eucalypti (Cooke & Massee) H.J. Swart
Notes – Cooke (1888) introduced Plectosphaera eucalypti from leaves of Eucalyptus in
Australia. Summerell (2006) cultured this fungus for the first time and provided sequence data.
Colonies on MEA, comprise flat, smooth, catenate, submerged spreading mycelium, with surface
cinnamon, smoke grey due to black ascomata forming from outer region of the colony.
Clypeosphaeriaceae G. Winter, Rabenh. Krypt.-Fl., Edn 2 (Leipzig) 1.2: 554 (1886)
Index Fungorum number: IF80613; Facesoffungi number: FoF01776; 53 species.
Hemibiotrophic or saprobic on woody or herbaceous plants. Sexual morph: Pseudoclypeus
comprising both host and fungal tissues, black. Ascomata immersed to erumpent, rarely superficial,
solitary or aggregated, globose to subglobose, coriaceous, brown to black, ostiolate, papillate.
Papilla short, narrow, internally lined with hyaline, filamentous periphyses. Peridium comprising
dark brown to light brown, thick-walled cells of textura angularis, inwardly hyaline. Paraphyses
numerous, hypha-like, septate, flexuose, embedded in a gelatinous matrix. Asci 8-spored,
unitunicate, cylindrical to broadly cylindrical, pedicellate, with a wedge-shaped, J-, or J+, apical
ring. Ascospores uniseriate to biseriate, hyaline to brown, ellipsoidal to fusiform, sometimes oval,
straight or curved, unicellular or septate, wall smooth or ornamented or striate, sometimes with
sheaths, appendages, rarely with germ slits or germ pores. Asexual morph: Undetermined.
Type genus – Clypeosphaeria Fuckel
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Figure 72 – Clypeophysalospora latitans (redrawn from Swart et al. 1981). a, b Ascomata on host
substrate. c Vertical section of ascoma. d Peridium. e, f Asci. g-j Ascospores. Scale bars: e, j = 10
µm.
Notes – Lindau (1897) resurrected Sphaeriales to accommodate Clypeosphaeriaceae which
has been accommodated in Xylariales (Krug 1978, Barr 1990b, 1994, Eriksson & Hawksworth
1993, Eriksson & Winka 1997, Hawksworth et al. 1995, Smith et al. 2003, Douanla-Meli & Langer
2012, Hernandez-Restrepo et al. 2015, Maharachchikumbura et al. 2015, 2016b). Senanayake et al.
(2015) excluded Clypeosphaeriaceae from Xylariales and transferred it to Amphisphaeriales.
Jaklitsch et al. (2016b) did not accept Amphisphaeriales because it lacked phylogenetic support in
their analysis. Furthermore, in their analysis, the generic type of Clypeosphaeria; C. mamillana
clusters as a basal clade within Xylariaceae. Therefore, they discontinued Clypeosphaeriaceae and
synonymised under Xylariaceae. However, in present study (Fig. 4) the family type; C. mamillana
form a distinct clade apart from Xylariaceae as a sister taxon to the Induratiaceae. Therefore, here
we maintain Clypeosphaeriaceae as a distinct family in Xylariales. However, most genera placed in
this family lack molecular data and the arrangement is reliant on ascospores being apiosporous, asci
having a J-, or J+, aprical ring and immersed ascomata.
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Ecological and economic significance of Clypeosphaeriaceae
Most taxa in Clypeosphaeriaceae have been reported as saprobes on wood worldwide (Kang
et al. 1999c, Catania & Romero 2012, Senanayake et al. 2015, Niranjan & Sarma 2018) or as
endophytic on Diospyros crassiflora (Ebenaceae fide Douanla-Meli & Langer 2012).
Genera included in Clypeosphaeriaceae
Aquasphaeria K.D. Hyde, Nova Hedwigia 61(1–2): 122 (1995)
Index Fungorum number: IF27561; 1 morphological species.
Type species – Aquasphaeria dimorphospora K.D. Hyde
Notes – Aquasphaeria was introduced as a freshwater species on submerged wood from
Queensland (Hyde 1995d). It was placed in Clypeosphaeriaceae based on morphology (Hyde
1995d). Aquasphaeria dimorphospora is illustrated in this entry. Hyde (1995d) introduced a new
genus Aquasphaeria with A. dimorphospora as the type species. Munk (1957) assumed this genus
to Lasiosphaeriaceae. Hyde (1995d) placed it to Clypeosphaeriaceae but Maharachchikumbura et
al. (2016b) excluded Aquasphaeria from Clypeosphaeriaceae and transferred it to Sordariomycetes,
genera incertae sedis. In this study, we include Aquasphaeria into Clypeosphaeriaceae base on
ascomata being immersed under a clypeus. Although this genus has been reported as ubiquitous
(Australia, Brunei, Seychelles, Malaysia, Thailand fide Hyde 1995d, Ho et al. 2001, Sivichai et al.
2002), there have been no recent collections.
Apioclypea K.D. Hyde, J. Linn. Soc., Bot. 116(4): 316 (1994)
Index Fungorum number: IF27441; 6 morphological species (Species Fungorum 2020),
molecular data available for 3 unnamed species in the genus.
Type species – Apioclypea livistonae K.D. Hyde
Notes – The genus was introduced by Hyde et al. (1994b) with A. livistonae as the type
species, as a saprobe on the dead rachis of Livistona (Arecaceae) in Papua New Guinea. Two
further species were introduced by Taylor & Hyde (2003). The genus is characterized by solitary,
globose or subglobose ascomata, immersed under a clypeus, with a J-, subapical, ascal ring and
hyaline apiospores with a mucilaginous sheath (Hyde et al. 1994b, Kang et al. 1999c, Taylor &
Hyde 2003).
Brunneiapiospora K.D. Hyde, J. Fröhl. & Joanne E. Taylor, Sydowia 50(1): 40 (1998)
Index Fungorum number: IF27879; 9 morphological species (Species Fungorum 2020),
molecular data available for an unnamed species.
Type species – Brunneiapiospora javensis K.D. Hyde, J. Fröhl. & Joanne E. Taylor
Notes – Nine species have been introduced to this genus including a recent species, B.
appendiculata from India (Hyde et al. 1998a, Crous et al. 2012a, Vitoria et al. 2012, Niranjan &
Sarma 2018). Brunneiapiospora species are characterized by solitary, subglobose ascomata,
immersed beneath a darkened clypeus and with a central ostiole, a J+, or J-, discoid, subapical,
ascal ring and hyaline to light brown apiospores with a mucilaginous sheath (Hyde et al. 1998a,
Kang et al. 1999c).
Clypeosphaeria Fuckel, Jb. nassau. Ver. Naturk. 23-24: 117 (1870)
Index Fungorum number: IF1125; 33 morphological species (Species Fungorum 2020), 3
species with sequence data.
Type species – Clypeosphaeria mamillana (Fr.) Lambotte
Notes – Clypeosphaeria is characterized by immersed to erumpent, dark pigmented ascomata,
cylindrical to broadly cylindrical asci, with J-, or J+, apical ring and ellipsoidal to fusiform,
unicellular to septate, smooth-walled to ornamented or striate, hyaline to dark brown ascospores,
sometimes with sheaths or appendages (Saccardo 1883c, Kang et al. 1999c). In this entry we
illustrate Clypeosphaeria mamillana.
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Figure 73 – Aquasphaeria dimorphospora (Material examined – AUSTRALIA, North Queensland,
on submerged wood, October 1991, K.D. Hyde, BRIP21482, holotype). a Material. b, c Ascomata
on host. d Close up of ascoma. e Section of ascoma. f Cell of peridium. g Neck. h Cylindrical asci.
i, j Immature ovoid ascus with paraphyses. k Mature ovoid asci. l Paraphyses. m J-, apical ring in
ascus. n-p Cylindrical ascospores. q, r Immature ovoid ascospores s Mature ovoid ascospores. Scale
bars: b-d = 500 μm, e = 200 μm, f, h-m = 50 μm, g = 100 μm, n-s = 20 μm.
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Figure 74 – Clypeosphaeria mamillana (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 Peridium. f Ascus apex with a J+,
subapical ring. g-h Asci. i-k Ascospores. Scale bars: c = 500 μm, d = 50 μm, e =10 μm, f, i‒k = 5
μm, g‒h = 20 μm.
Crassoascus Checa, Barrasa & A.T. Martínez, Mycotaxon 46: 300 (1993)
Index Fungorum number: IF26311; 3 morphological species (Species Fungorum 2020).
527
Type species – Crassoascus fusisporus Checa, Barrasa & A.T. Martínez
Notes – Crassoascus was introduced to accommodate C. fusisporus occurring on dead
branches of Erica arborea in Spain (Barrasa et al. 1993). The second species, C. canadensis was
found on wood of Salix in British Columbia (Barr 1994). The latest species is C. monocaudatus
which was reported from a dead branch of Podocarpus parlatorei from Tucumán, in Argentina
(Catania & Romero 2012). Crassoascus species are characterised by gregarious, semi-immersed,
black ascomata with ostioles, a J+, subapical, ascal ring and bright brown to dark brown, multiseptate, fusiform ascospores, with hyaline refractive cap-like appendages at each end (Barrasa et al.
1993, Kang et al. 1999c).
Palmaria K.D. Hyde, J. Fröhl. & Joanne E. Taylor, Fungal Divers. 86: 328 (2017)
Index Fungorum number: IF553238; 1 morphological species.
Type species – Palmaria montaneus (K.D. Hyde, J. Fröhl. & Joanne E. Taylor) K.D. Hyde, J.
Fröhl. & Joanne E. Taylor
Notes – Palmaria was introduced as Palmomyces by Hyde et al. (1998a) with P. montaneus
occurring on a leaf of Oraniopsis appendiculata in Queensland. The genus was recorded associated
with leaf spots on palms (Arecaceae) and only the sexual morph is known. Palmaria is
characterised by solitary, immersed, subglobose ascomata, cylindric-clavate asci with a J-,
subapical ring and hyaline, 1-septate, obclavate apiospores, with a mucilaginous sheath (Hyde et al.
1998a).
Cocoonihabitaceae W.Y. Zhuang & Z.Q. Zeng, Mycosystema 36(12): 1597 (2017)
Index Fungorum number: IF570496; Facesoffungi number: FoF06759; 1 species.
Cocoon‐inhabiting. Sexual morph: Ascomata superficial, solitary to aggregated, each seated
on host directly or on a small basal stroma, perithecial, coriaceous, pale orange to orange, not
changing colour in 100% lactic acid and 3% KOH. Peridium usually composed of 3 layers. Asci
cylindrical, with a thick cap at the apex penetrated by a narrow pore, short pedicellate. Ascospores
filiform, hyaline, multiseptate. Asexual morph: Undetermined (adapted from Zhuang & Zeng
2017).
Type genus – Cocoonihabitus W.Y. Zhuang & Z.Q. Zeng
Notes – Cocoonihabitaceae was introduced as a result of the separate branching of two strains
of a species, sister to species of Ophiocordycipitaceae and Flammocladiellaceae in a multi-gene
phylogram (Zhuang & Zeng 2017). The species has long cylindrical asci with thick cap at the apex,
penetrated by a narrow pore, characters which also exist in species belonging to Clavicipitaceae,
Cordycipitaceae and Ophiocordycipitaceae. However, the characteristics of the ascomata, absence
of stroma, ascospore features and habitats separated this new taxon from the other taxa. Therefore,
Cocoonihabitaceae was introduced (Zhuang & Zeng 2017).
Ecological and economic significance of Cocoonihabitaceae
Studies may likely unlock the significance of taxa accommodated in this family as has been
reported for those belonging to families such as Cordycipitaceae and Ophiocordycipitaceae (Yu et
al. 2004, Zhang et al. 2012b).
Genus included in Cocoonihabitaceae
Cocoonihabitus W.Y. Zhuang & Z.Q. Zeng, Mycosystema 36(12): 1597 (2017)
Index Fungorum number: IF570495; 1 species with sequence data.
Type species – Cocoonihabitus sinensis W.Y. Zhuang & Z.Q. Zeng
Notes – This monotypic genus was erected since the newly acquired strains of the freshly
collected fungus, namely, Cocoonihabitus sinensis, clustered independently in the phylogenetic
tree, apart from the other genera in other families (Zhuang & Zeng 2017). Cocoonihabitus sinensis
shares morphological resemblance to taxa belonging to some genera in Clavicipitaceae and
Ophiocordycepitaceae in terms of asci and ascospores. However, the ascomata of C. sinensis are
528
astromatic and seated directly on the substrate, contrary to the ascomata of the species in the other
genera of Clavicipitaceae and Ophiocordycepitaceae, which are mostly immersed or semiimmersed (Zhuang & Zeng 2017).
Figure 75 – Cocoonihabitus sinensis (redrawn from Zhuang & Zeng 2017). a Perithecia on
remaining leaf vein of a cocoon. b Cross-section of perithecium. c Ascus containing ascospores. d,
e Disarticulated cells of ascospores. Scale bars: a = 250 µm, b, c = 20 µm, d, e = 10 µm.
Coniocessiaceae Asgari & Zare, Mycol. Progr. 10(2): 195 (2011)
Index Fungorum number: IF518425; Facesoffungi number: FoF00671; 6 species.
Saprobic on grasses, soil and dung. Sexual morph: Ascomata less than 200 μm diam,
superficial, subglobose to pyriform, glabrous or pilose, ostiolate, with hypha-like ostiole. Peridium
membranaceous, thin, translucent or sometimes opaque, outer layer comprising pale-brown to dark
brown with cells of textura intricata. Paraphyses abundant or few, septate, tapering, hyphae-like,
thin-walled, filamentous, branched. Asci 4–spored, unitunicate, cylindrical to subcylindrical,
pedicellate, with or without an apical ring. Ascospores uniseriate, dark brown to black, ellipsoidal,
with narrowly rounded ends, smooth-walled, 1-celled, with or without germ-slit, if present full
length, straight. Asexual morph: Hyphomycetous, nodulisporium-like: Conidiophores
micronematous to macronematous, simple or branched, smooth-walled or verrucose, hyaline.
Conidiogenous cells integrated, terminal, discrete, elongating sympodially, with persistent
conspicuous denticles, hyaline. Conidia globose, subglobose to pyriform, smooth-walled or
verruculose, with rounded apex, hyaline, attenuated and truncated base and distinct projection at the
point of attachment to the conidiogenous cells (adapted from Maharachchimbura et al. 2016b).
Type genus – Coniocessia Dania García, Stchigel, D. Hawksw. & Guarro
Notes – Coniocessiaceae was established by Asgari & Zare (2011) to accommodate four
species of Coniocessia (C. anandra, C. cruciformis, C. maxima, C. minima) with the species C.
nodulisporioides. The family was placed in Xylariales based on morphology and molecular
analyses, and a hyaline asexual morph with polyblastic conidiogenesis (nodulisporium-like).
Phylogenetic analysis of Coniocessia species showed a distinct lineage close to Diatrypaceae,
which confirmed it as a family in Xylariales (Asgari & Zare 2011, Maharachchikumbura et al.
2016b).
Ecological and economic significance of Coniocessiaceae
Species of Coniocessiaceae are saprobic on grasses, soil and dung.
Genera included in Coniocessiaceae
Coniocessia Dania García, Stchigel, D. Hawksw. & Guarro, Mycol. Res. 110(11): 1284 (2006)
Index Fungorum number: IF29049; 5 species with sequence data.
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Type species – Coniocessia nodulisporioides (D. Hawksw.) Dania García, Stchigel, D.
Hawksw. & Guarro
Notes – García et al. (2006) introduced Coniocessia to accommodate C. nodulisporioides, by
synonymising Coniochaeta nodulisporioides, isolated from soil in Jordan by Hawksworth (1978).
The species was synonymised under Coniocessia, based on morphology and SSU and LSU
molecular data. Asgari & Zare (2011) introduced Coniocessiaceae and four new species in
Coniocessia based on morphological and ITS and LSU molecular data, as all species clustered into
a single monophyletic clade (Asgari & Zare 2011). Maharachchikumbura et al. (2016b) provided
an updated outline of the genus with five species.
Figure 76 – Coniocessia nodulisporioides (illustration based on García et al. 2006 and Asgari &
Zare 2011). a Masses of ascospores on host surface. b Ascomata. c Paraphyses. d Asci. e
Ascospores. f, g Conidiophores and conidiogenous cells. h Conidia. Scale bars: a = 100 μm, b = 50
μm, c-h = 10 μm.
Paraxylaria Wanas., E.B.G. Jones, Gafforov & K.D. Hyde, Fungal Divers. 89(1): 202 (2018)
Index Fungorum number: IF554226; 1 species with sequence data.
Type species – Paraxylaria rosacearum Wanas., Gafforov, E.B.G. Jones & K.D. Hyde
530
Notes – Paraxylaria was introduced to accommodate P. rosacearum and is a monotypic
genus saprobic on Rosaceae in Uzbekistan (Wanasinghe et al. 2018). The genus was introduced
based on morphology similar to Coniocessia. However, species differ as Paraxylaria have asci
with J+, apical rings with ascospores which lack germ-slits but have a close affinity in phylogenetic
analysis (Wanasinghe et al. 2018).
Coniochaetaceae Malloch & Cain, Can. J. Bot. 49: 878 (1971)
Index Fungorum number: IF80629; Facesoffungi number: FoF01332; 99 species.
Saprobic on dung, plant litter or in soil, water, or pathogens of plants and
immunocompromised humans, endophytic in leaves and lichen thalli. Sexual morph: Ascomata
perithecial or cleistothecial, solitary to gregarious, superficial, semi-immersed or immersed,
subglobose to globose or pyriform, dark brown to black, glabrous or hairy, ostiolate or lacking
ostiole. Ostiole periphysate, sometimes surrounded by a crown of setae. Peridium membranaceous
to pseudoparenchymatous, rarely coriaceous; composed of several layers of cells of textura
angularis or textura intricata, or less frequently cephalothecoid. Paraphyses numerous, filiform,
simple, septate, evanescent. Asci (4-), 8- to multi-spored, unitunicate, cylindrical to fusoid or
clavate, globose to subglobose, short pedicellate, with a truncate to rounded apex, with a J-, apical
ring, evanescent. Ascospores 1-seriate or irregularly arranged, initially hyaline, becoming brown to
dark brown or olive-greenish to dark olivaceous or black at maturity, ellipsoid to fusiform, broadly
ellipsoidal to globose, lenticular or cruciform, with rounded to apiculate ends, flattened on one or
both sides; 1-celled, with or without a germ slit, smooth-walled or pitted. Asexual morph:
Hyphomycetous or yeast-like. Colonies frequently characterized by pink or orange and a yeast-like
appearance. Conidiophores macronematous or semi-macronematous. Conidiogenous cells
phialidic, polyblastic, of various size and morphology, somewhat ampulliform, subulate or
indistinguishable from a normal hyphal cell, collarettes present, but usually indistinct, occasionally
somewhat flared. Conidia accumulating near the point of formation (in chains), hyaline, orange or
pink in mass, elliptical to oblong-elliptical to reniform, 1-celled, smooth-walled.
Type genus – Coniochaeta (Sacc.) Cooke
Notes – Coniochaetaceae was introduced by Malloch & Cain (1971) and its phylogenetic
relationships were clarified by García et al. (2006). The family was accepted in the newly
introduced order Coniochaetales (Huhndorf et al. 2004b). Hongsanan et al. (2017) provided
evidence through phylogenetic and MCMC trees to support the ordinal status. Additional notes for
Coniochaetaceae and two new species were provided by Friebes et al. (2016). For a key and type
herbarium study see Maharachchikumbura et al. (2016b).
Ecological and economic significance of Coniochaetaceae
Coniochaetaceae species are mainly saprobes on dead wood. In addition, some are plant
pathogens and opportunistic fungi causing serious infections in humans (Damm et al. 2010). They
are also producers of potent antibiotics and some strains were evaluated to have a high potential of
biological detoxification of lignocellulosic biomass (Segeth et al. 2003, López et al. 2004, Friebes
et al. 2016).
Genera included in Coniochaetaceae
Barrina A.W. Ramaley, Mycologia 89(6): 962 (1997)
Index Fungorum number: IF27745; 1 species with sequence data.
Type species – Barrina polyspora A.W. Ramaley
Notes – Ramaley (1997) introduced this monotypic genus to accommodate B. polyspora
which has ellipsoid-fusiform, aseptate, hyaline, smooth-walled, thin-walled ascospores and
ascoconidia. Based on its phialophora-like asexual morph, the genus is accepted in
Coniochaetaceae (Ramaley 1997, Kirk et al. 2013, Maharachchikumbura et al. 2015). However,
based on available sequences, this genus clusters in this family.
531
Coniochaeta (Sacc.) Cooke, Grevillea 16 (no. 77): 16 (1887)
Index Fungorum number: IF1209; 98 morphological species (Species Fungorum 2020), 55
species with sequence data.
Type species – Coniochaeta ligniaria (Grev.) Cooke
Figure 77 – Coniochaeta taeniospora (Material examined – ITALY, Province of Forlì-Cesena
[FC], Camposonaldo - Santa Sofia, on dead land branch of Quercus sp. (Fagaceae), 13 March 2017,
Erio Camporesi IT3275, MFLU 17-0832, HKAS 102311). a, b Stroma on the host. c Vertical
section of ascoma. d Cross section of ascoma. e Ostiole in section. f Paraphyses. g, h Peridium (h in
5% KOH). i-k Asci. l Ascus apex in Congo Red. m-p Ascospores (black arrow heads show germ
slits). Scale bars: a = 1 mm, b, c = 500 μm, d = 200 μm, e = 100 μm, i-k = 50 μm, g, h = 20 μm, l-p
= 10 μm, f = 5 μm.
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Notes – Phialidic asexual morphs proved to be an informative character to delimit
Coniochaetaceae, while the type of ascomata, which can be cleistothecial or perithecial, and the
ornamentation of the ascospore walls, proved to have little taxonomic value at the generic level
(García et al. 2006). Based on these conclusions, García et al. (2006) segregated several species
from Coniochaeta and moved Coniocessia and Coniolariella to Xylariales. Eleven new
Coniochaeta species supported by morphology and phylogeny have been introduced (Jiang et al.
2017, Coronado-Ruiz et al. 2018, Nasr et al. 2018, Samarakoon et al. 2018, Wanasinghe et al.
2018, Crous et al. 2019a, Harrington et al. 2019, Phookamsak et al. 2019). In this entry
Coniochaeta taeniospora is illustrated (Fig. 77).
Conioscyphaceae Réblová & Seifert, Persoonia 37: 63 (2015)
Index Fungorum number: IF813227; Facesoffungi number: FoF05190; 16 species.
Saprobic on wood and animal skin. Sexual morph: Ascomata perithecial, astromatic,
immersed, semi-immersed or superficial, subhyaline to pale orange or pale brown when fresh,
almost invisible when dry, papillate or with a cylindrical neck. Peridium coriaceous, two-layered.
Ostiole periphysate. Paraphyses filiform, septate, unbranched, longer than the asci. Asci 8-spored,
unitunicate, persistent, pedicellate, cylindrical to clavate, sigmoidal, with a tall and prominent J-,
refractive apical ring. Ascospores 2–4-seriate, hyaline to subhyaline, ellipsoid to fusiform or
fusiform-navicular, transversely septate, not constricted at the septa, lacking any gelatinous sheath
or appendages. Asexual morph: Hyphomycetous. Conidiophores micronematous, mononematous,
hyaline or pigmented. Conidiogenous cells holoblastic, sympodial, cyathiform or doliiform,
surrounded by hyaline, multi-layered, with conspicuous cup-like collarettes. Conidia brown,
unicellular, non-septate and varying in shape, thick-walled, smooth, with a pore at the point of
attachment to the conidiogenous cells, formed individually and in succession, apices of
conidiogenous cells proliferate percurrently, liberated after outer wall of the conidiogenous cell
ruptures at the apex (adapted from Réblová & Seifert 2004a, Zelski et al. 2015, Réblová et al.
2016c)
Type genus – Conioscypha Höhn.
Notes – Conioscyphaceae was introduced by Réblová et al. (2016c) under Conioscyphales
and it includes the single genus Conioscypha. Conioscypha lignicola, the generic type, was
described by Höhnel (1904) and it was reviewed by Shearer (1973) who introduced a second
species C. varia Shearer. The number of species gradually increased and C. tenebrosa is the latest
addition to Conioscyphales (Liu et al. 2019b). Most of the species have been isolated as saprobes
from submerged substrates (Shearer 1973, Matsushima 1975, 1993, 1996, Crous et al. 2014a,
2018b, Zelski et al. 2015, Chuaseeharonnachai et al. 2017, Hernández-Restrepo et al. 2017, Liu et
al. 2019b, Luo et al. 2019) or decaying wood in terrestrial biotopes (Munk 1957, Réblová & Seifert
2004a). Hongsanan et al. (2017) transferred Conioscyphales from Hypocreomycetidae (Réblová et
al. 2016c) into the newly introduced subclass Savoryellomycetidae with evidence from wellsupported phylogenetic and maximum clade credibility trees with stem age of 268 MYA.
Ecological and economic significance of Conioscyphaceae
Most species of Conioscypha are saprobic on decaying plant materials and are found in both
aquatic and terrestrial habitats, with rotten wood, leaves, twigs and bamboo stems being frequent
substrates. Conioscypha japonica is the sole species isolated from cuticular scrapings and hairs of a
living animal (Goh & Hyde 1998).
Genus included in Conioscyphaceae
Conioscypha Höhn., Annls mycol. 2(1): 58 (1904)
Index Fungorum number: IF7754; 16 morphological species (Liu et al. 2019b), 11 species
with sequence data.
Type species – Conioscypha lignicola Höhn.
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Notes – Conioscypha comprises species reported from decaying wood, leaves and bamboo
from both freshwater and terrestrial habitats, except for C. japonica which was isolated from dog
skin fragments and hair (Chuaseeharonnachai et al. 2017, Liu et al. 2019b). Conioscypha is
characterized by a unique mode of conidiogenesis with blastic conidia produced at inconspicuous
loci along the hyphae (Shearer 1973). Traditionally, the conidia were thought to be produced from
‘phialidic’ conidiogenous cells (Goh & Hyde 1998). Shearer & Motta (1973) described
Conioscypha conidiogenesis to be both ‘phialidic’ and ‘annelidic’ (Shearer & Motta 1973), but
Minter et al. (1983) did not agree with this observation. Cole & Samson (1979) reported conidial
development to be intermediate between the ‘phialidic’ and ‘annelidic’ process and after repetitive
basipetal conidial secession, the remains of the outer wall of conidia collect centripetally on the
conidiogenous cells to form ‘collarettes’. Based on in vitro experiments and molecular DNA data,
Réblová & Seifert (2004a) introduced Conioscyphascus, typified by Ca. varius, to accommodate
holomorphs with Conioscypha asexual morphs. Another sexual-asexual relationship was
established for C. peruviana (Zelski et al. 2015). Following abolishment of dual nomenclature and
adoption of one fungus, one name, Conioscyphascus was accepted as a synonym of Conioscypha
(Zelski et al. 2015, Réblová et al. 2016c). Although C. gracilis is the only species of the genus
known in its sexual state, the presence of typical conidia on the host near ascomata was repeatedly
observed (Réblová & Seifert 2004a, Zelski et al. 2015). Conioscypha varia Shearer is illustrated for
the sexual morph and C. tenebrosa is illustrated for the asexual morph, in this entry.
Conlariaceae H. Zhang, K.D. Hyde & Maharachch., Fungal Divers. 85: 90 (2017)
Index Fungorum number: IF553758; Facesoffungi number: FoF03336; 7 species.
Saprobic on submerged decaying wood in freshwater or grow on soil in terrestrial habitats.
Sexual morph: Ascomata perithecioid, gregarious, coriaceous or membranous, immersed to
erumpent or superficial, dark brown to black, globose to subglobose, smooth, ostiolate. Neck short
or elongated, cylindrical, straight or slightly flexuous. Peridium composed of several layers of cells
of textura angularis. Paraphyses cylindrical or globose, hyaline, septate. Asci 8-spored,
unitunicate, cylindrical, short pedicellate, apically rounded, with a distinct, refractive, massive,
barrel-shaped, apical ring or lacking apical structures. Ascospores uni- to bi-seriate, hyaline,
fusiform or ellipsoidal-fusiform, straight or slightly curved, aseptate to multi-septate, guttulate,
thin- or thick-walled, with or without appendages or appendages at one or both ends or surrounded
by an irregular gelatinous sheath. Asexual morph: Hyphomycetous. Colonies dark brown to black.
Mycelium mostly immersed, consisting of branched, septate, thin-walled, smooth, pale brown to
brown hyphae. Conidiophores micronematous or semi-macronematous, mononematous, septate or
aseptate, unbranched or irregularly branched, straight or flexuous, hyaline, becoming brown when
old. Conidiogenous cells holoblastic, determinate, doliiform, cylindrical. Conidia brown, muriform,
irregularly globose or subglobose, septate, constricted at the septa.
Type genus – Conlarium F. Liu & L. Cai
Notes – Conlariaceae was introduced in a new order Atractosporales by Zhang et al. (2017a)
for a single genus Conlarium which comprised C. aquaticum and C. dupliciascosporum. Asexual
morphs of the family were found in the culture of C. dupliciascosporum (Liu et al. 2012), on
natural decaying submerged wood (Zhang et al. 2017a) and dead wood or soil in terrestrial habitat
(Phookamsak et al. 2019, Xie et al. 2019). Another genus Riomyces is included in the family based
on our phylogenetic analyses (Fig. 3), and previous study (Luo et al. 2019). The phylogeny and
morphology warranted a new family (Zhang et al. 2017a).
Ecological and economic significance of Conlariaceae
Conlariaceae currently comprises two genera Conlarium and Riomyces with seven species
which are saprobic on submerged wood and dead wood or soil. They have so far been reported
from China, Costa Rica and Thailand (Ferrer et al. 2012, Liu et al. 2012, Zhang et al. 2017a,
Phookamsak et al. 2019, Xie et al. 2019). They are important plant decomposers involved in
nutrient cycling.
534
Figure 78 – Conioscypha varia (Material examined – CZECH REPUBLIC, Southern Bohemia,
Šumava Mts., Modrava, Modrava, brook Zhůřský potok close to Turner’s hut, associated with
Capronia pilosella, M.R. 1890/01, on decayed wood of a deciduous tree, 27 October 2001, M.
Réblová, PRM 900537, holotype). a, b Herbarium material. c Appearance of ascomata on host
substrate. d Peridium. e Transverse section through ostiole. f Paraphyses. g, h Asci. i Evagination
of ascal ring to allow discharge of spore. j J-, apical ring. k-m Ascospores. Scale bars: c = 200 µm,
d = 10 µm, e-m = 50 µm.
535
Figure 79 – Conioscypha tenebrosa (Material examined – CHINA, Guizhou Province, Dushan, on
decaying wood on the bank of a small freshwater stream, 6 July 2018, N.G. Liu, MFLU 19-0688,
holotype). a Colonies on natural substrate. b Mass of conidia. c-e Conidia and conidiogenous cells
with cup-shaped multi-collarette. f Conidium and conidiogenous cell originating from a hypha. g-i
Conidia. Scale bars: b = 50 μm, c-g, i = 30 μm, h = 10 μm.
Genera included in Conlariaceae
Conlarium F. Liu & L. Cai, Mycologia 104(5): 1180 (2012)
Index Fungorum number: IF564382; 6 species with sequence data.
Type species – Conlarium dupliciascosporum F. Liu & L. Cai
Notes – Conlarium was established by Liu et al. (2012) for the holomorph species C.
duplumascosporum based on its distinct lineage among related taxa. Conlarium aquaticum and C.
dupliciascosporum were found from submerged wood in freshwater, C. thailandense was from
dead wood in a terrestrial habitat, C. baiseense, C. nanningense and C. sacchari were from
536
sugarcane rhizosphere (Liu et al. 2012, Zhang et al. 2017a, Phookamsak et al. 2019, Xie et al.
2019). In this entry we illustrate Conlarium aquaticum and C. duplumascospora (Figs 80, 81).
Figure 80 – Sexual morph of Conlarium duplumascospora (Material examined – CHINA,
Guangdong Province, Zhaoqing Dinghu Mountain, on submerged wood in a stream, 29 December
2010, F. Liu, HMAS243129, holotype). a Gregarious ascomata on host substratum. b-d Asci. e
Paraphyses. f Ascospores. Scale bars: a = 500 µm, b, d = 20 µm, c, e = 10 µm, f = 5 µm.
Riomyces A. Ferrer, A.N. Mill., Sarmiento & Shearer, Mycologia 104(4): 876 (2012)
Index Fungorum number: IF561099; 1 species with sequence data.
Type species – Riomyces rotundus A. Ferrer, A.N. Mill., Sarmiento & Shearer
Notes – Riomyces was introduced by Ferrer et al. (2012) to accommodate a single species R.
rotundus, based on morphology and phylogenetic analyses of LSU sequence data. It was found on
submerged decaying wood in freshwater in Costa Rica (Fig. 3).
Cordanaceae Nann., Repert. mic. uomo: 498 (1934)
Index Fungorum number: IF80640; Facesoffungi number: FoF01673; 23 species.
Saprobic or pathogenic on wood of branches, twigs and leaves of various shrubs and trees,
bamboo, grasses and in soil of terrestrial habitats, occasionally in freshwater habitats. Sexual
morph: Ascomata perithecial, superficial, solitary or gregarious, globose to ovoid, setose or
glabrous, with basal stroma, papillate. Ostiole periphysate. Peridium comprising 3–5 layers of-
537
Figure 81 – Conlarium aquaticum (Material examined – THAILAND, Prachuap Khiri Khan, on
submerged wood in a small river, 30 July 2015, W. Dong 136A, MFLU 15-2703, holotype). a, b
Colonies on submerged wood. c-h Conidia. i-k Conidia with conidiogenous cells. Scale bars: c = 50
μm, d, f-k = 20 μm, e = 10 μm.
-cells of textura prismatica. Paraphyses septate, unbranched. Asci 8-spored, unitunicate,
cylindrical, with short pedicel, with or without a J-, apical ring. Ascospores uniseriate, pale brown
538
to brown, ellipsoid to fusiform, sometimes with pores at both ends, 1-septate. Asexual morph:
Hyphomycetous. Colonies effuse, dark brown to black. Conidiophores mononematous,
macronematous, brown, septate, erect, branched or unbranched. Conidiogenous cells terminal or
intercalary, polyblastic, denticulate. Conidia pale brown to brown, obovoid to ellipsoidal, smooth,
sometimes with germ pores at the ends, 1-septate or aseptate (adapted from Hughes 1955, Müller &
Samuels 1982b, Hernández-Restrepo et al. 2014).
Type genus – Cordana Preuss
Notes – Cordanaceae was introduced by Nannfeldt (1934) based on the type genus Cordana.
Initially, Hernández-Restrepo et al. (2015b) introduced Cordanales to accommodate Cordanaceae,
which was recognized as a monotypic, monophyletic sister group to Coniochaetales
(Maharachchikumbura et al. 2016b). However, in a study backed by a molecular clock evidence,
Cordanaceae was found to share a common ancestor with Coniochaetaceae in Coniochaetales 77
MYA (Hongsanan et al. 2017). While the family lineage had good statistical support, Cordanales
did not have enough support and was therefore combined under Coniochaetales (Hongsanan et al.
2017) of which the stem age was 176 MYA and had higher support as an order. Cordanaceae
differs from the closely related Coniochaetaceae by morphology of their basally stromatic
ascomata, septate ascospores and holoblastic-denticulate conidiogenesis. (Hernández-Restrepo et
al. 2015b).
Ecological and economic significance of Cordanaceae
Cordana species are widespread in temperate and tropical regions of the world, including
Africa, South and Central America, Spain, South East Asia and New Zealand (Fernández &
Huhndorf 2004, Hernández-Restrepo et al. 2014, Zelski et al. 2014, Ai et al. 2019). Species occur
on decaying plant matter, in soil or on other fungi (Hernández-Restrepo et al. 2014). They cause
leaf spots on banana and Canna species (Ellis 1976, Markovskaja 2003, Cai et al. 2004,
Hernández-Restrepo et al. 2014). The type, C. pauciseptata is a common saprobe reported on dead
wood.
Genus included in Cordanaceae
Cordana Preuss, Linnaea 24: 129 (1851)
Index Fungorum number: IF7777; 23 morphological species (Species Fungorum 2020), 11
species with sequence data.
Type species – Cordana pauciseptata Preuss
Notes – Cordana is a dematiaceous hyphomycetous genus introduced by Preuss (1851)
together with three species, C. polyseptata, C. pauciseptata and C. pedunculata, without
designation of a type species (Hernández-Restrepo et al. 2014). Saccardo (1877) and Hughes
(1955) attempted to designate C. pauciseptata as the lectotype, however in both occasions the
species was subjected to reclassification and the type material was reported as lost (HernándezRestrepo et al. 2014). Thus, Hernández-Restrepo et al. (2014) designated an epitype for this genus,
produced a phylogram based on ITS sequences and provided a dichotomous key to the species.
Based on cultural studies, Müller & Samuels (1982b) suggested Porosphaerella cordanophora
(Fig. 82) was the sexual morph of Cordana pauciseptata, and this was confirmed phylogenetically
by Hernández-Restrepo et al. (2014). ITS sequence data for eight species and LSU for some are
available in GenBank (Réblová & Winka 2000, Huhndorf et al. 2004b, Réblová & Seifert 2007,
Hernández-Restrepo et al. 2014, Zelski et al. 2014, Vu et al. 2019).
Cordycipitaceae Kreisel, Stud. Mycol. 57: 48 (2007)
Index Fungorum number: IF504360; Facesoffungi number: FoF01314; 313 species.
Parasites or pathogens of scale insects or mosses, or saprobes in leaf litter and upper soil
layers. Sexual morph: Stromata or subiculum, fleshy, pallid, or brightly coloured. Perithecia
superficial to completely immersed, oriented at right angles to the surface of the stroma. Asci
mostly 8–spored, cylindrical, with thickened ascus apex. Ascospores usually cylindrical, multi539
septate, disarticulating into part-spores or remaining intact at maturity. Asexual morph: beauverialike, engyodontium-like, lecanicillium-like, mariannaea-like, simplicillium-like.
Figure 82 – Cordana mercadiana (FMR 11828, a-c), drawings based on illustrations given in
Hernández-Restrepo et al. (2014). a, b Arrangement of conidia on conidiophore and denticulate
conidiogenous cells giving rise to conidia (based on SEM microphotographs provided in
Hernández-Restrepo et al. 2014). c Conidia. Sexual morph Porosphaerella cordanaphora, and its
Cordana asexual morph (d-j, redrawn from Müller & Samuels 1982b). d Asci with ascospores. e
Vertical section of ascoma. f J-, apical ring of asci with a mature ascospore. g Ascospores. h
Conidiophore with conidia clustered at apex. i Apical meristem protruding in conidiophore and
developing conidia. j Conidia. Scale bars: a = 50 µm, b-j = 10 µm.
540
Type genus – Cordyceps Fr.
Notes – Cordycipitaceae was first used by Kreisel (1969), while Wehmeyer (1976) used
Cordycipitoideae as subfamily for Clavicipitaceae based on the type genus Cordyceps.
Cordycipitaceae was validly segregated from Clavicipitaceae by Sung et al. (2007), based on
morphology analyses and multi-gene phylogenetic. Most of the species in the family are
entomogenous and produce superficial to partially immersed to completely immersed perithecia, on
a fleshy stroma or subicula, that are pallid or brightly coloured (Sung et al. 2007). Sung et al.
(2007) confined 11 genera to this family, while Kepler et al. (2017) proposed maintaining nine
generic names (Akanthomyces, Ascopolyporus, Beauveria, Cordyceps, Engyodontium, Gibellula,
Hyperdermium, Parengyodontium, and Simplicillium) and rejected 8 generic names (Evlachovaea,
Granulomanus, Isaria, Lecanicillium, Microhilum, Phytocordyceps, Synsterigmatocystis, and
Torrubiella). Kepler et al. (2017) also described two new generic names (Hevansia and
Blackwellomyces). The most recent addition is Samsoniella (Mongkolsamrit et al. 2018). In
addition, Index Fungorum (2020) records two generic names (Beejasamuha and Rotiferophthora),
which were not mentioned in papers, but are included here. Isaria and Microhilum were considered
as synonyms of Cordyceps, and Granulomanus was considered as a synonym of Gibellula (Kepler
et al. 2017). The asexual morphs in this family are confused.
Ecological and economic significance of Cordycipitaceae
Many species in this family are economically important, such as Cordyceps militaris,
Beauveria bassiana and Cordyceps cicadae, because many significant metabolites have been
extracted from these species such as nucleosides (adenine, adenosine, cordycepin) and
polysaccharides (CS-F10, CSP-1) (Kiho et al. 1993, Li et al. 2001, Rukachaisirikul et al. 2004, Yu
et al. 2004, Li et al. 2006). The bioactive compounds of Cordyceps militaris are well-documented
as Chinese traditional medicine as they benefit immune (Kuo et al. 2001), anti-cancer (Kuo et al.
1994, Bok et al. 1999, De Silva et al. 2012, 2013), anti-inflammatory (Won & Park 2005) and antiplasmodium (Pan et al. 2018) responses (Hyde et al. 2019b). Cordyceps militaris is cultivated in
many countries (Park et al. 2001, Shih et al. 2007, Wen et al. 2014) and is industrially profitable.
Genera included in Cordycipitaceae
Akanthomyces Lebert, Z. Wiss. Zool. 9: 449 (1858)
Index Fungorum number: IF7083; 24 morphological species (Species Fungorum 2020), 10
species with sequence data.
Type species – Akanthomyces aculeatus Lebert
Notes – This genus was established by Lebert (1858) with a single species Akanthomyces
aculeatus found in France (Hsieh et al. 1997a). The sexual morph was confirmed as cordyceps-like
(Hodge et al. 2003, Kepler et al. 2017) and thus may be a synonym. The genus infects Lepidoptera
adults and produces white, cream or flesh-colored, cylindrical, attenuated synnemata covered with a
hymenium of phialides and unicellular, hyaline conidia, in short or long chains (Hodge et al. 2003).
Species also occur on spiders producing synnematous or mononematous isaria-like phialides and
hyaline and long chains of conidia (Mongkolsamrit et al. 2018).
Amphichorda Fr., Syst. orb. veg. (Lundae) 1: 170 (1825)
Index Fungorum number: IF7134; 2 species with molecular data.
Type species – Amphichorda felina (DC.) Fr.
Notes – Species of the genus were isolated from dung, insects and soil (Seifert et al. 2011).
The genus is characterised by indeterminate synnemata, branched conidiophores, sympodial
conidiogenous and amerosporous, single conidia (Seifert et al. 2011). No sexual morph has been
reported.
Ascopolyporus Möller, Bot. Mitt. Trop. 9: 300 (1901)
541
Index Fungorum number: IF371; 7 morphological species (Species Fungorum 2020), 3
species with sequence data.
Type species – Ascopolyporus polychrous Möller
Notes – This genus is entomopathogenic and is strongly supported as monophyletic in
phylogenetic analyses (Kepler et al. 2017). The genus infects bamboo and has polypore-like, sessile
stroma (Hodge et al. 2003).
Beauveria Vuill., Bull. Soc. bot. Fr. 59: 40 (1912)
Index Fungorum number: IF7346; 40 morphological species (Index Fungorum 2020), 25
species with sequence data.
Type species – Beauveria bassiana (Bals.-Criv.) Vuill
Notes – Beauveria was introduced by Vuillemin (1912) by transferring “Botrytis bassiana”
and “Botrytis effusa” from Botrytis to Beauveria (Vuillemin 1912). Species of this genus are found
worldwide and are parasitic on insects. The sexual morph of the species are cordyceps-like (Li et al.
2001, Rehner et al. 2011). The genus infects insects and has basally-inflated conidiogenous cells
(Li et al. 2001, Rehner et al. 2011).
Beejasamuha Subram. & Chandrash., Can. J. Bot. 55(3): 247 (1977)
Index Fungorum number: IF7350; 1 morphological species.
Type species – Beejasamuha samala Subram. & Chandrash
Notes – Beejasamuha only has one hyphomycetous species, found on goat dung in India
(Subramanian & Chandrashekara 1977). The genus is characterised by flask-shaped conidiogenous
cells and blastoconidia (Subramanian & Chandrashekara 1977).
Blackwellomyces Spatafora & Luangsa-ard, IMA Fungus 8: 345 (2017)
Index Fungorum number: IF820864; 3 morphological species (Species Fungorum 2020), 2
species with sequence data.
Type species – Blackwelliella cardinalis (G.H. Sung & Spatafora) Spatafora & Luangsa-ard
Notes – The genus was introduced to accommodate two species of Cordyceps based on
morphological and phylogenetic analyses (Kepler et al. 2017). The asexual morph are
Clonostachys, Hirsutella, Isaria, and Mariannaea species (Kepler et al. 2017). The genus infects
insects and has irregularly septate ascospores that do not disarticulate into part-spores (Kepler et al.
2017).
Cordyceps Fr., Observ. mycol. (Havniae) 2: 316 (cancellans) (1818)
Index Fungorum number: IF1240; 180 morphological species (Species Fungorum 2020), 42
species with sequence data.
Type species – Cordyceps militaris (L.) Fr.
Notes – Cordyceps was validly published by Link (1833) and given as the group name by
Fries (1818) as Cordylia (Rogers 1954). Cordyceps includes about 564 species (Index Fungorum
2015) and was formally moved to Cordycipitoideae in 2007 (Sung et al. 2007). Sung et al. (2007)
moved some species from Cordyceps to several genera including Ophiocordyceps
(Ophiocordycipitaceae), Elaphocordyceps current name Tolypocladium (Ophiocordycipitaceae),
Metacordyceps (Clavicipitaceae), Claviceps (Clavicipitaceae), Tyrannicordyceps (Clavicipitaceae),
Epichloë (Clavicipitaceae), Podostroma (Hypocreaceae) and Podocrea (Hypocreaceae). Kepler et
al. (2017) suppressed the names Isaria, Microhilum, Phytocordyceps and Evlachovaea in favour of
Cordyceps. The genus infects insects and produces bright, stipitate, fleshy stromata, superficial to
embedded perithecia, ascospores which break or do not break into part spores (Sung et al. 2007).
The asexual morph of this genus produces evlachovaea-like, isaria-like, lecanicillium-like,
mariannaea-like, simplicillium-like phialides or conidia (Sung et al. 2007). A specimen of
Cordyceps militaris collected from China which shows the complete sexual and asexual
characteristics is illustrated here.
542
Figure 83 – Cordyceps militaris (Material examined – CHINA, Province of Liao-Ning, on dead
larva. 18 June 2014, Ting-Chi Wen TL2014091004, MFLU 15-3202). a Overview of stromata and
host. b Host. c Yellow, half superficial ascomata on stroma. d Cross section showing the stroma
and ascomata. e Ascomata. f Ostioles. g, h Asci. i Cap of ascus. j Part spores. k Conidiophores and
developing conidia. l Conidia. m Culture from above on PDA medium after 40 days. n Culture
from below on PDA medium after 40 days. Scale bars: d = 1000 µm, e = 500 µm, f = 100 µm, g, h
= 200 µm, i, l = 5 µm, j, k = 10 µm, m-n = 2 cm..
543
Coremiopsis Sizova & Suprun, Vestn. Moskov. Univ., Ser. biol. 12(2): 55 (1957)
Index Fungorum number: IF7781; 2 morphological species (Species Fungorum 2020).
Type species – Coremiopsis rosea Sizova & Suprun
Notes – The genus was isolated from soil and was considered as an uncertain status by
Kendrick (1974).
Engyodontium de Hoog, Persoonia 10(1): 53 (1978)
Index Fungorum number: IF8178; 4 morphological species (Species Fungorum 2020), 2
species with sequence data.
Type species – Engyodontium parvisporum (Petch) de Hoog
Notes – Engyodontium album was transfered to Parengyodontium by Tsang et al. (2016).
There is no sexual morph link to this genus (Tsang et al. 2016). The genus infects insects and has
stipitate synnemata, monophialidic conidiogenous cells and conidia in chains or slimey masses (de
Hoog 1978, Evans & Samson 1987).
Gibellula Cavara, Atti Ist. bot. R. Univ. Pavia, 2 Sér. 3: 347 (1894)
Index Fungorum number: IF8330; 24 morphological species (Species Fungorum 2020), 5
species with sequence data.
Type species – Gibellula pulchra (Sacc.) Cavara
Notes – The genus was recognized as spider pathogens with torrubiella-like sexual morphs
(Humber & Rombach 1987, Prathumpai et al. 2012, Kuephadungphan et al. 2014). The genus
infects spiders and has stipitate synnemata, and vesiculate conidiophores (Humber & Rombach
1987, Prathumpai et al. 2012).
Hevansia Luangsa-ard, Hywel-Jones & Spatafora in IMA Fungus 8: 348 (2017)
Index Fungorum number: IF820885; 8 morphological species (Kepler et al. 2017), 5 species
with sequence data.
Type species – Hevansia novoguineensis (Samson & B.L. Brady) Luangsa-ard, Hywel-Jones
& Spatafora
Notes – The genus was established with eight species and diagnosed by immersed perithecia
and an akanthomyces-like asexual morph (Kepler et al. 2017). The genus infects spiders and has
sessile stromata, immersed perithecia and monophialidic conidiogenous cells (Kepler et al. 2017).
Hyperdermium J.F. White, R.F. Sullivan, Bills & Hywel-Jones, Mycologia 92(5): 910 (2000)
Index Fungorum number: IF28464; 2 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Hyperdermium bertonii (Speg.) J.F. White, R.F. Sullivan, Bills & HywelJones
Notes – The members of this genus are parasitic on insects and plants (Sullivan et al. 2000).
The genus is characterised by bright, sessile stromata with 1-3 perithecia per stroma (Sullivan et al.
2000).
Leptobacillium Zare & W. Gams, Mycol. Progr. 15: 1001 (2016)
Index Fungorum number: IF815489; 1 species with sequence data.
Type species – Leptobacillium leptobactrum (W. Gams) Zare & W. Gams
Notes – This genus was separated from Verticillium as a saprotrophic or fungicolous taxon
without a sexual morph reported (Zare & Gams 2016). The genus is characterised by narrow, rodshaped conidia and chlamydospores (Zare & Gams 2016).
Parengyodontium C.C. Tsang, J.F.W. Chan, W.M. Pong, J.H.K. Chen, A.H.Y. Ngan, M. Cheung,
C.K.C. Lai, D.N.C. Tsang, S.K.P. Lau & P.C.Y. Woo, Medical Mycology 54: 708 (2016)
Index Fungorum number: IF815049; 1 species with sequence data.
544
Type species – Parengyodontium album (Limber) C.C. Tsang, J.F.W. Chan, W.M. Pong,
J.H.K. Chen, A.H.Y. Ngan, M. Cheung, C.K.C. Lai, D.N.C. Tsang, S.K.P. Lau & P.C.Y. Woo
Notes – The genus was emended to accommodate one species from Engyodontium and it
infects humans and produces numerous whorls of conidiogenous cells (Tsang et al. 2016).
Pseudogibellula Samson & H.C. Evans, Acta bot. neerl. 22(5): 524 (1973)
Index Fungorum number: IF9578; 1 morphological species.
Type species – Pseudogibellula formicarum (Mains) Samson & H.C. Evans
Notes – Pseudogibellula was introduced with a sing species, which parasitic on ants (Samson
& Evans 1973). The genus is characterised by producing singly conidia on sympodial
conidiogenous cells (Samson & Evans 1973). The sexual morph link to Torrubiella (Samson &
Evans 1973, White et al. 2003).
Samsoniella Mongkolsamrit, Noisripoom, Thanakitpipattana, Spatafora & Luangsa-ard, Mycologia
110(1): 230-257 (2018)
Index Fungorum number: IF823784; 3 species with sequence data.
Type species – Samsoniella inthanonensis Mongkolsamrit, Noisripoom, Thanakitpipattana,
Spatafora & Luangsa-ard
Notes – The genus was erected by Mongkolsamrit et al. (2018) and segregated from
Akanthomyces to accommodate species in a monophyletic clade that are easily recognized by their
oval to fusiform conidia, and bright red-orange stromata of the sexual morphs and synnemata of the
isaria-like asexual morphs.
Simplicillium W. Gams & Zare, Nova Hedwigia 73(1-2): 38 (2001)
Index Fungorum number: IF28570; 14 species with sequence data.
Type species – Simplicillium lanosoniveum (J.F.H. Beyma) Zare & W. Gams
Notes – The asexual morphs of this genus are cordyceps-like (White et al. 2003), but
molecular analyses has shown this as a separate genus from Cordyceps. The genus infects insects
and produces exclusively solitary phialides, adhering in globose slimy heads or imbricate chain
conidia (Zare & Gams 2001).
Coronophoraceae Höhn., Sber. Akad. Wiss. Wien, Math.-naturw. Kl., Abt. 1 116: 624 (1907)
Index Fungorum number: IF80647; Facesoffungi number: FoF01117; 17 species.
Saprobic on plant stems or wood in terrestrial habitats, or parasitic on other fungi. Sexual
morph: Ascomata perithecial, gregarious or solitary, superficial, black, semi-immersed becoming
erumpent through bark of host, ovoid to subglobose, carbonaceous, tuberculate, glabrous or with
hairs, laterally collapsing when dry, sometimes with a short neck with or without ostioles or
quellkörper. Peridium carbonaceous or membranaceous, composed of three layers, outer layer
comprising dark tissue, carbonaceous; middle layer comprising dark brown to brown cells of
textura angularis, membranaceous and the inner layer comprising hyaline cells of textura
prismatica, membranaceous. Paraphyses numerous, filamentous, septate, unbranched. Asci
polysporous, unitunicate, thin-walled, clavate to cylindrical, long pedicellate, apex blunt, without a
visible discharge mechanism. Ascospores crowded, hyaline, cylindrical to allantoid, slightly
curved, aseptate, smooth-walled, mostly with guttules. Asexual morph: Undetermined.
Type genus – Coronophora Fuckel
Notes – The monotypic family Coronophoraceae was introduced by Höhnel (1907b). As a
synonym of Nitschkiaceae, it had been placed in Coronophorales (Nannfeldt 1932, Müller & Arx
1973, Subramanian & Sekar 1990) or in Sordariales (Nannfeldt 1975, Barr 1990b). Mugambi &
Huhndorf (2010) used a tef1 and rpb2 combined sequence dataset, which included the type species
C. gregaria, to show that Coronophoraceae is distinct from Nitschkiaceae.
Ecological and economic significance of Coronophoraceae
545
Most Coronophora species are saprobic on wood and widespread in Europe and America and
involved in nutrient cycling (Fuckel 1870, Mugambi & Huhndorf 2010).
Figure 84 – Coronophora gregaria (Material examined – BELGIUM, on bark of Prunus cerasus
and Sorbus aucuparia, Autumn, collected by Libert MA, BR no. 5020094489111, isotype). a
Herbarium material. b Ascomata erumpent through bark of host. c Ascomata on decorticated wood.
d Ascoma in cross section (arrow head indicates ascogenous hyphae). e Peridium. f Munk pores. g
Paraphyses with asci. h, i Asci. j-o Ascospores. Scale bars: b-c = 2 mm, d = 200 µm, g = 100 µm, e
= 50 µm, h-i = 20 µm, f = 10 µm, j-o = 5 µm.
546
Genus included in Coronophoraceae
Coronophora Fuckel, Fungi rhenani exsic., fasc. 10: no. 961 (1864)
Index Fungorum number: IF1253; 17 morphological species (Species Fungorum 2020), 2
species with sequence data.
Type species – Coronophora gregaria Fuckel
Notes – Coronophora is characterized by tuberculate ascomata with polysporous asci and
tiny, allantoid, hyaline ascospores. Coronophora gregaria is illustrated in this entry. Sphaeria
gregaria is synonym as Coronophora gregaria and occurs on dead wood in Europe and America
(Fuckel 1870, Mugambi & Huhndorf 2010). The sequences of tef1 and rpb2 of C. gregaria
(ANM1555) were provided by Mugambi & Huhndorf (2010).
Coryneaceae Corda, Icon. fung. (Prague) 3: 36 (1839)
Index Fungorum number: IF80650; Facesoffungi number: FoF06868; 79 species.
Saprobic on dead wood or pathogenic on plants. Sexual morph: Stromata solitary, erumpent,
comprising pseudoparenchymatous cells. Ectostromatic disc well or poorly developed, brown to
black, comprising small cells of textura prismatica. Ascomata perithecial, arranged in valsoid
configuration, immersed, aggregated, globose to subglobose, coriaceous, brown to black, papillate,
ostiolate. Papilla upright, central, broad, sometimes converging, comprising brown cells of textura
porrecta. Peridium comprising outer, thick-walled, brown cells of textura angularis and inner,
thick-walled, hyaline, compressed cells of textura angularis. Paraphyses broad, cellular, septate,
attached to base, longer than asci. Asci 8-spored, unitunicate, ellipsoid to cylindrical, pedicellate,
rounded at apex with a J-, apical ring. Ascospores overlapping uni- to biseriate, hyaline or initially
hyaline, brown at maturity, irregularly fasciculate, ellipsoid, fusoid or elongate, 1–3-septate, often
distoseptate, with end-cells pale brown or hyaline, sometimes end-cells pointed, straight or curved
not constricted at the septa, guttulate, smooth-walled. Asexual morph: Coelomycetous.
Conidiomata acervular, solitary, erumpent through outer periderm layers of host or immersed,
scattered, surface tissues above somewhat dome-shaped. Conidiomatal wall composed of thinwalled, vertically arranged dark brown cells of textura angularis. Conidiophores branched at the
base or not, cylindrical to globose, septate or aseptate, hyaline or hyaline at the apex, pale brown at
the base. Conidiogenous cells terminal, hyaline, annellidic, cylindrical, sometimes with setulose
apical appendages. Conidia hyaline to dark brown, curved, broadly fusiform to cylindrical or
clavate, smooth-walled, 4–6-distoseptate, sometimes the apical and basal cell darker than other
cells with hyaline tip in apical cell.
Type genus – Coryneum Nees
Notes – Coryneaceae was introduced by Corda (1839a) to accommodate Coryneum which
was typified by C. umbonatum and comprises only one genus.
Ecological and economic significance of Coryneaceae
Some species of Coryneum cause coryneum-blight, a serious fungal disease that forms holes
in leaves, rough areas on fruits and concentric lesions on branches (Senanayake et al. 2018). Many
Coryneum species cause cankers in economically important trees and forest trees (Strouts 1972,
Gadgil & Dick 2007, Horst 2013, Senanayake et al. 2017a). Several Coryneum species occur on
chestnut and oak trees (Sutton 1975). Some saprobic species are associated with decaying wood
thus contributing to nutrient cycling.
Genus included in Coryneaceae
Coryneum Nees, Das System der Pilze and Schwamme 34 (1817)
Index Fungorum number: IF7798; 79 morphological species (Species Fungorum 2020), 10
species with sequence data, molecular data available for 2 unnamed species.
Type species – Coryneum umbonatum Nees
Notes – Rossman et al. (2015) synonymized Pseudovalsa under Coryneum. The asexual
morph is the most common morph seen. There are more than 70 species recorded under Coryneum.
547
However, molecular sequence data are lacking for most species. Wijayawardene et al. (2016b),
Senanayake et al. (2017a), Jiang et al. (2018) and Senwanna et al. (2018) introduced several species
providing phylogenetic analyses. Common host genera include Acer, Betula, Castanea, Eucalyptus,
Populus, Quercus, Rosa, Rhododendron, Salix, Sorbus, Sambucus, Tilia, and Vitis (Farr &
Rossman 2018). Coryneum species are common in temperate regions than in tropical regions.
Coryneum brachyurum, current name Coryneum lanciforme (Species Fungorum 2020) is illustrated
in this entry.
Figure 85 – Coryneum lanciforme, a-j sexual morph, k-q asexual morph (Material examined –
USA, Maryland, Beltsville, Narragansett Pkwy, on stem of Betula sp., 07 June 2002, Vasilyeva,
Larissa BPI 871006A (sexual morph); UK, Devon, Slapton, near Kingsbridge, slapton, Ley
National Nature Reserve, site J1, on twigs of Quercus petraea, 11 May 1994, B. C. Sutton, IMI
362542 (asexual morph)). a Herbarium packet. b Herbarium specimen. c Stromata on substrate. d
Transverse section of stroma. e-g Asci. h Peridium. i-j Ascospores. k, l Herbarium packet and
specimen. m Conidiomata on substrate. n Cross section of conidioma. o Conidia attached to
conidiophore. p-q Conidia. Scale bars: c = 1 mm, d = 100 µm, e-h = 20 µm, i-j = 10 µm, m = 500
μm, e = 100 μm, o-q = 10 μm.
Cryphonectriaceae Gryzenh. & M.J. Wingf., Mycologia 98: 246 (2006)
Index Fungorum number: IF510585; Facesoffungi number: FoF03455; 91 species.
548
Saprobic or pathogenic in forest trees and economic crops. Sexual morph: Ascostromata
scattered, immersed or erumpent, aggregated, oval to circular from above, comprising two layers,
with upper layer of yellowish orange to pale brown cells, purpling in KOH and inner layer of
hyaline cells, mixed with plant cells. Ascomata immersed, aggregated, several in one stroma,
globose to subglobose, fuscous black to umber, with long neck, sometimes ostiolar canal immersed
in stromatic tissues or superficial, with necks covered in umber stromatic tissue of texturaporrecta,
inner wall of the necks or ostiolar canal with hyaline, filamentous periphyses. Peridium comprising
inner layer of small, hyaline cells of textura angularis and outer layer of small, brown cells of
textura angularis. Paraphyses few, cellular, parenchymatous cells, attached at the base of asci and
dissolving at maturity. Asci 8-spored, unitunicate, cylindrical-fusoid to clavate, pedicellate, with
distinct, J-, refractive apical ring. Ascospores overlapping uniseriate to biseriate, hyaline,
sometimes brown,ellipsoid, fusoid to cylindrical, aseptate to multi-septate, not constricted at septa,
smooth-walled. Asexual morph: Coelomycetous. Conidiomata occurring as part of ascostromata, as
conidial locules or solitary structures, uni- to multi-loculate, pyriform, subglobose to pulvinate,
ostiolar necks absent or present; if present, with one to several attenuated necks, superficial or
semi-immersed, orange to fuscous-black. Conidiophores cylindrical, aseptate, hyaline, sometimes
reduced to conidiogenous cells. Conidiogenous cells lining the inner cavity of the conidiomata,
phialidic, sometimes within flattened base, ampulliform,conspicuous, with attenuated or truncate
apices, hyaline, smooth. Conidia minute, sometimes both micro- and macro-conidia present,
sigmoid, broadly ellipsoid to fusoid, obovoid-cylindrical to allantoid, aseptate, hyaline.
Type genus – Cryphonectria (Sacc.) Sacc. & D. Sacc.
Notes – Cryphonectriaceae comprises mostly tree and plant pathogens (Vermeulen et al.
2011) and rarely saprobes and endophytes. They cause diseases in economically important plants
and forest trees. Cryphonectriaceae was established by Gryzenhout et al. (2006b) to accommodate
the Cryphonectria-Endothia complex based on nrLSU sequence data.
Ecological and economic significance of Cryphonectriaceae
Cryphonectriaceae comprises highly virulent plant pathogens such as Cryphonectria
parasitica which is the causal agent of chestnut blight (Anagnostakis 1987, Heiniger & Rigling
1994). Some species are pathogenic on Myrtales tree species, especially forms causing stem canker
disease (Chen et al. 2016). Species in Cryphonectriaceaeare commonly associated with serious
cankers and leaf spots diseases (Crous et al. 2015a). Very few species are saprobes.
Genera included in Cryphonectriaceae
Amphilogia Gryzenhout, Glen & M. J. Wingf., Taxon 54(4): 1017 (2005b)
Index Fungorum number: IF510065; 2 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Amphilogia gyrosa (Berk. & Broome) Gryzenhout, Glen & M.J. Wingf.
Notes – Amphilogia was introduced to accommodate Cryphonectria gyrosa (CMW
10471/CRY 1551) based on its morphological distinctness such as superficial conical conidiomata,
conidia of variable size and ascospores with one to three septa (Gryzenhout et al. 2005). The
second species Amphilogia major, accommodated in this genus, was reported from New Zealand
(Gryzenhout et al. 2005).
Aurantioporthe G. Beier & Blanchette, Mycologia 107(1): 71 (2015)
Index Fungorum number: IF807278; 1 species with sequence data.
Type species – Aurantioporthe corni (Wehm.) G. Beier& R.A. Blanchette
Notes – Aurantioporthe is known from only its type species, A. corni. This species is the
causative agent of golden cankers disease in USA and Canada. Aurantioporthe differs from other
genera in Cryphonectriaceae in having ascogenous stromata, immersed to semi-immersed
conidiomata and uni- to multi-septate ascospores.
549
Aurantiosacculus Dyko & B. Sutton, Mycologia 71(5): 922 (1979)
Index Fungorum number: IF7296; 4 morphological species (Species Fungorum 2020), 2
species with sequence data.
Type species – Aurantiosacculus eucalypti (Cooke & Massee) Dyko & B. Sutton
Notes – Aurantiosacculus species are associated with leaf spots on Eucalyptus (Crous et al.
2012b). Aurantiosacculus produces leaf spots, with bright orange conidiomata with brown
furfuraceous tissue and aseptate conidia with swollen bases and thickened scars.
Aurapex Gryzenh. & M.J. Wingf., Mycologia 98(1): 112 (2006b)
Index Fungorum number: IF501067; 1 species with sequence data.
Type species – Aurapexpenicillata Gryzenh. & M.J. Wingf.
Notes – Aurapex is distinguished from other genera of Cryphonectriaceae by its orange
conidiomatal necks with black bases and a unique internal stromatal structure. Species of Aurapex
are serious canker pathogens of Eucalyptus species (Gryzenhout et al. 2006b).
Aurifilum Begoude, Gryzenh. & Jol. Roux, Antonie van Leeuwenhoek 98(3): 273 (2010)
Index Fungorum number: IF546909; 1 species with sequence data.
Type species – Aurifilum marmelostoma Begoude, Gryzenh. & Jol. Roux
Notes – The monotypic genus Aurifilum was introduced and typified by Aurifilum
marmelostoma which has broad convex conidiomata, darkened ostiolar openings at the apex of the
conidiomata, paraphyses or sterile cells, one septate, fusoid to ellipsoid ascospores and minute,
cylindrical conidia.
Capillaureum M.E.S. Oliveira, G.A. Silva & M.A. Ferreira, Mycol. Progr. 18(3): 392 (2019)
Index Fungorum number: IF827751; 1 species with sequence data.
Type species – Capillaureum caryovora M.E.S. Oliveira, G.A. Silva & M.A. Ferreira
Notes – Capillaureumwas isolated from Caryocar brasiliense, a tree used in the cosmetic and
food industries. This is a stem canker causing fungus. This genus is distinguished from other genera
in having a fuscous-black necks with septate, branched hyphal ostioles.
Celoporthe Nakab., Gryzenh., Jol. Roux & M.J. Wingf., Stud. Mycol. 55: 261 (2006)
Index Fungorum number: IF500886; 9 morphological species (Species Fungorum 2020), 9
species with sequence data.
Type species – Celoporthe dispersa Nakab., Gryzenh., Jol. Roux & M.J. Wingf.
Notes – This genus is distinguished from other genera in having immersed, umber to brown
to fuscous-black conidiomata with cylindrical conidia.
Chromendothia Lar.N. Vassiljeva, Mikol. Fitopatol. 27(4): 5 (1993)
Index Fungorum number: IF27570; 2 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Chromendothia citrina Lar.N. Vassiljeva
Notes – Chromendothia comprises three species. Chromendothia shares characters with other
taxonomic groups such as hypocreaceous fungi and some diaporthalean members. The genus is
characterised by characteristic brown ascospores and whitish yellow stromatic tissues. There are no
records of the asexual morph of this genus.
Chrysofolia Crous & M.J. Wingf., Persoonia, Mol. Phyl. Evol. Fungi 34: 207 (2015)
Index Fungorum number: IF812450; 2 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Chrysofolia colombiana Crous, Rodas & M.J. Wingf.
Notes – Chrysofolia is distinct in having erumpent conidiomata with bright, yellow-brown
furfuraceous margins (Crous et al. 2015c).
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Chrysomorbus S.F. Chen, Pl. Path. 67(1): 117 (2017)
Index Fungorum number: IF821021; 1 species with sequence data.
Type species – Chrysomorbus lagerstroemiae S.F. Chen & Q.L. Liu
Notes – Chrysomorbus is distinct from other genera having fusoid to oval conidia, convex to
globose conidiomata and lack of ostioles. Chrysomorbus lagerstroemiae is associated with cankers
on Lagerstroemia speciosa (Lythraceae, Myrtales).
Chrysoporthe Gryzenh. & M.J. Wingf., Stud. Mycol. 50(1): 129 (2004)
Index Fungorum number: IF500032; 8 species with sequence data.
Type species – Chrysoporthe cubensis (Bruner) Gryzenh. & M.J. Wingf.
Notes – Chrysoporthe was typified by C. cubensis and the sexual morph of this genus does
not commonly occur in nature. Chrysoporthe is distinguished from other genera in this family
having superficial, pyriform, fuscous-black conidiomata with oblong conidia.
Corticimorbus S.F. Chen & M.J. Wingf., Pl. Path. 65(8): 1258 (2016)
Index Fungorum number: IF815261; 1 species with sequence data.
Type species – Corticimorbus sinomyrti S.F. Chen, F.F. Liu & M.J. Wingf.
Notes – Corticimorbus sinomyrtiis a tree pathogen on Rhodomyrtus tomentosa forming stem
canker disease. The fungus is most similar to species in Chrysoporthe based on morphology.
However, this genus can be distinguished from the other genera having conical pycnidia and fusoid
to oval conidia.
Cryphonectria (Sacc.) Sacc. & D. Sacc., Syll. fung. (Abellini) 17: 783. (1905)
Index Fungorum number: IF9670; 14 morphological species (Species Fungorum 2020), 8
species with sequence data.
Type species – Cryphonectria parasitica D. Sacc.
Notes – Cryphonectria is typified by C. parasitica the causal agent of chestnut blight
(Anagnostakis 1987, Heiniger & Rigling 1994). Cryphonectria species are sometimes serious
canker pathogens but generally survive as saprobes (Roane 1986). Cryphonectria parasitica is
illustrated in this entry.
Cryptometrion Gryzenh. & M.J. Wingf., Australas. Pl.Path. 39(2): 166 (2010)
Index Fungorum number: IF514188; 1 species with sequence data.
Type species – Cryptometrion aestuescens Gryzenh. & M.J. Wingf.
Notes – Cryptometrion can be distinguished from other genera in Cryphonectriaceae based on
its orange, limited stromatic tissue, uniseptate, fusoid to ellipsoid ascospores and absence of
paraphyses among the conidiogenous cells in the asexual morph.
Diversimorbus S.F. Chen & Jol. Roux, Fungal Biol. 117: 300 (2014)
Index Fungorum number: IF564805; 1 species with sequence data.
Type species – Diversimorbus metrosiderotis S.F. Chen & Jol. Roux
Notes – Diversimorbus was isolated from South Africa associated with Myrtales’ tree
species. The fungus is pathogenic on its host (Chen et al. 2013). The genus is characterised by
pulvinate conidiomata without necks and fusoid to allantoid conidia.
Endothia Fr., Summa veg. Scand., Section Post. (Stockholm): 385 (1849)
Index Fungorum number: IF1810; 9 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Endothia gyrosa (Schwein.) Berk.
Notes – Endothia was introduced based on Sphaeria gyrosa, which causes a serious canker
disease (Snow et al. 1975). Endothia species cause hobnail canker on oak (Van Arsdel 1972). The
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genus is characterised by immersed, orange conidiomata without necks and with cylindrical
conidia.
Figure 86 – Cryphonectria parasitica (Material examined – USA, New York. Bronx Co. Bronx.
North of Botanical Museum, Bronx Park, on Castanea dentata (Fagaceae), 26 Nov. 1905, W.A.
Murrill, NY 01293321, holotype of Diaporthe parasitica). a Herbarium packet. b Herbarium
specimen. c Ascostromata on substrate. d Horizontal section of ascostroma. e Vertical cross section
of ascoma. f-i Asci. j-m Ascospores. n Horizontal cross section of conidiomata. o Vertical cross
section of conidioma. p, q Conidia attached to the conidiogenous cells and conidiophores. r
Conidia. Scale bars: c, d = 1 mm, f-i, p-r = 10 µm, e, q, o = 100 µm, n = 200 µm, j-m = 5 µm.
Foliocryphia Cheew. & Crous, Persoonia 23: 65 (2009)
Index Fungorum number: IF513846; 2 species with sequence data.
Type species – Foliocryphia eucalypti Cheew. & Crous
Notes – Foliocryphia is characterized by aseptate conidia in eustromatic conidiomata and
stromata which do not turn purple in 3% KOH, or yellow in lactic acid.
Holocryphia Gryzenh. & M.J. Wingf., Stud. Mycol. 55: 48 (2006b)
Index Fungorum number: IF500797; 4 species with sequence data.
Type species – Holocryphia eucalypti (M. Venter & M.J. Wingf.) Gryzenh. & M.J. Wingf.
Notes – Holocryphia eucalypti is an opportunistic canker pathogen of Eucalyptus and
Corymbia species (Chen et al. 2013). The genus is characterised by semi-immersed conidiomata,
septate conidiophores and cylindrical conidia.
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Immersiporthe S.F. Chen, M.J. Wingf. & Jol. Roux, Pl. Path. 62: 674 (2013)
Index Fungorum number: IF564804; 1 species with sequence data.
Type species – Immersiporthe knoxdaviesiana S.F. Chen, M.J. Wingf. & Jol. Roux
Notes – Immersiporthe knoxdaviesiana is an aggressive pathogen of Rapanea melanophloeos.
Immersiporthe is morphologically distinct in having orange pulvinate conidiomata, without
conidiomatal necks and with paraphyses.
Latruncellus M. Verm., Gryzenh. & Jol. Roux, Mycologia103(3): 562 (2011)
Index Fungorum number: IF518285; 1 species with sequence data.
Type species – Latruncellus aurorae M. Verm., Gryzenh. & Jol. Roux
Notes – This monotypic genus is associated with cankers on Galpinia transvaalica
(Lythraceae, Myrtales) in Swaziland. The genus is characterised by distinct conical conidiomata
with a distinct neck and subulate to flask-shaped conidiophores.
Luteocirrhus C. Crane & T.I. Burgess, IMA Fungus 4(1): 115 (2013)
Index Fungorum number: IF563390; 1 species with sequence data.
Type species – Luteocirrhus shearii C. Crane & T.I. Burgess
Notes – The monotypic genus Luteocirrhus was isolated from canker lesions in Banksia
species and Lambertia echinata. The genus is characterised by some semi-immersed conidiomata,
paraphyses within the locules and cylindrical conidia.
Mastigosporella Höhn. Sber. Akad. Wiss. Wien, Math.- naturw. Kl., Abt. 1 123: 135 (1914)
Index Fungorum number: IF8860; 5 morphological species (Species Fungorum 2020), 2
species with sequence data.
Type species – Mastigosporella hyalina (Ellis & Everh.) Höhn.
Notes – Mastigosporella hyalina is morphologically similar to Harknessia caudata, however
differs by colour of conidia. Additionally, the pattern of conidiogenesis in Harknessia and
Mastigosporella differ (Sutton 1971).
Microthia Gryzenh. & M.J. Wingf., Stud. Mycol. 55: 44 (2006)
Index Fungorum number: IF500792; 3 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Microthia coccolobae (Vizioli) Gryzenh. & M.J. Wingf.
Notes – Microthia comprises two species Microthiahavanensis and M. coccolobae. Microthia
is distinct in having small, pulvinate and semi-immersed stromata tending to be superficial on the
substrate.
Myrtonectria Marinc., D.B. Ali & J. Roux, Mycol. Progr. 17(8): 958 (2018)
Index Fungorum number: IF824022; 1 species with sequence data.
Type species – Myrtonectria myrtacearum Marinc., D.B. Ali & J. Roux
Notes – Myrtonectria myrtacearum appears to be an endophyte or cryptic pathogen that was
obtained from branches of Syzygium cordatum, root collars of Heteropyxisnatalensis and bark of
Tibouchina grandifolia. The genus is characterised by semi-immersed to superficial, dark greyish
brown, uniloculate conidiomata and aseptate, oblong conidia with a pointed base.
Rostraureum Gryzenh. & M.J. Wingf., Mycol. Res. 109(9): 1039 (2005b)
Index Fungorum number: IF28972; 2 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Rostraureum tropicale Gryzenh. & M.J. Wingf.
Notes – Rostraureum is distinguished by orange, superficial, rostrate, eustromatic
conidiomata (Gryzenhout et al. 2005).
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Ursicollum Gryzenh. & M.J. Wingf., Stud. Mycol. 55: 44 (2006a)
Index Fungorum number: IF500795; 1 species with sequence data.
Type species – Ursicollum fallax Gryzenh. & M.J. Wingf.
Notes – Ursicollum is typified by U. fallax which is distinct from other genera in
Cryphonectriaceae in having unique orange, pyriform to globose conidiomata with cylindrical to
attenuated necks (Gryzenhout et al. 2006a).
Wuestneia Auersw., Hedwigia 3: 159 (1864)
Index Fungorum number: IF5799; 12 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Wuestneia aurea (Fuckel) Auersw.
Notes – The type species of Wuestneia, W. aurea was placed in Cryphonectriaceae by
Rossman et al. (2007). However, most species of Wuestneia have been placed in Harknessia (Crous
et al. 2012d, Senanayake et al. 2017a). The genus is characterised by cylindrical asci with aseptate
ascospores and ampuliform conidiogeneous cells without conidiophores.
Cylindriaceae Crous & L. Lombard, Fungal Systematics and Evolution 1: 183 (2018)
Index Fungorum number: IF824770; Facesoffungi number: FoF06869; 10 species.
Associated with plant substrates, especially leaves. Sexual morph: Undetermined. Asexual
morph: Mycelium composed of septate, branched, hyaline, smooth-walled hyphae. Conidiophores
sporodochial or solitary, erect, hyphae and basal part of conidiophores subcylindrical, septate,
branched, becoming pale brown, smooth-walled. Conidiogenous cells terminal and intercalary,
subcylindrical, hyaline, smooth-walled, with several sympodial flat-tipped loci, unthickened, not
darkened. Ramoconidia subcylindrical, guttulate, hyaline, smooth-walled. Conidia aseptate,
arranged in long, branched chains, scars not thickened, slightly refractive, hyaline, smooth-walled
(adapted from Crous et al. 2018c).
Type genus – Cylindrium Bonord.
Notes – Cylindrium was considered as Hypocreales incertae sedis by Lombard et al. (2015).
Crous et al. (2018c) introduced Cylindriaceae in Hypocreales to accommodate this genus based on
phylogenetic analysis of ITS sequence data. We maintain Cylindriaceae in Amphisphaeriales based
on our phylogenetic anlysis of combined genes and divergence estimates (Figs 2, 4).
Figure 87 – Cylindrium candidum (redrawn from Bonorden 1851) Conidiophores, conidiogenous
cells and conidial chains.
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Ecological and economic significance of Cylindriaceae
Cylindrium purgamentum was isolated from leaf litter (Crous et al. 2016a). Other species
such as C. syzygii and C. algarvense were isolated from Syzygium and Eucalyptus leaves
(Cheewangkoon et al. 2009, Crous et al. 2011). There are no report of Cylindrium species as a leaf
pathogen.
Genus included in Cylindriaceae
Cylindrium Bonord., Handb. Allgem. mykol. (Stuttgart): 34 (1851)
Index Fungorum number: IF7865; 10 morphological species (Species Fungorum 2020), 3
species with sequence data.
Type species – Cylindrium elongatum Bonord.
Notes – This asexual morph genus is characterized by sporodochial conidiomata,
subcylindrical conidiogenous cells and aseptate hyaline conidia arranged in long chains (Bonorden
1851, Crous et al. 2018c). The taxonomic placement of Cylindrium was doubtful for years.
Cylindrium elongatum was accepted in Hypocreales incertae sedis by Lombard et al. (2015). Crous
et al. (2018c) revealed that Cylindrium is a member of Hypocreales and described Cylindriaceae
based on ITS sequence data. Cylindrium elongatum grouped within Amphisphaeriales in our
phylogenetic analysis (Fig. 4). Cylindrium candidum and C. algarvense are illustrated in this entry
(Figs. 87, 88).
Cytosporaceae Fr., Syst. orb. veg. (Lundae) 1: 118 (1825)
Index Fungorum number: IF82042; Facesoffungi number: FoF06870; 305 species.
Pathogenic or saprobic on plant tissues. Sexual morph: Ascomata perithecia or globose,
immersed to erumpent, solitary or 3–10 ascomata aggregated, black to brown, with long neck
swollen at the tips, ostiolate. Ostiole periphysate, with ostiolar neck. Peridium comprising outer, 4–
6 layers of dark brown with cells of textura angularis and 5–7 layers of, inner, small, hyaline, thinwalled, cells of textura angularis. Paraphyses few, hyaline. Asci 8-spored, unitunicate,clavate,
short-pedicellate, apex round, with or without apical ring. Ascospores uni- to biseriate, unicellular
or rarely bicellular, allantoid or ellipsoid, hyaline, smooth-walled. Asexual morph: Conidiomata
pyriform in section, brown to dark brown, divided into compartments by bending of peridium.
Peridium brown to pale brown, comprised 4–6 layers with cells of textura angularis.
Conidiophores reduced to conidiogenous cells. Conidiogenous cells arising from conidiomatal
wall, phialidic, simple or branched, hyaline, cylindrical. Conidia unicellular, allantoid, hyaline,
smooth-walled.
Type genus – Cytospora Ehrenb.
Notes – Cytosporaceae was introduced by Fries (1825) and placed into Diaporthales, which
comprises phytopathogens and saprobes (Wehmeyer 1975, Barr 1978, Eriksson 2001, Castlebury et
al. 2002). Maharachchikumbura et al. (2015, 2016b) listed 13 genera under Cytosporaceae (viz.
Amphicytostroma, Chadefaudiomyces, Cryptascoma, Cytospora, Ditopellina, Durispora,
Harpostroma, Hypospilina, Kapooria, Leptosillia, Maculatipalma, Pachytrype and Paravalsa).
Senanayake et al. (2017a) excluded seven genera and accepted five genera in Cytosporaceae (viz.
Cytospora, Pachytrype, Paravalsa, Waydora, Xenotypa) based on morphological characteristics.
Wijayawardene et al. (2018a) accepted six genera in Cytosporaceae in the outline of Ascomycota
(Cytospora, Hypophloeda, Pachytrype, Paravalsa, Waydora, Xenotypa). Senanayake et al. (2018)
re-examined and added two genera (viz. Cryptascoma, Hypophloeda) to Cytosporaceae based on
morphology. We place Hypophloeda in Diaporthales genera incertae sedis as it is atypical of
Cytosporaceae.
Ecological and economic significance of Cytosporaceae
Cytospora speies are important pathogens causing canker and dieback disease of a wide range
of plants (Adams et al. 2005, 2006, Hyde et al. 2016b, Norphanphoun et al. 2017, 2018).
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Figure 88 – Cylindrium algarvense (redrawn from Cheewangkoon et al. 2009). a Sporodochium on
OA. b Creeping hyphae. c Conidiophores, conidiogenous cells and conidial chains. d Conidia.
Scale bars: a = 200 µm, c = 10 µm, d = 20 µm.
Genera included in Cytosporaceae
Cryptascoma Ananthap., Trans. Br. mycol. Soc. 90(3): 479 (1988)
Index Fungorum number: IF25170; 2 morphological species (Species Fungorum 2020).
Type species – Cryptascoma bisetulum Ananthap.
Notes – Cryptascoma was introduced and typified by Ananthapadmanaban (1988) with C.
bisetula found on dead twigs in Tamil Nadu, India. This genus is distinguished by fusiform, 2celled, biseriate ascospores with protoplasmic extensions (Barr 1978, Monod 1983) and it was
assigned to Cytosporaceae by Ananthapadmanaban (1988).
Cytospora Ehrenb., Sylv. mycol. berol. (Berlin): 28 (1818)
Index Fungorum number: IF7904; 297 morphological species (Species Fungorum 2020), 117
species with sequence data.
Type species – Cytospora chrysosperma (Pers.) Fr.
Notes – Cytospora was introduced by Ehrenberg (1818). The genus contains important
pathogens causing canker and dieback disease of a wide range of plants (Adams et al. 2005, 2006,
Hyde et al. 2016b, Norphanphoun et al. 2017). Recent accounts of the genus have been published
with several new taxa (Norphanphoun et al. 2017, 2018, Tibpromma et al. 2017b, Senanayake et al.
2017a, Hyde et al. 2017b). Cytospora cedri is illustrated in this entry (Fig. 89).
Pachytrype Berl. Mycotaxon 48: 530 (1993)
Index Fungorum number: IF26475; 3 morphological species (Species Fungorum 2020), 2
species with sequence data.
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Type species – Pachytrype princeps (Penz. & Sacc.) M.E. Barr, J.D. Rogers & Y.M. Ju
Notes – Pachytrype was described by Barr et al. (1993) to accommodate two species,
Pachytrype princeps (= Diatrype princeps), species found from undetermined tree in Java; and
Pachytrype graphidioides (= Peroneutypella graphidioides), species found on Terminalia catappa
in the Philippines. Pachytrype rimosa was introduced from on a log in Costa Rica (Fernández et al.
2004). The genus is characterised by stroma showing multi- locules perithecial and entostroma with
perithecial beaks showing through cracks in stroma, cylindrical asci with cubical apical ring and
hyaline ascospores with ellipsoid to allantoid shaped (Fernández et al. 2004).
Figure 89 – Cytospora cedri (Material examined – ITALY, on branches of Rubus sp., Erio
Campesori IT3288, MFLU 17-0835). a Stromatal habit in wood. b Fruiting bodies on host surface.
c Surface of fruiting bodies showing the black ostioles. d Cross section of the stroma showing
conidiomata. e Peridium. f Ostiolar neck. g, h Conidiogenous cell containing conidia. i Conidia.
Scale bars: d = 100 µm, e, f = 20 µm, g-i = 5 µm.
Paravalsa Ananthap., Mycol. Res. 94(2): 275 (1990)
Index Fungorum number: IF25381; 1 morphological species.
Type species – Paravalsa indica Ananthap.
Notes – Paravalsa was introduced as monotypic genus to accommodate P. indica, which was
found on bark in Tamil Nadu (Ananthapadmanaban 1990). The genus is characterised by a lack of
557
stromata, solitary perithecia immersed under host tissues with prominent necks, 8-spored, clavate,
J- asci and 1-celled, allantoid ascospores (Ananthapadmanaban 1990).
Waydora B. Sutton, Trans. Br. mycol. Soc. 67(2): 248 (1976)
Index Fungorum number: IF10435; 1 species with sequence data.
Type species – Waydora typica (Rodway) B. Sutton
Notes – Waydora typica is found on capsules of Eucalyptus globulus in Tasmania (Sutton et
al. 1976). The genus is characterised by multilocular conidiomata, furfuraceous ostioles, septate
conidiophores which are branched at the base and above, enteroblastic phialidic conidiogenous
cells and straight or allantoid, aseptate conidia.
Xenotypa Petr., Sydowia 9(1-6): 499 (1955)
Index Fungorum number: IF5828; 1 morphological species.
Type species – Xenotypa aterrima (Fr.) Petr.
Notes – Xenotypa aterrima has stromata spreading around branches and often extending
between the cortical parenchyma and the periderm, splitting and pushing aside layers of bark
(Petrak 1955c). The genus is characterised by perithecia with papillate, cylindric-conical necks, 8spored, clavate asci and straight, hyaline, aseptate allantoid ascospores with rounded ends, often
with two rather indistinct oil droplets (Petrak 1955c).
Delonicicolaceae R.H. Perera, Maharachch. & K.D. Hyde, Cryptog. Mycol. 38(3): 334 (2017)
Index Fungorum number: IF553776; Facesoffungi number: FoF03604; 4 species.
Saprobic on fabaceous hosts. Sexual morph: Pseudostromata conspicuously pulvinate to
inconspicuous, embedded in host tissue or erumpent to rarely superficial, visible as raised, dark
spots on the host surface, more or less elevated patches on the wood or erumpent through the bark,
yellowish, brown to black; sometimes covered by yellow to yellow-green, bright turquoise, scurf.
Ascomata perithecial, immersed in pseudostroma, multi-loculate, aggregated, globose, subglobose
to conical or irregular, subhyaline to pale brown, ostiolate. Ostioles papillate. Peridium composed
of subhyaline to medium brown cells of textura angularis to textura prismatica, KOH-. Paraphyses
septate, unbranched or occasionally branched, hyaline. Asci 8-spored, unitunicate, arising from the
base or margins of the ascomata, clavate to cylindrical, straight, curved to sinuous, thin-walled, J-,
without a distinct apical ring. Ascospores bi-seriate to uniseriate, hyaline, ellipsoid or allantoid,
equilateral or inequilateral, with rounded apices, aseptate or 1-septate, not constricted at the septa,
thin-walled, smooth, without appendages or gelatinous sheath. Asexual morph: Undetermined.
Type genus – Delonicicola R.H. Perera, Maharachch. & K.D. Hyde
Notes – Perera et al. (2017) introduced Delonicicolaceae (Delonicicolales) to accommodate
Delonicicola and Liberomyces. Liberomyces was synonymised under Leptosillia and added to
Leptosilliaceae (Voglmayr et al. 2019a). A new genus Furfurella was added to Delonicicolaceae
based on morphological and molecular evidence by Voglmayr et al. (2019a). Vitale et al. (2018)
and Voglmayr et al. (2019a) accepted Delonicicolaceae within Xylariales. However, based on our
phylogenetic analysis (Fig. 4) we maintain Delonicicolaceae in Delonicicolales following the
classification of Perera et al. (2017).
Ecological and economic significance of Delonicicolaceae
Delonicicola siamense is a saprobic fungus occurring on decaying seed pods of Delonix regia
(Perera et al. 2017). There are no reports of pathogenic species in Delonicicola or Furfurella
(Perera et al. 2017, Voglmayr et al. 2019a).
Genera included in Delonicicolaceae
Delonicicola R.H. Perera, Maharachch. & K.D. Hyde, Cryptog. Mycol. 38(3): 334 (2017)
Index Fungorum number: IF553772; 1 species with sequence data.
Type species – Delonicicola siamense R.H. Perera, Maharachch. & K.D. Hyde
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Notes – The sexual morph, monotypic genus Delonicicola was established based on
Delonicicola siamense, which was isolated from Delonix regia seed pods in Thailand (Perera et al.
2017). Delonicicola species are saprobes, characterized by pseudostromatal immersed, papillate
ascomata, short pedicellate asci and 1-septate, hyaline ascospores (Perera et al. 2017). Delonicicola
siamense is illustrated in this entry.
Figure 90 – Delonicicola siamense (Material examined – THAILAND, Chiang Rai Province, Mae
Fah Luang University premises, on dried seed pods of Delonix regia (Boj.) Raf. (Fabaceae), 2 June
2015, R.H. Perera RHP 92, MFLU 17-0739, holotype). a Herbarium material. b Appearance of
ascomata on host substrate. c, d Vertical section through pseudostroma with ascoma. e Section of
the peridium. f Paraphyses. g-j Asci. k-o Ascospores. Scale bars: b = 2 mm, c = 200 μm, d = 100
μm, e-j = 20 μm, k-o = 5 μm.
Furfurella Voglmayr & Jaklitsch, Persoonia 42: 237 (2019)
Index Fungorum number: IF829925; 3 species with sequence data.
Type species – Furfurella stromatica Voglmayr & Jaklitsch
Notes – Furfurella was introduced by Voglmayr et al. (2019a) to accommodate F.
luteostiolata, F. nigrescens and F. stromatica (type species). Furfurella species are associated with
dead branches of fabaceous plants in the Mediterranean basin (Voglmayr et al. 2019a). The genus
is characterised by erumpent pseudostromata with bright yellow, yellow-green to turquoise scurfs,
clavate to cylindrical asci and allantoid, aseptate ascospores.
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Diaporthaceae Höhn., Am. J. Bot. 13: 638 (1926)
Index Fungorum number: IF80691; Facesoffungi number: FoF01383; 497 species.
Endophytic, pathogenic or saprobic on terrestrial and rarely submerged plants. Sexual morph:
Pseudostromata well or poorly developed, erumpent, pulvinate, slightly convex or flat, circular,
orbicular or irregular, coriaceous, sclerotioid, whitish or brownish to black, with or lacking black
zone or a crust containing of fungus tissue, solitary or having up to 10 ascomata in a stroma.
Ectostromatic disk subhyaline to hyaline or brown. Ascomata immersed to erumpent, perithecial,
globose or compressed, solitary or aggregated in a valsoid shape, black, coriaceous, ostiolate,
papillate. Papilla short or long, black, convergent, erumpent, conical or cylindrical, internal wall
concealed by hyaline periphyses, composed of vertically organized parenchymatous tissues.
Peridium covering exterior layer of thick-walled, flattened, dark-brown cells of textura angularis
and inner, thin-walled, hyaline cells of textura angularis. Paraphyses unbranched, septate,
cylindrical. Asci 8-spored, clavate, oblong-clavate to broadly fusoid, unitunicate, sessile, with a
distinct apical ring. Ascospores biseriate to partially biseriate, hyaline, dark brown, ellipsoid,
oblong to fusoid, unicellular or one septate, constricted at septum, with or without appendages at
both ends, occasionally narrowly rounded at ends, multi-guttulate, smooth-walled. Asexual morph:
Coelomycetous. Conidiomata globose, acervular or pycnidial, initially immersed, erumpent when
mature, black, solitary, coriaceous, scattered, elongated ostiolar necks, sometime becoming multiloculate with one to numerous noticeably demarcated black necks extending over the stroma,
frequently with yellowish conidial mass extruding from ostiole. Peridium comprising 3–4 layers of
light brown cells of textura intricata to textura angularis. Conidiophores dimorphic. Alpha
conidiophores subcylindrical, firmly aggregated, branched in middle area, comprising 2–3
associate cells, ampulliform, giving rise to septate, cylindrical to irregular conidiogenous cells or
paraphyses, straight to sinuous, smooth, 1–5-septate, hyaline to pale brown, cylindrical, branched
merely at the base, formed from the inner most layer cells of the conidiomata wall, sometime
terminal and lateral, apex with minute periclinal thickening and collarette. Beta conidiophores
hyaline, scattered between alpha conidiophores, subcylindrical, branched, 1–3-septate. Alpha
Conidiogenous cells phialidic, enteroblastic, cylindrical or subcylindrical, terminal and lateral,
slightly tapering towards the apex or sometimes apex with minute periclinal thickening and
collarette. Beta conidiogenous cells integrated, phialidic, terminal and lateral. Alpha conidia
hyaline, fusiform to ovate, subcylindrical to narrowly ellipsoid, straight or curved, occasionally
irregular, smooth-walled, abundant, 0–2-septate, apex obtuse, base truncate to sub-truncate, straight
to curved, occasionally slightly sigmoid, pale to medium brown, with many guttules, sometimes
short, bear appendages at both ends. Beta conidia hyaline, aseptate, smooth, subcylindrical, straight
to slightly curved, fusiform to hooked, base sub-truncate, occasionally widest in middle, tapering to
acutely rounded apex, truncate at base.
Type genus – Diaporthe Nitschke
Notes – Diaporthaceae was introduced and placed in Diaporthales by Höhnel (1917d) and it
encompasses numerous endophytic and phytopathogenic fungal species. Wehmeyer (1975)
confined two genera, Diaporthe and Mazzantia to this family. However, Barr (1978) synonymized
Diaporthaceae in Valsaceae. Castlebury et al. (2002) presented the distinct placement of
Diaporthaceae in Diaporthales, forming an analyzed LSU sequence data of diaporthoid taxa in a
well-supported clade. Previously, only Diaporthe (Phomopsis) and Mazzantia were accommodated
in Diaporthaceae based on phylogenetic analysis (Castlebury et al. 2002). Nevertheless,
Apioporthella and Leucodiaporthe were included in this family by Lumbsch & Huhndorf (2010).
Lamprecht et al. (2011) showed the phylogenetic placement of Stenocarpella and Phaeocytostroma
in Diaporthaceae using LSU sequences analysis. Based on combined gene analysis of LSU, SSU
and tef1 sequence data, Dai et al. (2014b) introduced Pustulomyces. Phylogenetic placement of
Phaeodiaporthe in Diaporthaceae was confirmed by Voglmayr & Jaklitsch (2014).
Maharachchikumbura et al. (2015) listed Allantoporthe, Apioporthella, Clypeoporthella,
Diaporthe,
Diaporthella,
Diaporthopsis,
Leucodiaporthe,
Mazzantia,
Mazzantiella,
Ophiodiaporthe and Pustulomyces as genera of Diaporthaceae based on an analysis of LSU
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sequence data. Based on greater usage of the name Mazzantia, Rossman et al. (2015) synonymized
Mazzantiella under Mazzantia. Clypeoporthella was established on C. brencklei Petr., and a newly
collected C. brencklei (BPI 843482) specimen was developed in culture and sequenced. It has a
Phomopsis asexual morph and DNA sequence data disclosed that C. brencklei clustered together
with Diaporthe. Thus, Clypeoporthella is regarded as a synonym of Diaporthe (Sogonov et al.
2008). Diaporthopsis was introduced to accommodate species that are similar to Diaporthe, with
unicellular ascospores and was typified by D. angelicae. Molecular analysis of LSU sequence data
showed that D. angelicae clustered within the Diaporthe. In addition, Diaporthopsis angelicae has
similar characters of Diaporthe such as stromata, perithecia, and centrum to species. Diaporthopsis
was synonymized under Diaporthe based on morphology and molecular data (Castlebury et al.
2002, Gomes et al. 2013). Diaporthella is comprised of aggregated perithecia within welldeveloped stromata and median, 1-septate ascospores. Diaporthella corylina is parasitic and causes
dieback of Corylus stems. Diaporthella corylina shows similar characters to Anisogramma
anomala morphologically. Anisogramma belongs in the Gnomoniaceae based on A. virgultorum
(Castlebury et al. 2002, Vasilyeva et al. 2007). Based on combined LSU, ITS, rpb2 and tef1 gene
analysis, Senanayake et al. (2017a) confirmed the phylogenetic placement of Diaporthella outside
of Diaporthaceae, and it does not show affinities with any families in Diaporthales. A recent study
by Senanayake et al. (2017a) accepted a few genera within this family in addition to
Maharachchikumbura et al. (2016b) by introducing three new genera: Chiangraiomyces,
Hyaliappendispora and Paradiaporthe based on morphology and phylogeny. Massariothea was
added to the family by Thambugala & Hyde (2018). Therefore 15 genera: Apioporthella,
Apiosphaeria, Chaetoconis, Chiangraiomyces, Diaporthe, Hyaliappendispora, Leucodiaporthe,
Massariothea, Mazzantia, Ophiodiaporthe, Paradiaporthe, Phaeocytostroma, Phaeodiaporthe,
Pustulomyces and Stenocarpella are accepted in Diaporthaceae (Senanayake et al. 2018).
Clypeoporthe, Cryptonectriella, Kensinjia, Lollipopaia and Skottsbergiella were listed in
Diaporthaceae by Wijawawardene et al. (2017). Clypeoporthe was introduced and is typified by C.
monocarpa. There are five species listed under this genus (Species fungorum 2020). However,
some species in this genus have eutypelloid configuration of ascomata in parenchymatous stromatic
tissues. Clypeoporthe was reduced to synonymy under Gnomonia by Monod (1983), while Kirk et
al. (2008) mentioned Clypeoporthe is the sexual morph of Phaeocytostroma. However, the latter is
not proven by culture or molecular data and it is necessary to obtain DNA sequence data to resolve
this genus. Therefore, Senanayake et al. (2017a, 2018) and Wijayawardhene et al. (2018) accepted
this genus in Gnomoniaceae. Cryptonectriella (Höhn.) Weese (≡ Cryptonectriopsis (Höhn.) Weese)
was introduced and is typified by C. biparasitica and a second species C. geoglossi (Species
Fungorum 2020). Weese (1919) accommodated this genus in Hypocreales. Kensinjia was
introduced and is typified by K. umbrina by Reid & Booth (1989). However this species was
synonymized under Cryptosporella as C. umbrina (Jenkins) Jenkins & Wehm., which is a genus in
Gnomoniaceae. The monotypic genus Lollipopaia was introduced and typified by L. minuta and
accommodated in Diaporthales genera incertae sedis (Inderbitzin and Berbee 2001). There are only
two nrSSU sequences and blast searches in GenBank show L. minuta is closely related to
economically important plant pathogens in Diaporthaceae. However our phylogenetic analysis
(unpublished) shows that L. minuta clusters away from Diaporthaceae; but within Diaporthales.
Lollipopaia minuta is somewhat different from taxa of Diaporthaceae in having solitary to
aggregated, carbonaceous ascomata with long, slender necks and long, filiform, multiseptate
ascospores. Senanayake et al. (2017a, 2018) accepted Lollipopaia within Diaporthales genera
incertae sedis based on morphology and here we follow this. Skottsbergiella was introduced and
typified by Skottsbergiella diaporthoides which has large perithecia immersed in massive,
externally crustose, pseudoparenchymatous stromata. Petrak (1971) assigned this genus to eutypoid
fungi based on its stromata. This genus is morphologically similar to Diaporthella, which is placed
in Diaporthales genera incertae sedis (Barr 1978). However, Skottsbergiella diaporthoides was
synonymized under Diaporthe diaporthoides (Barr 1978) and accommodated in Diaporthaceae.
This was followed by Senanayake et al. (2017a, 2018).
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Ecological and economic significance of Diaporthaceae
Species of Diaporthaceae includes a number of plant pathogens of major importance causing
diseases of a wide variety of plants (Dissanayake et al. 2017, Senanayake et al. 2018). Fruit
production is especially affected (Dissanayake et al. 2017, Senanayake et al. 2018).
Genera included in Diaporthaceae
Apioporthella Petr., Annls mycol. 27(5/6): 401 (1929)
Index Fungorum number: IF0260; 1 morphological species.
Type species – Apioporthella vepris (Lacroix) M.E. Barr
Notes – Apioporthella was described and typified by A. bavarica. Barr (1991) proposed two
new combinations as A. apiospora and A. vepris. The genus is characterised by an ectostromatic
disc which appears as a cushion of brown cells containing one or a few fused ostioles.
Apiosphaeria Höhn., Sber. Akad. Wiss. Wien, Math.-naturw. Kl., Abt. 1 118: 1218 (1909)
Index Fungorum number: IF7566; 1 species with sequence data.
Type species – Apiosphaeria guaranitica (Speg.) Höhn.
Notes – Apiosphaeria has been traditionally placed in Phyllachoraceae, based exclusively on
morphological studies, without supporting molecular evidence. Guterres et al. (2018) provided
molecular data for the link between sexual and asexual states of the fungus and elucidate the
phylogeny of A. guaranitica. Their results indicate a natural placement of Apiosphaeria within
Diaporthaceae, where it represents an ancient lineage of the crown group of Diaporthaceae.
Chaetoconis Clem., Gen. fung. (Minneapolis): 125 (1909)
Index Fungorum number: IF7566; 2 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Chaetoconis polygoni (Ellis & Everh.) Clem.
Notes – Chaetoconis comprises two species: C. polygoni and C. vaccinii (Index Fungorum
2020). De Gruyter et al. (2009) and Senanayake et al. (2017a) accommodated Chaetoconis in
Diaporthales genera incertae sedis and Diaporthaceae respectively, based on sequence analysis.
The genus is characterised by acropleurogenous conidia which form from the inner pycnidial wall
cells.
Chiangraiomyces Senan. & K.D. Hyde, Stud. Mycol. 86: 217 (2017)
Index Fungorum number: IF821544; 1 species with sequence data.
Type species – Chiangraiomyces bauhiniae Senan. & K.D. Hyde
Notes – Based on multigene phylogeny and morphology, Chiangraiomyces was introduced
by Senanayake et al. (2017a). The genus is characterised by immersed, solitary ascomata with
fusiform asci and a J-, funnel-shaped, apical ring. Oval to fusiform ascospores have two large
central guttules and two small marginal guttules.
Diaporthe Nitschke, Pyrenomyc. Germ. 2: 240 (1870)
Index Fungorum number: IF1497; 450 morphological species (Species Fungorum 2020), 261
species with sequence data.
Type species – Diaporthe eres Nitschke
Notes – Diaporthe is the type genus of Diaporthaceae, and it was established by Nitschke
(1867) to accommodate taxa with stromata in the Sphaeriales which are often with blackened zones
in the substrate, ellipsoid to fusiform ascospores and enclosed, unilocular pycnidia that contain
spermatia, stylospores and conidia (Wehmeyer 1933a). Diaporthe species are saprobes, pathogens
and endophytes and have a worldwide distribution (Dissanayake et al. 2017). Phomopsis was
earlier the asexual morph (Udayanga et al. 2012). However, these genera were linked based on
principle of significance and Diaporthe was nominated over Phomopsis to resolve nomenclatural
complications (Rossman et al. 2015). The genus contains 1067 epithets listed in Index Fungorum
562
(August 2019), but fewer than 200 species are accepted (Dissanayake et al. 2017). Diaporthe
asheicola, D. eucalyptorum, and D. salsuginosa are illustrated in this entry (Figs. 91, 92, 93, 94
respectively).
Figure 91 – Diaporthe asheicola (Material examined – RUSSIA, Rostov region, Rostov-on-Don
City Botanical Garden of Southern Federal University, on stem of Fraxinus pennsylvanica
(Oleaceae), 30 May 2015, T.S. Bulgakov T823, MFLU 15-2966). a, b Conidiomata on host surface.
b, c Vertical section of ascomata (semi-immersed to immersed on decaying wood). d Peridium. e
Paraphyses. f-l Immature and mature asci. m-q Ascospores with bipolar pad-like appendages. r
Germinating ascospore. Scale bars: b, d = 50 μm, c, e-r = 10 μm.
563
Figure 92 – Diaporthe eucalyptorum (Material examined – INDIA, Andaman and Nicobar Islands,
South Andaman, Manjery, (11˚52’68.2.0”N 92˚64’74.9”E), isolated from an unidentified twig, 17
May 2018, M. Niranjan & V.V. Sarma PUFNI 17624; living culture, NFCC-4374). a, b Ascomata
on host surface. c Section of ascomata. d, f Peridium. e Neck. g-k Asci. l, m Ascospores. n, o
Colonies on MEA (n-from above, o-below). Scale bars: c =100 μm, d-f = 50 μm, g-m = 10 μm.
564
Figure 93 – Diaporthe salsuginosa (Material examined – INDIA, Tamil Nadu, Parangipettai
mangroves (11.59°N 79.5°E), on decaying wood of Avicennia marina (Acanthaceae), 23 April
2018, B. Devadatha AMH-10013; living culture, NFCCI-4385). a Necks in crater-like depressions
on the host surface. b Vertical section of ascomata (semi-immersed to immersed on decaying
wood). b, d Ascomata. e Peridium. f Paraphyses. c, g-l Immature and mature asci. m-q Ascospores
with bipolar pad-like appendages. r Germinating ascospore. Scale bars: b-d = 50 μm, e-r = 10 μm.
565
Figure 94 –Diaporthe salsuginosa, from living culture (Material examined – INDIA, Tamil Nadu,
Parangipettai mangroves (11.59°N 79.5°E), on decaying wood of Avicennia marina (Acanthaceae),
23 April 2018, B. Devadatha AMH-10013; living culture, NFCCI-4385). a, b Front and reverse
view of culture. c Conidiomata producing hyaline conidial mass grown from the mature colonies.
d-f Vertical section through conidioma. g Peridium. h Conidiophores and conidiogenous cells i-l
Beta conidia. Scale bars: b-c = 100 μm, d = 50 μm, e-l = 10 μm.
566
Hyaliappendispora Senan., Camporesi & K.D. Hyde, Stud. Mycol. 86: 217 (2017)
Index Fungorum number: IF821548; 1 species with sequence data.
Type species – Hyaliappendispora galii Senan., Camporesi & K.D. Hyde
Notes – The genus is characterised by biguttulate, uniseptate, hyaline ascospores with long
filamentous apical and basal appendages and wall of the appendages makes a ring-like
ornamentation at the proximal end.
Leucodiaporthe M.E. Barr & Lar. N. Vassiljeva, Mycologia 99(6): 917 (2008)
Index Fungorum number: IF510561; 4 morphological species (Species Fungorum 2020).
Type species – Leucodiaporthe acerina M.E. Barr & Lar.N. Vassiljeva
Notes – This genus is distinct in the Diaporthales in having a pale to dark stromatic disk with
blackened marginal zones. Its hyaline ascospores occasionally have a gelatinous coat.
Massariothea Syd., Annls mycol. 37(3): 249 (1939)
Index Fungorum number: IF8855; 8 morphological species (Species Fungorum 2020), 2
species with sequence data.
Type species – Massariothea themedae Syd.
Notes – The genus is characterised by brown, distoseptate conidia with species commonly
reported on grasses (Poaceae) with a worldwide distribution (Thambugala et al. 2018).
Mazzantia Mont., Bull. Soc. bot. Fr. 2: 525 (1855)
Index Fungorum number: IF3036; 17 morphological species (Species Fungorum 2020), 2
species with sequence data.
Type species – Mazzantia galii (Fr.) Mont.
Notes – The genus is characterised by a sharply delimited, strongly melanized, sclerotial,
clypeal ectostroma, a whitish entostroma, and aseptate ascospores.
Ophiodiaporthe Y.M. Ju, H.M. Hsieh, C.H. Fu, C.Y. Chen & T.T. Chang, Mycologia 105(4): 866
(2013)
Index Fungorum number: IF801562; 1 species with sequence data.
Type species – Ophiodiaporthe cyatheae Y.M. Ju, H.M. Hsieh, C.H. Fu, Chi Y. Chen & T.T.
Chang
Notes – Based on its unique asexual characters and host preference, Ophiodiaporthe was
introduced and typified by O. cyatheae (Fu et al. 2013). The genus is characterised by its unique
host preference on Cyathea lepifera.
Paradiaporthe Senan., Camporesi & K.D. Hyde, Stud. Mycol. 86: 217 (2017)
Index Fungorum number: IF821546; 1 species with sequence data.
Type species – Paradiaporthe artemisiae Senan., Camporesi & K.D. Hyde
Notes – Paradiaporthe is morphologically comparable to Diaporthe. However,
phylogenetically, P. artemisiae is divergent from other genera in Diaporthaceae (Senanayake et al.
2017a). The genus is characterised by erumpent, solitary ascomata with prominent, wide papilla.
Phaeocytostroma Petr. Annls. mycol. 19(1/2): 45 (1921)
Index Fungorum number: IF9298; 6 morphological species (Species Fungorum 2020), 4
species with sequence data.
Type species – Phaeocytostroma ambiguum (Mont.) Petr.
Notes – Several molecular studies have established the taxonomic placement of
Phaeocytostroma within Diaporthaceae (Lamprecht et al. 2011, Verkley et al. 2014). The genus is
characterised by their aseptate, brown, eguttulate, cylindrical or ellipsoid, macroconidia with
truncate base and fusiform, short or elongate microconidia.
567
Phaeodiaporthe Petr. Annls mycol. 17(2/6): 99 (1920)
Index Fungorum number: IF3906; 1 morphological species.
Type species – Phaeodiaporthe appendiculata (G.H. Otth) Lar.N. Vassiljeva
Notes – Phaeodiaporthe was introduced based on P. keissleri and Petrak (1921) recognized it
as a synonym of Diaporthe appendiculata. Voglmayr & Jaklitsch (2014) epitypified P.
appendiculata and synonymized P. keissleri under P. appendiculata. The genus is characterised by
large brown ascospores.
Pustulomyces D.Q. Dai, Bhat & K.D. Hyde, Cryptog. Mycol. 35(1): 68 (2014)
Index Fungorum number: IF806063; 1 species with sequence data.
Type species – Pustulomyces bambusicola D.Q. Dai et al.
Notes – The monospecific genus Pustulomyces shows similar characters to Bambusicola.
However, phylogenetically of Pustulomyces bambusicola placed in Diaporthaceae (Dai et al.
2014b) and Bambusicola in Bambusicolaceae have distinct phylogenetically placements (Hyde et
al. 2013). The genus is similar to Bambusicola in Bambusicolaceae with cylindrical, phialidic,
conidiogenous cells and fusiform, straight to curved conidia.
Stenocarpella Syd. & P. Syd. Annls mycol. 15(3/4): 258 (1917)
Index Fungorum number: IF10085; 2 species with sequence data.
Type species – Stenocarpella macrospora (Earle) B. Sutton
Notes – Both Stenocarpella macrospora and S. maydis are well-known pathogens of corn, the
causal agents of stalk and ear rot and leaf spot (Latterell & Rossi 1983, De Silva et al. 2014). Based
on LSU sequence data, Stenocarpella was accommodated in Diaporthaceae (Crous et al. 2006a, b).
Lamprecht et al. (2011) and Wijayawardene et al. (2016b) confirmed its placement in
Diaporthaceae. The genus is characterised by brown, 0–3-septate conidia.
Diaporthosporellaceae C.M. Tian & Q. Yang, Mycoscience 59: 229–235 (2017)
Index Fungorum number: IF822663; Facesoffungi number: FoF05684; 1 species.
Pathogenic on branches and twigs of Cercis chinensis. Sexual morph: Ascomata perithecial,
immersed to erumpent, globose to subglobose, dark brown to black, penetrating through
ectostroma, convergent to disc. Paraphyses filiform, hyaline, unbranched, deliquescent at maturity.
Asci 8-spored, unitunicate, clavate to oblong-clavate, floating freely in the centrum, apical ring
become unclear in dried specimens. Ascospores irregularly uniseriate, allantoid or sub-reniform, 34-guttulate, hyaline, aseptate. Asexual morph: Coelomycetous. Conidiomata pycnidial, immersed,
erumpent at maturity, globose to subglobose, yellowish conidial mass release from ostiole.
Ectostromatic disc black, one ostiole for each disc. Conidiophores acropleurogenous, branched or
sympodially branched, cylindrical, aseptate. Conidiogenous cells enteroblastic, phialidic,
cylindrical, terminal, slightly tapering towards the apex. Conidia ellipsoidal, aseptate, hyaline,
slightly rounded at each end (adapted from Yang et al. 2018d, Senanayake et al. 2018).
Type genus – Diaporthosporella C.M. Tian & Q. Yang
Notes – Diaporthosporellaceae was introduced by Yang et al. (2018d) to accommodate
Diaporthosporella. Morphology and phylogeny of Diaporthosporellaceae is different from other
families of Diaporthales. Diaporthosporellaceae is characterized by unicellular ascospores,
branched or sympodially branched conidiophores, and hyaline, biguttulate conidia (Yang et al.
2018d, Senanayake et al. 2018). Diaporthosporellaceae comprises only one genus and one species.
Diaporthosporellaceae was found on branches and twigs of Cercis chinensis (Fabaceae) in China.
Sequence data are available in GenBank (Yang et al. 2018d, Fan et al. 2018).
Ecological and economic significance of Diaporthosporellaceae
Diaporthosporellaceae species play a major role as plant pathogens on branches and twigs of
Cercis chinensis (Yang et al. 2018d). Cercis chinensis is widely cultivated as an ornamental plant
568
(Na et al. 2009). Some parts of C. chinensis, such as stem, and root bark have been used to increase
blood circulation and to treat bruising and injuries (Na et al. 2009).
Figure 95 – Diaporthosporella cercidicola (Material examined – CHINA, Jiangsu Province,
Nanjing City, Xuanwu lake scenic area, on twigs and branches of Cercis chinensis, 11 Nov. 2015,
Q. Yang, BJFC-S1356, holotype). a Appearance on host. b-d Stromata on host. e Vertical section
through stroma. f Horizontal section through stroma. g Asci with ascospores and and paraphyses. h
Conidiophores with conidia. i Conidia. Scale bars: b-f = 200 µm, g-i = 10 µm.
Genus included in Diaporthosporellaceae
Diaporthosporella C.M. Tian & Q. Yang; Mycoscience 59: 229–235 (2017)
Index Fungorum number: IF822664; 1 species with sequence data.
Type species – Diaporthosporella cercidicola C.M. Tian & Q. Yang
Notes – Diaporthosporella was introduced by Yang et al. (2018d). Diaporthosporella
cercidicola is characterized by hyaline, allantoid or sub-reniform, aseptate ascospores and
acropleurogenous, branched or sympodially branched conidiophores, producing hyaline,
ellipsoidal, aseptate and biguttulate conidia (Yang et al. 2018d). Diaporthosporella comprises a
single species found on branches and twigs of Cercis chinensis in China (Yang et al. 2018d).
Diaporthosporella cercidicola is similar to Diaporthe eres Nitschke in having unitunicate, clavate
asci, hyaline, guttulate ascospores, but can be separated by phylogeny (Yang et al. 2018d). The
asexual morph produces a yellowish conidial mass and aseptate, hyaline conidia, characters that are
common to both Diaporthosporella cercidicola and Diaporthe eres (Udayanga et al. 2014, Yang et
al. 2018d). Diaporthosporella cercidicola is closely related to Juglanconidaceae according to
molecular phylogeny (Voglmayr et al. 2017).
569
Diaporthostomataceae X.L. Fan & C.M. Tian, Persoonia 40: 124 (2018)
Index Fungorum number: IF823983; Facesoffungi number: FoF05685; 1 species.
Pathogenic on twigs and branches of Machilus leptophylla. Sexual morph: Pseudostromata
immersed, slightly erumpent. Ectostromatic disc ovoid to ellipsoid, yellowish to dark grey. Central
column beneath the disc more or less conical. Stromatic zones lacking. Perithecia conical,
surrounding the ectostromatic disc. Ostiole single, dark grey to black. Paraphyses deliquescent.
Asci 8-spored, apical ring. Ascospores 2–3-seriate, hyaline, fusoid, bicellular, multiguttulate.
Asexual morph: Undetermined (adapted from Fan et al. 2018, Senanayake et al. 2018).
Type genus – Diaporthostoma X.L. Fan & C.M. Tian
Notes – Diaporthostomataceae was introduced by Fan et al. (2018) to accommodate the
monophyletic genus Diaporthostoma. The species of Diaporthostomataceae are characterized by
conical and separate perithecia and bicellular, fusoid, straight to curved ascospores with a median
septum (Fan et al. 2018, Senanayake et al. 2018). Multi-gene phylogenetic analysis revealed that
the family is monophyletic and sister to Diaporthosporellaceae (Senanayake et al. 2018).
Diaporthostomataceae species are morphologically distinct from Diaporthosporellaceae, in having
discrete perithecia and fusoid, straight to curved ascospores with a median septum (Fan et al. 2018,
Yang et al. 2018d). Diaporthostoma machili is the only species in Diaporthostomataceae and it was
collected on twigs and branches of Machilus leptophylla from China. Sequence data are available
in GenBank (Fan et al. 2018).
Ecological and economic significance of Diaporthostomataceae
Diaporthostomataceae species are plant pathogens on twigs and branches of Machilus
leptophylla causing cankers or dieback (Fan et al. 2018).
Genus included in Diaporthostomataceae
Diaporthostoma X.L. Fan & C.M. Tian, Persoonia 40: 124 (2018)
Index Fungorum number: IF823984; 1 species with sequence data.
Type species – Diaporthostoma machili X.L. Fan & C.M. Tian
Notes – Diaporthostoma was introduced by Fan et al. (2018) to accommodate
Diaporthostoma machili as the type species, which was collected from twigs and branches of
Machilus leptophylla in China. Diaporthostoma machili is characterized by scattered, conical
perithecia and fusoid, straight to curved ascospores with a median septum (Fan et al. 2018,
Senanayake et al. 2018). No asexual morph is known for this species. Diaporthostoma comprises
only the type species (Index Fungorum 2020). In this entry we illustrate Diaporthostoma machili.
Diatrypaceae Nitschke, Verh. naturh. Ver. preuss. Rheinl. 26: 73 (1869)
Index Fungorum number: IF80692; Facesoffungi number: FoF00679; 599 species.
Saprobic or pathogenic on woody plants. Sexual morph: Stromata eustromatic or
pseudostromatic, well-developed, immersed to erumpent, rarely superficial, mostly black or dark
brown, with somewhat carbonaceous outer layer, inner layer pale, loosely packed, parenchymatous.
Ascomata perithecial, immersed in stromatic tissues, mostly brown to black, globose to subglobose,
with ostiolar necks. Ostioles sulcate, inner layer covered with hyaline, periphyses. Peridium
consists of two layers, an inner hyaline layer and an outer layer of brown to black cells of textura
angularis. Paraphyses long, wide, branched, septate. Asci 8-spored or polysporous, rarely 1-spored
or 2-spored, unitunicate, cylindrical, clavate to pyriform, fusiform, with a very long pedicel, with a
truncate apex, with a J-, or J+, apical ring. Ascospores crowded, most hyaline to light brown, rarely
jet-black, allantoid, ellipsoidal, globose or filiform. Asexual morph: Coelomycetous, astromatic,
occurring on host asacervuli. Conidiomata acervuli sub cortical, erumpent, yellow to red, with
branched conidiophores and in culture as pycnidia, superficial, solitary or aggregated, yellow, dark
brown to black, subconical, globose to subglobose, and thick peridium, comprising brown, thickwalled cells of textura angularis with branched conidiophores, arising from pseudoparenchymatous
cells or interwoven hyphae. Conidiogenous cells in dense palisades, cylindrical, straight or curved,
570
apically distorted or annulated. Conidia filiform, curved, or rarely straight with flattened base and
blunt apex, hyaline (adapted from Maharachchikumbura et al. 2016b).
Figure 96 – Diaporthostoma machili (Material examined – CHINA, Zhejiang Province, Hangzhou
City, Linan, Tianmu Mountain, N30°19'18.21" E119°26'18.21", 354 m asl, on twigs and branches
of Machilus leptophylla, 20 April 2017, Q. Yang & Z. Du, CF 2017475, holotype). a, b
Pseudostromata on substrate. c Transverse section of pseudostroma. d Longitudinal section of
pseudostroma. e-h Asci. i-k Ascospores. Scale bars: a = 1 mm, b, c = 200 µm, d = 100 µm, e-h = 10
µm, i-k = 5 µm
Type genus – Diatrype Fr.
Notes – Diatrypaceae was introduced by Nitschke (1869) and is typified by Diatrype.
Diatrypaceae is a well-studied group. Recent studies of Diatrypaceae include de Almeida et al.
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(2016), Maharachchikumbura et al. (2016b), Senwanna et al. (2017) and Shang et al. (2017, 2018).
Hongsanan et al. (2017) provided a phylogenetic analysis and MCC tree with good support in
Xylariomycetidae for the family, where it occurs in Xylariales (Fig. 4). Neoeutypella was
introduced by Phookamsak et al. (2019) based on the phylogenetic analyses of a combined ITS and
tub2 sequence dataset and its unique morphology. Halocryptovalsa was introduced by Dayarathne
et al. (2019b) from marine habitats.
Ecological and economic significance of Diatrypaceae
Diatrypaceae are of great significance as pathogens of economic crops and forest trees
(Trouillas & Gubler 2010, Urbez-Torres et al. 2012, Senanayake et al. 2015). Species of
Anthostoma, Cryptosphaeria, Cryptovalsa, Diatrype, Diatrypella and Eutypella have been reported
as disease agents causing grapevine cankers in grape growing countries worldwide (Mostert et al.
2004, Trouillas et al. 2010b, 2011, Urbez-Torres et al. 2012, Paolinelli-Alfonso et al. 2015, Moyo
et al. 2017, Agustí-Brisach et al. 2019, Barna & Mihál 2019). Peroneutypa scoparia and Eutypella
sp. can produce ligninolytic enzymes and as endophytes or saprobes of decaying wood, they play
an important role in nutrient cycling ( de Errasti et al. 2014, Grassi 2014).
Genera included in Diatrypaceae
Allocryptovalsa Senwanna, Phookamsak & K.D. Hyde, Mycosphere 8(10): 1839 (2017)
Index Fungorum number: IF553857; 2 species with sequence data.
Type species – Allocryptovalsa polyspora Senwanna, Phookamsak & K.D. Hyde
Notes – Allocryptovalsa was introduced by Senwanna et al. (2017) to accommodate A.
polyspora and two combined species: A. cryptovalsoidea, which was transferred from Eutypella
and Allocryptovalsa rabenhorstii, which was transferred from Cryptovalsa. The genus is
characterised by stromata immersed in host tissues, polysporous asci and allantoid ascospores.
However, the combined species A. cryptovalsoidea is invalid according to the basionym of this
species, Eutypella cryptovalsoidea is invalid because Trouillas et al. (2011) designated holotype
with three collections when they introduced E. cryptovalsoidea. Therefore, the current status of
Allocryptovalsa cryptovalsoidea is further needed to be resolved. In this study, we introduce a new
species Allocryptovalsa truncata based on morphology and phylogeny support.
Allocryptovalsa truncata M. Niranjan & V.V. Sarma, sp. nov.
Fig. 97
Index Fungorum number: IF556619; Facesoffungi number: FoF06262
Etymology – Refers to the truncate shape of the ascal apex.
Holotypus – AMH-9983.
Saprobic on decaying twigs. Sexual morph: Stromata superficial, in irregular clusters,
beneath the periderm, outer black thin layer, inner white powdery coat around each ascoma.
Ascomata 280–400 × 470–620 μm, immersed in pseudostroma, perithecial, coriaceous, subglobose,
aggregated, raised individually, ostiolate, periphysate. Ostioles 220–250 × 190–220 μm. Peridium
outer brown and inner hyaline layers with cells of textura angularis. Paraphyses septate,
unbranched, guttulate, sparsely present. Asci (84–) 89–117(–122) × (10–)12–15(–16.5) μm ( = 103
× 13.9, n = 25), polysporous, unitunicate, clavate, apically obtuse, long pedicellate, apices truncate,
broader in the upper region, narrowing towards lower region, with a J-, apical ring, persistent.
Ascospores (7)7.2–10.7 × 2.1–2.7 μm ( = 8.7 × 2.4, n = 25), hyaline to brown, allantoid, smoothwalled. Asexual morph: Undetermined.
Culture characteristics – Colonies on malt extract agar (MEA) 46 mm diam. in 4 days at
28°C, white colony, radial, filamentous, raised in center, reverse center brown and white towards
edges.
Material examined – INDIA, Andaman and Nicobar Islands, South Andaman, Chidiyatapu
(11˚51’90.4” N 92˚65’23.8” E), decaying twig, 10 December 2017, M. Niranjan & V.V. Sarma,
AMH-9983, holotype; ex-type living cultures, NFCCI-4520.
GenBank numbers – ITS: MK990279, LSU: MK981538, SSU: MK981535.
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Notes – Allocryptovalsa truncata has narrower ascomata and smaller asci and ascospores
than the type species, A. polyspora. A unique feature of the new taxon is the ascal apices that are
apically, predominantly truncate in contrast to A. polyspora that has rounded apical ends. In the
phylogeny (Fig. 4), taxa from Anthostoma, Cryptosphaeria, Cryptovalsa, Diatrype and Eutypella
clustered with our new taxon and A. polyspora (strain MFLUC17-0364) nested with our taxon with
strong boostrap support (99% in ML).
Anthostoma Nitschke, Pyrenomyc. Germ. 1: 110 (1867)
Index Fungorum number: IF224; 36 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Anthostoma decipiens (DC.) Nitschke
Notes – The holomorphs of A. decipiens were studied by Rappaz (1992, 1993) and concluded
the genus belongs in Diatrypaceae. Phylogenetic analyses of sequence data suppoted these results
(Rocchi et al. 2010, Jaklitsch et al. 2014). The genus is characterised by dark brown to dark,
globose to subglobose ascomata, immersed in a stroma, with cylindrical, prominent ostioles,
cylindrical to clavate asci, with apically rounded to truncate apices, with a short pedicel and brown
to black brown ascospores (Nitschke 1867).
Cryptosphaeria Ces. & De Not., Comm. Soc. crittog. Ital. 1(4): 231 (1863)
Index Fungorum number: IF26092; 16 morphological species (Species Fungorum 2020), 5
species with sequence data.
Type species – Cryptosphaeria millepunctata Grev.
Notes – Cryptosphaeria has been widely accepted in Diatrypaceae (Nitschke 1867, 1870,
Clement & Shear 1931, Rappaz 1989, Trouillas et al. 2010b, 2015). The genus comprises
corticolous species and is characterised by widely effuse and poorly developed stromata, often
covered by a periderm, which is penetrated only by the separately emerging ostioles, generally 8spored, spindle-shaped, long pedicellate asci and sub-olivaceous to brown ascospores (Glawe 1984,
Rappaz 1987).
Cryptovalsa Ces. & De Not., Jb. nassau. Ver. Naturk. 23-24: 212 (1870)
Index Fungorum number: IF1340; 29 morphological species (Species Fungorum 2020), 4
species with sequence data.
Type species – Cryptovalsa protracta (Pers.) De Not.
Notes – Cryptovalsa was established by Cesati & De Notaris to accommodate C. protracta,
C. ampelina, C. nitschkei and C. effusa. The genus is characterised by eutypoid stromata that are
rather variable, when erumpent separately diatrypelloid, often immersed in wood, but sometimes
invading bark tissues. Asci are cylindrical or clavate, polysporous, with short or long pedicels.
Ascospores are crowded, allantoid and yellowish (Spooner 1981, Vasilyeva & Stephenson 2005).
Diatrypasimilis J.J. Zhou & Kohlm., Index Fungorum 305: 1 (2016)
Index Fungorum number: IF552473; 1 species with sequence data.
Type species – Diatrypasimilis australiensis J.J. Zhou & Kohlm.
Notes – Diatrypasimilis was established by Chalkley et al. (2010) based on conventional
taxonomic criteria and molecular phylogeny to accommodate a species from mangroves. However,
it was shown to be an invalid name and was legitimated as a nomenclatural novelty in Suh (2016).
The genus is characterised by carbonaceous, black stromata, 8-spored, cylindrical asci and
ellipsoidal, dark brown ascospores with a germ slit.
Diatrype Fr., Summa veg. Scand., Section Post. (Stockholm) 384 (1849)
Index Fungorum number: IF1504; 170 morphological species (Species Fungorum 2020), 17
species with sequence data.
Type species – Diatrype disciformis (Hoffm.) Fr.
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Figure 97 – Allocryptovalsa truncata (AMH-9983, holotype). a, d Ascomata on host. b Vertical
section. c Peridium. e Periphysate neck. f Germinating spores. g, k Ascospores. h-j Asci. l-m
Colonies on malt extract agar after 6 days (left) and 30 days (right) (l = from above, m = from
below). Scale bars: b =100 μm, c, e, h-j = 50 μm, k =20 μm, g = 10 μm.
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Figure 98 – Diatrype disciformis (Material examined – USA, New York, Ithaca, on twigs of Salix
sp., 21 May 1939, E.D. Hansing, NY 00305626). a Herbarium material. b Stromata on host surface.
c Ostiole appeared on stroma. d Vertical section of stroma. e Ostiole. f Peridium. g-i Asci. j
Ascospores. Scale bars: b = 1000 µm, c = 100 µm, d = 500 µm, e = 50 µm, f-h = 10 µm, i-j = 5 µm.
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Notes – Species of Diatrype are common inhabitants on dead stems and barks of various
plants throughout the world (Carmarán et al. 2006, Vasilyeva et al. 2009, de Almeida et al. 2016,
Senwanna et al. 2017). In this entry we illustrate a collection of Diatrype disciformis.
Diatrypella (Ces. & De Not.) De Not., Sfer. Ital.: 29 (1863)
Index Fungorum number: IF1505; 65 morphological species (Species Fungorum 2020), 15
species with sequence data.
Type species – Diatrypella verruciformis (Ehrh.) Nitschke
Notes – Diatrypella was established to accommodate members of stromatic Sphaeriales
which were characterized by ovoid and numerous ascospores. The genus is characterised by conical
to truncate, cushion-like or discoid stromata, usually delimited by a black zone in the host tissues,
with umbilicate or sulcate ostioles. Asci are cylindrical, polysporous, with long stalks. Ascospores
are allantoid, hyaline or yellowish.
Echinomyces Rappaz, Mycol. helv. 2(549): 547 (1987)
Index Fungorum number: IF25174; 2 morphological species (Species Fungorum 2020).
Type species – Echinomyces obesa (Syd.) Rappaz
Notes – Echinomyces was established by Rappaz (1987) to accommodate E. echidnus and E.
obesus. The genus is characterised by the carbonaceous, black stromata and pale yellow ascospores,
which are arc-like or full circles.
Endoxylina Romell, Bot. Notiser: 173 (1892)
Index Fungorum number: IF1814; 15 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Endoxylina stellulata Romell
Notes – Endoxylina was introduced and assigned to Diatrypales without assigning the
familial position by Romell (1892). Based on previous morphological literature and herbarium
studies, Hyde et al. (2017b) transferred Endoxylina to Diatrypaceae. The genus is characterised by
flask-shaped, immersed perithecia with long pedicels and asci containing cylindrical to curved, 1septate, brown ascospores (Romell 1892).
Eutypa Tul. & C. Tul., Select. fung. carpol. (Paris) 2: 52 (1863)
Index Fungorum number: IF1950; 66 morphological species (Species Fungorum 2020), 17
species with sequence data.
Type species – Eutypa lata (Pers.) Tul. & C. Tul.
Notes – Species of Eutypa are the causal agents of Eutypa dieback of grapevine, apricots and
cherries (Trouillas & Gubler 2004, Baumgartner et al. 2010, Camps et al. 2010, Trouillas & Gubler
2010, Blanco-Ulate et al. 2013, Camps et al. 2014). The genus is characterised by stromata which
are irregular in shape, as confluent bumps, with conspicuous, scattered, roundish to prominent
ostioles on the host surface. Asci are 8-spored, clavate, apically rounded to truncate, with indistinct
apical rings, and long pedicels. Ascospores are allantoid to ellipsoidal, aseptate and pale yellowish.
Eutypella (Nitschke) Sacc., Atti Soc. Veneto-Trent. Sci. Nat., Padova, Sér. 4 4: 80 (1875)
Index Fungorum number: IF1951; 111 morphological species (Species Fungorum 2020), 17
species with sequence data.
Type species – Eutypella cerviculata (Fr.) Sacc.
Notes – Species of Eutypella have been found associated with a wide range of canker
diseases (Vasilyeva & Stephenson 2006, Trouillas et al. 2011). The genus is characterised by
pustulate stromata with stout, converging ostiolar necks and asci with eight spores.
Halocryptovalsa Dayarathne & K.D. Hyde, Cryptog. Mycol. (2019)
MycoBank: MB824308; 2 species with sequence data.
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Type species – Halocryptovalsa avicenniae (Abdel-Wahab, Bahkali & E.B.G. Jones)
Dayarathne & K.D. Hyde
Notes – Halocryptovalsa was established by Dayarathne et al. (2019b) to accommodate
species resembling Cryptovalsa from marine environments namely Cryptovalsa avicenniae and a
new species Halocryptovalsa salicorniae. The genus is characterised by poorly developed stromata
and poly-spored asci, with a J-, cylindrical, conspicuous apical or subapical ring. Ascospores are
hyaline or yellow-brown to brown, allantoid, with small, fat globules at the end (Dayarathne et al.
2019b).
Halodiatrype M.C. Dayarathne & K.D. Hyde, Phytotaxa 7(5): 617 (2016)
Index Fungorum number: IF552254; 3 morphological species (Species Fungorum 2020), 2
species with sequence data.
Type species – Halodiatrype salinicola M.C. Dayarathne & K.D. Hyde
Notes – Halodiatrype was established by Dayarathne et al. (2016b) to accommodate H.
avicenniae and H. salinicola isolated from mangroves. The genus is characterised by 8-spored,
cylindrical to clavate, pedicellate asci, with apically rounded or flattened, lacking an apical ring and
deliquescing early. Ascospores are oblong to allantoid or sub-inequilateral, aseptate or septate, not
constricted at the septa, light brown, with one to few small guttules and slightly curved. The
asexual morphs are libertella-like (Dayarathne et al. 2016b, 2019b).
Leptoperidia Rappaz, Mycol. helv. 2(547): 544 (1987)
Index Fungorum number: IF25186; 4 morphological species (Species Fungorum 2020).
Type species – Leptoperidia macropunctata (Rehm) Rappaz
Notes – Leptoperidia was introduced to accommodate L. applanata, L. asperrima, L.
macropunctata and L. trifida. The genus is characterised by relatively small stroma, asci and
ascospores, perithecia with very thin and slightly melanized walls (Rappaz 1987).
Libertella Desm., Annls Sci. Nat., sér. 1 19: 275 (1830)
Index Fungorum number: IF8769; 33 morphological species (Species Fungorum 2020), 2
species with sequence data.
Type species – Libertella betulina Desm.
Notes – Libertella was introduced by Desmazières (1830) to accommodate L. betulina, L.
faginea and L. rosae. This genus was mostly reported as the asexual morph of Diatrypella,
however, some species were reported as asexual Eutypa, Eutypella, Diaporthe, and Polystigma
(Kirk et al. 2001). The genus is characterised by subcortical, erumpent, and yellow to red acervula
conidiomata and branched conidiophores that produce hyaline, 1-celled, filiform conidia (Barnett &
Hunter 1972, Sutton 1980, von Arx 1981b).
Monosporascus Pollack & Uecker, Mycologia 66(2): 348 (1974)
Index Fungorum number: IF3260; 4 morphological species (Species Fungorum 2020), 3
species with sequence data.
Type species – Monosporascus cannonballus Pollack & Uecker
Notes – Monosporascus was introduced by Pollack & Uecker (1974) with type species M.
cannonballus to accommodate a fungus obtained from necrotic muskmelon roots. The genus is
characterised by pyriform asci and the formation of one (rarely two) single large, sphaerical
ascospores (Pollack & Uecker 1974).
Neoeutypella M. Raza, Q.J. Shang, Phookamsak & L. Cai, Fungal Divers. 95 (1): 1–273 (2019)
Index Fungorum number: IF555373; 1 species with sequence data.
Type species – Neoeutypella baoshanensis M. Raza, Q.J. Shang, Phookamsak & L. Cai
Notes – Neoeutypella was introduced by Phookamsak et al. (2019) to accommodate two
fungal strains under the name Eutypella caricae and a new strain isolated from Pinus armandii
577
(Pinaceae). Neoeutypella is characterised by carbonaceous stromata, erumpent through host
epidermis, producing yellow pigments surrounding the stroma. Asci are 8-spored, spindle-shaped,
with long pedicellate, apically rounded, with refractive cytoplasmic strands, with a J+, subapical
ring.
Pedumispora K.D. Hyde & E.B.G. Jones, Mycol. Res. 96(1): 78 (1992)
Index Fungorum number: IF25433; 1 species with sequence data.
Type species – Pedumispora rhizophorae K.D. Hyde & E.B.G. Jones
Notes – Pedumispora was introduced by Hyde & Jones (1992) to accommodate a fungus
from mangrove habitats. A phylogenetic study based on nuclear LSU and ITS regions showed the
taxonomic position of Pedumispora was in Diatrypaceae (Klaysuban et al. 2014). The genus is
characterised by 8-spored, fusiform, pedicellate, unitunicate, apically truncate asci. Ascospores are
filiform, mutli-septate, curved, longitudinally striate, with tapering poles, with one or both ends
crook-like (Hyde & Jones 1992).
Peroneutypa Berl., Icon. fung. (Abellini) 3(3-4): 80 (1902)
Index Fungorum number: IF3834; 28 morphological species (Species Fungorum 2020), 11
species with sequence data.
Type species – Peroneutypa bellula (Desm.) Berl.
Notes – Peroneutypa was introduced by Berlese (1902) to accommodate P. bellula, P.
corniculata, and P. heteracantha without designating the type species. Rappaz (1987) proposed P.
bellula as the type species and synonymized Peroneutypa under Eutypella. However, based on
characteristics and phylogenetic data of Acero et al. (2004), Peroneutypa was reinstated by
Carmarán et al. (2006). The genus is characterised by valsoid stromata, with packed, long
prominent necks, sessile to long pedicels, small asci with truncate apices and allantoid ascospores
(Carmarán et al. 2006, 2014, Senwanna et al. 2017, Shang et al. 2017).
Quaternaria Tul. & C. Tul., Select. fung. carpol. (Paris) 2: 104 (1863)
Index Fungorum number: IF4632; 10 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Quaternaria persoonii Tul. & C. Tul.
Notes – Quaternaria was introduced by Tulasne & Tulasne (1863) and was typified by Q.
persoonii. Clements & Shear (1931) lectotypified the illegitimate name Q. quaternata to Q.
persoonii considered Quaternaria as a synonym of Eutypella (Tulasne & Tulasne 1863). Based on
analyses of LSU and ITS sequence data and the discussion of Gams (1994), Quaternaria was
considered to be an independent genus by Acero et al. (2004). The genus is characterised by
stromata that are cryptosphaeroid in appearance and develop within the bark parenchyma.
Distoseptisporaceae K.D. Hyde & McKenzie, Fungal Divers. 80(1): 402 (2016)
Index Fungorum number: IF551850; Facesoffungi number: FoF01754; 26 species.
Saprobic on wood and in terrestrial and freshwater habitats. Sexual morph: Ascomata
perithecial, dark brown to black, superficial, solitary, globose to ovoid, papillate. Ostiole
periphysate. Peridium leathery, carbonaceous, dark brown, composing two layers of textura
prismatica. Paraphyses numerous, hyaline, unbranched, septate, cylindrical. Asci 8-spored,
unitunicate, cylindrical, with a J-, apical ring. Ascospores biseriate, hyaline to light yellowishbrown, fusiform, straight, 3-septate, lacking a mucilaginous sheath or appendages. Asexual morph:
Hyphomycetous. Colonies effuse, hairy or velvety, olivaceous or black. Mycelium mostly
immersed, composed of branched, septate, smooth, pale brown hyphae. Conidiophores semimacronematous to macronematous, mononematous, septate, unbranched, single or in groups, erect,
straight or flexuous, smooth, olivaceous to brown, cylindrical, robust at the base, sometimes
elongating percurrently. Conidiogenous cells monoblastic, integrated, determinate, terminal,
cylindrical. Conidia acrogenous, solitary, olivaceous, brown or yellowish-brown to reddish-brown,
578
euseptate or distoseptate or muriform appearing cruciately divided by septa at right angles to one
another, obclavate or cylindrical, with rounded apex, indeterminately elongating percurrently,
truncate at base; basal cell with cross wall and basal scar; or transversal ellipsoid to subglobose,
sometimes with pores in the septa (adapted from Su et al. 2016b).
Type genus – Distoseptispora K.D. Hyde, McKenzie & Maharachch.
Notes – Distoseptisporaceae was introduced in Su et al. (2016b) to accommodate a single
genus Distoseptispora. Asexual morphs in Distoseptisporaceae share similar morphology with
species classified in Ellisembia and Sporidesmium in having holoblastic, euseptate or distoseptate
conidia and monoblastic, determinate or percurrently elongating conidiogenous cells
(Subramanian 1992, Shenoy et al. 2006, Yang et al. 2018b). However, many of the characters
used to delimit the genera do not appear phylogenetically significant (Shenoy et al. 2006, Su et al.
2016b).
Ecological and economic significance of Distoseptisporaceae
Distoseptispora contains saprobic lignicolous taxa found in terrestrial, but mostly in
freshwater habitats. They play important roles in decomposition of lignocelluloses in wood (Wong
et al. 1998a, Krauss et al. 2011, Hyde et al. 2016a).
Genus incuded in Distoseptisporaceae
Distoseptispora K.D. Hyde, McKenzie & Maharachch., Fungal Divers. 80(1): 402 (2016)
Index Fungorum number: IF551833; 26 species with sequence data.
Type species – Distoseptispora fluminicola McKenzie, H.Y. Su, Z.L. Luo & K.D. Hyde
Notes – The generic concept of Distoseptispora was emended by Yang et al. (2018b) in
respect to the characters of D. guttulata and D. suoluoensis, which have erect, macronematous
conidiophores, percurrently elongating conidiogenous cells and euseptate conidia. The type of
septum does not aid in identification of Distoseptispora species. Acrodictys martini was transferred
to Diaporthe martinii (Xia et al. 2017). It differs from all other species in the genus in having
muriform, transversally ellipsoid to subglobose brown conidia. Its placement in Distoseptispora
was resolved with molecular DNA data. Phylogenetic placement of the genus is shown in Fig. 4.
Distoseptispora clematidis was introduced by Phukhamsakda et al. (2020) from Clematis and Luo
et al. (2019) introduced D. appendiculata, D. lignicola, D. neorostrata and D. obclavata from
submerged wood in Thailand. In this entry we introduce a new species, Distoseptispora
rayongensis.
Distoseptispora rayongensis J. Yang & K.D. Hyde, sp. nov.
Fig. 99
Index Fungorum number: IF554770; Facesoffungi number: FoF04675
Etymology – Referring to the collecting site in Rayong Province, Thailand.
Holotype – MFLU 18-1045.
Saprobic on decaying twigs. Sexual morph: Undetermined. Asexual morph: Hyphomycetous.
Colonies effuse, dark brown to black, hairy, glistening. Mycelium partly superficial, partly
immersed in the substratum, composed of brown, septate, branched hyphae. Conidiophores
macronematous, mononematous, solitary, cylindrical, straight or slightly flexuous, septate, brown,
smooth, paler towards the apex, 75–125 × 3.5–5.5 µm (x̅ = 100 × 4.5 µm, n = 10), rounded at apex.
Conidiogenous cells integrated, terminal, monoblastic, cylindrical, brown. Conidia acrogenous,
solitary, obclavate or obspathulate, rostrate, mostly 9–13-euseptate, rarely 14–15-septate, pale
brown or pale olivaceous, becoming paler or hyaline towards the apex, guttulate, smooth, (36–)60–
106(–120) × 9–14.5 µm (x̅ = 80 × 11.5 µm, n = 20), with a darkened scar at the base, sometimes
with percurrent proliferation and forming another conidium from the conidial apex.
Culture characteristics – Conidia germinating on PDA within 24 h. Germ tubes produced
from the conidial base. Colonies on PDA slow growing, reaching 5–10 mm diam. after one month
at 25 °C in natural light, circular, aerial mycelium dense, brown; reverse dark brown, margin entire.
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Figure 99 – Distoseptispora rayongensis (MFLU 18-1045 holotype). a Colonies on the woody
substrate. b Conidiophore. c Conidiogenous cell with conidium. d, e Conidia and conidiophores. f-k
Conidia. l Germinated conidium. m, n Cultures, m from above, n from below. Scale bars: a = 100
µm, b, c, e-l = 30 µm, d = 50 µm.
580
Material examined – THAILAND, Rayong Province, Klaeng, 12°78′N, 101°66′E, on
decaying wood submerged in a freshwater stream, 24 April 2017, Y.Z. Lu, RAY-6, MFLU 181045, holotype; ibid. HKAS102138, isotype; ex-type living cultures MFLUCC 18-0415; ibid.
RAY-8, MFLU 18-1046, paratype; ex-paratype living culture MFLUCC 18-0417.
GenBank numbers – LSU: MH457137, SSU: MH457169, ITS: MH457172, tef1: MH463253,
rpb2: MH463255 (MFLUCC 18-0415); LSU: MH457138, SSU: MH457170, ITS: MH457173,
tef1: MH463254, rpb2: MH463256 (MFLUCC 18-0417).
Notes – Distoseptispora rayongensis clustered within Distoseptispora with high statistical
support in a sister clade to D. guttulata (Fig. 3). Distoseptispora rayongensis resembles D.
guttulata and D. suoluoensis in having conspicuous long conidiophores and euseptate conidia of
similar size. However, D. rayongensis can be distinguished from D. guttulata and D. suoluoensis in
conidial colour and shape. Percurrently elongating conidia are observed in D. rayongensis and D.
suoluoensis.
Dwiroopaceae K.V. Xavier, A.N. KC, J.Z. Groenew., Vallad & Crous, Fungal Systematics and
Evolution 4: 38 (2019)
Index Fungorum number: IF830873; Facesoffungi number: FoF06872; 3 species.
Saprobic or pathogenic on leaves, twigs, branches and fruits. Sexual morph: Undetermined.
Asexual morph: Coelomycetous. Conidiomata pycnidial, immersed to semi-immersed, erumpent,
solitary, uni- to multiloculate, scattered, globose, black, ostiolate or lacking ostioles. Peridium
thick-walled, comprising wall of 3–6 layers of pale brown cells of textura angularis.
Conidiophores lining the innermost wall layer of peridium, mostly reduced to conidiogenous
cells. Conidiogenous cells holoblastic, ampulliform to subcylindrical, hyaline to pale brown,
sometimes proliferating percurrently at apex. Conidia unicellular, solitary, truncate at base, of
three types: Macroconidia solitary, dark brown, aseptate, granular at surface, guttulate, thickwalled, broadly ellipsoid to obovoid, with longitudinal striations running along the length of
conidia. Mesoconidia hyaline to pale brown, ellipsoid, aseptate and microconidia ellipsoid,
hyaline, aseptate (adapted from Xavier et al. 2019).
Type genus – Dwiroopa Subram. & Muthumary
Notes – Castlebury et al. (2002) placed Harknessia and Dwiroopa in Diaporthales based on
LSU phylogeny. Subsequently, Crous et al. (2012d) established Harknessiaceae to include species
of Harknessia with wuestneia-like sexual morphs in Diaporthales. Dwiroopaceae was established
by Xavier et al. (2019) to accommodate a monotypic genus Dwiroopa including three species in
Diaporthales. Dwiroopaceae species share closer characters with taxa of Harknessiaceae and have
to sister relation with Harknessiaceae in a multigene phylogenetic analysis.
Ecological and economic significance of Dwiroopaceae
Species of Dwiroopaceae are saprobes and pathogens in terrestrial habitats. Dwiroopa ramya
is a saprobe on dead twigs in India (Subramanian & Muthumary 1986), while D. lythri is a
pathogen on a noxious weed purple loose strife in North America (Farr & Rossman 2001).
Dwiroopa punicae represents a severe pathogen of Punica granatum across the south eastern USA
causing up to 100% fruit damage (Xavier et al. 2019).
Genus included in Dwiroopaceae
Dwiroopa Subram. & Muthumary, Proc. Indian Acad. Sci., Pl. Sci. 96(3): 196 (1986)
Index Fungorum number: IF11077; 3 morphological species (Species Fungorum 2020), 2
species with sequence data.
Type species – Dwiroopa ramya Subram. & Muthumary
Notes – Members of Dwiroopa resemble Harknessia in having uni- to multiloculate, thickwalled, immersed to semi-immersed conidiomata, becoming erumpent, dark brown and aseptate,
thick-walled macroconidia with holoblastic conidiogenesis, with conidiogenous cells bearing
periclinal thickenings. Dwiroopa is distinguished from Harknessia in having distinct, holoblastic
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conidiogenesis, thick-walled conidiogenous cells and broadly spaced several longitudinal striations
in the macroconidia, lacking a basal appendage and showcasing a dark scar with minute marginal
frills at the conidial connection. Harknessia is distinct from Dwiroopa in having macroconidia that
are smooth-walled or have closely spaced longitudinal raised bands on only one side and usually
with a true basal appendage. Furthermore, Dwiroopa produces three different kinds of conidia viz.
macroconidia, mesoconidia and phialoconidia (Farr & Rossman 2003), which are not produced in
Harknessia. Farr & Rossman (2003) redescribed Dwiroopa ramya and designated it as a lectotype
and also introduced Harknessia lythri as Dwiroopa lythri in Dwiroopa based on morphological
similarities.
Figure 100 – Dwiroopa ramya (redrawn from Subramanian & Muthumary 1986). a Vertical
section of conidioma. b Conidia. c-f Stages in the development of macroconidia. g Paraphysis-like
structures. h Meso-conidia, note, developing and mature conidia. i Phialides with developing
phialoconidia. j Phialoconidia. Scale bars: a = 100 μm, b = 20 μm, c, d, f-i = 10 μm, e, j = 5 μm.
Erythrogloeaceae Senan., Maharachch. & K. D. Hyde, Stud. Mycol. 86: 217–296 (2017)
Index Fungorum number: IF821551; Facesoffungi number: FoF03478; 22 species.
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Foliicolous, associated with leaf spots. Sexual morph: Pseudostromata circular, erumpent,
consisting of an inconspicuous, usually orange ectostromatic disc, semi-immersed to superficial.
Ectostromatic disc flat or concave, orange. Perithecia conspicuous, umber to black, embedded in
pseudostromatic tissue, scattered, surrounding the ectostromatic disc, ostiolate. Ostioles cylindrical,
obscuring the disc, covered by orange to black crust. Paraphyses deliquescent. Asci 8-spored,
fusoid, 2–3-seriate, with an apical ring. Ascospores hyaline, fusoid to cylindrical, symmetrical to
asymmetrical, straight to curved, bicellular, with a median septum, constricted at the septum,
smooth, multiguttulate. Asexual morph: Coelomycetous. Conidiomata epiphyllous, subepidermal,
sometimes eustromatic, acervular or subglobose, brown to black or yellow-orange, amphigenous,
opening by irregular rupture, orange-brown wall composed of cells of textura angularis, exuding
slimy orange masses of conidia. Conidiophores reduced to conidiogenous cells. Conidiogenous
cells lining the inner cavity of conidioma, hyaline to olivaceous, smooth, subcylindrical to
ampulliform, tapering to a long, thin neck, at times apical part elongated into a long neck,
proliferating several times percurrently near apex, with flaring collarettes, or apex truncate, with
minute periclinal thickening. Conidia hyaline to olivaceous, smooth, guttulate or not, thin-walled,
ellipsoid, fusoid, ovoid to somewhat obclavate, straight to curved, apex subobtuse, obtusely
rounded, base truncate, with prominent marginal frill, or dimorphic, intermixed in same
conidiomata. Macroconidia broadly ellipsoid to obovoid, hyaline, smooth, granular to guttulate,
thick-walled, apex obtuse, base flattened. Microconidia hyaline, smooth, guttulate, fusoid-ellipsoid,
acutely rounded at apex, truncate at base (adapted from Senanayake et al. 2017a).
Type genus – Erythrogloeum Petr.
Notes – Erythrogloeaceae was erected based on morphology and phylogeny and it comprised
Chrysocrypta, Disculoides and Erythrogloeum (Senanayake et al. 2017a). Fan et al. (2018)
accommodated Dendrostoma within this family. Erythrogloeaceae is characterized by epiphyllous
acervuli, and subcylindrical to ampulliform conidiogenous cells. The sexual morph of this family is
reported only from Dendrostoma (Fan et al. 2018).
Ecological and economic significance of Erythrogloeaceae
Erythrogloeum hymenaeae causes leaf spots and anthracnose of Hymenaea sp. mainly during
the wet season (Ferreira et al. 1992). The second species, E. pini-acicola was described from
needles of Pinus oocarpa (Evans 1984). Chrysocrypta corymbiae is associated with leaf spots on
Corymbia. Disculoides forms leaf spots on Eucalyptus species.
Genera included in Erythrogloeaceae
Chrysocrypta Crous & Summerell, Persoonia 28: 165 (2012)
Index Fungorum number: IF800379; 1 species with sequence data.
Type species – Chrysocrypta corymbiae Crous & Summerell
Notes – Chrysocrypta, introduced by Crous et al. (2012b), is associated with leaf spots of
Corymbia species. This monotypic genus is similar to Foliocryphia. However, Chrysocrypta is
distinct in having dimorphic conidia.
Dendrostoma X.L. Fan & C.M. Tian, Persoonia 40: 124 (2018)
Index Fungorum number: IF823986; 14 species with sequence data.
Type species – Dendrostoma mali X.L. Fan & C.M. Tian
Notes – Dendrostoma is distinct in having diaporthe-type perithecia with clavate asci and
fusoid to cylindrical and bi-celled ascospores (Fan et al. 2018).
Disculoides Crous, Pascoe, I.J. Porter & Jacq. Edwards, Persoonia 28: 71 (2012)
Index Fungorum number: IF564737; 5 species with sequence data.
Type species – Disculoides eucalypti Crous, Pascoe, I.J. Porter & Jacq. Edwards
Notes – Disculoides species are foliar pathogens of Eucalyptus and Corymbia species. The
genus is characterized by acervular, amphigenous black conidiomata and conidiophores which are
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either reduced to conidiogenous cells. The conidiogenous cells are subcylindrical to ampulliform
and proliferate percurrently at the apex, with flaring collarettes. Conidia are hyaline to olivaceous
(Crous et al. 2012b, 2017a).
Erythrogloeum Petr., Sydowia 7(5-6): 378 (1953)
Index Fungorum number: IF8211; 2 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Erythrogloeum hymenaeae Gonz. Frag. & Cif.
Notes – Erythrogloeum comprises two species, E. hymenaeae and E. pini-acicola. Both
species are foliar pathogens of Hymenaeae and Pinus species. The sexual morph of this genus is
not reported. The genus is characterised by acervular conidiomata with side walls comprising dark
brown to black cells of textura angularis, lageniform to cylindrical conidiogenous cells and
monomorphic conidia (Senanayake et al. 2017a).
Figure 101 – Erythrogloeum hymenaeae (Material examined – COSTA RICA, San Jose, on leaves
of Hymenaea courbaril, Nov. 1929, H. Schmidt, F45468, syntype). a Herbarium specimen. b
Conidiomata on substrate. c, d Conidiophores, conidiogeneous cells and conidia. e Conidia. Scale
bars: b = 200 μm, c-d = 10 μm, e = 5 μm.
Etheirophoraceae Rungjindamati, Somrothipol, & Suetrong, Cryptog. Mycol. 35(2): 134 (2014)
Index Fungorum number: IF808178; Facesoffungi number: FoF01329; 5 species.
Saprobic on intertidal wood and bark in marine habitats. Sexual morph: Ascomata subglobose
to globose or pyriform, light brown to dark brown or black, immersed, oblique or vertical to the
host surface, clypeate, coriaceous, ostiolate, periphysate, papillate. Peridium composed several
layers of brown to dark brown cell layers of textura angularis. Paraphyses numerous, mostly
unbranched, attached to the apex of the ascomatal cavity, embedded in a gelatinous matrix. Asci 8spored, unitunicate, cylindrical to oblong, pedicellate, J-, persistent. Ascospores 1–2-seriate,
hyaline, ellipsoidal, 1 to many septate, constricted at the septa, with a filamentous appendage at one
or both ends. Appendages bristle-like, origin undetermined. Asexual morph: Undetermined
(adapted from Jones et al. 2014).
Type genus – Etheirophora Kohlm. & Volkm.-Kohlm.
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Notes – This family includes the genera Etheirophora (E. bijubata, E. blepharospora, E.
unijubata) and Swampomyces (S. armeniacus, S. triseptatus) (Jones et al. 2014). However, the
genera Etheirophora and Swampomyces are not congeneric and they form a sister clade with
Falcocladium species (Falcocladiaceae) in an unsupported clade in Hypocreomycetidae, order
incertae sedis (Maharachchikumbura et al. (2015). Subsequently, Jones et al. (2015) introduced
Torpedosporales to accommodate the families Etheirophoraceae, Juncigenaceae and
Torpedosporaceae. The order evolved with a stem age of 171–241 MYA (Hongsanan et al. 2017,
Hyde et al. 2017a).
Ecological and economic significance of Etheirophoraceae
Etheirophoraceae species are commonly found as saprobes on intertidal wood and bark in
marine habitats. Host-specificity of Keissleriella blephorospora (= Etheirophora blepharospora),
occurring on Rhizophora species in Hawaii has been reported and the species is involved in nutrient
cycling (Osorio et al. 2016).
Genera included in Etheirophoraceae
Etheirophora Kohlm. & Volkm.-Kohlm., Mycol. Res. 92(4): 414 (1989)
Index Fungorum number: IF25298; 3 morphological species (Species Fungorum 2020), 2
species with sequence data.
Type species – Etheirophora bijubata Kohlm. & Volkm.-Kohlm.
Notes – Kohlmeyer & Kohlmeyer (1989) introduced Etheirophora (type species E. bijubata)
to accommodate three marine species from tropical locations, including a species previously
referred to as Keissleriella blepharospora. The genus was assigned to Sphaeriales by Kohlmeyer &
Kohlmeyer (1989) and to Halosphaeriales by Hawksworth et al. (1995) and Kirk et al. (2001).
Schoch et al. (2006), based on molecular data and morphology, referred it to the TBM clade
comprising Bertia, Melanospora and Torpedospora, within Hypocreomycetidae, with affinities to
the Coronophorales. Etheirophora grouped with a range of unresolved taxa, Juncigena,
Swampomyces and Torpedospora and the asexual genera Glomerulispora (= Torpedospora) and
Moheitospora, in the TBM clade with high bootstrap support (Abdel-Wahab et al. 2010).
Subsequently the genus was referred to Etheirophoraceae (Torpedosporales) (Jones et al. 2014,
2015).
Swampomyces Kohlm. & Volkm.-Kohlm., Bot. Mar. 30(3): 198 (1987)
Index Fungorum number: IF6004; 2 species with sequence data.
Type species – Swampomyces armeniacus Kohlm. & Volkm.-Kohlm.
Notes – Abdel-Wahab et al. (2001) added S. triseptatus (Hyde & Nakagiri 1992), S.
clavatispora and S. aegyptiacus. Subsequently, S. aegyptiacus and S. clavatispora were transferred
to Fulvocentrum by Jones et al. (2014). Swampomyces armeniacus is illustrated in this entry.
Falcocladiaceae Somrith., E.B.G. Jones & K.L. Pang, Cryptog. Mycol. 35(2): 134 (2014)
Index Fungorum number: IF808179; Facesoffungi number: FoF01288; 5 species.
Saprobic on leaf litter, including leaves of Eucalyptus grandis and E. camaldulensis in
tropical, terrestrial habitats. Sexual morph: Undetermined. Asexual morph: Hyphomycetous.
Conidiomata hyaline, sporodochial or synnematal or penicillate, intermixed with setae, arising
from a stroma or microsclerotia or prostrate mycelium Setae cylindrical, thick and smoothwalled, hyaline, non-septate and terminating in variedly-shaped vesicles. Conidiophores
subcylindical, hyaline, septate, branched, forming up to three series of branches per conidioma
(primary, secondary and tertiary). Conidiogenous cells ampulliform, phialidic, arranged in 2–6
whorls, with elongate necks, with minute collarettes. Conidia trans- (1)-septate, hyaline, smoothwalled, falcate, guttulate, with short apical and basal appendages (adapted from Jones et al. 2014,
Maharachchikumbura et al. 2016b).
Type genus – Falcocladium S.F. Silveira
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Figure 102 – Swampomyces armeniacus (Material examined – BELIZE. Man-Of-War Cay
(Atlantic Ocean). 16° 53' 00"N, 88° 06' 15"W (16.883, -88.104); Driftwood; collected by J.
Kohlmeyer 4619 - NY taxon A, 23 November 1984, NY no.01271822, paratype). a, b Herbarium
material. c Ascomata on host substrate. d Ascoma cross section. e Peridium. f Paraphyses. g, h Asci
and paraphyses. i-n Ascospores. Scale bars: c, d = 100 µm, e, g, h = 50 µm, f, i-n = 20 µm.
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Notes – Crous et al. (2007a) used the BLASTn tool to compare ITS and LSU gene regions
with reference sequences, suggesting the placement of Falcocladium in Hypocreales and further
considered the genus to be polyphyletic. Jones et al. (2014) introduced the monotypic family
Falcocladiaceae which formed a monophyletic clade in Hypocreomycetidae, based on SSU and
LSU nuclear genes to accommodate members of Falcocladium. They suggested further taxon
sampling to determine its ordinal status. Based on an LSU and SSU combined gene phylogenetic
study, Falcocladiaceae was placed in Falcocladiales (Maharachchikumbura et al. 2015). In a
phylogenetic analysis by Réblová et al. (2016b), Falcocladiales showed close affinity to
Coronophorales and Melanosporales. Several phylogenetic studies showed similar results
(Maharachchikumbura et al. 2016b; Réblová et al. 2016b). Currently, the order includes one
hyphomycetous genus Falcocladium introduced by Crous et al. (1994a). Falcocladium species
can be distinguished based on morphology of vesicular apices of setae which ranges from
ellipsoidal, sphaeropedunculate to turbinate and conidial measurements (Somrithipol et al. 2007) .
No sexual morph has been recorded for the family.
Ecological and economic significance of Falcocladiaceae
Species in Falcocladium are saprobes on leaves of woody plants. Falcocladium africanum
occurs on leaves of Eucalyptus brassiana and E. tereticornis in Ghana and Sierra Leone (Crous et
al. 2018c), F. thailandicum occurs on leaves of E. camaldulensis in Thailand (Crous et al. 2007a),
F. turbinatum occurs on dead leaves of evergreen trees in Thailand (Somrithipol et al. 2007), F.
spltaeropeduncuiatuln occurs on leaves of E. pellita × brassiana in Brazil (Crous et al. 1997), F.
multivesiculatum occurs on leaf litter of E. grandis in Brazil (Crous et al. 1994a). This group plays
an important role in degrading plant leaves.
Genus included in Falcocladiaceae
FalcocladiumS.F. Silveira, Alfenas, Crous & M.J. Wingf., Mycotaxon 50: 447 (1994)
Index Fungorum number: IF25800; 5 species with sequence data.
Type species – Falcocladium multivesiculatum S.F. Silveira, Alfenas, Crous & M.J. Wingf.
Notes – Falcocladium includes five species, all of which occur on leaves, namely F.
africanum, F. multivesiculatum, F. sphaeropedunculatum, F. turbinatum, and F. thailandicum.
Falcocladium species can be distinguished based on their setal vesicle shape (ellipsoidal,
sphaerpedunculate or turbinate) and conidial dimensions. Phylogenetically they are distinct from
other genera in the Hypocreomycetidae and form a monophyletic clade (Jones et al. 2014). In this
entry we illustrate Falcocladium multivesiculatum. The species F. multivesiculatum has a wide host
range and mainly occurs on leaves, including Eucalyptus grandis and E. camaldulensis. The fungus
is distributed mainly in tropical forests (Somrithipol et al. 2007, Maharachchikumbura et al.
2016b). The taxon fits well within the species concept of Falcocladium with white sporodochia,
bearing thick-walled aseptate, stipe extensions and hyaline, 0–1-septate, falcate conidia, with short
apical and basal appendages (Crous et al. 1994a, Somrithipol et al. 2007).
Flammocladiellaceae Crous, L. Lombard & R.K. Schumach., Sydowia 67: 103 (2015)
Index Fungorum number: IF819623; Facesoffungi number: FoF06873; 2 species.
Fungicolous on Massaria spp. on corticated branches of Acer spp. Sexual morph: Ascomata
perithecial, pale luteous to yellow-orange, aggregated in clusters, on a single stromatic base,
covered in a dirty white crustose layer, turning pale luteous to dirty white, KOH-, ostiolate. Ostiole
papillate, periphysate. Peridium 3–4 layers of subhyaline, smooth-walled cells of textura angularis.
Asci 8-spored, unitunicate, fusoidal-ellipsoidal to subclavate, hyaline, smooth-walled. Ascospores
fasciculate, hyaline, fusoidal-ellipsoidal with obtuse ends, septate, warty, not to slightly constricted
at septa. Asexual morph: Conidiomata sporodochial, determinate, hyaline, becoming orange.
Conidiophores subcylindrical, septate, branched. Conidiogenous cells subcylindrical, terminal and
intercalary, hyaline, smooth-walled, proliferating sympodially at apex. Conidia slimy, solitary,
587
granular to guttulate, straight to gently curved, subcylindrical to narrowly obclavate, hyaline,
smooth-walled (adapted from Crous et al. 2015b).
Figure 103 – Falcocladium multivesiculatum (Material examined – BRAZIL, Aracruz, Espirito
Santo, from Eucalyptus grandis leaf litter, January 1993 Silvaldo F. Silveira, PREM 51541,
holotype). a, c Herbarium material. b Dried culture on MEA. d, e Ellpisoidal vesicles on setae. f-j
Conidia. Scale bars: d,e = 20 µm, f-j = 10 µm.
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Type genus – Flammocladiella Crous, L. Lombard & R.K. Schumach.
Notes – Flammocladiellaceae was introduced to accommodate the monotypic genus
Flammocladiella by Crous et al. (2015b). In the molecular analysis by Hongsanan et al. (2017) and
Sun et al. (2017), Flammocladiellaceae grouped within Clavicipitaceae, hence was not treated as a
separate family. Further, Hongsanan et al. (2017) synonymized Flammocladiellaceae under
Clavicipitaceae based on phylogenetic data and this was followed by Wijayawardene et al. (2018a).
However, in the molecular clock analysis by Hyde et al. (2017a) Flammocladiellaceae forms a
separate clade sister to Clavicipitaceae with low statistical support. Similar results were obtained
from phylogenetic analysis by Maharachchikumbura et al. (2016b) and Lechat & Fournier (2018).
Hence, we would like to treat Flammocladiellaceae as an accepted family within Hypocreales,
based on previous treatments and our phylogenetic analysis (Fig. 15).
Ecological and economic significance of Flammocladiellaceae
Flammocladiella decora is a fungicolous species occurring on immersed ascomata of
Massaria species (Lechat & Fournier 2018b). Currently, there is no data on the economic
significance of Flammocladiella species.
Genus included in Flammocladiellaceae
Flammocladiella Crous, L. Lombard & R.K. Schumach., Sydowia 67: 103 (2015)
Index Fungorum number: IF812530; 2 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Flammocladiella decora (Wallr.) Lechat & J. Fourn.
Notes – Flammocladiella was introduced by Crous et al. (2015b) based on a single species F.
aceris. Lechat & Fournier (2018b) proposed Flammocladiella decora to accommodate Nectria
decora. Flammocladiella decora produces long, subcylindrical conidia and a sexual morph, which
are indistinguishable from F. aceris (Lechat & Fournier 2018b). Considering morphological
similarities and phylogenetic analyses Lechat & Fournier (2018b) regarded F. aceris as a synonym
of F. decora. The genus is characterised by yellowish ascomata seated on ascomata of Massaria
species, aggregate in clusters, covered by a crustose layer, having a papillate, periphysate ostiolar
region, and a sporodochial asexual morph forming flame-like conidial masses, conidia
subcylindrical (Crous et al. 2015b, Lechat & Fournier 2018b).
Fuscosporellaceae J. Yang, J. Bhat & K.D. Hyde, Cryptogam. Mycol. 37(4): 457 (2016)
Index Fungorum number: IF552321; Facesoffungi number: FoF02421; 14 species.
Saprobic on dead wood mostly in freshwater habitats. Sexual morph: Ascomata immersed,
brown, solitary or in small groups, papillate or with a rostrate neck. Ostiole periphysate. Peridium
layered, with outer layer brown, inner layers hyaline. Paraphyses septate, hyaline. Asci 8-spored,
unitunicate, cylindrical, with a J-, apical ring at the apex. Ascospores uniseriate, fusiform, hyaline
or versicolorous, with brown middle cells and hyaline end cells, usually 5-septate, sometimes
slightly constricted at the septa and sometimes guttulate. Asexual morph: Hyphomycetous.
Colonies effuse or sporodochial, dark brown to black, scattered. Conidiophores macronematous
or semi-macronematous, mononematous, branched or unbranched, hyaline, pale brown to mid
brown. Conidiogenous cells monoblastic, globose to clavate, terminal, sometimes discrete.
Conidia clavate, obovoid or obpyriform, septate, smooth, brown to dark brown; basal cell pale
brown, trapeziform or triangular (adapted from Yang et al. 2016b).
Type genus – Fuscosporella J. Yang, J. Bhat & K.D. Hyde
Notes – The family concept was expanded to include Bactrodesmiastrum, Mucispora,
Fuscosporella, Parafuscosporella and two sexual genera Plagiascoma and Pseudoascotaiwania
(Boonyuen et al. 2016, Réblová et al. 2016c, Hernández-Restrepo et al. 2017, Hongsanan et al.
2017). Mucispora is characterized by macronematous conidiophores and obovoid dark conidia. The
characters shared by Bactrodesmiastrum, Fuscosporella and Parafuscosporella are sporodochial
conidiomata, monoblastic conidiogenous cells and brown septate conidia (Hernández-Restrepo et
589
al. 2013, 2015a, Yang et al. 2016b). Plagiascoma and Pseudoascotaiwania represent the only
known sexual morphs in the order characterized by cylindrical asci, with a J-, apical ring and
fusiform ascospores (Fallah et al. 1999, Réblová et al. 2016c). The understanding of
Fuscosporellales (Boonyuen et al. 2016, Réblová et al. 2016c, Yang et al. 2016b) requires further
collections and molecular data.
Figure 104 – Flammocladiella decora (CLL16020, holotype, redrawn from Lechat & Fournier
2018b). a Ascomata on bark. b Vertical section of peridium. c Ascospores. d, e Vertical sections of
ascomata. f Ascus. g Germinating spore. h Conidiophores. i Conidia. Scale bars: a = 1000 µm, b
=300 µm, c = 10 µm, d = 400 µm, e = 150 µm, f = 400 µm, g-i = 20 µm.
590
Ecological and economic significance of Fuscosporellaceae
Fuscosporellaceae contains saprobic lignicolous taxa found in terrestrial, but mostly in
aquatic habitats. They play important roles in decomposition of lignocelluloses in wood (Wong et
al. 1998a, Krauss et al. 2011, Hyde et al. 2016a).
Genera included in Fuscosporellaceae
Bactrodesmiastrum Hol.-Jech., Folia Geobot Phytotax 19 (1): 103 (1984)
Index Fungorum number: IF25786; 5 morphological species (Species Fungorum 2020), 3
species with sequence data.
Type species – Bactrodesmiastrum obscurum Hol.-Jech.
Notes – Bactrodesmiastrum is characterized by solitary or aggregated conidiophores, mostly
reduced to brown monoblastic conidiogenous cells, or arising from pulvinate to subpustulate
sporodochial conidiomata with moniliform or beaded hyphoid cells and brown, clavate, obovoid or
pyriform conidia often with black bands around the septa (Hernández-Restrepo et al. 2015a, Yang
et al. 2016b).
Fuscosporella J. Yang, J. Bhat & K.D. Hyde, Cryptog Mycol 37(4): 457 (2016)
Index Fungorum number: IF552289; 2 species with sequence data.
Type species – Fuscosporella pyriformis J. Yang, J. Bhat & K.D. Hyde
Notes – Fuscosporella is morphologically similar to Parafuscosporella, but differs in the
conidia which are produced in culture. Fuscosporella produced multi-celled, lamentous to helicoid
conidia in culture, while Parafuscosporella produce globose to obpyriform, uniseptate conidia in
culture (Yang et al. 2016b). Molecular data provided further evidence for their segregation.
Mucispora J. Yang, J. Bhat & K.D. Hyde, Cryptog Mycol 37(4): 466 (2016)
Index Fungorum number: IF552293; 2 species with sequence data.
Type species – Mucispora obscuriseptata J. Yang, J. Bhat & K.D. Hyde
Notes – Mucispora is similar to Monotosporella in having macronematous, mononematous
conidiophores, monoblastic conidiogenous cells and brown, ellipsoidal, clavate or obovoid conidia.
However, they are phylogenetically distinct. Phylogenetic studies showed Mucispora grouped in
Fuscosporellales (Yang et al. 2016b, 2017), while Monotosporella clustered within Pleurotheciales
(Hernández-Restrepo et al. 2017). In this entry we introduced a new species, Mucispora
infundibulata.
Mucispora infundibulata J. Yang & K.D. Hyde, sp. nov.
Fig. 105
Index Fungorum number: IF554769; Facesoffungi number: FoF04674
Etymology – Referring to the infundibular conidiogenous cells.
Holotype – MFLU 18-0142.
Saprobic on decaying, submerged twigs. Sexual morph: Undetermined. Asexual morph:
Colonies on substrate sparse, scattered, glistening, black. Mycelium mostly immersed, consisting of
septate, smooth, pale brown to hyaline hyphae. Conidiophores macronematous, mononematous,
solitary, straight, erect, smooth, mid brown, pale brown to hyaline and inflated at the apex,
guttulated, 1–2-septate, 50–65 × 4–6 µm ( x = 60 × 5 µm, n = 10), 10–12.5 µm wide at the apex,
with bulbous base. Conidiogenous cells monoblastic, integrated, terminal, cupulate or
infundibulate, pale brown to hyaline, guttulate. Conidia acrogenous, smooth, broadly ellipsoidal to
obovoid, 3-euseptate, constricted at the septa when young, with unobservable septa when mature,
truncate at the base, (22–)29–34 × (15–)19–21 µm ( x = 31 × 20 µm, n = 10).
Culture characteristics – Conidia germinating on PDA within 24 h. Germ tubes produced
from the conidial base. Colonies on PDA, reaching 5-10 mm diameter after one month at 25 °C in
natural light, circular, with brown, greyish-green, white and dark brown mycelium from inner to
outer circle; dark brown in reverse with entire margin. Hyphae subhyaline to pale brown,
sometimes constricted at the septa, (4–)6–10(–14) µm wide. Conidiophores reduced to a
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monoblastic conidiogenous cell. Conidiogenous cells 8–10.5 × 3.3–4.8 µm, integrated, subhyaline
to pale brown. Conidia (18–)23–35.5(–40) × (9.5–)12–17(–19) µm ( x = 31 × 14 µm, n = 20), pale
brown to mid brown, 1–3-septate, mostly 2-septate, globose to obovoid, with cells becoming bigger
towards the apical cell, smooth, constricted at the septa.
Figure 105 – Mucispora infundibulata (MFLU 18-0142, holotype). a-c Conidia and conidiophores.
d Germinated conidium. e, f Cultures, e from above, f from below. g-i Asexual morph in culture.
Scale bars: a-d, h = 30 µm, g, i = 50 µm.
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Material examined – THAILAND, Phang Nga Province, Bann Tom Thong Khang, on
decaying wood submerged in a freshwater stream, 17 December 2015, J. Yang, Site 7-21-3, MFLU
18-0142, holotype, HKAS 102139, isotype; ex-type living cultures MFLUCC 16-0866, GZCC 170021.
GenBank numbers – ITS: MH457174, LSU: MH457139, SSU: MH457171.
Notes – Mucispora infundibulata resembles Melanocephala australiensis in conidial shape
and size (Hughes 1979). Mucispora infundibulata was placed as a sister taxon to Mucispora
phangngaensis with strong support (Fig. 10).Conidiophores of Melanocephala australiensis are
initially 18–30 µm long, but can reach 100 µm by percurrent elongation, while conidiophores of
Mucispora infundibulata are mostly around 60 µm long. The obvious difference is the inflated
cupulate conidiogenous cells in Mucispora infundibulata compared to flared, percurrent
proliferating in Melanocephala australiensis.
Parafuscosporella J. Yang, J. Bhat & K.D. Hyde, Cryptog Mycol 37(4): 458 (2016)
Index Fungorum number: IF552292; 3 species with sequence data.
Type species – Parafuscosporella moniliformis J. Yang, J. Bhat & K.D. Hyde
Notes – Parafuscosporella resembles Fuscosporella based on their almost similar
morphology. However, the two genera are phylogenetically distinct (Fig. 10) and can be
distinguished by the morphology of conidia which are produced in culture. Among the three
accepted species, conidia of P. garethii are obpyramidal, coronate at the apex with conical
projections, while those of P. moniliformis and P. mucosa are obovoid to obpyriform (Boonyuen et
al. 2016, Yang et al. 2016b).
Plagiascoma Réblová & J. Fourn., Persoonia 37:69 (2016)
Index Fungorum number: IF813234; 1 species with sequence data.
Type species – Plagiascoma frondosum Réblová & J. Fourn.
Notes – Plagiascoma and Pseudoascotaiwania are monotypic sexual genera in
Fuscosporellaceae. The sexual morph of Plagiascoma frondosum (Fig. 106) resembles
Pseudoascotaiwania persoonii in having cylindrical asci with a J-, apical ring and uniseriate (in
fresh material), euseptate, fusiform ascospores. Ascospores of Plagiascoma frondosum are hyaline
and 3–5-septate, while those of Pseudoascotaiwania persoonii are 5-septate, rarely 6-septate and
have light brown central cells and hyaline end cells (Fallah et al. 1999, Réblová et al. 2016c).
Pseudoascotaiwania J. Yang, J. Bhat & K.D. Hyde, Cryptog Mycol 37(4): 471 (2016)
Index Fungorum number: IF552323; 1 species with sequence data.
Type species – Pseudoascotaiwania persoonii (Fallah, J.L. Crane & Shearer) J. Yang, J. Bhat
& K.D. Hyde
Notes – Pseudoascotaiwania, introduced with the type species P. persoonii, was transferred
from Ascotaiwania without known asexual morphs. The genus is characterised by 5-septate, rarely
6-septate ascospores with light brown, central cells and hyaline end cells.
Glomerellaceae Locq., Mycologia 98(6): 1083 (2007)
Index Fungorum number: IF80186; Facesoffungi number: FoF01100; 894 species.
Parasitic, endophytic and saprobic on plant leaves, stems and fruits. Sexual morph: Ascomata
solitary or gregarious, globose to subglobose, dark brown to black, ostiole periphysate. Peridium
composed of pale to medium brown flattened cells of textura angularis. Paraphyses composed of
hyaline, septate, branched at the base, rounded at the apex. Asci 8-spored, unitunicate, cylindrical to
subfusoid, short pedicellate, with a J-, refractive ring at the apex, Ascospores uni- to biseriate,
hyaline, aseptate, smooth-walled, cylindrical, oval, fusiform or rhomboid, straight or curved, one
end ± acute and one end rounded or both ends rounded. Asexual morph: Coelomycetous.
Conidiomata acervular, ranging from sporodochia-like aggregations to closed conidiomata that
open by rupture, conidiophores and setae formed on cushions of pale to medium brown, roundish to
593
angular cells. Setae may or may not be present, straight, pale to dark brown, sometimes hyaline
towards the tip, smooth-walled, verruculose to verrucose, 1–8-septate, base cylindrical, conical or
slightly inflated, tip ± rounded to ± acute. Conidiophores hyaline to pale brown, simple or septate,
branched or unbranched, smooth-walled, sometimes verruculose.
Conidiogenous cells
enteroblastic, hyaline to pale brown, smooth-walled, discrete, ampulliform, cylindrical, doliiform or
ellipsoidal, collarette usually distinct, periclinal thickening visible to conspicuous, sometimes
extending to form new conidiogenous loci ( percurrent) or surrounded by a gelatinous coating.
Conidia hyaline, smooth-walled, aseptate, cylindrical, clavate, fusiform, sometimes ellipsoidal to
ovoid, straight or curved, apex rounded to acute, sometimes with a filiform appendage, base
rounded to truncate, sometimes with a prominent hilum. Appressoria single or in small groups, pale
to dark brown, with a globose, elliptical, clavate, navicular or irregular outline and an entire,
undulate or lobate edge (adapted from Maharachchikumbura et al. 2016b).
Figure 106 – Plagiascoma frondosum (Redrawn from Réblová et al. 2016c). a, b Asci. c
Ascospores. d Paraphyses. Scale bars: a = 25 µm, b = 20 µm, c, d = 10 µm.
Type genus – Colletotrichum Corda
Notes – The ordinal name Glomerellales including Colletotrichum (= Glomerella) and three
other genera in a non-ranked group “ Eu-Glomérellales” was introduced by Chadefaud (1960), but
was not validly published. Earlier classification placed Colletotrichum in Phyllachoraceae
(Ainsworth 1971, Barr 1983, Hawksworth et al. 1983). However, due to its astromatic nature, it has
long been considered to be an outlier within Phyllachoraceae (Cannon 1991). The family name
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Glomerellaceae was invalidly published by Locquin (1984) and was ignored until preliminary
sequence-based studies and ontogenic research confirmed that Colletotrichum and Phyllachora did
not belong to the same order (Uecker 1994). In the 9th edition of the Dictionary of Fungi the family
name Glomerellaceae was adopted and placed with an uncertain position within the subclass
Sordariomycetidae (Kirk et al. 2001). Zhang et al. ( 2006) validated Glomerellaceae with a Latin
description, while placing it within Hypocreomycetidae. Kirk et al. (2008) placed this family as an
uncertain position in the subclass Hypocreomycetidae.
The phylogenetic position of
Glomerellaceae was further clarified and validated by Réblová et al. (2011) in a study using ITS,
LSU, SSU and rpb2 genes and two new families, Australiascaceae and Reticulascaceae, occupied a
common clade with Glomerellaceae. Maharachchikumbura et al. (2016b) provided evidence for the
phylogenetic position of Glomerellaceae within Glomerellales. This family is monotypic with the
single genus Colletotrichum.
Ecological and economic significance of Glomerellaceae
Colletotrichum includes many plant pathogens of major importance causing diseases of a
wide variety of plants (Cannon et al. 2012b, Jayawardena et al. 2016, Marin-Felix et al. 2017). Fruit
production is especially affected.
Genus included in Glomerellaceae
Colletotrichum Corda, Deutschl. Fl., 3 Abt. (PilzeDeutschl.) 3(12): 41 (1831)
Index Fungorum number: IF7737; 894 morphological species ( Species Fungorum 2 0 2 0 ) ,
<250 species with sequence data (Jayawardena et al. 2016).
Type species – Colletotrichum lineola Corda
Notes – Colletotrichum is the asexual morph of Glomerella, but Colletotrichum was selected
for conservation by Maharachchikumbura et al. (2015). In this entry we introduced a new species,
Colletotrichum orchidis.
Colletotrichum orchidis Jayaward., Camporesi & K.D. Hyde, sp. nov.
Fig. 108
Index Fungorum number: IF556214; Facesoffungi number: FoF05784
Etymology – Refers to the host genus.
Holotype – MFLU 16-2551.
Saprobic on aerial stem of Orchis sp. Sexual morph: Undetermined. Asexual morph:
Coelomycetous. Conidiomata 55–135 μm ( x = 95 μm, n = 10) diam., black, acervulus, oval,
solitary or gregarious, comprising dark brown, roundish cells from which setae and conidiophores
develop. Setae straight or ± bent, abundant, dark brown to light brown, becoming paler towards the
apex, smooth-walled, 3–5-septate, 105 μm long, base cylindrical, slightly inflated, 5. 6 μm diam,
apex acute to rounded. Chlamydospores not observed. Conidiophores simple, to 65 μm long,
hyaline to pale brown, smooth-walled. Conidiogenous cells 8–11 × 3.1–4.4 μm ( x = 10.5 ± 1.8 ×
3.6 ± 0.4 μm, n = 20) , hyaline, smooth-walled, cylindrical to slighty inflated, opening 0.5–1 μm
wide, collarette or periclinal thickening not observed. Conidia 15.9–20.3 × 1.9–3.2 μm ( x = 18.2 ±
1.5 × 2.5 ± 0.6 μm, n = 40), L/W ratio 7.3, hyaline, smooth or verruculose, aseptate, very variable
in size and shape, some strongly curved, strongly curved towards the often broadly rounded apex
than towards the truncate base, some small conidia almost straight, guttulate. Appressoria 8.2–13.1
× 4.5–8.2 μm ( x = 11.5 ± 3.2 × 4.9 ± 1.4 μm, n = 10), L/W ratio 2.3, solitary or in loose groups,
single-celled, olivaceous brown to dark brown, irregularly-shaped, but often globose or clavate,
smooth-walled.
Culture characteristics – Colonies on PDA flat with entire margin, aerial mycelium sparse,
short, pale olivaceous-grey, iron-grey acervuli can be observed mainly on the edge of the colony
after 7 days; reverse olivaceous green, reaching 60–75 mm in 7 d at 18 °C.
Material examined – ITALY, Province of Forlì-Cesena, near Premilcuore, on living dead
aerial stem of Orchis sp. (Orchidaceae), 5 October 2016, E. Camporesi IT3118 (MFLU 16-2551,
holotype), extype-living culture MFLUCC 17-1302; ibid cultures KUMCC 17-0119, JZB330118.
595
GenBank numbers – ITS: MK502144, gapdh: MK496857, CHS: MK496855, act:
MK496853, tub2: MK496859 (MFLUCC 17-1302), ITS: MK502143, gapdh: MK496858, CHS:
MK496856, act: MK496854, tub2: MK496860 (JZB330118).
Figure 107 – One of the 100 most phylogenetic tree generated by maximum parsimony analysis of
combined ITS, GAPDH, CHS, ACT and β-tubulin sequenced data of dematium species complex.
Seventeen strains are included in the analyses, which comprise 1780 characters including gaps (509
characters for ITS, 267 characters for GAPDH, 250 characters for CHS, 237 characters for ACT,
501 characters for β-tubulin) after alignment. Colletotrichum nigrum (CBS 169.49, Glomerellaceae,
Glomerellales) is used as the outgroup taxon and the tree is rooted with. Single gene analyses were
carried out and the topology of each tree had clade stability. Tree topology of the maximum
parsinmony analysis is similar to Bayesian analysis. The maximum parsimonious dataset consisted
of 1182 constant, 343 parsimony-informative and 255 parsimony-uninformative characters. The
parsimony analysis of the data matrix resulted in the maximum often equally most parsimonious
trees with a length of 939 steps (CI=0.804, RI=0.774, RC=0.623, HI=0.196) in the first tree.
Bootstrap support values for MP greater than 75% and Bayesian posterior probabilities greater than
0.90 are given near nodes respectively. Ex-type strains are in bold and black. The newly generated
sequences are indicated in blue.
Notes – Colletotrichum orchidis is only known from aerial stem of Orchis in Italy. The
conidia of this species resemble those of several species complexes in Colletotrichum (e.g.
dematium species complex, graminicola species complex, spaethianum species complex and
truncatum species complex) (Jayawardena et al. 2016). Based on DNA sequences, C. orchidis falls
within the C. dematium species complex clade and forms a separate branch as a sister taxon of C.
dematium ( Fig. 107) . A BLASTn search of NCBI GenBank with the ITS sequence of the new
species, showed 96% similarity to several Colletotrichum species with curved conidia. The closest
match in a BLASTn search in GenBank with the gapdh sequence of the new species showed 99%
similarity to C. dematium (2bp differences) and C. lineola (4bp differences). CHS sequence showed
a 98% similarity to C. dematium ( 6 bp differences) and C. insertae ( 7 bp differences) and act
sequences also showed a 98% similarity to C. dematium (4 bp differences). tub2 sequence showed
99% similarity to C. dematium (1bp differences) and C. lineola (1 bp difference). Colletotrichum
596
orchidis differs from C. dematium by smaller conidia and a lower L/W value (C. dematium 21.3 ×
3.5, L/W = 6.0).
Figure 108 – Colletotrichum orchidis (MFLUCC 17-1302, holotype). a-c Appearance of
conidiomata on host. d Setae. e Base of seta. f, h Conidiophores. g Conidiogenous cell. i-k Conidia.
l-n Appressoria. o 3 day old culture on PDA. p 10 day old culture on PDA. Scale bars: b, c = 200
µm, d, f, h = 20 µm, e, g, i-k, l-n = 10 µm.
Gnomoniaceae G. Winter, Rabenh. Krypt.-Fl., Edn 2 (Leipzig) 1.2: 570 (1886)
Index Fungorum number: IF80810; Facesoffungi number: FoF06299; 485 species.
Saprobic on bark and leaves of overwintered plants. Sexual morph: Stromata lacking, or
poorly to well-developed, scattered, erumpent, pustuliform with one or rarely two ascomata or
valsoid, broadly elliptic to rounded, large. Ectostromata well-developed, brown to black, thick
ectostromatic disc at perithecial necks. Ascomata immersed to erumpent, solitary or aggregated,
globose to subglobose, black, coriaceous, thin-walled, with one or more long, central or eccentric
necks with hyaline periphyses. Peridium comprising few layers of brown, thick-walled cells of
597
textura angularis. Paraphyses few, hyaline, septate, cellular. Asci 8–32-spored, unitunicate, oval,
fusiform to almost filiform, short pedicellate, with a distinct, J-, apical ring. Ascospores biseriate,
overlapping uniseriate to fasciculate, hyaline, oval, fusiform, ovoid to subulate, small, unicellular to
1-septate, rarely multiseptate, ends mostly rounded, rarely pointed, appendages absent or subulate,
navicular or whip-shaped, smooth-walled. Asexual morph: Coelomycetous. Conidiomata acervuli
or pycnidial, subcuticular, papillate or not, oblate to globose, black, thick-walled, with one chamber
containing whitish conidial mass. Conidiophores simple, filiform to fusiform, annellations visible
or invisible, densely branched. Conidiogenous cells usually phialidic, rarely with a few annellidic
scars, irregular in shape, lageniform to cylindrical, gradually tapering to ends for one quarter to
three quarters of their length, or abruptly narrowing to long neck at about half of the phialide
length, or abruptly narrowing at apex, straight or curved, sometimes asymmetric swollen nodes,
proliferating into other conidiogenous cells at basal or middle part. Conidia broadly ellipsoid to
oval, sometimes obovoid, allantoid, occasionally curved or sinuate to slightly angular, hyaline,
often unicellular (adapted from Senanayake et al. 2018).
Type genus – Gnomonia Ces. & De Not.
Notes – Gnomoniaceae was introduced to accommodate fungal species having upright
perithecia with or without long or short neck and presence or absence of stromatic tissues (Winter
1886). Species in this family are pathogens or endophytes in leaves of herbaceous or woody trees
(Rossman et al. 2007). Betulaceae, Fagaceae, and Salicaceae are the most common host families for
the gnomoniaceous taxa (Mejia et al. 2011). Gnomoniaceae comprises 34 genera (Senanayake et al.
2018, Minoshima et al. 2018).
Ecological and economic significance of Gnomoniaceae
Species in Gnomoniaceae mainly occur on living, fallen or attached, overwintered leaves and
petioles, leaf blades or herbaceous stems and rarely on woody substrates (Sogonov et al. 2008).
Members of this family are mostly reported from temperate regions. They occur as endophytes of
woody plants or pathogens causing disease to economically important hardwood trees (Danti et al.
2002, Green 2004, Moricca & Ragazzi 2008). Sycamore anthracnose by Apiognomonia veneta,
strawberry stem rot by Gnomoniopsis fructicola, foliar disease of birch and dieback of young
shoots by Discula betulae, and walnut anthracnose is caused and leaf blotch disease by
Ophiognomonia leptostyla are common diseases caused by members of this family (Maas 1998,
Green 2004, Green & Castlebury 2007, Pennycook 2007).
Genera included in Gnomoniaceae
Alnecium Voglmayr & Jaklitsch, Persoonia 33: 76 (2014)
Index Fungorum number: IF805342; 1 species with sequence data.
Type species – Alnecium auctum (Berk. & Broome) Voglmayr & Jaklitsch
Notes – Alnecium was introduced and typified by A. auctum (Voglmayr & Jaklitsch 2014)
which was collected from corticated dead branches of overwintered plants. Alnecium auctum is
considered as a saprobe (Voglmayr & Jaklitsch 2014). The genus is characterized by immersed
stroma host tissues and fusiform ascospores (Senanayake et al. 2017a).
Ambarignomonia Sogonov, Stud. Mycol. 62: 35 (2008)
Index Fungorum number: IF512170; 1 species with sequence data.
Type species – Ambarignomonia petiolorum (Schwein.) Sogonov
Notes – This is a pathogen on petioles and leaves of Liquidambar styraciflua.
Ambarignomonia is distinguished from other genera in Gnomoniaceae by the whitish powdery
collar surrounding the central neck (Sogonov et al. 2008).
Amphiporthe Petr., Sydowia 24(1-6): 257 (1971)
Index Fungorum number: IF169; 3 morphological species (Species Fungorum 2020), 2
species with sequence data and molecular data available for an unnamed species.
598
Type species – Amphiporthe hranicensis (Petr.) Petr.
Notes – Amphiporthe is one of the well-developed stromatic genus in Gnomoniaceae which
currently comprises three species (Senanayake et al. 2018). However only A. hranicensis and A.
raveneliana (Thüm. & Rehm) M.E. Barr have molecular data.
Anisomyces Theiss. & Syd., Annls mycol. 12(3): 270 (1914)
Index Fungorum number: IF202; 2 morphological species (Species Fungorum 2020).
Type species – Anisomyces papilloideoseptatus (Henn.) Theiss. & Syd.
Notes – Petrak (1947b) emended this genus with Anisomyces papilloideoseptatus as a
synonym of A. nectrioides. This genus is morphologically distinct from other genera in having
stromatic tissues surrounding the short beak and brown, apiosporous ascospores (Barr 1978).
Anisomyces also comprises A. theissenii.
Apiognomonia Höhn., Ber. dt. bot. Ges. 35(8): 635 (1917)
Index Fungorum number: IF257; 23 morphological species (Species Fungorum 2020), 7
species with sequence data.
Type species – Apiognomonia veneta (Sacc. & Speg.) Höhn.
Notes – The asexual morph of Apiognomonia has been reported as discula-like (Sogonov et
al. 2007). Apiognomonia is the causative agent of several serious diseases such as sycamore
cankers or plane tree anthracnose on some overwintered plant families (Sogonov et al. 2007).
Apioplagiostoma M.E. Barr, Mycol. Mem. 7: 101 (1978)
Index Fungorum number: IF258; 3 species with sequence data.
Type species – Apioplagiostoma populi (E.K. Cash & Waterman) M.E. Barr
Notes – Apioplagiostoma was introduced by Barr (1978) for Plagiostoma populi, Sphaerella
acerifera and Plagiostomella carpinicola, considering their morphological distinctness. However,
Plagiostomella carpinicola was transferred to Gnomonia. Apioplagiostoma populi is the causative
agent of bronze leaf disease of Populus (Senanayake et al. 2018).
Asteroma DC., Fl. franç., Edn 3 (Paris) 6: 162 (1815)
Index Fungorum number: IF7260; 62 morphological species (Species Fungorum 2020), 2
species with sequence data and molecular data available for an unnamed species.
Type species – Asteroma phyteumatis DC.
Notes – Asteroma is an asexual morph genus, characterized by filiform to fusiform conidia
(Senanayake et al. 2017a) and its sexual morph is not clearly determined. This genus lacks
sufficient molecular data to demarcate its species boundaries.
Bagcheea E. Müll. & R. Menon, Phytopath. Z. 22(4): 417 (1954)
Index Fungorum number: IF490; 3 morphological species (Species Fungorum 2020).
Type species – Bagcheea castaneae E. Müll. & R. Menon
Notes – This genus comprises three species Bagcheea taiwanensis and B. albomaculans,
together with its type species. Bagcheea is distinguishable from the other genera of Gnomoniaceae
by its conspicuous ascospores, which contain granular cytoplasm is divided into two parts, with the
wider space forming a diplastic polarity (Senanayake et al. 2017a).
Chadefaudiomyces Kamat, V.G. Rao, A.S. Patil & Ullasa, Revue Mycol., Paris 38(1-2): 19 (1974)
Index Fungorum number: IF929; 1 morphological species.
Type species – Chadefaudiomyces indicus Kamat, V.G. Rao, A.S. Patil & Ullasa
Notes – Based on the distinct characters of Gnomoniaceae, Senanayake et al. (2017a)
accommodated this genus within Gnomoniaceae. The fungus was from living leaves of Celastrus
paniculata (Ullasa et al. 1974).
599
Clypeoporthe Höhn., Sber. Akad. Wiss. Wien, Math.-naturw. Kl., Abt. 1 128(7-8): 584 (1919)
Index Fungorum number: IF431517; 8 morphological species (Species Fungorum 2020).
Type species – Clypeoporthe monocarpa Höhn.
Notes – Clypeoporthe was introduced and typified by C. monocarpa. There are eight species
listed under this genus (Species fungorum 2020). However, some species in this genus have
eutypelloid configuration of ascomata in parenchymatous stromatic tissues. Clypeoporthe was
reduced to synonymy in Gnomonia by Monod (1983) while Kirk et al. (2008) mentioned
Clypeoporthe is the sexual morph of Phaeocytostroma. However, this is not proved by cultural
examination or molecular data and it is necessary to obtain DNA sequence data to resolve this
genus. Until that Senanayake et al. (2017a, 2018), Wijayawardhene et al. (2018) accepted this
genus within Gonomoniaceae.
Cryptosporella Sacc., Michelia 1(no. 1): 30 (1877)
Index Fungorum number: IF1333; 33 morphological species (Species Fungorum 2020), 20
species with sequence data and molecular data available for an unnamed species.
Type species – Cryptosporella hypodermia (Fr.) Sacc.
Notes – Cryptosporella species are from temperate regions and are saprobic, endophytic and
occasionally pathogenic on hardwood trees (Mejia et al. 2011). Cryptosporella hypodermia is
illustrated.
Figure 109 – Cryptosporella hypodermia (Material examined – CZECH REPUBLIC, Moravia,
Hranice city, Canyon mountain, Ulmus sp., October 1925, F. Petrak, F48426, holotype). a, b
Conidiomata on substrate. c Vertical cross section of conidioma. d-e Conidiogenous cells and
conidia. f Conidia. Scale bars: a, b = 200 μm, c = 100 μm, d = 20 μm, e, f = 10 μm.
Dictyoporthe Petr., Sydowia 9(1-6): 556 (1955)
Index Fungorum number: IF25095; 4 morphological species (Species Fungorum 2020).
Type species – Dictyoporthe ahmadii Petr.
Notes – This genus is distinct from other genera of Diaporthaceae in having muriform
ascospores (Petrak 1955a).
600
Diplacella Syd., Annls mycol. 28(1/2): 101 (1930)
Index Fungorum number: IF1677; 2 morphological species (Species Fungorum 2020).
Type species – Diplacella paulliniae (Gonz. Frag. & Cif.) Syd.
Notes – Diplacella is an endophytic fungal genus collected from leaves when form leaf spots.
Currently there are two species in this genus as Diplacella mararyensis and D. paulliniae (Index
Fungorum 2020). Diplacella species lack sequence data. However, most characters of Diplacella
fits with gnomoniaceous species and Senanayake et al. (2018) accepted this genus within
Gnomoniaceae.
Ditopella De Not., Hedwigia 2: 179 (1863)
Index Fungorum number: IF1677; 12 morphological species (Species Fungorum 2020), 3
species with sequence data and molecular data available for an unnamed species.
Type species – Ditopella fusispora De Not.
Notes – Asci in Ditopella species contain more than eight ascospores (generally 32).
Senanayake et al. (2017a) and Tian et al. (2018) showed the phylogenetic placement of this genus
within Gnomoniaceae.
Ditopellopsis J. Reid & C. Booth, Can. J. Bot. 45(9): 1479 (1967)
Index Fungorum number: IF1679; 4 morphological species (Species Fungorum 2020),
molecular data available for an unnamed species.
Type species – Ditopellopsis clethrae J. Reid & C. Booth
Notes – Ditopellopsis is characterized by pseudoparenchymatous stroma surrounding the
upright perithecium, short and broad necks, and ascospores with medium septum (Reid & Booth
1967). Ditopellopsis clethrae has been reported from Alnus, Clethra, Gaultheria, and Sophora.
Gloeosporidina Petr., Annls mycol. 19(3-4): 214 (1921)
Index Fungorum number: IF8353; 6 morphological species (Species Fungorum 2020).
Type species – Gloeosporidina moravica Petr.
Notes – This genus produces subepidermal to epidermal acervuli conidiomata, enteroblastic
conidiogenous cells and distinctive tiny, aseptate, hyaline conidia (Sutton 1980).
Gnomonia Ces. & De Not., Comm. Soc. crittog. Ital. 1(fasc. 4): 231 (1863)
Index Fungorum number: IF2099; 111 morphological species (Species Fungorum 2020), 28
species with sequence data.
Type species – Gnomonia gnomon (Tode) J. Schröt.
Notes – Gnomonia is characterized by astromatic, solitary, thin-walled, immersed perithecia
with long necks and ascospores with a single median septum. The sexual morph of Gnomonia
gnomon is illustrated below.
Gnomoniella Sacc., Michelia 2(no. 7): 312 (1881)
Index Fungorum number: IF2100; 28 morphological species (Species Fungorum 2020), 6
species with sequence data.
Type species – Gnomoniella tubaeformis (Tode) Sacc.
Notes – Gnomoniella species are generally associated with leaves and petioles of
overwintered plants (Barr 1998), although two species are known from freshwater (Luo et al.
2019).
Gnomoniopsis Berl., Icon. fung. (Abellini) 1(3): 93 (1893)
Index Fungorum number: IF2102; 17 morphological species (Species Fungorum 2020), 20
species with sequence data and molecular data available for 66 unnamed species.
Type species – Gnomoniopsis chamaemori (Fr.) Berl.
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Figure 110 – Gnomonia gnomon (Material examined – FINLAND, Helsinki, Helsinki University
Botanical Garden, overwintered fallen leaves of Corylus avellana L. (Betulaceae), 19 April 2004,
Shchigel, Dmitry S., BPI 844273, epitype). a Packet of herbarium. b Herbarium specimen. c
Ascomata on substrate. d Cross section of ascoma. e Peridium. f-h Asci. i-l Ascospores. Scale bars:
c = 1 mm, d = 200 µm, e, f-h = 20 µm, i-l = 10 µm.
Notes – Some species in this genus are phytopathogens, forming leaf blotch and petiole blight
of strawberry (Fall 1951, Bolton 1954, Van Adriechem & Bosher 1958, Maas 1998). Gnomoniopsis
is characterized by small, black perithecia composed of single central, marginal or lateral neck and
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immersed in the host tissue or stromatic tissues, oval to fusiform asci and one-septate, oval to
fusiform ascospores (Sogonov et al. 2008).
Maculatipalma J. Fröhlich& K.D. Hyde, Mycol. Res. 99(6): 727 (1995)
Index Fungorum number: IF6033; 1 morphological species.
Type species – Maculatipalma frondicola J. Fröhl. & K.D. Hyde
Notes – Maculatipalma frondicola fits in Gnomoniaceae in having foliicolous life mode,
solitary, thin-walled ascomata, and cylindrical to fusiform asci (Fröhlich & Hyde 1995).
Mamianiella Höhn., Ber. dt. bot. Ges. 35(8): 635 (1917)
Index Fungorum number: IF2994; 2 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Mamianiella coryli (Batsch) Höhn.
Notes – Senanayake et al. (2017a) synonymized Mamiania and Anisogramma under
Mamianiella based on their morphological similarity. Mamianiella species mostly colonize living
leaves.
Marsupiomyces Senan. & K.D. Hyde, Stud. Mycol. 86: 265 (2017)
Index Fungorum number: IF821555; 2 species with sequence data.
Type species – Marsupiomyces epidermoideus Perera, Senan., Bulgakov & K.D. Hyde
Notes – Marsupiomyces comprises M. quercina and the type. Species of this genus have been
reported only from Fagaceae (Senanayake et al. 2017a).
Millerburtonia Cif., Mycopath. Mycol. appl. 6(1): 26 (1951)
Index Fungorum number: IF8948; 1 species with sequence data.
Type species – Millerburtonia oyedaeae Cif.
Notes – Millerburtonia was introduced to accommodate Chalcosphaeria oyedaeae (Ciferri
1950). This is a parasitic, leaf spot forming taxon on leaves of Oyedea everbesinoides.
Occultocarpon L.C. Mejía & Zhu L. Yang, Fungal Divers. 52(1): 101 (2012)
Index Fungorum number: IF519819; 1 species with sequence data.
Type species – Occultocarpon ailaoshanense L.C. Mejía & Zhu L. Yang
Notes – Occultocarpon is characterized by aggregated, stromatic perithecia with thin, central
to eccentric necks and oblong elliptical-elongated, one-septate ascospores (Mejía et al. 2012).
Ophiognomonia (Sacc.) Speg., Revta Fac. Agron. Vet. Univ. nac. La Plata 2(19): 231 (1896)
Index Fungorum number: IF3604; 52 morphological species (Species Fungorum 2020), 48
species with sequence data and molecular data available for 27 unnamed species.
Type species – Ophiognomonia melanostyla (DC.) Sacc.
Notes – Members of this genus are leaf-inhabiting endophytes, pathogens or saprobes on
several plant families such as Betulaceae, Fagaceae, Juglandaceae, Lauraceae, Malvaceae,
Platanaceae, Rosaceae, Salicaceae, and Sapindaceae (Walker et al. 2012).
Phragmoporthe Petr., Annls mycol. 32(5/6): 354 (1934)
Index Fungorum number: IF4033; 3 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Phragmoporthe ploettneriana (Henn.) Petr.
Notes – This genus is characterized by multi-septate ascospores and 8-spored asci (Sogonov
et al. 2008).
Phylloporthe Syd., Annls mycol. 23(3/6): 348 (1925)
Index Fungorum number: IF4071; 1 species with sequence data.
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Type species – Phylloporthe vernoniae Syd.
Notes – The monotypic genus Phylloporthe is a parasite on living leaves of Vernonia
triflosculosa. Phylloporthe is distinct in having strongly erumpent, almost superficial stromata, an
aparaphysate hamathecium and thin-walled ascospores (Senanayake et al. 2018).
Plagiostoma Fuckel, Jb. nassau. Ver. Naturk. 23-24: 118 (1870)
Index Fungorum number: IF4157; 52 morphological species (Species Fungorum 2020), 39
species with sequence data and molecular data available for 18 unnamed species.
Type species – Plagiostoma euphorbiae (Fuckel) Fuckel
Notes – Plagiostoma species occur as pathogens, endophytes or saprobes on stems, branches,
twigs and leaves of woody and herbaceous plants (Mejía et al. 2011). The genus is mainly
differentiated from other genera in the family by the neck characters of perithecia and ascospore
morphology (Mejía et al. 2011).
Pleuroceras Riess., Hedwigia 1(6): 25 (1854)
Index Fungorum number: IF4242; 20 morphological species (Species Fungorum 2020), 4
species with sequence data and molecular data available for 6 unnamed species.
Type species – Pleuroceras ciliatum Riess.
Notes – Pleuroceras is characterized by ascomata with eccentric, lateral necks and elongated
ascospores (Barr 1978). Species of this genus are found on overwintered leaves of hardwood trees
in temperate regions.
Sirococcus Preuss, Linnaea 26: 716 (1855)
Index Fungorum number: IF9927; 28 morphological species (Species Fungorum 2020), 11
species with sequence data.
Type species – Sirococcus strobilinus Preuss
Notes – Sirococcus is a phytopathogenic or endophytic fungal genus (Senanayake et al.
2018).
Spataporthe Bronson, Klymiuk, Stockey & Tomescu, Int. J. Pl. Sci. 174(3): 279 (2013)
Index Fungorum number: IF800815; 1 morphological species.
Type species – Spataporthe taylorii Bronson, Klymiuk, Stockey & Tomescu
Notes – Spataporthe is typified by S. taylorii and it is the only genus fossil in Diaporthales
(Bronson et al. 2013). Spataporthe is distinct from other genera in Gnomoniaceae in having
perithecial necks with a bell-shaped chamber, a peridium with outer, large pseudoparenchymatous
layer and inner, thin filamentous layer, and clavate asci with thinly tapered bases which are
detached from the hymenium to float freely in the perithecium (Bronson et al. 2013).
Tenuignomonia Minosh., D.M. Walker & Hirooka, Mycoscience 60: 39 (2018)
Index Fungorum number: IF824890; 1 species with sequence data.
Type species – Tenuignomonia styracis Minosh., D.M. Walker & Hirooka
Notes – Tenuignomonia is only known from its type species, Tenuignomonia styracis which
has been collected solely from Styraxo bassia. This genus is distinct from other genera in having
long perithecial necks and septate ascospores. Minoshima et al. (2018) proved that Tenuignomonias
tyracis has weak pathogenicity on Styraxo bassia and it may promote early defoliation.
Uleoporthe Petr., Annls mycol. 39(4/6): 279 (1941)
Index Fungorum number: IF5660; 1 morphological species.
Type species – Uleoporthe orbiculata (Syd. & P. Syd.) Petr.
Notes – This genus is a leaf epiphyllous taxon forming circular, erumpent to superficial
stromata. Morphologically this genus is closer to Phyloporthe in having a foliicolous life mode,
solitary, globose, astromatic ascomata and fusiform asci (Cannon 2001).
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Uniseta Ciccar., Nuovo G. bot. ital. 54: 711 (1948)
Index Fungorum number: IF10354; 1 morphological species.
Type species – Uniseta flagellifera (Ellis & Everh.) Ciccar.
Notes – Uniseta is morphologically distinct in having sessile, widely fusiform asci deeply
immersed in a hyaline, cellular mass and fusiform, slightly curved, guttulate ascospores
(Senanayake et al. 2018).
Valsalnicola D.M. Walker & Rossman, Persoonia 29: 149 (2012)
Index Fungorum number: IF801276; 1 species with sequence data.
Type species – Valsalnicola oxystoma (Rehm) D.M. Walker & Rossman
Notes – Valsalnicola is different from other genera in Gnomoniaceae in having allantoid, 1septate ascospores and a black line surrounding the ascomatal cavity (Crous et al. 2012c).
Vismaya V.V. Sarma & K.D. Hyde, Nova Hedwigia 73(1-2): 247 (2001)
Index Fungorum number: IF28571; 1 morphological species.
Type species – Vismaya chaturbeeja V.V. Sarma & K.D. Hyde
Notes – This genus has distinct characters such as immersed perithecial ascomata with long
protruding necks, four-spored asci and hyaline, 1-celled, verruculose ascospores with appendages at
both ends (Sarma & Hyde 2001).
Gondwanamycetaceae Réblová, W. Gams & Seifert, Stud. Mycol. 68(1): 188 (2011)
Index Fungorum number: IF515439; Facesoffungi number: FoF01329; 14 species.
Pathogenic on terrestrial and aquatic plants or parasitic on beetles or saprobic in compost.
Sexual morph: Ascomata perithecial, black, necks cylindrical to filiform, tapered towards the apex,
terminating in ostiolar hyphae. Peridium fragile, thin-walled. Paraphyses lacking. Asci 8 to multispored, evanescent. Ascospores hyaline, aseptate, fusiform to lunate or falcate or allantoid with or
without a gelatinous sheath. Asexual morph: Hyphomycetous. Conidiophores macronematous,
mononematous, mono-verticillate or penicillate, brown. Conidiogenous cells phialidic. Conidia
hyaline, 1-celled, smooth-walled, cylindrical to allantoid, aseptate, slimy (adapted from
Maharachchikumbura et al. 2016b).
Type species – Knoxdaviesia M.J. Wingf., P.S. van Wyk & Marasas
Notes – Gondwanamycetaceae was introduced for Gondwanamyces and its asexual morph
Custingophora by Réblová et al. (2011). The phylogenetic relationship of the asexual genera
Knoxdaviesia and Custingophora were documented in earlier studies of Viljoen et al. (1999) and
subsequently in Kolařík & Hulcr (2009) who suggested that Knoxdaviesia and Custingophora
should be treated as synonyms. Van der Linde et al. (2012) and de Beer et al. (2013a) conducted a
separate treatment of these genera and proposed using Knoxdaviesia, the oldest name, over
Gondwanamyces (Hawksworth 2011, Moubasher et al. 2017). The apparent absence of interascal
filaments in the ascomatal centrum and hyaline, allantoid ascospores, with a hyaline sheath, giving
the spore a fusiform to lunate or falcate appearance, are characteristic features of the sexual morph
of this family. The asexual morphs are characterized by conidiophores which are erect, darkly
pigmented, and paler towards the apex, and are either monoverticillate, sometimes with a terminal
vesicle or divergently penicillate, with whorls of phialides producing hyaline conidia while the
conidiogenous loci are located at the base of the shallow collarette (Kolařík & Hulcr 2009,
Moubasher et al. 2017). Réblová et al. (2011) and Maharachchikumbura et al. (2015, 2016b)
confirmed the placement of this family in Microascales based on analysis of combined SSU, LSU
and rpb2 data.
Ecological and economic significance of Gondwanamycetaceae
Gondwanamycetaceae comprises plant pathogens mostly occurring on infructescences of
Protea species and parasites on bark beetles such as Cossonus (Wingfield et al. 1988, Marais et al.
1998, Van der Linde et al. 2012). Knoxdaviesia was first observed in an infructescence of Protea
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spp. infected by insects (Wingfield et al. 1988, Marais et al. 1998) whereas, some described species
of Knoxdaviesia are associated with Scolytidae (bark beetles) (Bright & Torres 2006, Kolařík &
Hulcr 2008). Knoxdaviesia cecropiae was isolated from the body of Scolytidae unipunctatus and
Knoxdaviesia scolytodis from galleries in the sapwood of Curcuma angustifolia (Kolařík & Hulcr
2009). Knoxdaviesia serotectus was isolated from discoloured plant material of C. angustifolia in
Costa Rica (Kolařík & Hulcr 2009). These species are also vectors for mites that are associated
with insects which infest Euphorbia species (Van der Linde et al. 2012). Gondwanamyces species
produced lesions on healthy succulent branches of Euphorbia spp. hence causing decline in the
populations in South Africa (Van der Linde et al. 2012). Gondwanamycetaceae species are able to
grow on all monosaccharides that occur in Protea nectar (Aylward 2017). There are no reports of
severe effects of Gondwanamycetaceae species on these plant species or beetles. No control
methods have been reported. However, secondary metabolites are likely to be produced by
Custingophora species that enable Protea spp. to tolerate bacteria, arthropod and nematode
predation (Aylward 2017).
Genera included in Gondwanamycetaceae
Custingophora Stolk, Hennebert & Klopotek, Persoonia 5(2): 195 (1968)
Index Fungorum number: IF7851; 5 morphological species (Species Fungorum 2020), 2
species with sequence data.
Type species – Custingophora olivacea Stolk, Hennebert & Klopotek
Notes – Custingophora species have mononematous conidiophores that terminate in obovoid
conidiogenous cells with distinct collarettes and conidia in slimy droplets (de Beer et al. 2013a,
Moubasher et al. 2017).
Knoxdaviesia M.J. Wingf., P.S. van Wyk & Marasas, Mycologia 80(1): 26 (1988)
Index Fungorum number: IF11113; 9 morphological species (Species Fungorum 2020), 6
species with sequence data.
Type species – Knoxdaviesia proteae M.J. Wingf., P.S. van Wyk & Marasas
Notes – Knoxdaviesia was established for K. proteae, and the asexual morph of this species
was described as Ceratocystiopsis proteae (Wingfield et al. 1988). Subsequently, Marais et al.
(1998) introduced the sexual morph genus Gondwanamyces based on G. proteae to accommodate
two ophiostomatoid species Ceratocystiopsis proteae (Wingfield et al. 1988) and Ophiostoma
capense (Wingfield & van Wyk 1993). Gondwanamyces was characterized by black, globose to
subglobose perithecia and aseptate ascospores, with or without a sheath. The asexual morph
Knoxdaviesia formed phialoconidia holoblastically from obovoid conidiogenous cells. Sequence
data showed that Knoxdaviesia could be accommodated in Custingophora (Kolařík & Hulcr 2009,
Moubasher et al. 2017). Van der Linde et al. (2012) rejected the concept of synonymy of
Knoxdaviesia in Custingophora and de Beer et al. (2013a) erected Knoxdaviesia over
Gondwanamyces according to the one fungus one name protocol (Hawksworth 2011, Hawksworth
et al. 2011, Moubasher et al. 2017). In this entry, Knoxdaviesia proteae is illustrated.
Graphiaceae Z.W. de Beer, Seifert & M.J. Wingf., CBS Diversity Ser. 12: 8 (2013)
Index Fungorum number: IF622184; Facesoffungi number: FoF01099; 59 species.
Saprobic on plant debris, woody substrates, sometimes causing wounds on tree bark or
associated with beetles, occasionally isolated from soil, manure, polluted water, also reported to
cause fungaemia in an immunosuppressed child post stem-cell transplant. Sexual morph:
Undetermined. Asexual morph: Hyphomycetous. Conidiomata macronematous, synnematous,
determinate. Conidiophores compact, pigmented, septate, penicillately branched in two to three
levels, forming metulae at the apex. Conidiogenous cells in whorls of two to six, phialidic, with
annellidic extensions. Conidia produced in a transparent, slimy droplet, aseptate, cylindrical to
obovoid, sometimes slightly curved with age, truncate at base, often with a distinct basal frill,
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hyaline. Rarely synanamorphic forming obovoid, pigmented conidia (adapted from de Beer et al.
2013b, Maharachchikumbura et al. 2016b).
Figure 111 – Knoxdaviesia proteae (Material examined – SOUTH AFRICA, Cape Province,
Stellenbosch, from flower within the inflorescence, infested by insects, L.J. Strauss, 7 October
1985, PREM 48924 holotype) and asexual morph redrawn from Wingfield et al. (1988). a
Herbarium material. b, c Ascomata on host. d Long neck of ascomata. e, f Asci. g Ascospores. h-j
Conidiophores, conidiogenous cells and conidia. Scale bars: b, c = 100 μm. d = 50 μm, g = 20 μm,
e, f = 5 μm.
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Figure 112 – Graphium penicillioides (Material examined – CZECH REPUBLIC, PRM 842988,
epitype). a, b Herbarium material. c Colony on the surface of the substrate. d, e Synnemata on the
host surface. f, g Conidiogenous cells. h-k Conidia. Scale bars: c = 200 μm, d = 100 μm, e = 50 μm,
f-h = 20, i-k = 5.
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Type genus – Graphium Corda
Notes – Based on the phylogenetic distance between Graphium and other families of
Microascales, Graphiaceae was introduced by de Beer et al. (2013b) and accommodated in
Microascales (de Beer et al. 2013b, Maharachchikumbura et al. 2015, 2016b). The family is
characterized by hyaline, aseptate, cylindrical to obovoid, curved conidia. In rare cases, a
synanamorph with monoblastic, obovoid, pigmented chlamydospore-like conidia are formed.
Synnematous asexual morphs of Ophiostoma were treated as Graphium (Hedgcock 1906,
Upadhyay 1981, Seifert & Okada 1993). Phylogenetic studies showed that the asexual morphs of
Ophiostoma clustered distantly from Graphium sensu stricto species and thus they were
accommodated in Pesotum (Okada et al. 1998, 2000). However, species with annellidic
conidiogenous cells and those with Microascaceae affinities were retained in Graphium (Okada et
al. 1998, 2000, Jacobs et al. 2003). Graphium was referred to a separate family with nine accepted
species based on the description of G. penicillioides and available DNA sequence data (de Beer et
al. 2013b, Maharachchikumbura et al. 2015, 2016b). There are 14 species listed under Graphiaceae
based on the molecular data with the addition of new species such as, G. jumulu, G. scolytodis, G.
kuroshium (https://www.ncbi.nlm.nih.gov/taxonomy; Na et al. 2018) and many more
morphological species.
Hongsanan et al. (2017) list three families in the Microascales: Microascaceae,
Halosphaeriaceae and Graphiaceae with a stem age of 171–241 MYA, while Hyde et al. (2017a)
indicate there is grounds for the introduction of Graphidiales with a stem age of 166 MYA.
However, no formal designation was made (Hyde et al. 2017a).
Ecological and economic significance of Graphiaceae
Usually they cause wounds or colonize the outer bark of trees and are considered as
pathogens. Graphium basitruncatum causes invasive disease fungaemia in an immunosuppressed
child post stem-cell transplant (Kumar et al. 2007, El Feghaly et al. 2012).
Genus included in Graphiaceae
Graphium Corda, Icon. fung. (Prague) 1: 18 (1837)
Index Fungorum number: IF8398; 59 morphological species (Species Fungorum 2020), 5
species with sequence data.
Type species – Graphium penicillioides Corda
Notes – Graphium species are plant pathogens and also found in soil, plant debris, woody
substrates, manure, and polluted water (de Beer et al. 2013b, Maharachchikumbura et al. 2015,
2016b, Wijayawardene et al. 2017a). The genus was believed to have ophiostomatoid affinities
(Goidànich 1935, Upadhyay 1981, Seifert & Okada 1993, de Beer et al. 2013b). Cruywagen et al.
(2010), based on SSU and ITS sequence data, included eight described and seven undescribed
species, including G. penicillioides, in Graphium sensu stricto. de Beer et al. (2013b) provided a
description of Graphium sensu stricto and referred Graphium to the monophyletic family
Graphiaceae (Microascales). Phylogenetically Graphiaceae, are distantly placed from the
Microascaceae (Zhang et al. 2006, Spatafora et al. 2006, Schoch et al. 2009, Maharachchikumbura
et al. 2015, 2016b). Members of Graphium are characterized by distinctive, erect, black synnemata,
each bearing a single, terminal, 1-celled, hyaline conidium produced from phialidic conidiogenous
cell with annellidic extensions.
Graphostromataceae M.E. Barr, J.D. Rogers & Y.M. Ju, Mycotaxon 48: 533 (1993)
Index Fungorum number: IF81957; Facesoffungi number: FoF00624; 101 species.
Saprobic on trunks, branches, and twigs of angiosperm plants, endophytic and pathogenic.
Sexual morph: Stromata erumpent, widely effuse, irregular in outline, bipartite with ectostroma and
entostroma, ectostroma develops first, young ectostroma surface buff, with matted mycelium,
ectostroma develops beneath, hard and brittle. Ascomata perithecial, immersed in the enostroma,
bottle-shaped, more rarely pyriform or obpyriform, black, opaque, monostichous, arranged in
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diatrypoid configuration in entostroma, compressed, varying in size with carbonaceous, doliiform
ascomatal embedded necks. Peridium comprising hyaline, multi-layered, thin, indistinct, flattened
cells. Paraphyses sparse, elongate, tapering from wide base. Asci 8-spored, unitunicate, narrowly
clavate, short-pedicellate, rounded to subtruncate at the apex, with J+, discoid, apical ring,
inconspicuously bluing in Melzer’s reagent. Ascospores uni-multiseriate, unicellular, allantoid and
hyaline or brown and ellipsoid, at times with appendages, with or without germ slits, without
dehiscent perispores. Stromatal pigments absent. Asexual morph: in culture nodulisporium-type,
most often periconiella-like or xylocladium-like (reduced form of Periconiella) (adapted from
Wendt et al. 2018).
Type genus – Graphostroma Piroz.
Notes – Barr et al. (1993) introduced Graphostromataceae to accommodate Graphostroma
based on species having ascospores typical of the Diatrypaceae and conidiogeneous structures
typical of the Hypoxylon. Maharachchikumbura et al. (2016b) synonymized Graphostromataceae
with Xylariaceae and treated Graphostroma as a genus in Xylariaceae. In recent multi-gene
phylogenetic analyses reveal that Graphostromataceae is close to Barrmaeliaceae and Xylariaceae
(Daranagama et al. 2018, Voglmayr et al. 2018, Wendt et al. 2018). Hongsanan et al. (2017) and
Hyde et al. (2017a) provided a 63 MYA stem age for Graphostromataceae and accepted the family
based on molecular clock analyses. Daranagama et al. (2015) and Senanayake et al. (2015)
confirmed the monophyletic association of Biscogniauxia, Graphostroma and camillea-like genera
that share common characters. Based on phylogeny and morphology, Theissenia was excluded
from the family and accepted in Hypoxylaceae (Wendt et al. 2018, this study). Five accepted
genera are contained in the family, Biscogniauxia, Camillea, Graphostroma, Obolarina, and
Vivantia.
Ecological and economic significance of Graphostromataceae
Graphostromataceae species are mainly saprobes on terrestrial wood. Among the genera in
the family, Biscogniauxia consists of known pathogens and endophytes. Biscogniauxia
mediterranea, a causal organism charcoal canker is widely distributed on European oaks that
probably initially invades the host via the leaves and can be detected as endophyte in apparently
healthy plants, but can turn pathogenic in water-stressed host plants (Edwards et al. 2003).
Raimondo et al. (2016) described Biscogniauxia rosacearum, a charcoal canker fungus on Pyrus
sp., Prunus sp., and Cydonia sp. In addition, Biscogniauxia nummularia, B. nothofagi and B.
uniapiculata are canker causing species (Granata & Sidoti 2004, Nugent et al. 2005). Interestingly,
ambrosia beetles (Platypus cylindrus; Coleoptera, Platypodidae) act as vectors for B. mediterranea
(Inácio et al. 2011). Amand et al. (2012) described bioactive compounds viz. xylaranone and
xylaranol B isolated from endophytic B. nummularia from Cephalotaxus harringtonia which have
antigerminative activity on Raphanus sativus seeds might act as latent pathogens. Obolarina
dryophila was described as an inner bark endophyte from Salix alba that is propagated by insects
because these authors found viable ascospores from the gut of Gasterocercus depressirostris
(Coleoptera, Curculionidae) beetles (Pažoutová et al. 2010). Even though little is still known about
the lifestype of most Graphostromataceae species, the above cited examples indicate that these taxa
may play a very important role in the ecology of their hosts, and they should be studied and
monitored further, in particular with regard to the challenges that agriculture and forestry are now
facing during the imminent global climate change.
Genera included in Graphostromataceae
Biscogniauxia Kuntze, Revis. gen. pl. (Leipzig) 2: 398 (1891)
Index Fungorum number: IF582; 53 morphological species (Species Fungorum 2020), 23
species with sequence data.
Type species – Biscogniauxia nummularia (Bull.) Kuntze
Notes – Biscogniauxia species develop within the bark and emerge via the overlying bark. A
periconiella-like asexual morph is associated with several Biscogniauxia species and sometimes
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vary from nodulisporium-like to periconiella-like (Stadler et al. 2013). The genus appears
paraphyletic together with morphologically dissimilar genera, such as Camillea, Obolarina and
Graphostoma (Pažoutová et al. 2010, Raimondo et al. 2016, Daranagama et al. 2018). A case could
be made to merge all these genera, were it not for the fact that over 90% of the species remain to be
studied by multi-gene genealogies and of over 80% only the less relevant rDNA sequences are
extant. A new species, Biscogniauxia mangiferae, is illustrated in this entry.
Biscogniauxia mangiferae Samarak. & K.D. Hyde, sp. nov.
Fig. 113
Index Fungorum number: IF556894; Facesoffungi number: FoF06780
Etymology – Named after the host genus on which the fungus occurs.
Holotype – MFLU 18–0827.
Saprobic on dead branch of Mangifera indica L. (Anacardiaceae). Sexual morph: Stromata
0.5–3.5 cm long, 0.25–1.5 cm width, applanate, dull brown to black, carbonaceous. Perithecia 185–
225 μm (x̅ = 205 μm) diam., 310–430 μm (x̅ = 360 μm) high, ovoid to obpyriform, laterally
compressed. Ostiole pointed, raised. Paraphyses 3.9–6.5 μm (x̅ = 5.4 μm, n = 20) wide, hyaline,
septate, round apex, guttulate. Asci 80.7–94.4 × 6.7–8.6 μm (x̅ = 88.6 × 7.8 μm, n = 25), 8-spored,
unitunicate, cylindrical, pedicellate, with J+, apical ring bluing in Melzer’s reagent, wedge-shaped
with 2.6–5 μm (x̅ = 4.1 μm, n = 25) high, 2.5–3.7 μm (x̅ = 3.1 μm, n = 25) broad. Ascospores 8.8–
11.7 × 5.3–6.5 μm (x̅ = 10.3 × 6 μm, n = 35), uniseriate, unicellular, ellipsoid, rounded ends, dark
brown to black at mature, smooth, with straight germ slit spore-length. Asexual morph:
Undetermined.
Material examined – THAILAND, Doi Phu Kha, Naan 55000, dead branch of Mangifera
indica (Anacardiaceae), 4 August 2017, M.C. Samarakoon, SAMC049 (MFLU 18-0827, holotype);
ibid. (HKAS 102330, isotype).
GenBank numbers – ITS: MN337232, LSU: MN336236, rpb2: MN366247, tub2:
MN509782.
Notes – Biscogniauxia nummularia (type species), B. capnodes, B. petrensis and the strain
isolated in this study clustered with high statistical support (98% ML). Our strain is sister to B.
petrensis (100% ML). Biscogniauxia petrensis was described by Zhang et al. (2017c) from a karst
cave ecosystem with its asexual morph. Morphological comparisons are not possible and ITS
comparison shows 5.7% bp differences among these two strains. Biscogniauxia capnodes differs
from our strain by its obovoid to tubular perithecia, discoid apical ring and large ascospores.
Biscogniauxia reticulospora described from undetermined corticated wood (Thailand) differs from
our strain by its comparatively larger ascospores (27–34 × 14–15.5 μm) with ornamentations and
reticulations (Ju & Rogers 2001), as compared to ascospores in our strain (8.8–11.7 × 5.3–6.5 μm).
Camillea Fr., Summa veg. Scand., Section Post. (Stockholm): 382 (1849)
Index Fungorum number: IF777; 44 morphological species (Species Fungorum 2020), 4
species with sequence data.
Type species – Camillea leprieurii (Mont.) Mont.
Notes – Camillea species are characterized by long, erect, hard, carbonaceous, cylindrical,
black stromata with light coloured ascospores (Læssøe et al. 1989). Many of the species of
Camellea remain to be cultured and sequenced.
Graphostroma Piroz., Can. J. Bot. 52(10): 2131 (1974)
Index Fungorum number: IF2132; 1 species with sequence data.
Type species – Graphostroma platystomum (Schwein.) Piroz.
Notes – Pirozynski (1974) erected Graphostroma for Sphaeria platystoma with
nodulisporium-like asexual morph. The combination of nodulisporium-like asexual morph and a
diatrypaceae-like sexual morph having allantoid ascospores are considered as characteristic of
Graphostromataceae with the type species Graphostroma platystomum (Barr et al. 1993).
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Phylogenetic analyses by Daranagama et al. (2018), and Wendt et al. (2018) confirmed that
Graphostroma clustered with Biscogniauxia and Camillea in Graphostromataceae.
Figure 113 – Biscogniauxia mangiferae (MFLU 18-0827, holotype). a, b Stroma on the host. c, d
Cross section of stromata showing the ascoma and ostiole. e-g Asci. h, i J+, apical ring in Melzer’s
reagent. j Paraphyses. k-n Ascospores. Scale bars: b, c = 1000 μm, d = 500 μm, e-h = 20 μm, i, j =
10 μm, k-n = 10 μm.
Obolarina Pouzar, Česká Mykol. 40(1): 7 (1986)
Index Fungorum number: IF25727; 2 morphological spcies with sequence data.
Type species – Obolarina dryophila (Tul. & C. Tul.) Pouzar
Notes – Obolarina species are endophytic, saprotrophic, or rarely parasitic on wood of
Quercus sp. (Daranagama et al. 2018). Mirabolfathy et al. (2013) introduced Obolarina persica
from Iran, where it is widely associated with dying Quercus brantii. The phylogenetic position of
Obolarina was determined in Graphostromataceae (Mirabolfathy et al. 2013, Daranagama et al.
2018, Wendt et al. 2018).
612
Vivantia J.D. Rogers, Y.M. Ju & Cand., Mycol. Res. 100(6): 672 (1996)
Index Fungorum number: IF27648; 1 morphological spcies.
Type species – Vivantia guadalupensis J.D. Rogers, Y.M. Ju & Cand.
Notes – Daranagama et al. (2018) and Wendt et al. (2018) tentatively placed Vivantia in
Graphostromataceae based on nodulisporium-like asexual morph and bipartite stromata. However,
this species is only known from a single collection made in the Caribbean many decades ago and
fresh collection and sequence data are needed.
Halosphaeriaceae E. Müll. & Arx, Can. J. Bot. 50: 1951 (1972)
Index Fungorum number: IF80832; Facesoffungi number: FoF01294; 171 species.
Saprobic on algae, immersed or submersed on phanerogams, wood, bark, leaves, and other
cellulosic plant remains, grains of sand, or on calcareous shell fragments, rarely parasitic or
symbiotic, found in marine (oceans, mangroves and estuaries) and freshwater habitats. Sexual
morph: Ascomata subglobose, cylindrical or pyriform, hyaline or dark; sometimes subiculate, rarely
stromatic; superficial or immersed. Ostioles papillate to long cylindrical; ostiolar canal with
periphyses or pseudoparenchyma; rarely without ostioles. Peridium soft or subcarbonaceous,
composed of flattened, thick- or thin-walled cells. Centre of immature ascomata consisting of
polygonal, thin-walled, pseudoparenchymatic cells, sometimes with pits, at maturation separating
to form catenophyses or compressed by the asci and dissolving. Paraphyses absent. Asci fusiform,
clavate or rarely subglobose, with or without apical structures, thin-walled, 1-layered, persistent or
swelling and deliquescing at or before ascospore maturity. Hymenial layer at base of venter, flat or
convex. Mature ascospores filling the venter of the ascoma, released singly through the ostiole or
rarely within the ascus, which swells after dispersal. Ascospores overlapping 2-3 seriate, hyaline or
light brown, 1- multi-celled, mostly with characteristic ornamentations, appendages or gelatinous
sheaths, or both. Asexual morph: Hyphomycetous. Hyphae hyaline or brown, superficially or
immersed, septate, branched. Conidiophores simple or micronematous, hyaline or mid to dark
brown, paler towards the apex, unbranched, walls smooth, thin or thick-walled, terminal,
integrated, monoblastic, determinate or arise as erect lateral branches from the cells of the
vegetative hyphae, wide at the base, tapering only slightly above or arising singly or in groups of 24 from a small stroma or flexuous, initially short and simple becoming longer and many times
septate (up to 10), old conidiophore apices become displaced laterally as the conidiophore
elongates and persists as short conoid denticles. Conidiogenous cells erect, some are with a distinct
collarette, produce conidia terminally in a sympodial fashion or monoblastic, integrated, terminal,
determinate. Conidia solitary, hyaline to bright reddish brown, holoblastic arising singly from the
apices of the conidiogenous cells or develop as a small swelling at the end of the collarette, thinwalled, smooth, fusoid, (2) 7-11 septate, sigmoid with a flat detachment scar or initially oval
gradually the upper half widens and becomes turbinate and finally triangular-shaped, each corner of
the triangle with a long, hair-like divergent process or helicoids, semi-contorted, mostly coiled
0.75-1 times, 3-4 (5) septate, constricted at the septa; cells increase in size and pigments from base
to apex: apical cell conspicuously swollen, subglobose, darker than the others, basal cell cylindrical
and tapering (adapted from Maharachchikumbura et al. 2016b).
Type genus – Halosphaeria Linder
Notes – Halosphaeriaceae which was discussed by Müller & von Arx (1962) and Eriksson
(1984) currently comprises 163 species in 64 genera (Pang 2002, Jones et al. 2009, 2015, 2017,
2019, Maharachchikumbura et al. 2015, Wijayawardene et al. 2017a, 2018a). The Halosphaeriaceae
was validated by Kohlmeyer (1972) and assigned as a single family to Halosphaeriales
(Hawksworth & Eriksson 1986). Halosphaeriaceae was previously polyphyletic and the genera
Lulworthia and Lindra were referred to a new family Lulworthiaceae (Campbell et al. 2003, Pang
et al. 2003, Koch et al. 2007). Even though Hibbett et al. (2007) and Schoch et al. (2007) referred
Halosphaeriaceae to Microascales, some other researchers included it in Halosphaeriales (Zhang et
al. 2006, Tang et al. 2007, Jones et al. 2009). Recent studies by Jones et al. (2015) and
Maharachchikumbura et al. (2015, 2016b) established the placement of the Halosphaeriaceae as
613
one of the families in the Microascales. Perithecial ascomata, necks (usually with periphyses), the
presence of catenophyses that easily deliquesce, unitunicate, thin-walled asci that deliquesce early,
with or lacking an apical ring and appendaged ascospores are the most common characters of this
family (Jones 1995, Campbell et al. 2003, Pang et al. 2003, Koch et al. 2007), while some genera
have been introduced based on ascospore appendage morphology and ontogeny, such as
Bovicornuta, Kohlmeyeriella, Ondiniella, and Marinospora (Jones et al. 1983, 1984, Jones 1995)
but subsequently confirmed by molecular data. Halosphaeriaceous species are primarily marine,
found on wood (especially mangrove wood) or seagrasses and are cosmopolitan in distribution
(Jones 2011, Jones & Pang 2012, Jones et al. 2013). Some species are found in freshwater habitats,
which was belong to the genera Aniptodera, Lignincola, Luttrellia, Magnisphaera, Naïs,
Natantispora, Oceanitis, Panorbis and Phaeonectriella and the majority are lignicolous (Pang &
Jheng 2012, Cai et al. 2014). Around 75% of halosphaeriaceous species have been sequenced
(Jones et al. 2017).
Ecological and economic significance of Halosphaeriaceae
Halosphaeriaceous taxa play a significant role in the decomposition of complex organic
matter in the marine ecosystem by secreting wood-modifying enzymes (cellulases, laccases and
lignases) that degrade lignocelluloses (Jones & Hyde 1988). Many species are able to cause soft rot
decay of wood e.g. Halosphaeria appendiculata (Mouzouras 1986). Hence, contribute to the
generation of particulate and dissolved organic matter available in the food web of the oceanic
environment (Jones & Pang 2012). Several studies have shown that mangrove associated
halosphaeriaceous fungi are capable of producing hydrolytic and oxidative enzymes, thus
indicating their probable role in recycling of lignocellulose in mangroves (Bucher et al. 2004). A
number of halosphaeriaceous species have been shown to produce bioactive secondary metabolites,
e.g. Corollospora maritima the source of corollosporine (Liberra et al. 1998).
Genera included in Halosphaeriaceae
We do not provide notes on each genus as these can be found at marinefungi.org (Jones et al.
2019). A monograph on genera of Halosphaeriaceae is also planned by Dayarathne et al. (2020).
Alisea J. Dupont & E.B.G. Jones, Mycol. Res. 113(12): 1358 (2009)
Index Fungorum number: IF513289; 1 species with sequence data.
Type species – Alisea longicolla J. Dupont & E.B.G. Jones
Amphitrite S. Tibell, Svensk Mykologisk Tidskrift 37(2): 45 (2016)
Index Fungorum number: IF817480; 1 morphological species.
Type species – Amphitrite annulata S. Tibell
Aniptodera Shearer & M.A. Mill., Mycologia 69(5): 893 (1977)
Index Fungorum number: IF198; 19 morphological species (Species Fungorum 2020), 3
species with sequence data.
Type species – Aniptodera chesapeakensis Shearer & M.A. Mill.
Aniptosporopsis K.L. Pang, C.L. Lu, W.T. Ju & E.B.G. Jones, Bot. Mar.: 459 (2016)
Index Fungorum number: IF818199; 1 species with sequence data.
Type species – Aniptosporopsis lignatilis (K.D. Hyde) K.L. Pang, C.L. Lu, W.T. Ju & E.B.G.
Jones
Anisostagma K.R.L. Petersen & Jørg. Koch, Mycol. Res. 100: 209 (1996)
Index Fungorum number: IF27558; 1 morphological species.
Type species – Anisostagma rotundatum K.R.L. Petersen & Jørg. Koch
614
Antennospora Meyers, Mycologia 49: 501 (1957)
Index Fungorum number: IF219; 2 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Antennospora quadricornuta (Cribb & J.W. Cribb) T.W. Johnson
Appendichordella R.G. Johnson, E.B.G. Jones & S.T. Moss, Can. J. Bot. 65(5): 941 (1987)
Index Fungorum number: IF25087; 1 morphological species.
Type species – Appendichordella amicta (Kohlm.) R.G. Johnson, E.B.G. Jones & S.T. Moss
Arenariomyces Höhnk, Veröff. Inst. Meeresf. Bremerhaven 3: 28 (1954)
Index Fungorum number: IF303; 5 morphological species (Species Fungorum 2020), 4
species with sequence data.
Type species – Arenariomyces trifurcatus Höhnk
Bathyascus Kohlm., Revue Mycol. 41(2): 190 (1977)
Index Fungorum number: IF519; 5 morphological species (Species Fungorum 2020).
Type species – Bathyascus vermisporus Kohlm.
Carbosphaerella I. Schmidt, Feddes Repert. 80(2-3): 108 (1969)
Index Fungorum number: IF820; 2 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Carbosphaerella pleosporoides I. Schmidt
Ceriosporopsis Linder, Farlowia 1: 408 (1944)
Index Fungorum number: IF915; 9 morphological species (Species Fungorum 2020), 2
species with sequence data.
Type species – Ceriosporopsis halima Linder
Chadefaudia Feldm.-Maz., Revue Générale de Botanique 64: 150 (1957)
Index Fungorum number: IF927; 6 morphological species (Species Fungorum 2020).
Type species – Chadefaudia marina Feldm.-Maz.
Corallicola Volkm.-Kohlm. & Kohlm., Mycotaxon 44(2): 418 (1992)
Index Fungorum number: IF25426; 1 morphological species.
Type species – Corallicola nana Volkm.-Kohlm. & Kohlm.
Corollospora Werderm., Notizbl. Bot. Gart. Berlin-Dahlem: 248 (1922)
Index Fungorum number: IF1251; 27 morphological species (Species Fungorum 2020), 16
species with sequence data.
Type species – Corollospora maritima Werderm.
Notes – Corollospora cinnamomea was introduced by Koch (1986) and differs from other
species of the genus by brown rugose 1-septate ascospores and specialized pseudoparenchyma in
the ostiolar canal. The peridium is thin and consists only of two (rarely three) cell layers in contrast
to the other Corollospora spp. that have three or more layers (Nakagiri & Tubaki 1986). Sequence
data available for this species and its phylogenetic placement within Halosphaeriaceae is confirmed
(Jones et al. 2017).
Cucullosporella K.D. Hyde & E.B.G. Jones, Mycotaxon 37: 200 (1990)
Index Fungorum number: IF25506; 1 species with sequence data.
Type species – Cucullosporella mangrovei (K.D. Hyde & E.B.G. Jones) K.D. Hyde & E.B.G.
Jones
615
Figure 114 – Corollospora cinnamomea (Material examined – THAILAND, Phuket, Patong beach,
on driftwood, 30 Oct. 1983, J. Koch., CP1012566, isotype). a Herbarium material. b, c Ascomata
on sand grain. d Outer apperence of ascomata. e Squash mount of ascomta (microslide). f Peridium.
g-i Asci. j, k Ascospores. Scale bars: b, c = 200 µm, d = 100 µm, e, g-i = 50 µm, f, j, k = 20 µm.
616
Figure 115 – Corollospora marina (redrawn from Haythorn et al. 1980). a Conidiophore with a
developing conidium and scars left by the detached conidia. b, c Single conidia produced from a
simple conidiophore. Scale bar: 50 µm.
Ebullia K.L. Pang, Mycoscience 56: 40 (2015)
Index Fungorum number: IF803444; 1 species with sequence data.
Type species – Ebullia octonae (Kohlm.) K.L. Pang
Gesasha Abdel-Wahab & Nagahama, Nova Hedwigia 92(3-4): 501 (2011)
Index Fungorum number: IF518670; 3 species with sequence data.
Type species – Gesasha mangrovei Abdel-Wahab & Nagah.
Haiyanga K.L. Pang & E.B.G. Jones, Raffles Bull. Zool., Suppl. 19: 8 (2008)
Index Fungorum number: IF535842; 1 species with sequence data.
Type species – Haiyanga salina (Meyers) K.L. Pang & E.B.G. Jones
Haligena Kohlm., Nova Hedwigia 3: 87 (1961)
Index Fungorum number: IF2204; 1 species with sequence data.
Type species – Haligena elaterophora Kohlm.
Halosarpheia sensu stricto Kohlm. & E. Kohlm., Trans. Br. Mycol. Soc. 68(2): 208 (1977)
Index Fungorum number: IF2208; 8 morphological species (Species Fungorum 2020), 4
species with sequence data.
Type species – Halosarpheia fibrosa Kohlm. & E. Kohlm.
Halosphaeria Linder, Farlowia 1: 412 (1944)
Index Fungorum number: IF2209; 4 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Halosphaeria appendiculata Linder
617
Halosphaeriopsis T.W. Johnson, J. Elisha Mitchell Scient. Soc. 74: 44 (1958)
Index Fungorum number: IF2210; 1 species with sequence data.
Type species – Halosphaeriopsis mediosetigera (Cribb & J.W. Cribb) T.W. Johnson
Havispora K.L. Pang & Vrijmoed, Mycologia 100(2): 293 (2008)
Index Fungorum number: IF506744; 1 species with sequence data.
Type species – Havispora longyearbyenensis K.L. Pang & Vrijmoed
Iwilsoniella E.B.G. Jones, Syst. Ascomyc. 10(1): 8 (1991)
Index Fungorum number: IF25574; 1 morphological species.
Type species – Iwilsoniella rotunda E.B.G. Jones
Kitesporella Jheng & K.L. Pang, Bot. Mar. 55: 462 (2012)
Index Fungorum number: IF563899; 1 morphological species.
Type species – Kitesporella keelungensis J.S. Jheng & K.L. Pang
Kochiella Sakay., K.L. Pang & E.B.G. Jones, Fungal Divers. 46: 96 (2011)
Index Fungorum number: IF518769; 1 species with sequence data.
Type species – Kochiella crispa (Kohlm.) Sakay., K.L. Pang & E.B.G. Jones
Lautisporopsis E.B.G. Jones, Yusoff & S.T. Moss, Mycotaxon 67: 1 (1998)
Index Fungorum number: IF27443; 1 morphological species.
Type species – Lautosporopsis circumvestita (Kohlm.) E.B.G. Jones, Yusoff & S.T. Moss
Lignincola Höhnk, Veröff. Inst. Meeresf. Bremerhaven 3: 216 (1955)
Index Fungorum number: IF2864; 4 morphological species (Species Fungorum 2020), 2
species with sequence data.
Type species – Lignincola laevis Höhnk
Limacospora Jørg. Koch & E.B.G. Jones, Can. J. Bot. 73(7): 1011 (1995)
Index Fungorum number: IF27610; 1 morphological species.
Type species – Limacospora sundica (Jørg. Koch & E.B.G. Jones) Jørg. Koch & E.B.G.
Jones
Luttrellia Shearer, Mycologia 70(3): 692 (1978)
Index Fungorum number: IF2954; 4 morphological species (Species Fungorum 2020).
Type species – Luttrellia estuarina Shearer
Magnisphaera J. Campbell, J.L. Anderson & Shearer, Mycologia 95(3): 546 (2003)
Index Fungorum number: IF28707; 2 species with sequence data.
Type species – Magnisphaera spartinae (E.B.G. Jones) J. Campb., J.L. Anderson & Shearer
Marinospora A.R. Caval., Nova Hedwigia 11: 548 (1966)
Index Fungorum number: IF3002; 2 species with sequence data.
Type species – Marinospora calyptrata (Kohlm.) A.R. Caval.
Moana Kohlm. & Volkm.-Kohlm., Mycol. Res. 92 (4): 418 (1989)
Index Fungorum number: IF25325; 1 morphological species.
Type species – Moana turbinulata Kohlm. & Volkm.-Kohlm.
Morakotiella Sakay., Mycologia 97(4): 806 (2005)
Index Fungorum number: IF28969; 1 species with sequence data.
618
Type species – Morakotiella salina (C.A. Farrant & E.B.G. Jones) Sakay.
Naïs Kohlm., Nova Hedwigia 4: 409 (1962)
Index Fungorum number: IF92064; 2 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Naïs inornata Kohlm.
Natantispora J. Campbell, J.L. Anderson & Shearer, Mycologia 95(3): 543 (2003)
Index Fungorum number: IF28703; 3 species with sequence data.
Type species – Natantispora lotica (Shearer) J. Campb., J.L. Anderson & Shearer
Nautosphaeria E.B.G. Jones, Trans. Br. Mycol. Soc. 47(1): 97 (1964)
Index Fungorum number: IF3428; 1 species with sequence data.
Type species – Nautosphaeria cristaminuta E.B.G. Jones
Neptunella K.L. Pang & E.B.G. Jones, Mycol. Progr. 2(1): 35 (2003)
Index Fungorum number: IF28749; 1 species with sequence data.
Type species – Neptunella longirostris (Cribb & J.W. Cribb) K.L. Pang & E.B.G. Jones
Nereiospora E.B.G. Jones, R.G. Johnson & S.T. Moss, J. Linn. Soc. Bot. 87(2): 204 (1983)
Index Fungorum number: IF25825; 2 species with sequence data.
Type species – Nereiospora comata (Kohlm.) E.B.G. Jones, R.G. Johnson & S.T. Moss
Nimbospora J. Koch, Nordic J. Bot. 2(2): 166 (1982)
Index Fungorum number: IF3509; 2 species with sequence data.
Type species – Nimbospora bipolaris K.D. Hyde & E.B.G. Jones
Nohea Kohlm. & Volkm.-Kohlm., Syst. Ascomyc. 10: 121 (1991)
Index Fungorum number: IF25883; 3 morphological species (Species Fungorum 2020), 2
species with sequence data.
Type species – Nohea umiumi Kohlm. & Volkm.-Kohlm.
Oceanitis Kohlm., Revue Mycol. 41(2): 193 (1977)
Index Fungorum number: IF532552; 4 species with sequence data.
Type species – Oceanitis scuticella Kohlm.
Ocostaspora E.B.G. Jones, R.G. Johnson & S.T Moss, Bot. Mar. 26: 353 (1983)
Index Fungorum number: IF25829; 1 species with sequence data.
Type species – Ocostaspora apilongissima E.B.G. Jones, R.G. Johnson & S.T. Moss
Okeanomyces K.L. Pang & E.B.G. Jones, J. Linn. Soc. Bot. 146(2): 228 (2004)
Index Fungorum number: IF28838; 1 species with sequence data.
Type species – Okeanomyces cucullatus (Kohlm.) K.L. Pang & E.B.G. Jones
Ondiniella E.B.G. Jones, R.G. Johnson & S.T. Moss, Bot. Mar. 27: 136 (1984)
Index Fungorum number: IF25830; 1 species with sequence data.
Type species – Ondiniella torquata (Kohlm.) E.B.G. Jones, R.G. Johnson & S.T. Moss
Ophiodeira Kohlm. & Volkm.-Kohlm., Can. J. Bot. 66 (10): 2062 (1988)
Index Fungorum number: IF25267; 1 species with sequence data.
Type species – Ophiodeira monosemeia Kohlm. & Volkm.-Kohlm.
619
Panorbis J. Campbell, J.L. Anderson & Shearer, Mycologia 95(3): 544 (2003)
Index Fungorum number: IF28704; 1 species with sequence data.
Type species – Panorbis viscosus (I. Schmidt) J. Campb., J.L. Anderson & Shearer
Paraaniptodera K.L. Pang, C.L. Lu, W.T. Ju & E.B.G. Jones, Bot. Mar. 60(4): 460 (2017)
Index Fungorum number: IF818212; 1 species with sequence data.
Type species – Paraaniptodera longispora (K.D. Hyde) K.L. Pang, C.L. Lu, W.T. Ju &
E.B.G. Jones
Phaeonectriella R.A. Eaton & E.B.G. Jones, Nova Hedwigia 19(3-4): 779 (1971)
Index Fungorum number: IF3928; 3 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Phaeonectriella lignicola R.A. Eaton & E.B.G. Jones
Pileomyces K.L. Pang & Jheng, Bot. Stud. 53: 536 (2012)
Index Fungorum number: IF519625; 1 species with sequence data.
Type species – Pileomyces formosanus K.L. Pang & J.S. Jheng
Praelongicaulis E.B.G. Jones, Abdel-Wahab, & K.L. Pang, Fungal Divers. 73(1): 54 (2015)
Index Fungorum number: IF812594; 1 species with sequence data.
Type species – Praelongicaulis kandeliae (Abdel-Wahab & E.B.G. Jones) E.B.G. Jones,
Abdel-Wahab, & K.L. Pang
Pseudolignincola Chatmala & E.B.G. Jones, Nova Hedwigia 83(1-2): 225 (2006)
Index Fungorum number: IF29053; 1 species with sequence data.
Type species – Pseudolignincola siamensis Chatmala & E.B.G. Jones
Remispora Linder, Farlowia 1(3): 409 (1944)
Index Fungorum number: IF4674; 6 morphological species (Species Fungorum 2020), 5
species with sequence data.
Type species – Remispora maritima Linder
Saagaromyces K.L. Pang & E.B.G. Jones, Mycol. Progr. 2(1): 35 (2003)
Index Fungorum number: IF28748; 4 species with sequence data.
Type species – Saagaromyces abonnis (Kohlm.) K.L. Pang & E.B.G. Jones
Sablicola E.B.G. Jones, K.L. Pang & Vrijmoed, Can. J. Bot. 82(4): 486 (2004)
Index Fungorum number: IF28801; 1 species with sequence data.
Type species – Sablicola chinensis E.B.G. Jones, K.L. Pang & Vrijmoed
Thalassogena Kohlm. & Volkm.-Kohlm., Syst. Ascomyc. 6: 223 (1987)
Index Fungorum number: IF25208; 1 morphological species.
Type species – Thalassogena sphaerica Kohlm. & Volkm.-Kohlm.
Thalespora Chatmala & E.B.G. Jones, Nova Hedwigia 83(1–2): 228 (2006)
Index Fungorum number: IF29054; 1 species with sequence data.
Type species – Thalespora appendiculata Chatmala & E.B.G. Jones
Tinhaudeus K.L. Pang, S.Y. Guo & E.B.G. Jones, Fungal Divers. 75: 160 (2015)
Index Fungorum number: IF812936; 1 species with sequence data.
Type species – Tinhaudeus formosanus K.L. Pang, S.Y. Guo & E.B.G. Jones
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Tirispora E.B.G. Jones & Vrijmoed, Can. J. Bot. 72(9): 1373 (1994)
Index Fungorum number: IF27405; 2 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Tirispora unicaudata E.B.G. Jones & Vrijmoed
Toriella Sakay., K.L. Pang & E.B.G. Jones, Fungal Divers. 46(1): 99 (2011)
Index Fungorum number: IF518774; 1 species with sequence data.
Type species – Toriella tubulifera (Kohlm.) Sakay., K.L. Pang & E.B.G. Jones
Trailia G.K. Sutherl., Trans. Br. Mycol. Soc. 5: 149 (1915)
Index Fungorum number: IF5516; 1 morphological species.
Type species – Trailia ascophylli G.K. Sutherl.
Trichomaris Hibbits, G.C. Hughes & Sparks, Can. J. Bot. 59(11): 2123 (1981)
Index Fungorum number: IF5559; 1 morphological species.
Type species – Trichomaris invadens Hibbits, G.C. Hughes & Sparks
Tubakiella Sakay., K.L. Pang & E.B.G. Jones, Fungal Divers. 46: 97 (2011)
Index Fungorum number: IF518772; 1 species with sequence data.
Type species – Tubakiella galerita (Tubaki) Sakay., K.L. Pang & E.B.G. Jones
Tunicatispora K.D. Hyde, Aust. Syst. Bot. 3: 712 (1990)
Index Fungorum number: IF25558; 1 morphological species.
Type species – Tunicatispora australiensis K.D. Hyde
Hansfordiaceae Crous, Fungal Systematics and Evolution 3: 84 (2019)
Index Fungorum number: IF829455; Facesoffungi number: FoF06206; 13 species.
Saprobic on leaf and stem tissues bearing ascomata. Sexual morph: Undetermined. Asexual
morph: Mycelium superficial to immersed on stems or twigs. Conidiophores superficial or setiform,
solitary, erect, multi-septate with lateral branches, surface smooth, straight to flexuous, light brown.
Conidiogenous cells hyaline, cylindrical or clavate, with subdenticulate apical loci. Conidia
solitary, dry, aseptate, hyaline to pale brown, globose, ellipsoid to fusoid, smooth-walled or finely
roughened (adapted from Crous et al. 2019b).
Type genus – Hansfordia S. Hughes.
Notes – Hansfordiaceae was introduced by Crous et al. (2019b) and belongs in Xylariales.
The family comprises a single genus, Hansfordia. Hansfordiaceae is closely related to the families
Sporocadaceae, Phlogicylindriaceae and Beltraniaceae.
Ecological and economic significance of Hansfordiaceae
Mitchell & Taber (1986) and Alderman (2010) suggested that Hansfordia pulvinata as a
possible biological control agent against plant pathogenic fungi. Alderman (2010) accepted Dicyma
pulvinata (= Hansfordia pulvinata), as a potential biocontrol agent of Cereosporidim personatum in
peanut.
Genus included in Hansfordiaceae
Hansfordia S. Hughes, Mycol. Pap. 43: 15 (1951)
Index Fungorum number: IF8423; 13 morphological species (Species Fungorum 2020), 2
species with sequence data.
Type species – Hansfordia ovalispora S. Hughes
Notes – Hansfordia was introduced by Hughes (1951a) with 12 species based on
morphology. Crous et al. (2019b) introduced H. pruni and H. pulvinata to this genus based on
morphology and LSU and ITS sequence data. The main characters to differentiate species of
621
Hansfordia is the denticulate nature of cylindrical conidiogenous cells with regular verticils
inserted directly on the stipe (Hughes 1951a).
Figure 116 – Hansfordia ovalispora (redrawn asexual morph of H. ovalispora from Hughes
1951a). a, b, d Conidiophores with conidia. c Single conidiophore immature conidia. e-g Lateral
branches. Scale bars: b, c = 10 µm.
Harknessiaceae Crous, Persoonia, Mol.Phyl. Evol. Fungi 28: 55 (2012)
Index Fungorum number: IF564740; Facesoffungi number: FoF01387; 69 species.
Saprobic or pathogenic on leaves or twig litter. Sexual morph: Ascomata perithecial, solitary
or aggregated, immersed, globose, coriaceous, brown, ostiolate, papillate. Papilla emergent to
depressed, comprising 3–5 layers of brown-walled cells of textura angularis. Paraphyses septate,
branched. Asci 8-spored, unitunicate, cylindrical to clavate, short pedicellate, with J-, apical ring.
Ascospores uniseriate to biseriate, hyaline, ellipsoid to fusoid, aseptate, thick-walled, guttulate,
smooth-walled. Asexual morph: Coelomycetous. Conidiomata eustromatic, pycnidial, scattered or
aggregated, immersed, globose, coriaceous, with single or several locules, dark brown to black,
ostiolate. Peridium comprising thin-walled, almost hyaline to brown cells of textura angularis.
Ostiole wide, centrally located, surrounded by brown cells. Conidiophores lining the inner cavity,
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or reduced to basal layer; sometimes septate, branched; sometimes reduced to conidiogenous cells.
Conidiogenous cells holoblastic, integrated, lageniform, subcylindrical to cylindrical, hyaline to
pale yellow, smooth, producing macroconidia and sometimes microconidia from same
conidiogenous cell; sometimes proliferating sympodially one or several times. Macroconidia with a
basal appendage, hyaline when young, brown at maturity, unicellular, although basal appendage
separated by a thick, smooth-walled septum, with or without light and dark longitudinal bands,
sometimes longitudinally striate, guttulate; basal appendage cellular, cylindrical to subcylindrical,
hyaline, thin-walled, devoid of contents; apical appendage present or absent, if present elongate.
Microconidia hyaline, oval to ellipsoid, aseptate, smooth-walled (adapted from Senanayake et al.
2018).
Type genus – Harknessia Cooke
Notes – Harknessiaceae was initially proposed by Castlebury et al. (2002), subsequently
formally introduced by Crous et al. (2012d) to accommodate Harknessia with their wuestneia-like
sexual morphs. Species of Harknessiaceae have pycnidial conidiomata with brown, furfuraceous
margins, brown conidia with hyaline, tube-like basal appendages, longitudinal striations, and
rhexolytic secession (Crous et al. 2012d, Senanayake et al. 2018). Harknessiaceae species seem to
have a cosmopolitan distribution, since they have been recorded worldwide, and are commonly
associated with leaf spots and branches of various hosts (Sankaran et al. 1995, Farr & Rossman
2019). They have also been recoded as endophytes and saprobes in leaves and twigs of various
angiosperm plants and noxious weeds (Crous et al. 2012d).
LSU analysis support this family which was earlier placed in Diaporthales (Crous et al.
2012d). Crous et al. (2012d) introduced six novel species of Harknessia on Eucalyptus and
supported by a multi-gene analysis (ITS, calM and tub2) for these species.
Ecological and economic significance of Harknessiaceae
Most members in the family are associated with leaf spots, leaf tip-dieback or leaf scorch and
stem cankers and they are suspected to be pathogens mainly in Eucalyptus (Crous et al. 1989,
Marincowitz et al. 2008). However, many species have been isolated from asymptomatic plant
tissues and are assumed to be saprobes. Therefore, their pathogenicity is still ambiguous. Normally
they are species with less economical significance (Park et al. 2000).
Genera included in Harknessiaceae
Harknessia Cooke, Grevillea 9 (no.51): 85 (1881)
Index Fungorum number: IF8449; 68 morphological species (Species Fungorum 2020), 38
species with sequence data.
Type species – Harknessia eucalypti Cooke
Notes – Harknessia species have been recorded in wide variety of habitats and normally they
appear as leaf spots, wherein leaves with tip-dieback or leaf scorch and stem cankers occur (Crous
et al. 1989, Wijayawardene et al. 2016b). Some species have been recorded from leaf and twig litter
(Marincowitz et al. 2008). However, the pathogenicity of Harknessia species on host plants is not
proven yet, since nobody has resolved it experimentally. Harknessia species with hyaline conidia
and apical appendages were transferred to Mastigosporella by Höhnel (1917c), and species with
brown conidia with apical and basal appendages were placed to Apoharknessia (Lee et al. 2004),
whereas species with very thick conidial walls and longitudinal slits were included in Dwiroopa
(Farr & Rossman 2003). Harknessia has enough molecular support to accommodate in a distinct
lineage in Harknessiaceae. Currently, there are 74 Harknessia epithets in Index Fungorum (2020).
Yuan & Mohammed (1997) introduced Wuestneia epispora which was identical to the
asexual morph of Harkenessia eucalypti. According to the cultural characters conidia of W.
epispora are shorter (11–22 × 11–15 µm) than those of H. eucalypti (Crous et al. 1993). However,
the average of conidial size (25.4 × 13.9 µm) by Yuan & Mohammed (1997) is closely similar with
the averages of H. eucalypti (23 × 14 µm) (Nag Raj 1993). Therefore, Lee et al. (2004) treated this
culture as authentic to H.eucalypti.
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Figure 117 – Harknessia eucalypti (Material examined – USA, California, on leaves of Eucalyptus
globulus. (Myrtaceae), Harknessia 1280, isotype K (M) 195744). a Herbarium specimen. b
Conidiomata on host substrate. c Cross section of conidioma. d Peridium. e, f Conidia attached to
conidiophores. g-k conidia are attached to conidiogenous cells. l Conidium with basal appendage.
Scale bars: c = 100 μm, d-f = 20 μm, g-l = 10 μm.
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Mebarria J. Reid & C. Booth, Can. J. Bot. 67(3): 898 (1989)
Index Fungorum number: IF25321; 1 morphological species.
Type species – Mebarria thujina (Nag Raj & DiCosmo) J. Reid & C. Booth
Notes – Nag Raj & DiCosmo (1981) introduced Cryptosporella thujina and Reid & Booth
(1989) synonymized as Mebarria thujina accommodating it in a new genus Mebarria. The asexual
morph of this species was recorded as Harknessia thujina (Index Fungorum 2020). However,
molecular data needs to confirm sexual and asexual connection of these two species.
Helminthosphaeriaceae Samuels, Cand. & Magni, Mycologia 89(1): 144 (1997)
Index Fungorum number: IF81913; Facesoffungi number: FoF01142; 140 species.
Saprobic on wood or decorticated branch in terrestrial and aquatic habitats, some
fungicolous. Sexual morph: Ascomata perithecial, gregarious or scattered, dark brown to black,
solitary, superficial or immersed, ovoid, globose to subglobose, rough, tuberculate, smooth or with
setae, papillate or ostiole indistinct, the apex collapsing when dry. Ostioles periphysate. Peridium
composed of two layers, outer layer comprising brown cells of textura angularis or prismatica,
carbonaceous or membranaceous; inner layer comprising hyaline cells of textura prismatica, thin,
membranaceous. Paraphyses numerous, septate, persistent or deliquescing, swollen, filiform or
cylindrical. Asci 8-spored, unitunicate, thin or thick-walled, cylindrical to clavate, pedicellate, apex
truncate, J-, apical ring refractive or indistinct. Ascospores 2-seriate, hyaline or brown or becoming
dark colored in part, allantoid, clavate, cylindrical to ellipsoid, 0–3-septate, smooth-walled, with or
without guttules or pores. Asexual morph: Hyphomycetous. Conidiophores macronematous,
mononematous, scattered or gregarious, brown, straight, septate, unbranched, smooth-walled.
Conidiogenous cells mono- or polyblastic, terminal, intercalary, integrated, percurrently elongating;
conidiogenesis tretic. Conidia solitary, acrogenous, brown, aseptate or septate, obclavate, globose
to fusiform to cylindrical, straight, subtruncate to obtuse at the apex, truncate or swollen at the base,
sometimes with guttules.
Type genus – Helminthosphaeria Fuckel
Notes – Helminthosphaeriaceae was introduced by Samuels et al. (1997) for taxa with black,
setose ascomata and cylindrical asci containing brown ascospores and included the only genus
Helminthosphaeria (Fuckel 1870). Tengiomyces was added to the family by Réblová (1999a) based
on similar characters to Helminthosphaeria, viz. setose ascomata. Echinosphaeria, Hilberina,
Ruzenia and Synaptospora, many of which are Lasiosphaeria segregates, were included based on
analysis of LSU sequence data and characters (Miller & Huhndorf 2004, Miller et al. 2014).
Endophragmiella was accepted as the asexual morph of Echinosphaeria (Hughes 1979).
Helminthosphaeriaceae forms a well-supported clade in Chaetosphaeriales, with
Chaetosphaeriaceae as a sister clade (Maharachchikumbura et al. 2015). Currently seven genera are
accepted in the family based on morphological and phylogenic analyses of LSU, SSU, tef1 and
rpb2 sequence data (Fig. 8). Although we include Tengiomyces here, it lacks molecular data, which
is needed to confirm its placement.
Ecological and economic significance of Helminthosphaeriaceae
Members of Helminthosphaeriaceae are usually saprobic on wood or/and other fungi in
terrestrial, marine or freshwater habitats and are distributed worldwide, but more taxa are reported
from Europe, such as Echinosphaeria canescens (on driftwood of half buried in sand),
Helminthosphaeria clavariarum (on Clavulina sp.) (Hughes 1979, Réblová 1999a, Miller &
Huhndorf 2004, Miller et al. 2014, Hernández-Restrepo et al. 2017).
Genera included in Helminthosphaeriaceae
Echinosphaeria A.N. Mill. & Huhndorf, Mycol. Res. 108(1): 29 (2004)
Index Fungorum number: IF28829; 14 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Echinosphaeria canescens (Pers.) A.N. Mill. & Huhndorf
625
Notes – Echinosphaeria, typified by Echinosphaeria canescens, has setose ascomata and
cylindrical asci with reniform ascospores. This genus is closely related to Helminthosphaeria and
Synaptospora within Helminthosphaeriaceae (Miller & Huhndorf 2004) (Fig. 8).
Endophragmiella B. Sutton, Mycol. Pap. 132: 58 (1973)
Index Fungorum number: IF8167; 84 morphological species (Species Fungorum 2020), 2
species with sequence data.
Type species – Endophragmiella pallescens B. Sutton
Notes – Endophragmiella pallescens has branched, septate, flexuous conidiophores and
ellipsoid, septate conidia (Sutton 1973). This genus was accepted in Helminthosphaeriaceae and is
closely related to Helminthosphaeria (Hernández-Restrepo et al. 2017).
Helminthosphaeria Fuckel, Jb. nassau. Ver. Naturk. 23-24: 166 (1870)
Index Fungorum number: IF2263; 18 morphological species (Species Fungorum 2020), 9
species with sequence data.
Type species – Helminthosphaeria clavariarum (Desm.) Fuckel
Notes – Helminthosphaeria was included in Melanosporaceae, in Sordariales based on its 1septate ascospores with apical pores (Parguey-Leduc 1960). Lundqvist (1972) excluded
Helminthosphaeria from Melanosporaceae due to its distinct filamentous pseudoparaphyses and
asci lacking sheaths. Barr (1990b) reevaluated Sordariaceae and included Helminthosphaeria.
Eriksson & Hawksworth (1993) placed Helminthosphaeria in Sordariales. Samuels et al. (1997)
monographed the genus, including the asexual morph Diplococcium, and introduced
Helminthosphaeriaceae (Goh & Hyde 1998, Réblová 1999a, Hernández-Restrepo et al. 2017).
Figure 118 – Helminthosphaeria pilifera (diplococcium-like, redrawn from Réblová 1999a).
Conidiophore with conidia of Diplococcium asexual morph. Scale bar = 20 µm.
Hilberina Huhndorf & A.N. Mill., Mycol. Res. 108(1): 31 (2004)
Index Fungorum number: IF28830; 17 morphological species (Species Fungorum 2020), 5
species with sequence data.
Type species – Hilberina caudata (Fuckel) Huhndorf & A.N. Mill.
626
Notes – Hilberina caudata has setose ascomata, cylindrical asci with rings and geniculate
ascospores with one end tapering to a distinct point (Miller & Huhndorf 2004). Sequence data of
five members in Hilberina were provided by Miller et al. (2014) and reveals that Hilberina was
closely related to Synaptospora, Ruzenia and Helminthosphaeria within Helminthosphaeriaceae
(Fig. 8).
Figure 119 – Ruzenia spermoides (Material examined – SWEDEN, Uppland, Bladaker; on dead
wood of Tilia cordata; 20 September 1992, S-F 242341). a Material label. b Material. c Ascoma on
host. d Ascoma cross section. e Peridium. f Asci. g-l Ascospores. Notes: figs f-l stained in KOH
reagent. Scale bars: d = 200 µm, e-f = 50 µm, g-l = 10 µm.
Ruzenia O. Hilber, The Genus Lasiosphaeria and Allied Taxa (Kelheim): 7 (2002)
Index Fungorum number: IF530833; 1 species with sequence data.
Type species – Ruzenia spermoides (Hoffm.) O. Hilber
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Notes – Ruzenia spermoides has glabrous ascomata, cylindrical asci and allantoid ascospores
with guttules (Miller & Huhndorf 2004). It is closely related to Synaptospora based on multi-gene
analysis (Miller et al. 2014) (Fig. 8).
Synaptospora Cain, Beih. Sydowia 1: 4 (1957)
Index Fungorum number: IF5328; 5 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Synaptospora petrakii Cain
Notes – Synaptospora petrakii has setose ascomata, cylindrical asci and cylindrical
ascospores breaking ascoconidia (Cain 1957, Huhndorf et al. 1999). It is closely related to Ruzenia
within Helminthosphaeriaceae (Miller et al. 2014, Fig. 8).
Tengiomyces Réblová, Mycotaxon 70: 408 (1999)
Index Fungorum number: IF28271; 1 morphological species.
Type species – Tengiomyces indicus (Varghese & V.G. Rao) Réblová
Notes – Tengiomyces has setose ascomata, cylindrical asci and versicolored ascospores with
2–3 septa. This genus is similar to Helminthosphaeria based on setose ascomata and versicoloured
ascospores and it is associated with spadicoides-like asexual morphs (Réblová 1999a).
Hispidicarpomycetaceae Nakagiri, Mycologia 85(4): 649 (1993)
Index Fungorum number: IF81958; Facesoffungi number: FoF01098; 1 species.
Parasitic on a marine alga. Sexual morph: Ascomata superficial, solitary to gregarious,
ostiolate, apapillate. Peridium composed of three-layers: outer layer with thick-walled, short hyphal
projections, brown to dark brown, middle layer of light brown to dark brown, thick-walled hyphal,
flattened cells of textura epidermoidea, and an inner layer of pale brown, thin-walled, flattened
cells of textura epidermoidea, lining the whole ascomatal venter. Paraphyses septate branched.
Asci 8-spored, unitunicate, thin-walled, clavate to pyriform, deliquescing early and lacking an
apical ring. Ascospores hyaline, ellipsoid to elliptic-fusiform, unicellular, lacking appendages or a
sheath, overlapping. Reproductive state of brown hyphae, Spermodochia composed of a mass
spermatiosphores. Spermatiosphores, verticilliate, asymmetrical penicilliate-like, brown at the base,
paler and hyaline towards the apex. Stipes septate, branched, brown at the base. Spermatia
cylindrical, unicellular, hyaline. Trichogynes septate, cylindrical, brown.
Type genus – Hispidicarpomyces Nakagiri
Notes – Hispidicarpomycetaceae is a monotypic family, which was introduced to include a
marine ascomycete, Hispidicarpomyces galaxauricola (Nakagiri 1993). Previously,
Hispidicarpomycetaceae was referred to Spathulosporales by Nakagiri (1993). However,
phylogenetic studies by Inderbitzin et al. (2004) placed Spathulospora (S. adelpha, S. antartica) in
Lulworthiales. Consequently, the affinity of Spathulospora within the Lulworthiales raises the
question as to the phylogenetic position of Hispidicarpomycetaceae. Maharachchikumbura et al.
(2015) accepted the taxonomic placement of Hispidicarpomycetaceae within Spathulosporales.
Cultures and sequences are unavailable for this family and consequently a generic revision is
needed for the family.
Ecological and economic significance of Hispidicarpomycetaceae
Biotrophic on the marine alga Galaxaura falcata (Rhodophyta), which was found along the
Japanese coast (Nakagiri 1993). Infection of the host alga by H. galaxauricola probably occurs in
June to July when its ascospores are abundantly released (Nakagiri 1993). Spermodochia and
ascocarps are produced from the base up to around the first to third branches of the algal frond. It is
believed that the absorption of photosynthetic nutrients from the epithelial cells can be occurring
due to dense distribution of hyphae beneath the epithelial layer. However, there was no evidence of
hyphal invasion into a host cell, though close contact of the hyphae with algal cells was often
observed (Nakagiri 1993).
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Genus included in Hispidicarpomycetaceae
Hispidicarpomyces Nakagiri, Mycologia 85(4): 639 (1993)
Index Fungorum number: IF26466; 1 morphological species.
Type species – Hispidicarpomyces galaxauricola Nakagiri
Notes – Hispidicarpomyces was introduced by Nakagiri (1993) is a monotypic genus which
shares similar morphological features with Spathulospora. There is no sequence data available for
this genus, therefore, further taxon sampling is essential to determine its phylogenetic position
within the Ascomycota (Jones et al. 2015, 2019).
Hispidicarpomyces galaxauricola is similar to members of Spathulospora in morphology and
parasitism on marine red algae (Kohlmeyer 1973, Kohlmeyer & Kohlmeyer 1979). However, they
are different in that H. galaxauricola has true hyphae growing throughout the algal frond and, in
contrast, those of Spathulospora spp. are reduced-hyphoid (Kohlmeyer 1973). Further, intracellular
invasion into the algal cell by an infection peg and formation of intracellular crusts can be seen in
Spathulospora species, but H. galaxauricola showed only intercellular infection (Nakagiri 1993).
Hypocreaceae De Not., G. bot. ital. 2(1): 48 (1844)
Index Fungorum number: IF80892; Facesoffungi number: FoF01904; 606 species.
Biotrophic, hemibiotrophic, saprobic or hypersaprobic on plants, other fungi, myxomycetes
and lichens, some are mycoparasites, rarely coprophillus, occuring in terrestrial and aquatic habitats
worldwide. Sexual morph: Stromatic tissue well-developed, or weakly developed and existing only
as subiculum of interwoven pallid or brightly pigmented hyphae, when present soft and fleshy,
pallid or brightly pigmented to light brownish, immersed erumpent, effuse, tuberculate or pulvinate,
occasionally stipitate, then fertile region clavate. Ascomata solitary or often arranged in groups,
perithecial or rarely cleistothecial, immersed in or erumpent to superficial on substrate, pallid,
brightly pigmented, or shades of light brown or blue to violet (appearing black), rarely brown,
globose, ovoid, obpyriform or spheroid, collabent at times, apex papillate, with periphysate ostiole,
surface glabrous or warted or bearing hyaline or pallid hyphal appendages or rarely thick-walled
setae. Peridium externally composed of pseudoparenchymatous cells, sometimes with thick,
sclerotial walls, internally composed of compressed rows of cells, pallid to brightly pigmented or
brown, blue or violet. Paraphyses apical (periphysoids), usually deliquescent, occasionally visible
at maturity, as cellular remnants among asci or as remnants of apical fringe. Asci basal to
peripheral, mostly 8-spored, occasionally polysporous or less than eight, cylindric, oblong or
inflated, apical ring often lacking, when present shallow, refractive, J-. Ascospores uniseriate,
biseriate or in fascicle, hyaline, yellowish, pinkish to greenish or occasionally brown, 1-celled or
one to several septate, occasionally with longitudinal septa, disarticulating into part-spores at times
or budding to form conidia within ascus, ellipsoid, fusoid, allantoid, elongate or globose, with
smooth, verruculose or longitudinally striate cell wall. Asexual morph: Hyphomycetous or
coelomycetes. Primirily phialidic. Conidiomata non-exsistent to sporodochial or synnematal,
conidiogenous cells enteroblastic phialidic, thick-walled structures present at times. Conidia
hyaline to bright colored, less commonly produce aleurioconidia and ampulloconidia.
Type genus – Trichoderma Pers.
Notes – Hypocreaceae is characterized by often disarticulating ascospores and perithecia that
are mostly immersed in a stroma or seated on a subiculum. Hypocreaceae was recognized within
Hypocreales and divided into six subfamilies by Lindau (1897). Rogerson (1970) recognized
Hypocreales as the only family in Hypocreales. However, Kreisel (1969) accepted Hypocreaceae
and Nectriaceae as two families in Hypocreales. Rossman et al. (1999) defined Hypocreaceae in a
more restricted sense and recognized Nectriaceae, Hypocreaceae and Bionecriaceae as three
separate families within the Hypocreales. Molecular analyses of Rossman et al. (2001) based on
LSU data supported this family. Members of Hypocreaceae are characterized by their brightly
coloured and fleshy perithecial stromata (Rogerson 1970).
Ecological and economic significance of Hypoceaeceae
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Trichoderma species play an important ecological role as producers of bioactive compounds,
biocontrol agents of plant diseases and as pathogens of animals and mushrooms (Schuster &
Schmoll 2010). For example, Trichoderma harzianum, and T. viride are widely used as
biofungicides for the biocontrol of plant pathogens (Wraight et al. 1998, Harman et al. 2004,
Kaewchai et al. 2009, Thiruvudainambi et al. 2010, Stocco et al. 2016, Hyde et al. 2019b). They
have been also utilized for production of antibiotics (Harman et al. 2006). Trichoderma polysporum
produces antifungal Cyclosporine A (CsA), a calcineurin inhibitor, used as an immunosuppressant
(Dreyfuss et al. 1976). Root colonization by Trichoderma species enhances root growth and
development, crop productivity, resistance to abiotic stresses and the uptake and use of nutrients,
hence it is becoming widely used in horticulture (Harman 2000, Harman et al. 2004). Fungicolous
specis of Hypoceaeceae, such as species of Mycogone and Hypomyces are mushroom pathogens
(Rossman et al. 1999). Mycogone perniciosa, Hypomyces rosellus and H. aurantius are the causal
agents of dry bubble, wet bubble and cobweb disease of the button mushroom, Agaricus bisporus
(Baars et al. 2013, Tanovic et al. 2006).
Genera included in Hypoceaeceae
Arachnocrea Z. Moravec, Bull. trimest. Soc. mycol. Fr.72 : 161 (1956)
Index Fungorum number: IF292; 3 morphological species (Species Fungorum 2020), 2
species with sequence data.
Type species – Arachnocrea stipata (Lib.) Z. Moravec
Notes – Arachnocrea species occur on decaying wood, herbaceous substrates or polypores
(fungicolous) (Moravec 1956, Rossman et al. 1999). The genus is characterized by white to yellow
orange, papillate, perithecial ascomata, cylindrical asci with hylaine, fusiform, 1-septate ascospores
disarticulating into monomorphic part ascospores and verticillium-like asexual morphs (Põldmaa
1999, Rossman et al. 1999, Rossman 2000).
Dialhypocrea Speg., Boln Acad. nac. Cienc. Córdoba 23(3−4): 475 (1919)
Index Fungorum number: IF1490; 1 species with sequence data.
Type species – Dialhypocrea puiggariana Speg.
Notes – Dialhypocrea puiggariana is associated with decaying branches (Rossman et al.
1999). The genus is characterized by stromatic, perithecial ascomata, cylindrical asci and hyaline,
1-septate ascospores disarticulating into part ascospores (Rossman et al. 1999). No asexual morph
is linked to this genus.
Escovopsioides H.C. Evans & J.O. Augustin, PLoS ONE 7(12): e51392, 6 (2012)
Index Fungorum number: IF800474; 1 species with sequence data.
Type species – Escovopsioides nivea H.C. Evans & J.O. Augustin
Notes – Augustin et al. (2013) introduced the monotypic genus Escovopsioides based on
mycoparasitic species E. nivea isolated from of leaf-cutting ant Acromyrmex subterraneus
subterraneus. The genus is characterized by conidiophores with terminal and intercalary, globose
vesicles and phialides produced on vesicles and, hyaline conidia occuring in long chains (Augustin
et al. 2013). No sexual morph is linked to this genus.
Escovopsis J.J. Muchovej & Della Lucia, Mycotaxon 37: 192 (1990)
Index Fungorum number: IF11249; 14 species with sequence data.
Type species – Escovopsis weberi J.J. Muchovej & Della Lucia
Notes – Escovopsis, typified by E. weberi was introduced by Muchovej & Della Lucia (1990)
to replace Phialocladus (1972). Taxa are mycoparasites found only from the fungus gardens of
attine ants (Reynolds & Currie 2004, Augustin et al. 2013, Montoya et al. 2019). The genus is
characterized by conidiophores with or without terminal variously-shaped vesicles and synchronous
or sympodial conidiogenesis with pigmented conidia occuring in chains (Muchovej & Della Lucia
1990, Montoya et al. 2019). No sexual morph is linked to this genus.
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Figure 120 – Hispidicarpomyces galaxauricola (Material examined – JAPAN, O-ura, Shimoda, 2
June 1990, IFO H-12135, holotype). a Herbarium material of Hispidicarpomyces galaxauricola. b
Microslides. c Ascomata on host surface. d Section through ascoma. e Peridium. f Short hyphal
projections arising from outer layer of peridium. g-j Ascospores. k, l Branching spermatiophores. m
Penicilliate phialides of spermatiophores. Scale bars: c = 500 μm, d = 50 μm, e, f, k = 20 μm, l-m =
10 μm, g-j = 5 μm.
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Hypocreopsis P. Karst., Bidr. Känn. Finl. Nat. Folk 23: 251 (1873)
Index Fungorum number: IF2436; 10 morphological species (Species Fungorum 2020), 3
species with sequence data.
Type species – Hypocreopsis riccioidea (Bolton) P. Karst.
Notes – Hypocreopsis species occur on decaying woody substrates and resupinate
basidiomycetes (Rossman et al. 1999, Stasinska 2004, Johnston et al. 2007). The genus is
characterized by globose, white to pale yellow, perithecial ascomata, immersed in a welldeveloped, radially shreaded, often lobate, reddish brown to grey stromata, cylindrical asci and
ellipsoid to fusiform, 1–3 septate, hyaline, minutly to coarsely warted ascospores (Rossman et al.
1999, Stasinska 2004, Johnston et al. 2007). No asexual morph is linked to this genus.
Hypomyces (Fr.) Tul.&C. Tul.,Annls Sci. Nat., Bot., sér. 4 13: 11 (1860)
Index Fungorum number: IF2446; 115 morphological species (Species Fungorum 2020), 54
species with sequence data.
Type species – Hypomyces lactifluorum (Schwein.) Tul. & C. Tul.
Notes – Hypomyces species are fungicolous, mostly on mushrooms (Agaricales, Boletales),
bracket fungi (Poriales) and discomycetes (Helotiales, Pezizales) (Rossman et al. 1999, Põldmaa
2011, Zeng & Zhuang 2016, Wei & Kirschner 2017). Arnold (1970) introduced the fungicolous
genus Sibirina based on S. fungicola. Sibirina was synonymyzed under Cladobotryum by Rogerson
& Samuels (1993), and was followd by Põldmaa (2003). However the name Hypomyces was
protected against Cladobotryum by Rossman et al. (2013). Hence here we consider Sibrina as a
synonym of Hypomyces by following Gams (2017) and Vu et al. (2019). The genus is characterized
by brightly or lightly coloured, perithecial ascomata in a concolorous subiculum, cylindrical asci
with a thickened apex and, fusiform, aseptate or 1-septate, usually warted ascospores with welldeveloped apiculus at both ends (Põldmaa et al. 2000, Rossman & Seifert 2011). Asexual morphs
are mainly cladobotryum-like, while several other asexual morphs are reported for the genus (i.e.
acremonium-, dactylaria-, mycogone-, papulaspora-, sepedonium-, stephanoma-, trichotheciumand verticillium-like) (Rogerson & Samuels 1993, 1994, Rossman et al. 1999, 2013, Rossman
2000, Wijayawardene et al. 2017b).
Kiflimonium Summerb., J.A. Scott, Guarro & Crous, Microorganisms 6(3, 88): 17 (2018)
Index Fungorum number: IF826819; 1 species with sequence data.
Type species – Kiflimonium curvulum (W. Gams) Summerb., J.A. Scott, Guarro & Crous
Notes – The monotypic genus Kiflimonium was introduced by Summerbell et al. (2018) based
on K. curvulum. Kiflimonium species are mostly isolated from the soil (Summerbell et al. 2018).
The genus is characterized by simple conidiophores arising laterally from somatic hyphae, bearing
1–4 phialidic conidiogenous cells, and 0–1 septate, lunate to falcate, hyaline conidia (Summerbell
et al. 2018).
Lichenobarya Etayo, Diederich & Lawrey, Bryologist 118: 88 (2015)
Index Fungorum number: IF811832; 1 species with sequence data.
Type species – Lichenobarya usneae (Etayo) Etayo, Diederich & Lawrey
Notes – Lawrey et al. (2015) established Lichenobarya for the lichenicolous species N.
usneae based on DNA sequence data. The genus is characterized by superficial, obpyriform,
perithecial, brown ascomata, immersed in a poorly-developed and inconspicuous subiculum,
elongate asci with or without slightly-thickened apex and long, filiform, multi-septate ascospores
(Lawrey et al. 2015). There is no report of a asexual morph for this genus (Lawrey et al. 2015).
Mycogone Link, Mag. Gesell. naturf. Freunde, Berlin 3(1–2): 18 (1809)
Index Fungorum number: IF9026; 28 morphological species (Species Fungorum 2020), 2
species with sequence data.
Type species – Mycogone rosea Link
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Notes – Mycogone species are fungicolous parasites of mushrooms (Agaricales) (Veihmeyer
1914). The genus is characterized by monoblastic, branched or verticillate conidiophores; hyaline,
aseptate or rarely 2–3 septate phialoconidia and 1-septate aleurioconidia (Gray & Morgan-Jones
1980, Gams 1983). No sexual morph is linked to this genus.
Protocrea Petch, J. Bot., Lond. 75: 219 (1937)
Index Fungorum number: IF4383; 6 morphological species (Species Fungorum 2020), 3
species with sequence data.
Type species – Protocrea farinosa (Berk. & Broome) Petch
Notes – Petch (1937) established Protocrea to accomodate Hypocrea farinosa, H. delicatula
and H. stipata. Protocrea species occur on bark, decaying woody substrates or fungicolous on
effused basidiomycetes (Rossman et al. 1999, Jaklitsch et al. 2008). The genus is characterized by
perithecial ascomata, partly or completely immersed in the subiculum, cylindrical asci, fusiform to
ellipsoid, 1-septate ascospores, disarticulating at the septum inside the asci and gliocladium-like
asexual morphs (Rossman et al. 1999, Jaklitsch et al. 2008).
Rogersonia Samuels & Lodge, Sydowia 48(2): 250 (1996)
Index Fungorum number: IF27816; 1 morphological species.
Type species – Rogersonia striolata Samuels & Lodge
Notes – The monotypic genus Rogersonia was introduced by Samuels & Lodge (1996) to
accommodate a single species R. striolata, isolated from decaying wood. The genus is
characterized by perithecial ascomata, partly to completely immersed in the subiculum, cylindrical
asci and aseptate, broadly ellipsoidal, striate ascospores (Samuels & Lodge 1996). No sexual morph
is described for this genus.
Sepedonium Link, Mag. Gesell. naturf. Freunde, Berlin 3(1–2): 18 (1809)
Index Fungorum number: IF9876; 23 morphological species (Species Fungorum 2020), 4
species with sequence data.
Type species – Sepedonium mycophilum (Pers.) Nees
Notes – Link (1809) introduced the asexual morph genus Sepedonium based on S.
mycophilum (Uredo mycophila) as the type of the genus. Sepedonium species are parasitic on
basidiomycetes, mostly on Boletales species (Sahr et al. 1999). The genus is characterized by
hyaline, septate conidiophores and occurrence of two or more synanamorphs producing
aleurioconidia, phialoconidia and ampulloconidia (Sahr et al. 1999). No sexual morph has been
linked to this genus.
Sphaerostilbella (Henn.) Sacc. & D. Sacc., Syll. fung. (Abellini) 17: 778 (1905)
Index Fungorum number: IF5122; 12 morphological species (Species Fungorum 2020), 5
species with sequence data.
Type species – Sphaerostilbella lutea (Henn.) Sacc. & D. Sacc.
Notes – Sphaerostilbella species are mainly fungicolous, occurring on aphyllophorales
basidiomycetes and rarely on wood (Rossman et al. 1999, Põldmaa et al. 2000, Põldmaa & Samuels
2004, Lombard et al. 2015). The genus is characterized by obipyriform ascomata with dark red to
purple, or in shades of yellow, naviculate to ellipsoid, hyaline, 1-septate ascospores and a
gliocladium-like asexual morph (Rossman et al. 1999, Põldmaa et al. 2019).
Sporophagomyces K. Põldmaa & Samuels, Can. J. Bot. 77(12): 1765 (1999)
Index Fungorum number: IF28375; 3 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Sporophagomyces chrysostomus (Berk. & Broome) K. Põldmaa & Samuels
Notes – Sporophagomyces species are fungicolous, associated with aphyllophorous
basidiomycetes (Põldmaa et al. 2000). The genus is characterised by buff to yellow, obypyriform
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ascomata, cylindrical asci with or without thickened apices and (0-) or 1-septate fusiform to
naviculate or lanceolate ascospores (Põldmaa et al. 2000). Asexual morphs are acremonium-like,
comprising simple conidiogenous cells immersed directly on mycelial hyphae, bearing 3-septate
conidia (Põldmaa et al. 2000).
Stephanoma Wallr., Fl. crypt. Germ. (Norimbergae) 2: 269 (1833)
Index Fungorum number: IF10087; 6 morphological species (Species Fungorum 2020).
Type species – Stephanoma strigosum Wallr.
Notes – Stephanoma species are mycoparasites on ascomycetes or sometimes associated with
orchid roots (Wallroth 1833, van Zinderen-Bakker 1934, Butler & McCain 1968, Rakvidhyasastra
& Butler 1973). The genus is characterised by hyphomycetous asexual morphs producing globose
to cuboid, hyaline or golden brown to brown conidia and ovoid, hyaline conidia on verticillate
branches (Wallroth 1833, Butler & McCain 1968). No sexual morph has been linked to this genus.
Trichoderma Pers., Neues Mag. Bot. 1: 92 (1794)
Index Fungorum number: IF10282; 378 morphological species (Species Fungorum 2020),
318 species with sequence data.
Type species – Trichoderma viride Pers.
Notes – Trichoderma is typified by T. viride (Persoon 1794). Trichoderma species are
associated with decaying woody substrates and other fungi (Rossman et al. 1999, Jaklitsch &
Voglmayr 2015). Barrasa et al. (1985) introduced the monotypic genus Aphysiostroma based on
coprophilous taxon, A. stercorarium. However this genus was synonymized under Trichoderma by
Jaklitsch & Voglmayr (2015) based on DNA sequence data. Doi (1972) introduced
Pseudohypocrea to accommodate Hypocrea citrinella. Pseudohypocrea species occur mainly on
decaying wood, bark and dead leaves (Zhu & Zhuang 2014). However, based on morphological
and molecular data Pseudohypocrea was synonymised under Trichoderma by Zeng & Zhuang
(2017). Trichoderma erinaceum is illustrated in this entry (Fig. 121).
Verticimonosporium Matsush., Microfungi of the Solomon Islands and Papua-New Guinea
(Osaka): 68 (1971)
Index Fungorum number: IF10401; 3 morphological species (Species Fungorum 2020), 2
species with sequence data.
Type species – Verticimonosporium diffractum Matsush.
Notes – Matsushima (1971) introduced the asexual morph genus Verticimonosporium based
on V. diffractum. The genus is characterised by swollen, phialidic, conidiogenous cells which are
discrete or in clusters and aseptate, hyaline conidia (Matsushima 1971, 1975, Wang et al. 2005).
Hyponectriaceae Petr., Annls mycol. 21(3/4): 305 (1923)
Index Fungorum number: IF80898; Facesoffungi number: FoF01775; 229 species.
Saprobic or pathogenic on dead plant matter, in terrestrial and aquatic habitats. Sexual
morph: appearing as black, shiny spots on host surface or small black lines arising from cracks in
bark. Pseudostromata present or lacking, if present, superficial, forming clypeus over the ascomata.
Ascomata solitary or aggregated, immersed, erumpent or rarely somewhat superficial, globose to
ovoid, upright or horizontal, brown to black. Papilla short, ostiolate, with or without periphyses.
Peridium comprising two layers, outer layer of brown cells of textura angularis, inner layer of
hyaline cells of textura angularis, in some genera cells are of textura globosa. Paraphyses sparse,
septate, tapering towards the apex, often deliquescing at maturity. Asci 8-spored, unitunicate,
oblong, cylindrical or ellipsoidal, short pedicellate, with a J+, or J-, apical ring. Ascospores
overlapping biseriate, or fasciculate, hyaline, yellow to light brown, fusoid, isthmoid, elongate
filiform, obovoid, or oblong, asymmetric or symmetric, unicellular or septate, smooth-walled or
verruculose, with or without a mucilaginous sheath. Asexual morph: Undetermined.
Type genus – Hyponectria Sacc.
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Notes – The most recent treatments and updated accounts of Hyponectriaceae were by
Maharachchikumbura et al. (2016b) which we follow here. Most of the genera included in this
family lack molecular data, but have characteristics typical of Hyponectria. This includes ascomata
visible as tiny black spots on the leaf surface and J+, or J-, apical rings and unicellular ascospores.
The family is therefore, likely to be a dustbin for numerous unrelated genera and is in need of fresh
collections and sequence data.
Figure 121 – Trichoderma erinaceum (Material examined – INDIA, Andaman and Nicobar
Islands, North Andaman, Diglipur, Nabagram (13˚13’18.1”N 92˚55’59.3”E), found on Baccaurea
ramiflora decaying twig, 17 May 2018, M. Niranjan & V.V. Sarma, AMH-10060; living culture,
NFCCI-4431; INDIA, South Andaman, Port Blair, Wright Mayo (11˚47’39” N 92˚42’36”E), on
Pterocarpus dalbergioides, 22 October 2015, T38F5, T39F2, T52F2, isotypes; INDIA, Mannarghat
(11˚43’14” N 92˚39’33”E), on Pterocarpus dalbergioides, 04 January 2017, Niranjan. M & Sarma.
V.V., T184F1, T193Fl). a, b Stromata on host. c, d Vertical section of ascoma. e, f Asci. g, h
Culture (NFCCI-4431) on malt extract agar plates (g-from above, h-reverse). Scale bars: c = 50 μm,
d = 20 μm, e, f = 10 μm.
Ecological and economic significance of Hyponectriaceae
Species are saprobes, endophytes, and plant pathogens (Boddy & Griffith 1989, Hyde &
Pearce 1993, Olatinwo et al. 2003, Tubajika 2006, Schaechter 2011, Hussain et al. 2014,
Promputtha et al. 2007, Uppala 2019). Physalospora has also been reported as lichenicolous
635
(Hoffmann & Hafellner 2000). Physalospora vaccinia causes a common and widely distributed
fruit disease (blotch rot, fruit rot), and other species are plant pathogens, such as P. piricola (apple
ring rot) (Olatinwo et al. 2003, Tubajika 2006, Kang et al. 2019, Uppala 2019). Extracts of an
endophytic Physalospora sp. exhibited biological activity and showed significant antibacterial,
antifungal, and herbicidal activities (Hussain et al. 2014).
Genera included in Hyponectriaceae
Apiothyrium Petr., Sydowia 1(1-3): 1 (1947)
Index Fungorum number: IF269; 2 morphological species (Species Fungorum 2020).
Type species – Apiothyrium arcticum Petr.
Notes – Apiothyrium was introduced to accommodate A. arcticum occurring on dead leaves
of Diapensia lapponica (Diapensiaceae) in Finland and later A. tasmanicum on dead needles of
Athrotaxis cupressoides (Cupressaceae) in Tasmania, Australia (Petrak 1947a, Swart 1988b).
Apiothyrium species have solitary, immersed ascomata, visible as dots on the host surface, with a
central ostiole, irregular stromata, cylindric-clavate asci with a J+, apical ring, and hyaline, oblong
to clavate, apiospores with a mucilaginous sheath (Petrak 1947a, Swart 1988b).
Arecomyces K.D. Hyde, Sydowia 48(2): 227 (1996)
Index Fungorum number: IF27749; 10 morphological species (Species Fungorum 2020).
Type species – Arecomyces frondicola K.D. Hyde
Notes – Most species of Arecomyces are found on palms (Arecaceae, Hyde 1996a, Hyde &
Fröhlich 2003, Vitoria et al. 2011). The genus has been reported from Australia, Brazil, Brunei, and
Ecuador (Hyde 1996a, Hyde & Fröhlich 2003, Vitoria et al. 2011). Arecomyces is characterised by
solitary or gregarious, immersed, raised darkened discs or minute black dots on the host surface, a
pseudostroma, central ostioles, (2–-)4–8 spored, broadly cylindrical asci with J-, apical rings, and
hyaline, ellipsoidal-fusiform or ovoid, unicellular ascospores surrounded by a mucilaginous sheath
(Hyde 1996a, Vitoria et al. 2011).
Arwidssonia B. Erikss., Svensk bot. Tidskr. 68: 199 (1974)
Index Fungorum number: IF340; 2 morphological species (Species Fungorum 2020).
Type species – Arwidssonia empetri (Rehm) B. Erikss.
Notes – Arwidssonia has immersed ascomata opening by 3–5 lobes, cylindrical to slightly
saccate asci, with J+, apical rings and hyaline, (1–)3(–5)-septate ascospores (Wang & Hyde 1999).
In this study, we examined the holotype of Arwidssonia empetri and accept it in Hyponectriaceae
based on its morphology (Barr 1990b, Wang & Hyde 1999, Maharachchikumbura et al. 2016b,
Wijayawardene et al. 2018a).
Cesatiella Sacc., Michelia 1(no. 2): 250 (1878)
Index Fungorum number: IF917; 3 morphological species (Species Fungorum 2020).
Type species – Cesatiella australis Sacc. & Speg.
Notes – The type species was found on what was probably a dead branch of Olea europaea
(Oleaceae) in Italy (Rossman et al. 1999). Cesatiella has solitary, immersed, cylindric asci with a
J+, apical ring and hyaline, fusoid, oblong to ellipsoid, 3-septate ascospores, without a
mucilaginous sheath (Saccardo 1878).
Chamaeascus L. Holm, K. Holm & M.E. Barr, Blyttia 51(3-4): 121 (1993)
Index Fungorum number: IF26460; 1 morphological species.
Type species – Chamaeascus arcticus L. Holm, K. Holm & M.E. Barr
Notes – The species is a saprobe on dried leaves of Carex (Cyperaceae) found in Norway.
Chamaeascus has solitary, immersed ascomata without papilla, which are darker around the pore,
ellipsoidal asci without an apical ring and bacilliform to lunate, 1-celled ascospores, with obtuse or
slightly pointed ends (Wang & Hyde 1999).
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Figure 122 – Trichoderma viride (Material examined – CZECH REPUBLIC, South Bohemia,
Frymburk, on partly decorticated logs of Pinus sylvestris L., leg. W. Jaklitsch, W.J. 2753, 3
October 2004, WU 24013, epitype). a Herbarium material. b Stromata on host substrate. c, d
Transverse sections through stromata with perithecia. e Peridium in section. f Ostiole in transvers
section. g-i Asci (i in Melzer’s reagent). j Hair on surface of stroma. k-n Ascospores. Scale bars: b
= 2 mm, c = 200 µm, d = 100 µm, e, f = 50 µm, g-j = 20 µm, k-n = 5 µm.
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Figure 123 – Arwidssonia empetri (Material examined – NORWAY: Hedmark: 50 km W of
Tynset: Tatasjöhöi; on dead attached leaves of Empetrum nigrum, 23 August 1985, P.F. Cannon, IMI
312725, holotype). a Material label. b Specimen. c, d Ascomata on the upper leaf surface. e Close
up of ascomata. f Section of ascoma. g Paraphyses. h Peridium. i J+, reaction of apical ascal ring
stained with Melzer’s reagent. j-m Asci. n-s Ascospores. Scale bars: e = 1000 µm, d = 500 μm, e =
200 μm, f = 50 μm, g-h, j-s =10 μm, i-m = 20 μm.
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Discosphaerina Höhn., Sber. Akad. Wiss. Wien, Math.-naturw. Kl., Abt. 1 126(4-5): 353 (1917)
Index Fungorum number: IF1667; 21 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Discosphaerina discophora Höhn.
Notes – Discosphaerina cytisi is the only species in this genus with sequence data (Vu et al.
2019). Species occur on leaves and stems of various plants worldwide, as saprobes, endophytes,
and plant pathogens (Höhnel 1917b, Petrak 1924, 1927, 1934, 1941, Gonzales-Menendez et al.
2014, Godeas et al. 1985, Holm et al. 1999). Discosphaerina has gregarious, often two or three
confluent, semi-immersed ascomata formed under a clypeus, cylindric-clavate asci with a J+, apical
ring and hyaline, ellipsoid, aseptate ascospores (Höhnel 1917b, 1919).
Exarmidium P. Karst., Bidr. Känn. Finl. Nat. Folk 23: 29, 222 (1873)
Index Fungorum number: IF1959; 14 morphological species (Species Fungorum 2020).
Type species – Exarmidium hysteriiforme P. Karst.
Notes – The type species occurs on branches of Juniperus (Cupressaceae) in Finland.
Exarmidium has solitary, aggregated, dark-brown or black ascomata, immersed under a clypeus,
elongate to clavate asci, with a poorly defined apical ring, J+ in some species and hyaline or faintly
yellowish, ascospores with 3 or more transverse septa (Barr & Boise 1985, Aptroot 1998).
Frondicola K.D. Hyde, Bot. J. Linn. Soc. 110(2): 100 (1992)
Index Fungorum number: IF26302; 1 morphological species.
Type species – Frondicola tunitricuspis K.D. Hyde
Notes – Frondicola tunitricuspis, illustrated in this entry, was found on decaying fronds of
Nypa fruticans (Arecaceae). Frondicola has solitary ascomata, visible as dark, raised, circular spots
with central ostiolar dots, immersed under a clypeus, cylindrical asci with J-, apical ring and
hyaline, unicellular, ellipsoidal ascospores with a mucilaginous sheath (Hyde 1992a).
Hyponectria Sacc., Michelia 1(no. 2): 250 (1878)
Index Fungorum number: IF2448; 30 morphological species (Species Fungorum 2020); 1
species and 3 unnamed species with sequence data.
Type species – Hyponectria buxi (Alb. & Schwein.) Sacc.
Notes – Hyponectria was revised and described in Maharachchikumbura et al. (2016b).
Hyponectria has solitary or mostly gregarious, immersed ascomata with a central ostiole, visible as
orange to brown dots on the host surface, cylindric-clavate to clavate, short pedicellate asci with a
J-, apical ring and hyaline, ellipsoidal or oblong, aseptate ascospores, without a mucilaginous
sheath (Maharachchikumbura et al. 2016b). Hyponectria buxi is illustrated in this entry.
Lichenoverruculina Etayo, Herzogia 24(2): [379] (2011)
Index Fungorum number: IF563512; 1 morphological species.
Type species – Lichenoverruculina sigmatospora (Speg.) Etayo & Sharuddin
Notes – Lichenoverruculina is lichenicolous with immersed perithecia under the thallus of
Heterodermia sp. The genus has abundant paraphyses, 32-spored, cylindrical asci and fusoidsigmoid, 2-celled, hyaline ascospores (Spegazzini 1889, Etayo & Rosato 2008).
Micronectria Speg., Anal. Soc. cient. argent. 19(1): 45 (1885)
Index Fungorum number: IF3175; 4 morphological species (Species Fungorum 2020).
Type species – Micronectria guaranitica Speg.
Notes – Micronectria guaranitica forms a sexual morph on leaves of Luehea divaricata while
no asexual morph is known (Spegazzini 1885). This species has been reported from India, Sri
Lanka, and Ethiopia, (Petch 1920, Ciccarone 1951, Ananthanarayanan 1964). Micronectria appears
as scattered, often aggregated, brownish-black, raised spots, immersed, flask-shaped ascomata,
639
oblong-clavate asci without an apical ring and hyaline, 4–5-septate, filiform ascospores, pointed at
both ends (Spegazzini 1885, Ananthanarayanan 1964).
Papilionovela Aptroot, Mycol. Res. 101(3): 266 (1997)
Index Fungorum number: IF27748; 1 morphological species.
Type species – Papilionovela albothallina Aptroot
Notes – Papilionovela albothallina is a saprobe found on branches of an unidentified large
tree in lowland rain forest in Papua New Guinea (Aptroot 1997). Papilionovela has solitary, dark
brown discoid ascomata, cylindrical asci without apical rings, and hyaline, 1(–3)-septate, ellipsoid
ascospores, ornamented with gelatinous wings (Aptroot 1997).
Pellucida Dulym., Sivan., P.F. Cannon & Peerally, Mycol. Res. 105(2): 250 (2001)
Index Fungorum number: IF28461; 1 morphological species.
Type species – Pellucida pendulina Dulym., Sivan., P.F. Cannon & Peerally
Notes – Pellucida pendulina occurs on native plants in Mauritius (Dulymamode et al. 2001).
The genus was placed in Hyponectriaceae and no close relatives have been identified
(Dulymamode et al. 2001). Pellucida has immersed, white to buff, globose, ascomata on the
abaxial and adaxial leaf surfaces, externally only visible as minute depressed areas representing the
ostioles, cylindrical asci with a J+, apical ring, and hyaline, filiform, multiseptate, ascospores,
parallel or helically arranged in the ascus (Dulymamode et al. 2001).
Phragmitensis M.K.M. Wong, Poon & K.D. Hyde, Bot. Mar. 41(4): 379 (1998)
Index Fungorum number: IF27890; 2 morphological species (Species Fungorum 2020).
Type species – Phragmitensis marina M.K.M. Wong, Poon & K.D. Hyde
Notes – Phragmitensis marina occurs on dead culms of Phragmites australis in a saline
estuarine habitat in Hong Kong (Wong et al. 1998b), while P. ellipsoidea was found in freshwater
(Wong et al. 1999). Phragmitensis has aggregated, immersed, pseudostromata, containing several
obpyriform ascomata, clavate asci, lacking an apical ring and hyaline unicellular, cymbiform,
ellipsoidal to reniform ascospores, with distal end tapered more than the proximal end and
surrounded by a mucilaginous sheath (Wong et al. 1998b, 1999).
Physalospora Niessl, Verh. nat. Ver. Brünn 14: 170 (1876)
Index Fungorum number: IF4079; 132 morphological species (Species Fungorum 2020), 3
species and 5 unnamed species with sequence data.
Type species – Physalospora alpestris Niessl
Notes – Physalospora species have solitary, immersed, clypeate, black ascomata, cylindrical
asci, and hyaline, fusiform ascospores, sometimes becoming pale brown at maturity and with or
without a mucilaginous sheath (Wang & Hyde 1999, Sivanesan & Shivas 2002).
Rachidicola K.D. Hyde & J. Fröhl., Sydowia 47(2): 217 (1995)
Index Fungorum number: IF27632; 1 morphological species.
Type species – Rachidicola palmae K.D. Hyde & J. Fröhl.
Notes – Rachidicola palmae occurs on living and dead fronds of palms (Arecaceae) in Brunei
(on rachis of Oncosperma horridum), Hong Kong (on rachis of Calamus sp.) and Malaysia (on
Daemonorops sp.). Rachidicola has solitary, dark-brown, raised ascomata, immersed under a
clypeus, with a central ostiole, cylindric-clavate asci with a non-refractive apical ring and 2–3seriate, hyaline, straight or slightly curved, 3-celled ascospores surrounded by a mucilaginous
sheath. (Hyde & Fröhlich 1995).
Xenothecium Höhn., Sber. Akad. Wiss. Wien, Math.-naturw. Kl., Abt. 1 128: 589 (1919)
Index Fungorum number: IF5827; 1 morphological species.
Type species – Xenothecium jodophilum Höhn.
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Figure 124 – Frondicola tunitricuspis (Material examined – BRUNEI, on bark of Nypa fruticans
Wurmb (Arecaceae), 14 June 1987, K.D. Hyde, BRIP17110, holotype). a Material label. b
Ascomata on host substrate. c Close up of ascoma. d Section of ascoma. e Peridium. f Paraphyses.
g-j Asci. k-o Ascospores. Scale bars: c = 500 μm, d = 100 μm, e = 20 μm, f, k-o = 10 μm, g-j = 50
μm.
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Figure 125 – Hyponectria buxi (Material examined – UK, Surrey, Mickleham, on leaves of Buxus
sp., 1927, E.W. Mason no. 365, IMI 16895, IMI52229, IMI49466). a Herbarium specimen. b
Ascomata on the upper leaf surface. c Ascomata on the lower leaf surface. d Close up of ascoma. e
Close up of ascomata under microscope. f J-, reaction of apical ring in Melzer’s reagent. g Cells of
peridium. h Section of ascomata. i-k Asci. l-q Ascospores. Scale bars: b-c = 500 μm, d = 200 μm, e,
h = 50 μm, f, i-k = 10 μm, g = 20 μm, l-q = 5 μm.
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Notes – Xenothecium jodophilum has short-pedicellate, cylindrical asci, with a J+, apical ring
and uniseriate, 1-celled, hyaline ascospores, lacking a mucilaginous sheath (Höhnel 1919).
Hypoxylaceae DC. Fl. franc¸., Edn 3 (Paris) 2: 280 (1805)
Index Fungorum number: IF81885; Facesoffungi number: FoF02979; 422 species.
Saprobic or endophytic in wood, leaves and fruits, sometimes symbiotically associated with
insect vectors in terrestrial and aquatic habitats (Pažoutová et al. 2010, 2013). Sexual morph:
Stromata (if present) erect, glomerate, pulvinate, discoid, effused-pulvinate, hemispherical,
spherical or peltate, or sometimes rudimentary or lacking, arising singly or aggregated into groups,
with one to several ascomata, surface colour variable, mostly black-brown, ostiolate, mostly with
extractable stromal pigments, unipartite, attached to the surface by broad bases, conical-dome
shaped, raised areas. Ascomata variable in size and shape, globose-subglobose or elongate
cylindrical-pyriform, embedded in the stroma, monostichous, interior sometimes zonate or filled
with liquid. Ostiole papillate or umbilicate or at the same level as the stromal surface, with or
without discs. Paraphyses hyaline, filamentous, septate, embedded in a gelatinous matrix, usually
longer than the asci. Asci 4–8-spored, unitunicate, cylindrical to clavate, rarely globose, pedicellate
or apedicellate, apically rounded, with or without a J+, or J-, apical ring, or with apical thickenings.
Ascospores uniseriate-biseriate (crowded in Phylacia and Pyrenomyxa), uni-bicellular, ellipsoidalsubglobose, brown to black, rarely hyaline, mostly with a germ slit, straight, spiral or sigmoid,
sometimes with rostrate, hyaline dwarf cell. Perispore dehiscent or sometimes lacking, smooth or
with patterns. Asexual morph: Conidiophores mostly nodulisporium-like, hyaline to light brown,
smooth, branched, mononematous or infrequently synnematous, usually macronematous.
Conidiogenous cells cylindrical, usually hyaline, one to several on each branch of the conidiophore,
swollen apex. Conidia hyaline, roughened or smooth, ellipsoidal (adapted from Wendt et al. 2018).
Type genus – Hypoxylon Bull.
Notes – The term Hypoxylaceae was not used in modern classifications until 2017, and
treated under Xylariaceae (as ‘Hypoxyloideae’). Wendt et al. (2018) validated the family in the
Xylariales based on multi-locus phylogeny, morphology and chemotaxonomy studies. Molecular
clock evidence confirmed the familial position of Hypoxylaceae within Xylariales (Hongsanan et
al. 2017). Daranagama et al. (2018) accepted 18 genera including Alloanthostomella,
Neoanthostomella and Pseudoanthostomella in the Hypoxylaceae. However, according to the
phylogenetic studies by Voglmayr et al. (2018) and Wendt et al. (2018), these three genera cluster
as a separate clade in Xylariaceae sensu stricto; thus they are placed in Xylariales genera incertae
sedis. Herein, we accept 19 genera in Hypoxylaceae.
Ecological and economic significance of Hypoxylaceae
Species of Hypoxylaceae are mainly saprobes and endophytes, however, none are serious
pathogens although there have been several reports on Hypoxylaceae species causing disease
symptoms in water-stressed trees (Stadler 2011). The endophytic stages of Hypoxylaceae species
have a rather interesting ecology and may even be beneficial to their host plants, because they
protect them from pathogens, including other fungi, as well as invertebrates. Daldinia and
Hypoxylon have been intensively studied for chemical profiling (Helaly et al. 2018).
Genera included in Hypoxylaceae
Annulohypoxylon Y.M. Ju, J.D. Rogers & H.M. Hsieh, Mycologia 97(4): 855 (2005)
Index Fungorum number: IF500298; 62 morphological species (Wendt et al. 2018), 35
species with sequence data.
Type species – Annulohypoxylon truncatum (Starbäck) Y.M. Ju, J.D. Rogers & H.M. Hsieh
Notes – Annulohypoxylon was introduced to accommodate Hypoxylon sect. Annulata sensu
Ju & Rogers (1996). Kuhnert et al. (2017) provided a concise revision of the genus and described
several additional species based on molecular phylogenetic and chemotaxonomic evidence. Based
on a multigene genealogy, Jackrogersella was segregated from Annulohypoxylon (Kuhnert et al.
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2017). Jackrogersella is characterised by papillate ostioles and its species produce azaphilones of
the cohaerin type as predominant stromatal metabolites, while stromata of Annulohypoxylon sensu
stricto mostly have ostioles encircled by an annulated disc and lack azaphilones but contain
binaphthalenes as major stromatal metabolites (Wendt et al. 2018).
Anthocanalis Daranag., Camporesi & K.D. Hyde, Fungal Divers. 73: 211 (2015)
Index Fungorum number: IF809515; 1 species with sequence data.
Type species – Anthocanalis sparti Daranag., Camporesi & K.D. Hyde
Notes – The monotypic genus Anthocanalis was introduced to accommodate an
anthostomella-like species characterized by ascomata with a reduced clypeus, shiny papilla, a
peridium with hyaline cells of textura angularis, light brown cells of textura irregularis and
limoniform ascospores with a prominent sheath only at the ventral side. Combined ITS, LSU, tub2
and rpb2 phylogenetic analyses suggested that the genus is closely related to Rhopalostroma,
Ruwenzoria and Thamnomyces in Hypoxylaceae (Daranagama et al. 2018, Wendt et al. 2018).
However, the available LSU, ITS, rpb2 and tub2 DNA sequences of A. sparti as well as the
reported conidiogeneous structures are similar to Daldinia eschscholtzii, which needs further
clarification.
Chlorostroma A.N. Mill., Lar.N. Vassiljeva & J.D. Rogers, Sydowia 59(1): 142 (2007)
Index Fungorum number: IF29126; 3 morphological species (Daranagama et al. 2018).
Type species – Chlorostroma subcubisporum A.N. Mill., Lar.N. Vassiljeva & J.D. Rogers
Notes – Chlorostroma has character of green stromata, J-, ascal apical ring and more or less
cubical ascospores. All the species have been described based on stromata of Hypoxylon (Miller et
al. 2007, Nordén et al. 2014). Læssøe et al. (2010) and Wendt et al. (2018) placed Chlorostroma in
Hypoxylaceae based on highly similar secondary metabolite profiles to Hypoxylon aeruginosum.
Daldinia Ces. & De Not., Comm. Soc. crittog. Ital. 1(fasc. 4): 197 (1863)
Index Fungorum number: IF1408; 53 morphological species (Species Fungorum 2020), 43
species with sequence data.
Type species – Daldinia concentrica Ces. & De Not.
Notes – The genus was revisited and redefined by Stadler et al. (2014) and later included in
Hypoxylaceae because of its affinities with Hypoxylon. Most species are characterized by welldefined concentric zones in the stromatal interior (Stadler et al. 2014). Multigene phylogeny studies
showed that the genus comprises two well-supported clades within the family (Daranagama et al.
2018, Wendt et al. 2018). Daldinia eschscholtzii is illustrated in this entry (Figs. 126, 127).
Durotheca Læssøe, Srikitik., Luangsa-ard & M. Stadler, IMA Fungus 4(1): 62 (2013)
Index Fungorum number: IF803610; 6 morphological species (De Long et al. 2019), 6
species with sequence data.
Type species – Durotheca depressa Læssøe & Srikitik.
Notes – Durotheca species have narrow, undulating stromata with ostioles in deep
depressions and ascospores with a germ slit or inconspicuous germ slit. Daranagama et al. (2018)
and Wendt et al. (2018) treated the genus in Xylariales, incertae sedis due to chemotaxonomic
affinities with Chlorostroma and Hypoxylon aeruginosum. Based on multigene phylogeny, De
Long et al. (2019) described two species D. crateriformis and D. guizhouensis confirming their
taxonomic placement in Hypoxylaceae.
Entonaema A. Möller, Botanische Mittheilungen aus den Tropen 9: 306 (Möller 1901)
Index Fungorum number: IF1837; 6 morphological species (Daranagama et al. 2018), 2
species with sequence data.
Type species – Entonaema liquescens A. Möller
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Figure 126 – Daldinia eschscholtzii (Material examined – INDIA, Andaman and Nicobar Islands,
South Andaman, Manjery, Near Amber Sunset Point (11˚51’68.6”N 92˚65’38.8”E), on Tamarindus
indica 10 December 2017, M. Niranjan, PUFNI 1761 (AMH-10066); living culture, NFCC 4365.
INDIA, South Andaman, Manjery (11°52'68"N 92°64'74"E), on unidentified twig, 11 December
2017, M. Niranjan (T101F1); INDIA, Middle Andaman, Mayabunder (12˚49’15” N 92˚51’02”E),
on unidentified twig, 7 January 2017 (T95F2). a, b Stromata on decaying host. c Vertical section of
stromata. d Ascospores. e Germinating spore. f Stromata giving the purple colour in 10% KOH. g
Peridium of textura oblita. h Textura angularis. i Paraphyses. j-m Immature and mature asci. Scale
bars: d = 200 μm, g, h, l, n = 50 μm, j, k m = 20 μm, d, e, g = 10 μm.
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Figure 127 – Daldinia eschscholtzii (NFCC 4365). a, b Colonies on MEA (a-from above, bbelow). c Germinating spore. d-h Conidiophores with attached conidia. i Conidia. Scale bars: d-f =
20 μm, g-i = 10 μm.
Notes –Entonaema is characterized by hollow, gelatinous stromata similar to Daldinia
(Stadler et al. 2008). Based on the stromatic sexual morph, nodulisporium-like asexual morph,
chemotaxonomy and multigene phylogeny, Entonaema has been placed in Hypoxylaceae (Wendt et
al. 2018).
Hypomontagnella Sir, L. Wendt & C. Lamb. Mycological Progress: 1–15 (2018)
Index Fungorum number: IF827251; 5 morphological species (Lambert et al. 2019), 2
species with sequence data.
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Type species – Hypomontagnella monticulosa (Mont.) Sir, L. Wendt & C. Lamb.
Notes – Lambert et al. (2019) erected Hypomontagnella to accommodate several species
previously described as Hypoxylon species. Hypomontagnella differs from Annulohypoxylon and
Jackrogersella by smooth perispores or with transversally striate ornamentations. It differs from
species of Hypoxylon in having woody to carbonaceous stromata lacking coloured granules,
papillate ostioles usually with a black annulate disc, and without apparent KOH-extractable
pigments in mature stromata (Lambert et al. 2019). Cultures of all Hypomontagnella species
studied so far produce the strongly antifungal polyketides of the sporothrolide type, which have
never been found in any other species of Xylariales and are otherwise only known from spurious
endophytes. Hypomontagnella monticulosa is illustrated in this entry (Fig. 128).
Hypoxylon Bull., Histoire des champignons de la France. I: 168 (1791)
Index Fungorum number: IF2456; 232 morphological species (Species Fungorum 2020), 95
species with sequence data.
Type species – Hypoxylon fragiforme (Pers.) J. Kickx f.
Notes – Hypoxylon is an extensively studied, species rich genus with more than 200 taxa, but
with more than 1000 epithets (Kuhnert et al. 2014, Wijayawardene et al. 2017a). Miller (1961)
followed stromatal and ascospore morphology for species delimitation and Ju & Rogers (1996)
widely used morphology of asexual morph and stromatal pigments. Currently, the genus has further
been restricted with a modern phylogenetic concept based on molecular data. Hypoxylon species
are widely distributed and mainly inhabit dead wood as saprobes (Ju & Rogers 1996, Wendt et al.
2018). Some Hypoxylon species occur as endophytes and facultative parasites on diseased hosts (Ju
& Rogers 1996, Kuhnert et al. 2014). Many secondary metabolites have been investigated from
Hypoxylon species (Helaly et al. 2018).
Jackrogersella L. Wendt, Kuhnert & M. Stadler, Mycol. Prog. (2018)
Index Fungorum number: IF819742; 6 morphological species (Wendt et al. 2018), 3 species
with sequence data.
Type species – Jackrogersella multiformis L. Wendt, Kuhnert & M. Stadler
Notes – Jackrogersella was introduced to accommodate species previously described as
Annulohypoxylon, and characterized by papillate ostioles and lacking prominent ostiolar disks.
Species in Jackrogersella also differ from Annulohypoxylon by containing cohaerin/multiformin
type azaphilones rather than binaphthalele derivatives as predominant secondary metabolites.
Based on the phylogeny of ITS, LSU, tub2 and rpb2, the genus is well supported in Hypoxylaceae
(Wendt et al. 2018). One species, which is for the time being retained as A. michelianum, occupies
a separate clade.
Natonodosa Heredia, R.F. Castañeda & D.W. Li, Mycological Progress 19: 23 (2020)
Index Fungorum number: IF819412; 1 species with sequence data (Heredia et al. 2020).
Type species – Natonodosa speciosa Heredia, R.F. Castañeda & D.W. Li
Notes –Natonodosa is characterized by effuse, hairy, pigmented colonies. Conidiophores are
septate, greyish brown to yellowish brown, paler towards the apex, with integrated, terminal and
intercalary, doliiform, cylindrical or subcylindrical conidiogenous cells, with several subhyaline
minute denticles and acropleurogenous, simple, fusiform to navicular, unicellular, dry, smooth and
hyaline conidia. The sexual morph is undetermined (Heredia et al. 2020).
Phylacia Lév., Annls Sci. Nat., Bot., sér. 3 3: 61 (1845)
Index Fungorum number: IF4047; 10 morphological species (Species Fungorum 2020), 2
species with sequence data.
Type species – Phylacia globosa Lév.
Notes – Phylacia is characterized by cleistocarpous pyrenomycetes, lacking an ostiolar canal
and opening by regular rupture of the ascomatal apex (Rodrigues & Samuels 1989, Medel et al.
647
2006). Earlier the genus was accepted in Phylaciaceae (Speer 1980b). Based on morphology
(particularly the asexual morph), the strikingly similar metabolite profiles of the cultures to
Daldinia (Bitzer et al. 2008) as well as inferred from molecular phylogenetic data, Phylacia was
accommodated in Hypoxylaceae by Wendt et al. (2018). The asexual morph and stromatal
pigments produced by Phylacia species also have close affinities to Thamnomyces and
Rhopalostroma (Stadler et al. 2004, 2010).
Pyrenomyxa Morgan, J. Cincinnati Soc. Nat. Hist. 18: 42 (1895)
Index Fungorum number: IF22327; 3 morphological species (Species Fungorum 2020), 2
species with sequence data.
Type species – Pyrenomyxa invocans Morgan
Notes – Stromata of Pyrenomyxa are similar too species of the H. rubiginosum complex when
collected in the field, but their ascal morphology is different, since they feature subglobose asci in
which phaseoliform, laterally compressed ascospores are arranged in a similar manner to the slices
of an orange fruit. The ascospores are not forcibly discharged as in most other Hypoxylaceae
genera aside from Phylacia, Rhopalostroma and Thamnomyces, but they are released from the
mature stromata in powdery masses.
A comparison of morphological features and secondary metabolite profiles of Pyrenomyxa
invocans, Hypoxylon piceum and Pulveria porrecta were evaluated by Stadler et al. (2005) and it
was concluded that they are congeneric with Pyrenomyxa which is the older name and takes
preference. The molecular data and the secondary metabolite profiles (rubginosins and hypomiltins
as prevailing stromatal metabolites) clearly suggest that this genus has affinities with Hypoxylon.
Pyrenopolyporus Lloyd, Mycological Writings 5 (50): 706 (1917)
Index Fungorum number: IF4599; 5 morphological species (Species Fungorum 2020), 4
species with sequence data.
Type species – Pyrenopolyporus hunteri Lloyd
Notes – Pyrenopolyporus is characterized by massive stromata with vinaceous shades on the
surface, long tubular perithecia and umbilicate ostioles. Superficially, these stromata actually
resemble a polyporaceous basidiomycete. Based on morphology, phylogeny and chemotaxonomy,
Wendt et al. (2018) resurrected Pyrenopolyporus and transferred it to Hypoxylaceae. The genus has
a close affinity to Daldinia, and the stromata resemble those of D. placentiformis and D. korfii,
which lack conspicuous internal stromatal concentric zones. However, Pyrenopolyporus species
deviate from those of the latter genus by producing a characteristic virgariella-like asexual morph
in culture, by having characteristic, irregular ascospores and by producing drastically different
secondary metabolite profiles in their stromata as well as their cultures (Wendt et al. 2018). For
example, the cultures of Pyrenopolyporus produce cochliodinol, whereas those of Daldinia produce
chromones and other small polyketides.
Rhopalostroma D. Hawksw., Kew Bull. 31(3): 422 (1977)
Index Fungorum number: IF4722; 11 morphological species (Species Fungorum 2020), 4
species with sequence data.
Type species – Rhopalostroma indicum D. Hawksw. & Muthappa
Notes – Rhopalostroma species are characterized by pedicellate, melanized stromata with
often abruptly expanded convex heads. The globose to subglobose stromatal morphology of
Rhopalostroma is close to Phylacia. Rhopalostroma possesses a single layer of ascomata in the
stroma, asci with apical ring and ascospores with a germ slit, which differs from Phylacia with
multilayers and compact ascomata in the stroma, lack of ascal apical ring and lack of germ slit in
the ascospores. Rhopalostroma species are restricted to subtropical Africa and South Asia. Based
on morphological affinities with Thamnomyces and Phylacia, phylogenetic analyses and secondary
metabolite profiles, the genus is accepted in Hypoxylaceae (Hawksworth 1977, Stadler et al. 2010,
Daranagama et al. 2014, 2018, Wendt et al. 2018).
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Figure 128 – Hypomontagnella monticulosa (Material examined – THAILAND, Naan, Doi Phu
Kha, dead branch of Leucaena leucocephala (Fabaceae), 04 August 2017, MC. Samarakoon
SAMC007, MFLU 18-0822, HKAS 102355, living culture MFLUCC 18-0362). a Mature stroma
on bark. b Stromatal surface showing papillate and ostiolar discs (indicated by white arrows). c
Stromata in vertical section. d-f Asci. g Ascospores (sigmoid germ slits indicated by white arrows).
h Ascus tip in Lugol’s reagent. i, j Perispores. k, l KOH-extractable pigments. m Surface of colony.
n Reverse of colony. o-q Conidiophores and conidia. Scale bars: a = 2 mm, b, c = 500 μm, d-g = 20
μm, i, j = 10 μm, o-q = 5 μm.
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Rostrohypoxylon J. Fourn. & M. Stadler, Fungal Divers. 40: 24 (2010)
Index Fungorum number: IF512543; 1 species with sequence data.
Type species – Rostrohypoxylon terebratum J. Fourn. & M. Stadler
Notes – Rostrohypoxylon is a monotypic genus characterized by its erumpent effuse stromata
featuring stout, strongly protruding ostiolar necks and KOH extractable pigments. Multigene
phylogenies confirmed that Rostrohypoxylon has a close affinity to Annulohypoxylon, but it
appeared in a sister clade to the species of the latter genus and has different stromal HPLC profiles.
Therefore, the genus is accepted as a separate genus in Hypoxylaceae (Fournier et al. 2010,
Daranagama et al. 2018, Wendt et al. 2018).
Ruwenzoria J. Fourn., M. Stadler, Læssøe & Decock, Mycol. Progr. 9(2): 171 (2010)
Index Fungorum number: IF515155; 1 species with sequence data.
Type species – Ruwenzoria pseudoannulata J. Fourn., M. Stadler, Læssøe & Decock
Notes – Ruwenzoria is characterized by a blackish, slightly nodulose and cracked stromatal
surface, ostioles with an umbilicate opening at the centre of a raised disc and J-, ascal apical ring.
Based on phylogenetic and chemotaxonomic analysis, Ruwenzoria is accepted in Hypoxylaceae
and has close affinity to Daldinia and Thamnomyces (Stadler et al. 2010, 2014, Daranagama et al.
2018, Wendt et al. 2018).
Thamnomyces Ehrenb., Horae Phys. Berol.: 79 (1820)
Index Fungorum number: IF5402; 11 morphological species (Stadler et al. 2010), 1 species
with sequence data.
Type species – Thamnomyces chamissonis Ehrenb.
Notes – Thamnomyces is characterized by erect, black, unbranched to dendroid stromata with
ascomata embedded on the tips of branches or laterally on the unbranched, wiry stromata. Its
species are mostly known from the neotropics, except for Thamnomyces camerunensis, which was
only found from Western Africa. Morphology, multigene phylogenies and chemical profiling have
revealed that the genus is close to Daldinia, Phylacia and Rhopalostroma (Stadler et al. 2010), and
accordingly is included in Hypoxylaceae (Daranagama et al. 2018, Wendt et al. 2018).
Theissenia Maubl., Bull. Soc. mycol. Fr. 30(1): 52 (1914)
Index Fungorum number: IF5418; 1 species with sequence data.
Type species – Theissenia pyrenocrata (Theiss.) Maubl.
Notes – Theissenia has a conspicuous central bulge or umbo in the perithecium and early
dissolving asci. Ju et al. (2007) accepted the genus in Xylariaceae based on its nodulisporium-like
asexual morph and act and tub2 phylogeny. Læssøe et al. (2013) accepted it in Graphostromataceae
because of its nodulisporium-like asexual morph and close relationship to Biscogniauxia.
Daranagama et al. (2018) and Wendt et al. (2018) accepted the genus in Xylariales genera incertae
sedis. A recent study by De Long et al. (2019) placed the genus in Hypoxylaceae.
Thuemenella Penz. & Sacc., Malpighia 11(11–12): 518 (1898)
Index Fungorum number: IF5459; 4 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Thuemenella cubispora (Ellis & Holw.) Boedijn
Notes – Thuemenella is characterized by having bright yellow stromata, green ascospores and
a nodulisporium-like asexual morph. Several recent studies have transferred this genus to
Hypoxylaceae based on its nodulisporium-like asexual morph and ITS and tub2 similarity of a
sequence derived from Thuemenella cubispora with those of Hypoxylon rubiginosum (Miller et al.
2007, Wendt et al. 2018).
Induratiaceae Samarak., Thongbai, K.D. Hyde & M. Stadler, Fungal Divers. (in press)
MycoBank number: MB833443; Facesoffungi number: FoF06905; 29 species.
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Saprobic on dead wood, leaves, endophytic on leaves, stems and barks. Sexual morph:
Stromata scattered, solitary or rarely pair-wise, globose to subglobose, barely raised above the
host surface, visible as light brown-black, carbonaceous spots, immersed to semi-immersed in
host tissue, attached to substrate with a broad base, with a single ascomata per stroma. Clypeus
black, thick-walled, short, comprising dark fungal hyphae and host epidermal cells. Ostiole
papillate, centric. Peridium wide at base, composed of two layers; inner layer hyline, thin walled
cells of textura angularis; outer layer yellowish brown, thick walled cells of textura angularis.
Paraphyses hyaline, smooth, thin-walled, two types; type 1 branched, with short, lateral bridges,
aseptate, type 2 infrequently branched, septate. Asci 8-spored, unitunicate, cylindrical, short
pedicellate, apically rounded with a J+, apical ring. Ascospores uniseriate, naviculate to
ellipsoidal or long fusiform, mostly hyaline, smooth-walled, constricted apiosporous; apical cell
conical shape, guttulate with remnant at the top; base cell rarely brown, guttulate with remnant at
the base, produce germ tube during the spore germination. Culture charcateristics: White to pale
orange, straight, right angle hyphae, wavy or cottony hyphal growth, rope-like with cauliflowerlike hyphal bodies, strong odour. Asexual morph: Conidiophores nodulisporium-like (Induratia),
abundant, and irregularly branched in the upper part. Conidiogenous cells terminal, solitary or
sometimes two celled at the ends of branches, cylindrical, pale brown, bearing inconspicuous
denticles. Conidia narrowly ellipsoidal to subglobose, hyaline, smooth, with a flat, wide, basal
scar. idriella-like (Emarcea) (Samarakoon et al. 2019b).
Type genus – Induratia Samuels, E. Müll. & Petrini
Notes – Samarakoon et al. (2020) introduced Induratiaceae to accommodate genera Emarcea
and Induratia (= Muscodor) with apiosporous ascospores. Two novel species collected from
northern Thailand revealed their phylogenetic affinity to earlier described Muscodor, Emarcea
species and one Induratia sp. SMH 2155. Based on morphology, ITS-LSU-rpb2-tub2 phylogeny
and gas chromatography coupled with mass spectrometry (GC-MS), Induratiaceae was established
in Xylariales (Samarakoon et al. 2020).
Ecological and economic significance of Induratiaceae
Induratiaceae consists of mainly endophytic species (which were previously referred to as
Muscodor spp.) and a few known saprobic species. Strobel et al. (2001) described at least 28 VOCs
associated with Induratia species (as “Muscodor”) that can induce lethal effects against a broad
range of fungal and bacterial plant and human pathogens, as well as acari, insects and nematodes
(Daisy et al. 2002, Atmosukarto et al. 2005, Mercier & Manker 2005, Strobel 2006, Kudalkar et al.
2012, Hutchings et al. 2017).
Genera included in Induratiaceae
Emarcea Duong, Jeewon & K.D. Hyde, Stud. Mycol. 50(1): 255 (2004)
Index Fungorum number: IF500070; 3 morphological species (Species Fungorum 2020), 2
species with sequence data.
Type species – Emarcea castanopsidicola Duong, Jeewon & K.D. Hyde
Notes – Emarcea castanopsidicola was introduced from dead leaves of Castanopsis
diversifolia in northern Thailand (Duong et al. 2004). The genus is characterized by immersed,
globose to subglobose, coriaceous, solitary ascomata, immersed under a blackened clypeus and
hyaline apiospore with mucilage at the ends and idriella-like asexual morphs. Emarcea differs from
Induratia by its long fusiform ascospores, obclavate apical cells and similar by its apiospores
without germ slits (Li et al. 2009, Daranagama et al. 2018). The ITS, LSU, rpb2 and tub2
multigene phylogeny revealed that Emarcea is basal to Induratia with high statistical support.
Induratia Samuels, E. Müll. & Petrini, Mycotaxon 28(2): 484 (1987)
Index Fungorum number: IF25403; 26 morphological species (Samarakoon et al. 2020;
mostly previously classified as Muscodor spp.), 25 species with sequence data.
Type species – Induratia apiospora Samuels, E. Müll. & Petrini
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Figure 129 – Induratia thailandica (Material examined – THAILAND, Chang Wat Chiang Mai,
Amphoe Mae Taeng, Tambon Pa Pae, on dead wood, 1 September 2017, M.C. Samarakoon,
SAMC015, MFLU 18-0784, holotype). a-c Ascomata on the host surface. d Vertical section of
ascoma. e, f Section of peridium (f in Congo Red). g, k-m Asci (g in Congo Red). h Paraphyses. i
Apical ring J+ in Melzer’s reagent. j Ascus apex in Congo Red. n-r Ascospores. Scale bars: d = 200
µm, e-g, k-m = 20 µm, h, n-r =10 µm, i, j = 5 µm.
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Notes – Muscodor was introduced by Worapong et al. (2001) and the species identification is
mainly based on their characteristic volatile organic compounds (VOCs), inconclusive hyphal
characters such as coiling, ropyness and by right angle branching patterns and inadequate
phylogenetic analyses (Worapong et al. 2001, González et al. 2009). Due to the lack of taxon
sampling, incomplete phylogenetic comparisons and undetermined morphology the taxonomic
placement of Muscodor was considered doubtful (Stadler et al. 2013, Wendt et al. 2018).
Therefore, Lumbsch & Huhndorf (2010) did not accept the genus, while Seifert & Gams (2011),
Maharachchikumbura et al. (2016b) and Daranagama et al. (2018) accepted the genus and placed it
in Xylariales genera incertae sedis. Wendt et al. (2018) proposed to reject the genus from the
family because of poor taxonomic standards. Samarakoon et al. (2020) found evidence that the
Muscodor is connected to Induratia, whose sexual morph was described by Samuels et al. (1987).
Based on a polyphasic taxonomic approach including a multi gene genealogy, as well as volatile
metbolite profiles and dual culture experiments, Samarakoon et al. (2020) erected nine new species
and several new combinations of Induratia mainly resolving the taxonomic confusions of
previously published Muscodor species into a new family Induratiaceae. Induratia thailandica is
illustrated in this entry (Fig 129).
Iodosphaeriaceae O. Hilber, The Genus Lasiosphaeria and Allied Taxa (Kelheim): 7 (2002)
Index Fungorum number: IF82138; Facesoffungi number: FoF06189; 8 species.
Saprobic on dead twigs and leaves of various hosts. Sexual morph: Ascomata superficial,
solitary, black, and easily removed from the substrate, covered with dark brown, setae-like, brown
hairs, comprising agglutinated mycelial strands, with a stellate arrangement, arising from cells at
the perithecial surface. Ostioles pore-like opening, central, with periphyses. Peridium outer region
comprising pigmented, brown cells of textura angularis, inner region comprising hyaline, flattened
cells. Paraphyses numerous, hypha-like, septate, flexuose, slightly tapered towards the apex. Asci
8-spored, unitunicate, narrowly clavate, short pedicellate or apedicellate, apex rounded, with a J+,
subapical ring. Ascospores biseriate, hyaline, allantoid, unicellular, smooth-walled, lacking sheaths
or appendages. Asexual morph: ceratosporium-like conidia have been observed on the surface of
perithecia, but may not be related.
Type genus – Iodosphaeria Samuels, E. Müll. & Petrini
Notes – Iodosphaeriaceae was introduced to accommodate Iodosphaeria and placed in
Amphisphaeriales (Hilber & Hilber 2002). Maharachchikumbura et al. (2016b) and Samarakoon et
al. (2016) placed this family within Xylariales. Iodosphaeriaceae was placed in Xylariomycetidae,
families incertae sedis (Hyde et al. 2017b, Hongsanan et al. 2017, Wijayawardene et al. 2018a).
We accept Iodosphaeriaceae in Amphisphaeriales based on multigene analysis of combined LSU,
ITS, rpb2 and tub2 sequence data (Fig. 5).
Ecological and economic significance of Iodosphaeriaceae
Most Iodosphaeria species are saprobes which occur worldwide (Samuels et al. 1987, Barr
1994, Hyde 1995d, Candoussau et al. 1996, Hsieh et al. 1997b, Taylor & Hyde 1999, Catania &
Romero 2012, Li et al. 2015a, Marasinghe et al. 2019), but have never been reported as a
pathogenic on hosts. They are probably endophytes that become saprobes at leaf senescence.
Genus included in Iodosphaeriaceae
Iodosphaeria Samuels, E. Müll. & Petrini, Mycotaxon 28(2): 486 (1987)
Index Fungorum number: IF25402; 8 morphological species (Species Fungorum 2020), 2
species with sequence data.
Type species – Iodosphaeria phyllophila (Mouton) Samuels, E. Müll. & Petrini
Notes – Iodosphaeria was placed in Amphisphaeriaceae by Samuels et al. (1987) and also
accepted by Eriksson et al. (2001). However, Barr (1994) and Hyde (1995d) suggested this genus
should be placed in Lasiosphaeriaceae. Kang et al. (1998, 1999a) and Jeewon et al. (2003b)
suggested that Iodosphaeria should be excluded from Amphisphaeriaceae and the genus was
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transferred to Trichosphaeriaceae (Réblová 1999d). Hilber & Hilber (2002) introduced
Iodosphaeriaceae and treated. Iodosphaeria aquatica was excluded from Iodosphaeria based on
morphology being similar to Pseudohalonectria lignicola (Hsieh et al. 1997b). Iodosphaeria
tongrenensis was introduced by Li et al. (2015a) from China with morphology and molecular
evidence. Iodosphaeria honghense was introduced from China by Marasinghe et al. (2019) and is
illustrated here.
Figure 130 – Iodosphaeria honghense (Material examined – CHINA, Yunnan Province, Honghe,
on dead stems, 15 June 2018, Junfu Li, MFLU 19-0719, holotype). a, b Appearance of ascomata on
host substrate. c Close up of ascoma. d Fruiting body under the light microscope. e Section of
ascoma. f Periphyses (septate at red arrows). g Peridium. h Associated ceratosporium-like conidia,
but probably not related. i Unbranched brown hairs with septa (red arrows). j, k Paraphyses. l-n
Asci. o Ascus apical ring (stained in Melzer´s reagent). p-v Ascospores. Scale bars: b, c = 500 μm,
d = 200 μm, e, h = 100 μm, f-h, l-n = 50 μm, j-k, o-v = 10 μm.
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Jobellisiaceae Réblová, Mycologia 100(6): 899 (2008)
Index Fungorum number: IF508692; Facesoffungi number: FoF03334; 8 species.
Saprobic on wood in terrestrial and freshwater habitats. Sexual morph: Ascomata perithecial,
basally immersed to superficial, astromatic, globose to subglobose, lageniform to obpyriform,
brown to black or yellowish, glabrous or slightly rugose, with a papilla or with upright neck.
Peridium three-layered, comprising cells of textura angularis or textura prismatica or textura
intricata, some with an orange, middle wall layer. Paraphyses numerous, septate. Asci 8-spored,
unitunicate, cylindrical to clavate, short pedicellate, with a J-, distinct, refractive, apical ring.
Ascospores uniseriate or overlapping uniseriate, oblong to ellipsoidal, fusoid to fusiform, straight or
slightly curved, reddish-brown or greenish-brown to brown, darker at the median septum, 1-septate,
with germ pores at one or both ends. Asexual morph: Undetermined (adapted from Réblová 2008
and Maharachchikumbura et al. 2016b).
Type genus – Jobellisia M.E. Barr
Notes – Jobellisiaceae was introduced by Réblová (2008) to accommodate a single genus
Jobellisia based on LSU sequence data in the Sordariomycetes incertae sedis. In the phylogenetic
analysis, Maharachchikumbura et al. (2015) introduced a new order Jobellisiales to accommodate
this family, which was accepted by Maharachchikumbura et al. (2016b). With the use of molecular
clock evidence, Jobellisiales fell in the ordinal time frame (146 MYA) (Hyde et al. 2017a).
However, Hongsanan et al. (2017) stated that the placement of this order is unstable as sometimes it
clustered with Pleurostomataceae.
Ecological and economic significance of Jobellisiaceae
Jobellisiaceae species are saprobic lignicolous taxa in terrestrial and freshwater habitats
(Maharachchikumbura et al. 2016b). They play an important role in nutrient cycling as
decomposers, especially on submerged woody debris or dry wood.
Genus included in Jobellisiaceae
Jobellisia M.E. Barr, Mycotaxon 46: 60 (1993)
Index Fungorum number: IF26310; 8 morphological species (Luo et al. 2019, Species
Fungorum 2020), 3 species with sequence data.
Type species – Jobellisia luteola (Ellis & Everh.) M.E. Barr
Notes – Jobellisia was introduced by Barr (1993) with two new combinations, J. luteola (type
species) and J. nicaraguensis. Sequences of three species are available in GenBank (J. fraterna, J.
guangdongensis and J. luteola), and have been used in phylogenetic analyses
(Maharachchikumbura et al. 2015, 2016b, Hongsanan et al. 2017). Jobellisia guangdongensis is
illustrated in this study (Fig. 131).
Juglanconidaceae Voglmayr & Jaklitsch, Persoonia 38: 142 (2017)
Index Fungorum number: IF819587; Facesoffungi number: FoF03489; 5 species.
Saprobic on dead corticated twigs and branches of Juglandaceae attached to trees. Sexual
morph: Pseudostromata comprising an inconspicuous ectostromatic disc which are erumpent from
bark. Central column beneath the disc, nearly conical. Perithecia surrounding the central column,
with nearly globose base, with long lateral ostioles that emerge at the margin or within an
ectostromatic disc. Asci 8-spored, clavate to fusoid, with a J-, apical ring. Ascospores overlapping
1–2-seriate, hyaline, broadly fusiform, bicelled, slightly curved, guttulate, with or without
gelatinous appendages. Asexual morph: Coelomycetous. Conidiomata acervular, with
ectostromatic disc and conical central column. Conidiophores aseptate or septate, smooth, hyaline
to brownish, occasionally branched. Conidiogenous cells annellidic. Conidia initially hyaline,
becoming brown with age, various in shape, pip-shaped, narrowly ellipsoid, elongate to
suballantoid, unicellular, often truncate with scar at the base, with gelatinous sheath and irregular
verrucae on the inner surface of the conidial wall (adapted from Voglmayr et al. 2017).
Type genus – Juglanconis Voglmayr & Jaklitsch
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Figure 131 – Jobellisia guangdongensis (Material examined – CHINA, Yunnan Province, on
submerged wood in a stream, 25 August 2019, G.N. Wang DQ09, MFLU 19-2827). a Appearance
of black ascoma on host. b Structure of peridium. c, e-i Asci. d Paraphyses. j, k Ascospores. Scale
bars: b = 40 μm, c, d, h, i = 30 μm, e = 10 μm, f, g = 20 μm, j, k = 5 μm.
Notes – Juglanconidaceae was erected by Voglmayr et al. (2017) to accommodate four
Melanconium species on hosts of Juglandinae, viz. three species (Juglanconis appendiculata, J.
juglandina, J. oblonga) on various Juglans species, and one species (J. pterocaryae) from
Pterocarya spp. (Voglmayr et al. 2019b). Du et al. (2017) introduced Melanosporellaceae to
accommodate Melanosporella typified by M. juglandium. It was shown that the strains Juglanconis
juglandina (MC1, ME16, ME22, ME23, MC3) from Austria and Spain and strains Melansporella
juglandium (CFCC 51727, CFCC 51728, CFCC 51729) from China clustered together in the
multigene phylogenetic analyses and share close morphological features (Du et al. 2017, Fan et al.
2018). Thus, Senanayake et al. (2018) treated Melanosporellaceae as a synonym of
Juglanconidaceae. Juglanconidaceae seems specific to Juglandaceae tree species and cause black
pustular dieback (Senanayake et al. 2018).
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Ecological and economic significance of Juglanconidaceae
All species of Juglanconis have been reported on Juglandaceae (walnut family), such as
Juglans ailanthifolia, J. cinerea, J. nigra, J. regia and Pterocarya rhoifolia. Juglanconis species
have a wide distribution across Austria, China, the Czech Republic, France, Germany, Greece,
Japan, Russia, Spain and the USA (Fan et al. 2018). Juglanconis juglandina and J. oblonga were
regarded as the main causal agent of canker and dieback disease of Juglans regia in China (Fan et
al. 2018).
Genus included in Juglanconidaceae
Juglanconis Voglmayr & Jaklitsch, Persoonia 38: 142 (2017)
Index Fungorum number: IF819582; 5 species with sequence data.
Type species – Juglanconis juglandina (Kunze) Voglmayr & Jaklitsch
Notes – Juglanconis was established by Voglmayr et al. (2017) to accommodate three
combined species including J. juglandina (WU 35965, neotype), J. oblonga (MBT374386,
lectotype), J. pterocaryae (TFM FPH2623, holotype) and a newly introduced species J.
appendiculata (WU 35954, holotype). These species were previously placed in Melanconium but
they are morphologically different from Melanconium sensu stricto in having irregular
ornamentation on the inner surface of the conidial wall (Voglmayr et al. 2017). Fresh collections
and updated morphological descriptions of Juglanconis juglandina and J. oblonga collected from
Juglans regia in China were also provided by Fan et al. (2018). Subsequently, two new species
Juglanconis japonica occurring on Pterocarya rhoifolia and J. pterocaryae occurring on
Pterocarya fraxinifolia were introduced to this genus (Voglmayr et al. 2019b). To date, five species
are accepted in Juglanconis and more fresh collections are needed (Voglmayr et al. 2019b).
Currently Juglanconis species have calM, histone, ITS, LSU, ms204, rpb1, rpb2, tef1 and tub2
gene sequence data available for multilocus analyses (Voglmayr et al. 2019b).
Figure 132 – Morphology of Juglanconis juglandina (Material examined – CHINA, Gansu
Province, Qingyang City, Shishe village, on twigs and branches of Juglans regia, 14 July 2013,
X.L. Fan, BJFC-S908). a, b Habit of acervuli on branches. c Transverse section through acervulus.
d Longitudinal section through acervulus. e, f Conidiophores, conidiogenous cells and conidia.
Annellides of conidiogenous cells are marked by black arrow in f. Scale bars: a-d = 0.5 mm, e, f =
20 μm.
Juncigenaceae E.B.G. Jones, Abdel-Wahab & K.L. Pang, Cryptog. Mycol. 35(2): 133 (2014)
Index Fungorum number: IF808177; Facesoffungi number: FoF01665; 8 species.
Saprobic intertidal wood, mangrove and herbaceous wood and roots, bark, leaves in marine
habitats. Sexual morph: Ascomata perithecial, globose, subglobose, ovoid to pyriform, immersed,
erumpent to superficial, subcoriaceous to coriaceous, olivaceous-brown, brown to dark-brown to
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black, hyaline to yellow-orange to reddish-brown, ostiolate, periphysate, papillate or hyaline to
apricot coloured long neck surrounded by dense brown, septate hyphae. Peridium comprising
several cell layers of ellipsoidal to subglobose cells forming textura angularis, textura
epidermoidea or both, or textura prismatica or textura globulosa. Paraphyses numerous, narrow,
branched or unbranched, persistent, connected to the apex and base of the peridium or
catenophyses. Asci 8-spored, unitunicate, thin-walled, persistent, clavate, cymbiform, cylindrical to
fusiform, short pedicellate, with or without apical ring. Ascospores 1-3 seriate, hyaline, ellipsoidal,
clavate to fusiform, unicellular, or 1–4-septate, with or without equatorial and polar or subpolar
appendages. Asexual morph: hyphomycetous. Hyphae septate, branched, hyaline to brown.
Conidiogenous cells non-specialized, short, lateral, solitary, helicoid, septate, and light to dark
brown. Conidia brown, single, helicoid, septate, constricted at the septa (adapted from AbdelWahab et al. 2010, Jones et al. 2014, Maharachchikumbura et al. 2015, Poli et al. 2019).
Type genus – Juncigena Kohlm., Volkm.-Kohlm. & O.E. Erikss.
Notes – Juncigenaceae was typified by Juncigena by Jones et al. (2014) and included the
genera Fulvocentrum, Marinokulati and Moheitospora. They formed a phylogenetically stable
monophyletic clade in a SSU and LSU based phylogeny. Juncigenaceae is sister to
Etheirophoraceae, Falcocladiaceae and Torpedosporaceae in the subclass Hypocreomycetidae.
Jones et al. (2014) introduced Fulvocentrum to accommodate F. aegyptiacum and F.
clavatisporium, which were previously introduced under Swampomyces sensu stricto. Likewise, the
marine ascomycete Chaetosphaeria chaetosa did not group in Chaetosphaeria sensu stricto
(Chaetosphaeriales) and was transferred to a new genus Marinokulati (Jones et al. 2014). In Jones
et al. (2014), Juncigenaceae was placed in the subclass Hypocreomycetidae, order incertae sedis.
This was supported by Maharachchikumbura et al. (2015). Jones et al. (2015) placed
Etheirophoraceae, Juncigenaceae and Torpedosporaceae in Torpedosporales. Abdel-Wahab et al.
(2018) introduced Khaleijomyces as sister to Juncigena and Poli et al. (2019) introduced
Elbamycella as a separate lineage in Juncigenaceae.
Ecological and economic significance of Juncigenaceae
Members in this family have mainly been found on driftwood collected from the intertidal
zone, submerged sea grasses and brown alga (Abdel-Wahab et al. 2010, Poli et al. 2019). They are
likely saprobes involved in nutrient cycling.
Genera included in Juncigenaceae
Elbamycella A. Poli, E. Bovio, V. Prigione & G.C. Varese, MycoKeys 55: 21 (2019)
Index Fungorum number: IF830648; 1 species with sequence data.
Type species – Elbamycella rosea A. Poli, E. Bovio, V. Prigione & G.C. Varese
Notes – This genus was recorded from submerged marine environments (sea grass Posidonia
oceanica and with the brown alga Padina pavonica) (Poli et al. 2019). The genus is well-supported
by morphology and phylogeny. This is the first record of Juncigenaceae found in submerged
environment (Poli et al. 2019). The genus is morphologically distinct from other genera with
subpolar appendaged spores (Poli et al. 2019).
Fulvocentrum E.B.G. Jones & Abdel-Wahab, Cryptog. Mycol. 35(2): 131 (2014)
Index Fungorum number: IF808181; 3 species with sequence data.
Type species – Fulvocentrum aegyptiacum (Abdel-Wahab et al.) E.B.G. Jones & AbdelWahab
Notes – Found on decayed attached branches of Avicennia marina in the intertidal zone. The
genus has immersed, dark brown, ostiolate ascomata with numerous unbranched paraphyses in a
gel and short pedicellate, apically thickened asci, with 3-septate, ellipsoidal, hyaline ascospores
(Jones et al. 2014). The genus is distinct in having, short pedicellate, apically thickened asci with 3septate, ellipsoidal hyaline ascospores (Jones et al. 2014). In this entry, Fulvocentrum aegyptiacum
is illustrated to demonstrate the sexual morph. Fulvocentrum aegyptiacus is commonly found in
658
intertidal wood from Avicennia marina in Red sea mangroves Egyptian coast (Abdel-Wahab 2005)
and Arabian Gulf mangroves of the Saudi Arabia coast (Abdel-Wahab 2005).
Figure 133 – Juglanconis juglandina (a-j), Juglanconis appendiculata (k-n) redrawn from
Voglmayr et al. (2017). a, b Ectostromatic discs. c Transverse sections below ectostromatic disc. d
Vertical section through pseudostroma, showing perithecia and entostroma. e Apical ring of asci. f
Asci. g-n Ascospores. Scale bars: a = 1 mm, b, c = 0.5 mm, f =20 µm, e, g-n = 10 µm.
659
Figure 134 – Juglanconis juglandina (a-h), Juglanconis oblonga (i, j), Juglanconis pterocaryae (k)
redrawn from Voglmayr et al. (2017). a Conidiomata on the host. b, c Transverse and vertical
sections of conidiomata. d, e Conidiophore with conidia. f-k Conidia. Scale bars: a = 1 mm, b, c =
0.5 mm, d-k = 10 µm.
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Figure 135 – Fulvocentrum aegyptiacus (Material examined – EGYPT, Red Sea Coast, Safaga
mangrove, on decayed attached branches of Avicennia marina in the intertidal zone, January 1999,
M.A. Abdel- Wahab, IMI 386146, holotype). a Herbarium packet. b Herbarium material. c
Ascomata on host. d Section through ascoma. e Peridium. f Paraphyses. g-j Asci. k-m Ascospores.
Scale bars: c, d = 100 µm, f-j = 50 µm, e, k-m = 10 µm.
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Juncigena Kohlm., Volkm.-Kohlm. & O.E. Erikss., Bot. Mar. 40(4): 291 (1997)
Index Fungorum number: IF27750; 2 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Juncigena adarca Kohlm., Volkm.-Kohlm. & O. E. Erikss.
Notes – Juncigena Kohlmeyer et al. (1997). was initially included under Magnaporthaceae by
Eriksson (1999). Further protein coding and rDNA based phylogeny by Schoch et al. (2006)
revealed Juncigena adarca clusters with Swampomyces sp. in Coronophorales. Jones et al. (2009)
included Juncigena under Hypocreales incertae sedis. With resolving the TBM
(Torpedospora/Bertia/Melanospora) clade with terrestrial Falcocladium, Jones et al. (2014)
referred Juncigena under Juncigenaceae. Kohlmeyer et al. (1997) revealed that Cirrenalia adarca
was the asexual morph of Juncigena adarca. Abdel-Wahab et al. (2010) transferred C. adarca to
Moheitospora based on phylogenetic and morphological evidence. The genus currently comprises
Juncigena adarca and J. fruticosae. Juncigena is distinctive from other genera in having the
cylindrical, short pedicellate asci with an apical ring (Jones et al. 2014). Réblová et al. (2016b)
referred Moheitospora fruticosae to Juncigena as the two genera are congeneric, and Juncigena has
priority.
Figure 136 – Juncigena fruticosae (Material examined – EGYPT, Mediterranean coast, on decayed
driftwood in the intertidal zone, June 2006, M. A. Abdel-Wahab, IMI 391650, holotype). a
Herbarium packet. b Herbarium material. c Hyphae, conidiophores and conidia on host. d
Conidiophores. f-k Conidiophores and variously-shaped mature conidia. Scale bars: c = 200 µm, d
= 100 µm, e, f = 50 µm, g-k = 20 µm.
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Khaleijomyces M.A. Abdel-Wahab, Phytotaxa 340(3): 280 (2018)
Index Fungorum number: IF823891; 1 species with sequence data.
Type species – Khaleijomyces marinus M.A. Abdel-Wahab
Notes – The genus differs from other genera in the family, in having ostiolar necks with
periphyses and not having apical rings or thickenings in the asci (Abdel-Wahab et al. 2018). The
ascomata differ in being yellow-orange to reddish brown. Species are found on intertidal driftwood
(Abdel-Wahab et al. 2018).
Marinokulati E.B.G. Jones & K.L. Pang, Cryptog. Mycol. 35(2): 132 (2014)
Index Fungorum number: IF808184; 1 species with sequence data.
Type species – Marinokulati chaetosa (Kohlm.) E.B.G. Jones & K.L. Pang
Notes – While resolving the TBM (Torpedospora/Bertia/Melanospora) clade with the marine
species Chaetosphaeria chaetosa and terrestrial Falcocladium, Jones et al. (2014) revealed that the
former species was significantly different from Chaetosphaeria and Juncigena. Hence,
Marinokulati was introduced to accommodate the previously known Chaetosphaeria chaetosa. The
genus is characterised by apically thickened asci and 3-septate, hyaline, fusiform to ellipsoidal
ascospores with polar and equatorial appendages.
Junewangiaceae J.W. Xia & X.G. Zhang, Scientific Reports 7 (no. 7888): 12 (2017)
Index Fungorum number: IF818897; Facesoffungi number: FoF06602; 9 species.
Saprobic on terrestrial or freshwater habitats. Sexual morph: Ascomata scattered or solitary,
immersed with neck superficial, uniloculate, globose, subglobose or ellipsoidal, glabrous, brown,
coriaceous. Ostiole long, cylindrical, periphysate. Peridium comprising several layers of brown,
thick-walled, compressed cells of textura porrecta. Paraphyses numerous, cylindrical, hyaline,
septate, unbranched, persistent. Asci 8-spored, unitunicate, long cylindrical, with a tapering pedicel,
apically rounded, with an indistinct, J-, apical ring. Ascospores fusiform, hyaline, 3-septate, with
filamentous bipolar appendages. Asexual morph: Hyphomycetous. Colonies on natural substrate
effuse, brown. Mycelia partly superficial, partly immersed, composed of hyaline to brown,
branched, septate hyphae. Conidiophores micronematous or macronematous, when
macronematous, erect, cylindrical, unbranched, straight or slightly flexuous, thick-walled, septate,
brown, paler towards the apex. Conidiogenous cells blastic, integrated, terminal, cylindrical,
subhyaline to brown, smooth or verrucose. Conidia acrogenous, solitary, dry, smooth, ellipsoidal,
oval, clavate or obovoid, brown to dark brown (adapted from Xia et al. (2017).
Type genus – Junewangia W.A. Baker & Morgan-Jones
Notes – Junewangiaceae was established by Xia et al. (2017) to accommodate a single genus,
Junewangia, based on combined of LSU, SSU, ITS and tub2 sequence data. Although
Dictyosporella and Junewangia formed a monophyletic clade with strong support, Song et al.
(2018) did not transfer Dictyosporella to Junewangiaceae. Luo et al. (2019) accepted three genera,
i.e. Dictyosporella, Junewangia and Sporidesmiella in Junewangiaceae in their comprehensive
study of freshwater Sordariomycetes. However, in our phylogenetic analyses, Acrodictyaceae and
Junewangiaceae and Sporidesmiella formed a monophyletic clade, while Acrodictyaceae and
Junewangiaceae are sister groups. Therefore, we suggest Junewangiaceae contains Dictyosporella
and Junewangia, while Sporidesmiella is treated as Diaporthomycetidae genus incertae sedis.
Ecological and economic significance of Junewangiaceae
Junewangiaceae species are mostly saprobic on decaying wood in both aquatic and terrestrial
habitats. The main role of Junewangiaceae species in the ecosystem is participating in nutrient
cycling as decomposers and recyclers. Thus, they are important for ecological balance.
Genera included in Junewangiaceae
Dictyosporella Abdel-Aziz, Fungal Divers. 75: 143 (2015)
Index Fungorum number: IF551480; 3 species with sequence data.
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Type species – Dictyosporella aquatica Abdel-Aziz
Notes – Dictyosporella was introduced by Ariyawansa et al. (2015) to accommodate an
asexual morph species, D. aquatica, which is characterized by micronematous conidiophores and
brown to black, muriform conidia. Based on phylogenetic analyses of LSU sequence data,
Dictyosporella was assigned to Annulatascaceae (Ariyawansa et al. 2015). Zhang et al. (2017a)
introduced a sexual morph species, D. thailandensis, and treated Dictyosporella as
Diaporthomycetidae, genera incertae sedis. Luo et al. (2019) transferred Dictyosporella to
Junewangiaceae based on multi-gene phylogenetic analyses.
Junewangia W.A. Baker & Morgan-Jones, Mycotaxon 81: 307 (2002)
Index Fungorum number: IF28602; 6 morphological species (Species Fungorum 2020), 4
species with sequence data (Xia et al. 2017, Song et al. 2018).
Type species – Junewangia sphaerospora W.A. Baker & Morgan-Jones
Notes – Baker et al. (2002) introduced Junewangia with J. sphaerospora as the type species.
At the same time, they transferred four Acrodictys species which have subglobose, uniformly
pigmented conidia with angular or oblique septa to Junewangia. Junewangia is morphologically
similar with Acrodictys. However, conidia of Acrodictys is conspicuous, cuneiform or funnelshaped basal cells, while the basal cells of Junewangia are inconspicuous (Xia et al. 2017).
Kathistaceae Malloch & M. Blackw., Can. J. Bot. 68(8): 1719 (1990)
Index Fungorum number: IF81991; Facesoffungi number: FoF06874; 6 species.
Saprobic on herbivore dung and insects. Sexual morph: Ascomata globose to subglobose,
with a long necks composed of parallel hyphae, straight curved ostiolar setae, with sphaerical
spore-bearing structures (sporidiomata), cylindrical unicellular extension present. Peridium thinwalled, pseudoparenchymatous, of cells of textura angularis in surface view and thick-walled in
cross section. Paraphyses lacking. Asci 8-spored, unitunicate, ellipsoidal to fusoid, thin-walled,
evanescent at maturity, arranged in a basal fascicle. Ascospores clavate to falcate, hyaline or pale
brown, 0–1 or transversely multi septate. Asexual morph: Conidiomata superficially similar to the
ascomata, sphaerical, hyaline, without hyphal attachments, single wall layer thick at base, with or
without ostiole, when present ostiole comprising with long neck. Conidiomatal wall composed of
flattened cells of textura angularis. Conidiogenous cells enteroblastic, phialidic, hyaline, and
smooth. Conidia sphaerical or cylindrical, produced in the centre of the conidiomata, hyaline,
smooth, 0–1 or multi septate, escaping in a continuous chain through the ostiolar neck (adapted
from Maharachchikumbura et al. 2016b).
Type genus – Kathistes Malloch & M. Blackw.
Notes – Malloch & Blackwell (1990) established Kathistaceae to accommodate Kathistes
with the type species K. calyculata, and two more species, K. fimbriata and K. analemmoides.
Kathistaceae has sufficient taxonomic distance from Pyxidiophoraceae. Therefore, it was reported
as a family in Ophiostomatales (Malloch & Blackwell 1990). Maharachchikumbura et al. (2016b)
provided an updated outline of Kathistaceae with the genera Kathistes, Mattirolella, and
Termitariopsis.
Ecological and economic significance of Kathistaceae
Kathistaceae species are saprobic on herbivore dung and insects (Malloch & Blackwell
1990).
Genera included in Kathistaceae
Kathistes Malloch & M. Blackw., Can. J. Bot. 68(8): 1712 (1990)
Index Fungorum number: IF25513; 3 morphological species (Species Fungorum 2020), 2
species with sequence data.
Type species – Kathistes calyculata Malloch & M. Blackw.
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Figure 137 – Dictyosporella thailandensis (Material examined – THAILAND, Prachuap Khiri
Khan, on submerged bamboo in a small river, 30 July 2015, W. Dong, MFLU 15-2706, holotype).
a, b Appearance of necks on host. c Vertical section of ascoma. d Peridium. e Paraphyses. f, g Asci.
h Ascus with a apical ring. i Ascospore mounted in India ink. j-o Ascospores. Scale bars: a, b = 50
μm, c = 25 μm, d, e, j-o = 10 μm, f, g = 20 μm, h = 5 μm.
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Figure 138 – Junewangia sphaerospora (redrawn from Baker et al. 2002). This drawing shows the
mycelia, conidiophores, conidiogenous cells and conidia of Junewangia sphaerospora. Scale bars:
15 μm.
Notes – Kathistes was introduced as the type genus in Kathistaceae by Malloch & Blackwell
(1990), which was originally described with three species (K. analemmoides, K. calyculata, K.
fimbriata). The genus is characterized by long-necked, brown to dark ascomata, transparent at the
base, and germinating ascospores produced by budding yeast-like cells (Malloch & Blackwell
1990).
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Figure 139 – Kathistes spp. (a-i: Kathistes calyculata, j, k: K. fimbriata, l-r: K. analemmoides,
redrawn from Malloch & Blackwell 1990). a Ascoma with dark neck and hyaline base. b Asci. c
Ascospores. d Ascospores after 4 h on agar Germ tubes have developed by this time. e-g
Ascospores after 19, 27, and 42 h on agar, respectively. Yeast cells have begun to develop. h Yeast
cells after several days. i Sporidioma showing single-celled wall at base and sporidiolae filling
hollow darkened neck cell. j Ascoma with dark neck and hyaline base. k Three ascospores
illustrating differences in septation, size, and pigmentation. l Ascomata. m Asci. n Ascospores from
an extruded spore mass. One ascospore has begun producing yeast cells. o-q Three ascospores,
each drawn at 2, 3, 4, and 18 h after being placed on agar r Sporidioma showing single wall layer of
base and darkened hollow neck cell.
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Mattirolella S. Colla, Boll. Lab. Zool. Portici 22: 44 (1929)
Index Fungorum number: IF8867; 2 morphological species.
Type species – Mattirolella silvestrii S. Colla
Notes – Mattirolella was described by Colla (1929) with the type species M. silvestrii parasite
on integument of the termite Rhinotermes marginales in British Guiana. Khan & Kimbrough
(1974) introduced Mattirolella crustosa, a species found on Isoptera (termites) in Panama.
Termitariopsis M. Blackw., Samson & Kimbr., Mycotaxon 12(1): 98 (1980)
Index Fungorum number: IF10184; 1 morphological species.
Type species – Termitariopsis cavernosa M. Blackw., Samson & Kimbr.
Notes – Termitariopsis was described by Blackwell et al. (1980) as a monotypic genus and is
a species found on insects (Neivamyrmex opacithorax) in Kansas. Although Wijayawardene et al.
(2017a) listed Termitariopsis as Ascomcota genera incertae sedis, we maintain Mattirolella and
Termitariopsis in Kathistaceae based on similar morphology (Blackwell et al. 1980).
Koralionastetaceae Kohlm. & Volkm.-Kohlm., Mycologia 79(5): 764 (1987)
Index Fungorum number: IF82025; Facesoffungi number: FoF01391; 13 species.
Saprobic on coralline covered rocks, sponges or on algae, found in marine habitats. Sexual
morph: Ascomata black, subglobose, ovoid or ellipsoidal, superficial, ostiolate, papillate or
epapillate, periphysate, subiculate, or without subiculum. Paraphyses simple, septate. Asci 8spored, unitunicate, clavate to ellipsoidal or fusiform, pedicellate, deliquescent. Ascospores
overlapping, hyaline, ellipsoidal to fusiform, multi-septate near the apices, or filiform and evenly
multi-septate, thick or thin walled, germinating apically, lacking appendages. Reproductive
structures: Spermatia enteroblastic, subglobose.
Type genus – Koralionastes Kohlm. & Volkm.-Kohlm.
Notes – Koralionastetaceae currently comprises 13 species (five Koralionastes species and
eight Pontogeneia species) (Wijayawardene et al. 2017a). The family was introduced by
Kohlmeyer & Volkmann-Kohlmeyer (1987) to include a single genus, Koralionastes. Pontogeneia
was introduced by Kohlmeyer (1975) for five perithecial ascomycetes. Koralionastetaceae is
characterized by thick-walled ascospores that germinate into hyphae bearing phialidic antheridia
with enteroblastic spermatia. Earlier the ordinal and higher classification of Koralionastetaceae was
unresolved and Koralionastes was placed under Ascomycota genera incertae sedis (Eriksson 2006).
Based on a phylogenetic study with SSU and LSU sequence data, the genera Koralionastes and
Pontogeneia were assigned to Koralionastetales which is a sister group to Lulworthiales (Campbell
et al. 2009). The subclass Lulworthiomycetidae was introduced to accommodate Lulworthiales and
Koralionastetales based on combined sequence data by Maharachchikumbura et al. (2015).
Ecological and economic significance of Koralionastetaceae
Koralionastetaceae members are coral and algal inhabiting fungi and play a major role in
heterotrophic conversion of reef biomass to nutrients (Raghukumar & Ravindran 2012).
Koralionastes have been reported to have a unique association with crustaceous sponges,
developing their ascomata on or within these hosts (Kohlmeyer & Volkmann-Kohlmeyer 1990a;
Wang 2006) and is the only marine fungal group which have been identified in marine sponges
(Wang et al. 2008). Pontogeneia are algae inhabiting fungi. For an example, Pontogeneia rostrata
are known from red algae (Momota & Nakaoka 2017). Furthermore, studies by Yu & Suh (2011)
showed that Pontogeneia rostrata plays an important role in the sandy shore ecosystem as a
trophic link between primary producers and higher consumers.
Genera included in Koralionastetaceae
Koralionastes Kohlm. & Volkm.-Kohlm., Mycologia 79(5): 765 (1987)
Index Fungorum number: IF25184; 5 morphological species (Wijayawardene et al. 2017a;
Species Fungorum 2020), 1 species with sequence data.
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Figure 140 – Koralionastes ovalis (Material examined – BELIZE, back reef of South water Cay,
sub tidal coral slab (0.2-0.5 m deep), November 1986, J. Kohlmeyer; NY 1317811, NY 1317823,
NY 1317828, NY 1317838, slides from holotype), Pontogeneia padinae (Material examined –
Intertidal zone at the Marine Laboratory of the University of Arizona, on living plants near the base
among rhizoids of the holdfast, 22 Jun 1974, J. J. Kohlmeyer, NY 985934, holotype). a Microslides
of Koralionastes ovalis. b Ascomata of Pontogeneia padinae on the base of an alga. c, d Section
through ascoma. e Ascoma neck region of Koralionastes ovalis. f, g Peridium. h Ascus of
Koralionastes ovalis. i Paraphyses of Koralionastes ovalis. j, k Ascospores of Koralionastes ovalis.
l, m Ascospores of Pontogeneia padinae. Scale bars: c, d = 200 μm, h = 100 μm, e = 50 μm, f, g, l,
m = 20 µm, f-i = 10 µm.
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Type species – Koralionastes ovalis Kohlm. & Volkm.-Kohlm.
Notes – Koralionastes was introduced to accommodate Koralionastes angustus, K. ellipticus
and K. ovalis (the type species) by Kohlmeyer & Volkmann-Kohlmeyer (1987). Kohlmeyer &
Volkmann-Kohlmeyer (1990a) introduced K. giganteus, and K. violaceus found on dead coralline
covered rocks among crustose sponges and their ascospores have double walls and are ellipsoidal
or fusiform (Campbell et al. 2009).
Pontogeneia Kohlm., Bot. Jb. 96(1–4): 201 (1975)
Index Fungorum number: IF4335; 8 morphological species (Wijayawardene et al. 2017a;
Species Fungorum 2020), 2 species with sequence data.
Type species – Pontogeneia padinae Kohlm.
Notes – Pontogeneia was introduced by Kohlmeyer (1975) to include five perithecial
ascomycetes parasitizing marine algae, belonging to the Chlorophyta, Phaeophyta or Rhodophyta
(Kohlmeyer & Kohlmeyer 1979, Kohlmeyer & Demoulin 1981, Momota & Nakaoka 2017).
Lamproconiaceae Norph., T.C. Wen & K.D. Hyde, Phytotaxa 270(2): 94 (2016)
Index Fungorum number: IF552187; Facesoffungi number: FoF02248; 14 species.
Pathogenic and saprobic on dead herbaceous branches. Sexual morph: Stromata
prosenchymatous around perithecia, delimited externally by greenish-blackened dense
pseudoparenchymatous zone, interior whitish, composed of interwoven hyphae mixed with
substrate cells, 3–5 ascomata in a stromata. Ascomata perithecial, small, aggregated, scattered,
globose to subglobose, light brown to dark brown, coriaceous, ostiolate, papillate. Papilla
converging and erumpent through stroma surface as single, large opening, wide at the top,
narrowing towards the base, dark brown region around base of papilla. Peridium comprises light
brown, compressed, cells of textura angularis. Asci 8-spored, unitunicate, cylindrical, short
pedicellate, with a J-, apical ring. Ascospores uniseriate, broadly ellipsoid, 1-septate, not or lightly
constricted at the septa, hyaline, smooth-walled. Asexual morph: Conidiomata pycnidial, solitary,
partly immersed in host tissue, uniloculate, multilocular or convoluted, dark blue (Lamproconium),
dark blackish brown (Hercospora), erumpent in the centre. Pycnidium thick-walled, thin at inner
layer, hyaline (Lamproconium), dark brown (Hercospora), comprising wall cells of textura
angularis (Lamproconium) or textura intricate (Hercospora). Ostiole absent, dehiscence irregular.
Paraphyses interspersed within conidiophores. Conidiophores filiform or cylindrical, pale-bluish or
hyaline, septate, branched, smooth-walled, formed at the base of conidiomatal wall. Conidiogenous
cells holoblastic, cylindrical to subcylindrical, each forming a single conidium at the conidiophore
apex, or annellidic, colourless to olivaceous, smooth-walled. Conidia fusiform, ellipsoid, thickwalled, with granular, contents granular, aseptate, bluish to glistening dark blue (Lamproconium),
hyaline (Hercospora), smooth-walled, produced in mucilage but without a distinct mucilaginous
envelope or appendage (adapted from Norphanphoun et al. 2016).
Type genus – Lamproconium (Grove) Grove
Notes – Lamproconiaceae was established by Norphanphoun et al. (2016) to accommodate
Lamproconium and Hercospora, based on morphology and phylogenetic analyses.
Lamproconiaceae forms a robust clade basal to Sydowiellaceae and Stilbosporaceae in the
combined ITS and LSU phylogeny and the conidia differ from those of Sydowiellaceae and
Stilbosporaceae (Norphanphoun et al. 2016, Senanayake et al. 2017b).
Ecological and economic significance of Lamproconiaceae
Species in Lamproconiaceae are pathogens and saprobes on branches and bark of various
trees (Norphanphoun et al. 2016). Lamproconium desmazieresi causes canker on branches or twigs
of lime trees (Tilia spp.).
Genera included in Lamproconiaceae
Hercospora Fr., Syst. orb. veg. (Lundae) 1: 119 (1825)
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Index Fungorum number: IF2301; 13 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Hercospora tiliae (Pers.) Tul. & C. Tul.
Notes – Hercospora was introduced by Fries in 1825 to accommodate Sphaeria tiliae and S.
atrovirens. Tulasne & Tulasne (1863) accepted Hercospora tiliae (= S. atrovirens) as the type
species of Hercospora. The characters of the genus were described in Petrak (1938) and Sutton
(1980). Phylogenetic studies based on LSU sequence data, placed H. tiliae in Diaporthales, genera
incertae sedis, where it grouped with Melanconis desmazierii (Castlebury et al. 2002, Rossman et
al. 2007, Voglmayr et al. 2012, Voglmayr & Jaklitsch 2014). Norphanphoun et al. (2016) placed
Hercospora in a new family Lamproconiaceae based on sequence data which clustered with
Melanconis desmazieri, a species synonymized under Lamproconium desmazierii.
Figure 141 – Hercospora tiliae (redrawn from Sutton 1980 and Norphanphoun et al. 2016). a
Ascomata. b, c Asci. d Ascospores. e Conidioma. f Conidiophores and developing conidia. g
Conidia. Scale bars: a = 200 μm, b, c = 40 μm, d, f = 20 μm, e = 100 μm, g = 10 μm.
Lamproconium (Grove) Grove, British Stem- and Leaf-Fungi (Coelomycetes) (Cambridge) 2: 321
(1937)
Index Fungorum number: IF8703; 1 species with sequence data.
Type species – Lamproconium desmazieri (Berk. & Broome) Grove
Notes – Lamproconium was introduced by Grove (1918) as a section of Melanconium which
was represented by M. desmazieri. Melanconium desmazieri was reported as the asexual morph of
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Melanconis desmazieri, from Tilia sp. (Petrak 1938). Grove (1937) re-circumscribed the species
and found that M. desmazieri differed from the type species in having bluish to glistening dark
blue, not brownish-black, 1-septate conidia. Therefore, Grove (1937) introduced Lamproconium as
a new genus (Sutton 1980) and it was placed in Diaporthales genera incertae sedis by Cannon &
Minter (2014). Based on phylogenetic study and fresh collections of L. desmazieri (Norphanphoun
et al. 2016) introduced Lamproconiaceae as new family for accommodate Lamproconium and
Hescospora.
Figure 142 – Lamproconium desmazieresi (Material examined – RUSSIA, Rostov region:
Krasnosulinsky district, Donskoye forestry, artificial forest, on dead branches of Tilia cordata Mill.
(Tiliaceae), 21 May 2014, T. Bulgakov, MFLU 14-0780, reference specimen). a, b Conidiomata on
host. c Cross section of the conidioma. d Conidiogenous cells with attached. e-i Immature
conidium. j Mature conidia. Scale bars: b = 1 mm, c = 500 µm, d = 40 µm, e-j = 20 µm.
Lasiosphaeriaceae Nannf., Nova Acta R. Soc. Scient. upsal., Ser. 4 8(no. 2): 50 (1932)
Index Fungorum number: IF80930; Facesoffungi number: FoF01145; 315 species.
Saprobic on wood, rotting vegetation, soil, dung of the herbivore, in freshwater, marine or
terrestrial habitats, many coprophilous. Sexual morph: Ascomata perithecial or cleistothecial,
gregarious or scattered, yellow, brown to black, solitary, superficial, erumpent or immersed,
globose, subglobose to ovoid, carbonaceous, coriaceous or membranaceous, ornamented,
tuberculate or smooth, with setae or hair, papillate or indistinct or absent, ostiolate, when present
periphysate, the apex collapsing when dry. Peridium thick, composed of two layers, outer layer
comprising brown cells of textura angularis or globulosa, carbonaceous, coriaceous or
membranaceous; inner layer comprising hyaline cells of textura prismatica or porrecta, thin,
membranaceous. Paraphyses numerous, hyaline, septate, filiform or cylindrical. Asci 4- or 8spored, unitunicate, thin or thick-walled, cylindrical to clavate, pedicellate, with a J-, apical ring.
Ascospores uni-seriate to irregular, hyaline, brown or black, allantoid, clavate, cylindrical, ellipsoid
to dumbbell-like, curved or not, concolorous or versicolorous, ornamented or smooth-walled, with
or without germ pore, appendage present or absent, with or without guttules. Asexual morph:
Hyphomycetous. Conidiophores macronematous or mononematous, scattered or gregarious, brown,
straight, septate, branched, smooth-walled. Conidiogenous cells enteroblastic or holoblastic,
phialidic, hyaline to dark brown, subglobose to ampulliform, proliferating percurrently, with
collarette. Conidia solitary, globose, subglobose to cylindrical, hyaline to brown, smooth, aseptate
(adapted from Maharachchikumbura et al. 2016b).
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Type genus – Lasiosphaeria Ces. & De Not.
Notes – Lasiosphaeriaceae has black ascomata and cylindrical to clavate asci with brown to
hyaline ascospores, and is typified by Lasiosphaeria (Cesati & De Notaris 1863). Wijayawardene
et al. (2018a) accepted 32 genera in Lasiosphaeriaceae, but the family is in need of revision with
DNA sequence data analyses. Many of the genera included in this family lack sequence data and
require confirmation with fresh collections and phylogenetic analyses. Species of
Lasiosphaeriaceae cluster in several clades within Sordariales (Huhndorf et al. 2004b). Wang et al.
(2019a) made progress towards resolving the family and proposed Cladorrhinum, Podospora and
Triangularia in Podosporaceae based on multi-gene analysis and the family is sister to
Chaetomiaceae. In this study, the lasiosphaeriaceous complex is composed by Lasiosphaeriaceae
sensu stricto (Lasiosphaeriaceae II) and Lasiosphaeriaceae sensu lato (Lasiosphaeriaceae I and III)
(Fig. 23).
Ecological and economic significance of Lasiosphaeriaceae
Species of Lasiosphaeriaceae are found worldwide, usually as saprobes on herbs and woody
plants and on dung, in freshwater, marine or terrestrial habitats (Shearer 1989, Guarro et al. 1996,
Huhndorf et al. 2004b, Kruys et al. 2015, Maharachchikumbura et al. 2016b). A few species are
model organisms for genetic research. Podospora anserina has been extensively studied as an
experimental model, viz. mitochondrial mutants, protein products and genome sequences (Stahl et
al. 1978, Cummings et al. 1979, Coustou et al. 1997, Espagne et al. 2008, Scheckhuber & Osiewacz
2008).
Genera included in Lasiosphaeriaceae
Anopodium N. Lundq., Bot. Notiser 117: 356 (1964)
Index Fungorum number: IF211; 2 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Anopodium ampullaceum N. Lundq.
Notes – Anopodium was introduced based on A. ampullaceum (type species) and A. epile. The
genus is characterized by subglobose ascomata, clavate asci and brown ascospores with a pedicel at
one end (Lundqvist 1964a). The genus is phylogenetically related to Cercophora sulphurella with
good support (Kruys et al. 2015).
Apiosordaria Arx & W. Gams, Nova Hedwigia 13: 201 (1967)
Index Fungorum number: IF262; 24 morphological species (Species Fungorum 2020), 11
species with sequence data.
Type species – Apiosordaria verruculosa (C.N. Jensen) Arx & W. Gams
Notes – Apiosordaria is characterized by pyriform ascomata, cylindrical asci and ascospores
with verrucose or pitted walls (von Arx & Gams 1966, Guarro et al. 1984). Echinopodospora and
Lacunospora were considered as synonyms of Apiosordaria based on similar characters, such as
verrucose or pitted ascospores (Krug et al. 1983, Stchigel et al. 2000).
Apodospora Cain & J.H. Mirza, Can. J. Bot. 48(5): 891 (1970)
Index Fungorum number: IF275; 6 morphological species (Species Fungorum 2020), 3
species with sequence data.
Type species – Apodospora simulans Cain & J.H. Mirza
Notes – Apodospora simulans has ellipsoid to subglobose ascomata, cylindrical asci and
verrucose ascospores with a gelatinous sheath. The members of Apodospora nested in
Lasiosphaeriaceae sensu lato based on multi-gene analysis (Wang et al. 2019a; Fig. 23).
Apodus Malloch & Cain, Can. J. Bot. 49(6): 872 (1971)
Index Fungorum number: IF277; 2 species with sequence data.
Type species – Apodus deciduus Malloch & Cain
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Notes – Apodus deciduus has cleistothecial ascomata, clavate asci and brown ascospores. Cai
et al. (2006c) proposed to transfer this genus to Lasiosphaeriaceae from Sordariaceae based on
phylogenetic analysis.
Arnium Nitschke, Bot. Ztg. 31: 450 (1873)
Index Fungorum number: IF313; 29 morphological species (Species Fungorum 2020), 9
species with sequence data.
Type species – Arnium lanuginosum Nitschke
Notes – Arnium has pyriform to ellipsoid ascomata, cylindrical asci and oblong to oval
ascospores (Nitschke 1873, Kruys et al. 2015). Kruys et al. (2015) found that species of Arnium
were dispersed into several groups and interspersed in Lasiosphaeriaceae sensu lato (Fig. 23).
Bellojisia Réblová, Mycologia 100(6): 897 (2008)
Index Fungorum number: IF508691; 1 species with sequence data.
Type species – Bellojisia rhynchostoma (Höhn.) Réblová
Notes – The monotypic Bellojisia is characterized by brown to black ascomata with a long
neck, cylindrical asci and reniform to navicular ascospores and phylogenetic analysis of LSU
sequences revealed that this genus is related to Anopodium within Lasiosphaeriaceae sensu stricto
(Réblová 2008; Fig. 23).
Biconiosporella Schaumann, Veröff. Inst. Meeresf. Bremerhaven 14(1): 24 (1972)
Index Fungorum number: IF568; 1 morphological species.
Type species – Biconiosporella corniculata Schaumann
Notes – Biconiosporella corniculata is characterized by membranaceous ascomata,
cylindrical asci and broadly fusiform ascospores with a tubercule at the center (Schaumann 1972).
Bombardia (Fr.) P. Karst., Bidr. Känn. Finl. Nat. Folk 23: 20 (1873)
Index Fungorum number: IF616; 15 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Bombardia bombarda (Batsch) J. Schröt.
Notes – Miller & Huhndorf (2005) reported that this genus is similar to Bombardioidea based
on the bombardioid peridium and these species were also related in phylogenetic analysis.
Bombardioidea C. Moreau, Symb. bot. upsal. 20(no. 1): 274 (1972)
Index Fungorum number: IF619; 4 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Bombardioidea bombardioides (Auersw.) C. Moreau
Notes – Bombardioidea bombardioides is characterized by coriaceous ascomata, cylindrical
asci and ellipsoid to ovoid ascospores with a gelatinous sheath (Krug & Scott 1994). Huhndorf et
al. (2004b) accepted Bombardioidea as member of Lasiosphaeriaceae based on the molecular
analysis.
Camptosphaeria Fuckel, Jb. nassau. Ver. Naturk. 23-24: 140 (1870)
Index Fungorum number: IF781; 4 morphological species (Species Fungorum 2020).
Type species – Camptosphaeria sulphurea Fuckel
Notes – Huhndorf et al. (2004b) accepted this genus in Lasiosphaeriaceae based on the
cylindrical asci with a subapical globule similar to Lasiosphaeria.
Cercophora Fuckel, Jb. nassau. Ver. Naturk. 23-24: 244 (1870)
Index Fungorum number: IF906; 63 morphological species (Species Fungorum 2020), 30
species with sequence data.
Type species – Cercophora mirabilis Fuckel
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Notes – Cercophora has globose to subglobose ascomata, cylindrical asci with a subapical
globule and oblong ascospores with appendages at each end (Fuckel 1870). Members of
Cercophora nested in the lasiosphaeriaceous complex based on multi-gene analysis (Huhndorf et
al. 2004b, Kruys et al. 2015; Fig. 23).
Corylomyces Stchigel, M. Calduch & Guarro, Mycol. Res. 110(11): 1362 (2006)
Index Fungorum number: IF510040; 1 species with sequence data.
Type species – Corylomyces selenosporus Stchigel, M. Calduch & Guarro
Notes – Corylomyces is monotypic and characterized by tomentose ascomata, cylindrical asci
and lunate to reniform ascospores (Stchigel et al. 2006). It is closely related to Zopfiella tabulata
within Lasiosphaeriaceae based on multi-gene analysis (Wang et al. 2019a).
Diffractella Guarro, P.F. Cannon & Aa, Syst. Ascom. 10: 107 (1991)
Index Fungorum number: IF25499; 1 morphological species.
Type species – Diffractella curvata (Fuckel) Guarro, P.F. Cannon & Aa
Notes – The monotypic Diffractella has cleistothecial ascomata with alveolate-reticulate
ornamentation, globose asci and reniform to fusiform ascospores with a cap-like gelatinous
appendage at each end (Guarro et al. 1991).
Diplogelasinospora Cain, Can. J. Bot. 39: 1669 (1961)
Index Fungorum number: IF1609; 4 species with sequence data.
Type species – Diplogelasinospora princeps Cain
Notes – Diplogelasinospora was transferred from Sordariaceae to Lasiosphaeriaceae based
on multi-gene analysis and it was found to be closely related to Bombardia and Bombardioidea
(Cai et al. 2006c, Crous et al. 2014a, Wang et al. 2019a).
Emblemospora Jeng & J.C. Krug, Can. J. Bot. 54(16): 1971 (1976)
Index Fungorum number: IF1768; 2 morphological species (Species Fungorum 2020).
Type species – Emblemospora monotrema Jeng & J.C. Krug
Notes – Jeng & Krug (1976) introduced Emblemospora ditrema and E. monotrema (type
species) characterised by pyriform ascomata, clavate asci and ellipsoid ascospores with plate-like
wrinkles.
Eosphaeria Höhn., Annls mycol. 15(5): 362 (1917)
Index Fungorum number: IF1843; 1 morphological species.
Type species – Eosphaeria uliginosa (Kunze) Höhn.
Notes – The monotypic genus Eosphaeria is characterized by verrucose ascomata, clavate
asci and curved ascospores. The curved ascospores are similar to those found in Lasiosphaeria
(Kruys et al. 2015). In this entry we illustrate a specimen of E. uliginosa collected in the USA. We
have not compared this with the type specimens of Eosphaeria, but its morphological features fit
the description of E. uliginosa (Höhnel 1917).
Episternus Górz & Boroń, Phytotaxa 376(1): 49 (2018)
Index Fungorum number: IF811848; 1 species with sequence data.
Type species – Episternus onthophagi Górz & Boroń
Notes – The monotypic Episternus was found on the mesosternum, legs and dung of
Onthophagus beetles in Poland (Górz & Boroń 2018).
Fimetariella N. Lundq., Bot. Notiser 117: 239 (1964)
Index Fungorum number: IF1990; 9 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Fimetariella rabenhorstii (Niessl) N. Lundq.
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Notes – Fimetariella is characterized by ovoid to subglobose ascomata, cylindrical asci and
oval ascospores with a gelatinous sheath (Lundqvist 1964b). The ascospores are similar to
Bombardioidea (Kruys et al. 2015).
Figure 143 – Eosphaeria uliginosa (Material examined – USA, Maryland, of 728 Lead Mountain
(Humpback), along road near parking area, 44°51’28’’N, 68°05’52’’W, beech-maple forest, on
soil, 2 September 2014, RC Harris, NY 01818643). a Material label. b Material. c Ascomata on the
host. d Ascoma cross section. e Peridium. f Asci with paraphyses. g-l Ascospores. Scale bars: c = 1
mm, d = 200 µm, e, f = 100 µm, g-l = 20 µm.
Immersiella A.N. Mill. & Huhndorf, Mycol. Res. 108(1): 31 (2004)
Index Fungorum number: IF28831; 2 species with sequence data.
Type species – Immersiella immersa (P. Karst.) A.N. Mill. & Huhndorf
Notes – Immersiella caudata and I. immersa (type species) were introduced within
Sordariales, characterised by globose to subglobose ascomata, cylindrical to clavate asci and
geniculate ascospores with appendages at each end (Miller & Huhndorf 2004). The genus is closely
related to Arnium hirtum, within Lasiosphaeriaceae based on phylogenetic analysis (Kruys et al.
2015).
Jugulospora N. Lundq., Symb. bot. upsal. 20(no. 1): 256 (1972)
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Index Fungorum number: IF2538; 1 species with sequence data.
Type species – Jugulospora rotula (Cooke) N. Lundq.
Notes – The monotypic Jugulospora is characterized by ovoid to subglobose ascomata,
cylindrical asci and broadly ellipsoid to ovoid ascospores with plate-like wrinkles (Lundqvist
1972). It is closely related to Strattonia carbonaria within Lasiosphaeriaceae sensu lato based on
multi-gene analysis (Miller & Huhndorf 2005; Fig. 23).
Lasiosphaeria Ces. & De Not., Comm. Soc. crittog. Ital. 1(4): 229 (1863)
Index Fungorum number: IF2656; 63 morphological species (Species Fungorum 2020), 6
species with sequence data.
Type species – Lasiosphaeria ovina (Pers.) Ces. & De Not.
Notes – Lasiosphaeria has globose ascomata, cylindrical asci with a subapical globule and
geniculate ascospores. This genus was included in Lasiosphaeriaceae based on its pale brown,
geniculate ascospores (Barr 1990b). It is closely related to Anopodium based on multi-gene analysis
(Maharachchikumbura et al. 2016b). Fig. 23 reveals that Lasiosphaeria groups with Anopodium
and Bellojisia within Lasiosphaeriaceae sensu stricto.
Mammaria Ces., Bot. Ztg. 12: 190 (1854)
Index Fungorum number: IF8836; 2 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Mammaria echinobotryoides Ces.
Notes – Mammaria is an asexual genus which has brown to vinaceous brown mycelium and
broadly ellipsoid conidia with a longitudinal germ slit (Rabenhorst 1854). Mammaria
echinobotryoides was obtained in culture of Cercophora natalitia (Barr 1990b, del Valle Catania et
al. 2011). Pseudocercophora was found in Singapore and introduced as the sexual morph of
Mammaria, as they share similar conidial characters (Subramanian & Sekar 1986). However,
Mammaria is recommended as the current name, if the nomenclature principle of priority is
followed (Réblová et al. 2016b).
Periamphispora J.C. Krug, Mycologia 81(3): 476 (1989)
Index Fungorum number: IF25331; 1 morphological species.
Type species – Periamphispora phacelodes J.C. Krug
Notes – The monotypic Periamphispora is characterised by tuberulate ascomata surrounded
by hairs, clavate asci and ellipsoid to oval ascospores with lumpy ornamentation and a gelatinous
sheath. It is similar to Emblemospora in having ornamented ascospores (Kruys et al. 2015).
Ramophialophora M. Calduch, Stchigel, Gené, Guarro, Stud. Mycol. 50(1): 84
(2004)
Index Fungorum number: IF500021; 5 morphological species (Species Fungorum 2020), 4
species with sequence data.
Type species – Ramophialophora vesiculosa M. Calduch, Stchigel, Gené & Guarro
Notes – Ramophialophora has macronematous, mononematous conidiophores and aseptate,
brown conidia in slimy masses. Wijayawardene et al. (2020) proposed to transfer this genus to
Lasiosphaeriaceae from Sordariales genera incertae sedis based on phylogenetic analysis.
Rinaldiella Deanna A. Sutton, Y. Marín, Guarro & E.H. Thomps., Persoonia 32: 301 (2014)
Index Fungorum number: IF807137; 1 species with sequence data.
Type species – Rinaldiella pentagonospora Deanna A. Sutton, Y. Marín, Guarro & E.H.
Thomps.
Notes – Rinaldiella pentagonospora resembles species of Apiosordaria, with versicolored
ascospores with ornamented walls (Crous et al. 2014a). The genus is closely related to Echria in
Lasiosphaeriaceae sensu lato, based on multi-gene analysis (Wang et al. 2019a; Fig. 23).
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Schizothecium Corda, Icon. fung. (Prague) 2: 29 (1838)
Index Fungorum number: IF4908; 31 morphological species (Species Fungorum 2020), 7
species with sequence data.
Type species – Schizothecium fimicola Corda
Notes – Schizothecium is characterized by pyriform ascomata, cylindrical asci and versicolor
ascospores (Corda 1838). It is closely related to Jugulospora within Lasiosphaeriaceae based on
multi-gene analysis (Stchigel et al. 2006, Kruys et al. 2015).
Strattonia Cif., Sydowia 8(1-6): 245 (1954)
Index Fungorum number: IF5278; 10 morphological species (Species Fungorum 2020), 4
species with sequence data.
Type species – Strattonia tetraspora (R. Stratton) Cif.
Notes – Strattonia tetraspora has pyriform to subglobose ascomata and cylindrical asci with
ellipsoid ascospores (Ciferri 1954). It is closely related to Jugulospora within Lasiosphaeriaceae
based on phylogenetic analysis (Wang et al. 2019a).
Thaxteria Sacc., Syll. fung. (Abellini) 9: 687 (1891)
Index Fungorum number: IF5407; 4 morphological species (Species Fungorum 2020).
Type species – Thaxteria kunkelii Giard
Notes – Thaxteria has globose ascomata, clavate asci and allantoid ascospores (Saccardo
1891).
Tripterosporella Subram. & Lodha, Curr. Sci. 37: 246 (1968)
Index Fungorum number: IF5611; 2 morphological species (Species Fungorum 2020).
Type species – Tripterosporella coprophila Subram. & Lodha
Notes – Tripterosorella coprophila has cleistotecial ascomata, clavate asci and triangular
ascospores.
Zopfiella G. Winter, Rabenh. Krypt.-Fl., Edn 2 (Leipzig) 1.2: 56 (1884)
Index Fungorum number: IF5876; 23 morphological species (Species Fungorum 2020), 18
species with sequence data.
Type species – Zopfiella tabulata (Zopf) G. Winter
Notes – Huhndorf et al. (2004b) suggested that Zopfiella was placed in Chaetomiaceae. Cai et
al. (2006c) transferred this polyphyletic genus to Lasiosphaeriaceae and we found that species of
Zopfiella are widely distributed in the lasiosphaeriaceous complex (Fig. 23).
Zygopleurage Boedijn, Persoonia 2(3): 316 (1962)
Index Fungorum number: IF5889; 3 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Zygopleurage zygospora (Speg.) Boedijn
Notes – Zygopleurage has pyriform ascomata, cylindrical asci and ascospores consisting of 2
ellipsoid cells connected by a long cylindrical cell (Boedijn 1962, Lundqvist 1969). Sequence data
of Z. zygospora (Huhndorf et al. 2004b) revealed that Zygopleurage was closely related to Arnium
cirriferum based on phylogenetic analysis (Kruys et al. 2015).
Zygospermella Cain, Mycologia 27(2): 227 (1935)
Index Fungorum number: IF5894; 2 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Zygospermella setosa (Cain) Cain
Notes – Zygospermella is characterized by pyriform ascomata, clavate to cylindrical asci and
ascospores with appendages at each end (Lundqvist 1969). It is closely related to Cercophora
sordarioides based on tub2 and LSU sequence analysis (Kruys et al. 2015).
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Lautosporaceae Kohlm., Volkm.-Kohlm. & O.E. Erikss., Bot. Mar. 38(2): 169 (1995)
Index Fungorum number: IF81975; Facesoffungi number: FoF01764; 2 species.
Saprobic on submerged mangrove wood and culms of salt marsh plants in marine habitats.
Sexual morph: Ascomata fusiform to ellipsoidal, ostiolate, brown, coriaceous, solitary. Paraphyses
simple, septate, persistent. Asci 4-spored, cylindrical, short pedicellate, thick-walled, unitunicate,
with an ocular chamber. Ascospores uni- or biseriate, fusiform, muriform, distoseptate, hyaline,
outer wall very thick. Asexual morph: Undetermined (adapted from Jones et al. 2015).
Type genus – Lautospora K.D. Hyde & E.B.G. Jones
Notes – Lautosporaceae comprises two marine Lautospora species, collected on driftwood of
Sonneratia griffithii (L. gigantea) and culms of Juncus roemerianus (L. simillima) (Hyde & Jones
1989, Kohlmeyer et al. 1995, respectively). The family was not assigned to any order, but referred
to Ascomycota incertae sedis. Jones et al. (2009) included the family in Dothideomycetidae
incertae sedis, based on the thick-walled nature of the ascus. A recent collection of L. simillima on
mangrove wood in Thailand by Suetrong, enabled the extraction of DNA. Based on LSU sequence
data (Suetrong, unpublished data), L. simillima was referred to an unnamed clade in
Sordariomycetes and in a basal clade to Diaporthales (Jones et al. 2015). Lautosporaceae was
therefore excluded from the class Dothideomycetes (Hyde et al. 2013). Lautospora simillima
grouped with the neotropical ascomycete Mirannulata sameulsii with weak support, and shares few
morphological features with this genus (Huhndorf et al. 2003, Jones et al. 2015). The
Lautospora/Mirannulata clade formed a sister group to Vertexicola caudatus and Rhamphoria
delicatula (Annulatascaceae), hence Jones et al. (2015) emended the diagnosis of Lautosporaceae
family.
Ecological and economic significance of Lautosporaceae
Lautosporaceae species are saprobic on submerged mangrove wood of Sonneratia griffithii,
and the salt marsh sedge Juncus roemerianus (Kohlmeyer et al. 1995, Huhndorf et al. 2003, Jones
et al. 2015).
Genus included in Lautosporaceae
Lautospora K.D. Hyde & E.B.G. Jones, Bot. Mar. 32(3): 479 (1989)
Index Fungorum number: IF25368; 2 morphological species (Species Fungorum 2020).
Type species – Lautospora simillima Kohlm., Volkm.-Kohlm. & O.E. Erikss.
Notes – Lautospora species can be distinguished by immersed ascomata, 4-spored,
unitunicate asci with an ocular chamber and ring, cellular paraphyses and large ascospores which
are muriform, hyaline and thick-walled (Hyde & Jones 1989). Kohlmeyer et al. (1995) introduced
Lautosporaceae to include this genus and assigned it to Ascomycotina family incertae sedis. More
taxon sampling and DNA sequencing are recommended for this genus including the type L.
gigantea to confirm the phylogenetic placement. Lautospora simillima is as an obligate marine
fungus, with its ascomata occurring singly, or at the most, in pairs at the base of very soft culms,
between 3 and 12 cm above the rhizome. However, we could not observe the ascomata on
microslides of the holotype. Lautospora simillima is a second species assigned to the genus,
occurring on the culms of Juncus roemerianus and no asexual morph has been reported for this
species (Kohlmeyer et al. 1995, Wijayawardene et al. 2017a).
Leptosilliaceae Voglmayr & Jaklitsch, Persoonia 42: 240 (2019)
Index Fungorum number: IF829929; Facesoffungi number: FoF06183; 9 species.
Endophytic, pathogenic or lichenized. Sexual morph: Ascomata perithecial, superficial to
partly immersed, scattered or collapsed, black. Ostioles central, papillate, base of canal sulcate,
sometimes with hyaline periphyses. Peridium melanized, KOH-, comprises cells of textura
angularis or prismatica. Paraphyses septate, occasionally branched, embedded in a gelatinous
matrix. Asci 8-spored, clavate to cylindrical, curved or sinuous, with J+, apical ring. Ascospores bi
or tri-seriate or fasciculate, hyaline, falcate, lunate, sinuous, sigmoid to hook-shaped, aseptate or
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septate, not constricted, thin and smooth-walled, with rounded to subacute apices, without
appendages or gelatinous sheath. Asexual morph: Coelomycetous. Conidiomata superficial to
partly immersed, scattered, aggregated or confluent, globose to pyriform, uni or irregularly
pluriloculate, black. Conidiomatal wall more or less melanized, textura globulosa or angularis.
Conidiophores hyaline, short with arising from the inner layer of the peridium. Conidiogenous cells
cylindrical to lageniform. Conidiogenesis enteroblastic phialidic or holoblastic with sympodial
proliferation to both types. Conidia falcate or filiform, aseptate, hyaline, allantoid, thin-walled
(adapted from Voglmayr et al. 2019a) .
Type genus – Leptosillia Höhn.
Notes – Leptosilliaceae was introduced by Voglmayr et al. (2019a) which belongs to
Xylariales and is morphologically similar to Delonicicolaceae, but differs in ascomata structure.
Only one genus, Leptosillia is recognized in the family based on molecular and morphology data
(Voglmayr et al. 2019a). Voglmayr et al. (2019a) clearly showed Leptosillia should be placed in the
Xylariales with strong evidence from ITS and LSU analyses. However, in this study we accept this
family under Delonicicolales, based on high statistical support (Figs. 1, 2, 4).
Ecological and economic significance of Leptosilliaceae
Most Leptosilliaceae species are facultative lichenised taxa. Leptosilliaceae can be
endophytes in woody hosts in tropical or subtropical areas (Voglmayr et al. 2019a). Leptosillia
pistaciae is a serious canker pathogen of Pistacia vera (Vitale 2018). In addition, Vitale (2018) and
Voglmayr et al. (2019a) mentioned that some Leptosillia species isolated as endophytes within
living plant tissues and may actually represent latent pathogens.
Genus included in Leptosilliaceae
Leptosillia Höhn., Mitt. bot. Inst. tech. Hochsch. Wien 5: 111 (1928)
Index Fungorum number: IF2797; 9 morphological species (Species Fungorum 2020), 6
species with sequence data.
Type species – Leptosillia notha Höhn.
Notes – When establishing Cresporhaphis, a synonym of Leptosillia, Aguirre-Hudson (1991)
described its association with an unidentified globose chlorococcoid photobiont. Various species
were included, as probably lichenized and Calatayud & Aguirre-Hudson (2005) considered
Cresporhaphis ulmi as not lichenised. Voglmayr et al. (2019a) introduced seven species as new
combinations in Leptosillia. Species of Leptosillia have superficial to partly immersed, subglobose
to pyriform, ostiolate ascomata, with a peridium of textura angularis or prismatica, and clavate to
cylindrical asci and lunate ascospores. The asexual morph has superficial conidiomata with cells of
textura globulosa to angularis in the conidiomatal wall, short hyaline conidiophores, cylindrical to
lageniform conidiogenous cells and lunate or filiform conidia (Voglmayr et al. 2019a).
Leptosporellaceae S. Konta & K.D. Hyde, Mycosphere 8(10): 1956 (2017)
Index Fungorum number: IF553956; Facesoffungi number: FoF03840; 11 species.
Saprobic or endophytic on various host plants. Sexual morph: Ascomata solitary, superficial,
comprising black, subglobose, carbonaceous, dome-shaped, raised, blister-like areas, within the
plant tissues, flattened at the base, ostiole central. Peridium outer cells merging with the host
epidermal cells, composed of dark brown to black cells of textura angularis. Paraphyses numerous,
hyaline, hypha-like, septate, longer than asci. Asci 8-spored, unitunicate, cylindrical, longpedicellate, with a J-, wedge-shaped, subapical ring. Ascospores fasciculate, hyaline or paleyellowish in mass, spiral, filiform, straight or curved, aseptate, ends rounded, with or without polar
mucilaginous appendages. Asexual morph: Undetermined (adapted from Konta et al. 2017).
Type genus – Leptosporella Penz. & Sacc.
Notes – The family was introduced by Konta et al. (2017) in Chaetosphaeriales with support
from the analysis of combined LSU and ITS sequence data. In this study, a new species,
Leptosporella elaeidis, is introduced (Fig. 8).
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Figure 144 – Lautospora simillima (Material examined – USA, North Carolina, on senescent
culms of Juncus roemerianus, J. Kohlmeyer, J.K. 5532 IMS), microslides from holotype). a
Microslides of Lautospora simillima. b Apex of an asci in methylene blue. c, d Asci. e An ascus in
methylene blue. f Paraphyses. g, h Ascospores. i Germinating ascospore on agar. Scale bar: c-e =
200 μm, d-i = 100 μm, b = 50 μm.
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Figure 145 – Sexual morph of Leptosillia muelleri (redrawn from Voglmayr et al. 2019a). a, b
Ascomata on the substrate. c Paraphyses. d, e Asci. f Ascus apex in Lugol’s reagent. g Dead
ascospores. h-n Vital ascospores. Scale bars: a, b = 100 µm, c = 5 µm, d-f = 10 µm, g-n = 5 µm.
Ecological and economic significance of Leptosporellaceae
Species of Leptosporellaceae are saprobes and endophytes on various plants, especially
monocotyledons. Appressoria have been re-observed in culture, indicating the genus may be
endophytic (Konta et al. 2016).
Genus included in Leptosporellaceae
Leptosporella Penz. & Sacc., Malpighia 11(9-10): 406 (1897)
Index Fungorum number: IF2804; 11 morphological species (Species Fungorum 2020), 4
species and 2 unnamed species with sequence data.
Type species – Leptosporella gregaria Penz. & Sacc.
Notes – A phylogenetic tree for this genus was provided in Konta et al. (2017). The genus is
characterised by solitary, superficial, carbonaceous, dome-shaped ascomata, with a central ostiole,
cylindrical asci with a J-, subapical ring and spiral, filiform, hyaline (pale-yellowish in mass),
aseptate ascospores, with or without polar mucilaginous appendages (Konta et al. 2017). The
asexual morph and appressoria have not been reported. Leptosporella arengae is illustrated for this
genus with updated the information of an appressoria-like structure that was produced at the hyphal
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tip when ascospores were germinated. Appressoria were 4–5 μm high × 4–5 μm diameter (x̅ = 5
μm, n = 5), rare, globose to subglobose, irregular, and hyaline. A new species, Leptosporella
elaeidis is introduced herein.
Figure 146 – Asexual morph of Leptosillia muelleri (redrawn from Voglmayr et al. 2019a). a, b
Conidiophores, conidiogenous cells and conidia from pycnidia on natural substrates. c, d
Conidiophores, conidiogenous cells and conidia from pycnidia in pure culture. e-i Conidia from
natural substrate (e - dead, f-I - vital). Scale bars: a-d = 10 µm, e-i = 5 µm.
Leptosporella elaeidis Konta & K.D. Hyde, sp. nov.
Fig. 148
Index Fungorum number: IF556361; Facesoffungi number: FoF06026
Etymology – Named after the host genus, Elaeis.
Holotype – MFLU 19–0669.
Saprobic on rachis and petioles of Elaeis guineensis Jacq. Sexual morph: Ascomata (included
neck) 242–600 μm high × 625–1000 μm diameter (x̅ = 444 × 787 μm, n = 10), solitary, superficial,
comprising black, carbonaceous, dome-shaped, raised blister-like areas, subglobose, flattened at the
base, ostiolate. Ostiole central, carbonaceous, black, with periphyses. Peridium 63–113 μm wide ( x
= 83 μm, n = 10), outer cells merging with the host epidermal cells, composed of dark brown to
black cells of textura angularis. Paraphyses numerous, 2–7 µm diameter (x̅ = 4 μm, n = 10),
branched, septate, longer than asci. Asci 125–210 × 9–14 μm (x̅ = 163 × 11 μm, n = 20), 8-spored,
unitunicate, cylindrical, long-pedicellate, with a J-, wedge-shaped, subapical ring. Ascospores 124–
136 × 2–4 μm (x̅ = 128 × 3 μm, n = 20), fasciculate, yellowish in mass, spiral, filiform, straight or
curved, C-shaped or sigmoid, aseptate, rounded at the apex, pointed at the base, without polar
mucilaginous appendages, smooth-walled. Asexual morph: Undetermined.
Material examined – THAILAND, Suratthani Province, on dead rachis and petioles of Elaeis
guineensis Jacq. (Arecaceae), 21 July 2017, Sirinapa Konta SRWD14b, MFLU 19-0669, holotype.
GenBank numbers – ITS: MK659767, LSU: MK659772, SSU: MK659774, tef1: MN883560.
Notes – The ascospores of Leptosporella elaeidis failed to germinate and therefore the DNA
was obtained directly from fruiting bodies. Phylogenetic analysis placed our collection with L.
arengae, 61% ML (Fig. 8). Leptosporella elaeidis differs from L. arengae in having larger
ascomata, asci and ascospores, and lack of polar mucilaginous appendages. This is the first record
of L. elaeidis on Elaeis guineensis and the fourth record of Leptosporella species from a palm tree
(Arecaceae).
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Figure 147 – Leptosporella arengae (Material examined – THAILAND, Phang-Nga Province, on
dead rachis of Arenga pinnata (Wurmb) Merr. (Arecaceae), 5 December 2014, Sirinapa Konta
PHR07a, MFLU 15-0305, holotype). a Appearance of ascomata on host substrate. b Close up of
ascomata. c Yellowish ascospore mass. d Section of ascoma. e Peridium. f Paraphyses. g, h Asci. i,
j Ascospores. k J-, reaction of the apical ring. l Germinated ascospores (appressoria-like at red
arrow). m-o Appressoria. p Colony on MEA. Scale bars: b = 1,000 μm, c = 500 μm, d = 200 μm, e,
g-j, k = 50 μm, f, k = 20 μm, m-o = 5 μm.
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Figure 148 – Leptosporella elaeidis (MFLU 19-0669, holotype). a, b Appearance of ascomata on
host substrate. c, d Close up of ascomata. e Yellowish ascospore mass. f, g Sections of ascoma. h
Neck with periphyses. i Peridium. j Paraphyses (branch and septate at red arrow). k-m Asci. n-o J-,
reaction of apical ring. p-u Ascospores. Scale bars: b, c = 1000 μm, d, e = 500 μm, f, g = 200 μm, h,
j-n, p-u = 100 μm, i = 50 μm, o = 20 μm.
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Linocarpaceae S. Konta & K.D. Hyde, Mycosphere 8(10): 1962 (2017)
Index Fungorum number: IF553959; Facesoffungi number: FoF03843; 56 species.
Saprobic and endophytic fungi on monocotyledons and rarely dicotyledons. Sexual morph:
Ascomata solitary, superficial comprising black, dome-shaped (Linocarpon), slightly raised or
flattened circular areas, or immersed (Neolinocarpon) with a black shiny papilla. Peridium
composed of dark brown to black cells of textura angularis. Paraphyses septate, longer than asci,
wider at the base, tapering towards the apex. Asci 8-spored, unitunicate, cylindrical, with a J-,
apical ring, developing from the base and periphery of the ascomata. Ascospores parallel or spiral
in asci, hyaline or pale-yellowish in mass, filiform, straight or curved, unicellular with refringent
bands, with or without polar appendages. Asexual morph: A phialophora-like spp. was found in
Linocarpon appendiculatum and L. elaeidis cultures (Hyde 1992a), but has not been recovered in
other species (adapted from Konta et al. 2017).
Type genus – Linocarpon Syd. & P. Syd.
Notes – A recent treatment and updated account of Linocarpaceae by Konta et al. (2017)
placed this family in Chaetosphaeriales which was confirmed by Hyde et al. (2019a) . The first
record of appressorium structures were from Neolinocarpon (Hyde et al. 2019a).
Ecological and economic significance of Linocarpaceae
Linocarpaceae species are saprobes and endophytes on monocotyledons and rarely
dicotyledons. It is likely that they change from endophytes to saprobes as leaves scenesce and are
involved in nutrient cycling.
Genera included in Linocarpaceae
Linocarpon Syd. & P. Syd., Annls mycol. 15(3/4): 210 (1917)
Index Fungorum number: IF2882; 43 morphological species (Species Fungorum 2020); 10
species and 9 unnamed species with sequence data.
Type species – Linocarpon pandani Syd. & P. Syd.
Notes – The most recent update of this genus is by Konta et al. (2017) who we follow here.
Linocarpon calami Konta, sp. nov.
Fig. 149
Index Fungorum number: IF556362; Facesoffungi number: FoF06024
Etymology – Named after the host genus, Calamus.
Holotype – MFLU 15-0277.
Saprobic on petioles of Calamus. Sexual morph: Ascomata 147–187 μm high × 565–732 μm
diameter (x̅ = 165 × 637 μm, n = 10), solitary or aggregated, superficial, comprising black, domeshaped, raised, blister-like areas, subglobose in section, flattened at the base, with a central ostiole.
Peridium 33–70 μm diameter ( x̅ = 47 μm, n = 10) , outer cells merging with the host epidermal
cells, composed of dark brown to black cells of textura angularis. Paraphyses 3–6 µm diameter (x̅
= 4 μm, n = 10), numerous, septate, longer than asci. Asci 120–160 × 13–21 μm (x̅ = 140 × 16 μm,
n = 20), 8-spored, unitunicate, cylindrical, long-pedicellate, with a J-, wedge-shaped, subapical ring.
Ascospores 78–95 × 2–4 μm (x̅ = 91 × 5 μm, n = 20), parallel when immature, becoming spiral
when mature, hyaline, filiform, straight or curved, aseptate, containing numerous refringent
septum-like bands, ends rounded, without polar mucilaginous appendage at the apex, smoothwalled. Asexual morph: Undetermined.
Material examined – THAILAND, Phang-Nga Province, on dead petioles of Calamus
(Arecaceae), 6 December 2014, Sirinapa Konta DNH02a, MFLU 15-0277, holotype.
Notes – A new species is introduced based on morphology and comparison with known
Linocarpon species as well as species reported in Thailand. Linocarpon calami fits with the generic
concept of Linocarpon in having superficial, dome-shaped, cylindrical asci, with J-, apical ring and
filiform ascospores with refringent bands. Comparisons of Linocarpon species known in Thailand,
shows that L. calami differs from L. arengae in having smaller ascomata, asci and ascospores, and
ascospores without polar mucilaginous appendage, while, Linocarpon calami differs from L. cocois
686
in having larger ascospores containing numerous refringent septum-like bands. Therefore,
Linocarpon calami is introduced as a new species and the first geological recorded from the host
genus Calanus (Arecaceae) in Thailand.
Figure 149 – Linocarpon calami (MFLU 15-0277, holotype). a, b Appearance of ascomata on host
substrate. c Close up of ascoma. d Off white to yellowish ascospores mass. e, f Section of ascoma.
g Peridium. h J-, reaction of apical ring. i Paraphyses (septate at the red arrow). j, k Asci (k mature
asci with immature asci and paraphyses). l-o Ascospores. Scale bars: b = 1000 μm, c = 500 μm, d-f
= 200 μm, g-h = 20 μm, i-o = 50 μm.
Neolinocarpon K.D. Hyde, Bot. J. Linn. Soc. 110(2): 104 (1992)
Index Fungorum number: IF26303; 13 morphological species (Species Fungorum 2 0 2 0 ); 7
species and 3 unnamed species with sequence data.
Type species – Neolinocarpon globosicarpum K.D. Hyde
Notes – Hyde (1992a) introduced Neolinocarpon with four new species to accommodate N.
globosicarpum which is saprobic on Nypa fruticans in Brunei. Two new species were introduced
with molecular data (Konta et al. 2017). An appressorium has been reported from N. rachidis
687
( MFLU 15-2347) ( Hyde et al. 2019a) . We provide an illustration of Neolinocarpon rachidis to
show the appressorium characters of Neolinocarpon (Hyde et al. 2019a).
Figure 150 – Neolinocarpon rachidis (Material examined – THAILAND, Prachaupkhirikhan
Province, on dead of petiole, Cocos nucifera (Arecaceae) , 30 July 2015, Sirinapa Konta PJK04i,
MFLU 15-2347). a Appearance of ascomata on host substrate. b Close up of ascomata. c Yellowish
ascospore mass. d Section of ascoma. e Papilla. f Peridium. g Paraphyses. h-i Asci. j J-, a reaction
of apical ring. k-m Ascospores. n Germinated ascospore (appressoria at red arrow). o-r
Appressoria. s Culture on MEA. Scale bars: a = 1000 μm, b, c = 500 μm, d = 100 μm, e, h-n = 50
μm, f = 20 μm, g = 10 μm, o-r = 5 μm.
688
Lopadostomataceae Daranagama & K.D. Hyde, Fungal Divers 73: 1 (2015)
Index Fungorum number: IF91005; Facesoffungi number: FoF00071; 36 species.
Saprobic on dead wood. Sexual morph: Pseudostromata/stromata immersed to erumpent or
nearly superficial, gregarious, scattered, densely compact, with abrupt margins, appearing as
darkened areas, dark brown to dull black, coalescing in black linear rows, waxy, roughened,
flattened at the top, with inconspicuous ascomatal mounds, embedded in carbonized encasement.
Ascomata cylindrical-subglobose or flask-shaped, multi-peritheciate, single to multi-layered,
clustered into valsoid groups, sometimes with long ostiolar necks (Lopadostoma), lined with
whitish material. Ostioles umbilicate or at the same level as stromal surface. Peridium amorphous,
outwardly comprised of dark brown to black cells, inwardly hyaline, flattened cells. Paraphyses
numerous, long, rarely branched, apically free. Asci (4–)8-spored, unitunicate, cylindrical,
pedicellate, apex rounded, with J+, discoid to wedged-shaped, apical ring, J+. Ascospores uniseriate
or partially biseriate, initially hyaline, turning light brown to nearly black, unicellular, oblong,
narrowly ellipsoidal, smooth-walled, without any patterns, germ slit full length, straight. Asexual
morph: Coelomycetous or Hyphomyceteous, libertella-like or sometimes nodulisporium or
geniculosporium-like synanamorph (adapted from Senanayake et al. 2015, Daranagama et al.
2018).
Type genus – Lopadostoma (Nitschke) Traverso
Notes – Lopadostomataceae was introduced by Senanayake et al. (2015) in Xylariales to
accommodate Creosphaeria and Lopadostoma. The family is typified by Lopadostoma turgidum.
In earlier considerations, Creosphaeria and Lopadostoma, which have libertella-like asexual
morphs were treated as a basal lineage of Xylariaceae (Tang et al. 2009b). ITS and LSU combined
phylogenetic analyses revealed that the Lopadostomataceae is a monophyletic sister clade to
Coniocessiaceae, hence it was treated as a separate family (Senanayake et al. 2015). At present, the
family consists with four genera including Creosphaeria, Jumillera, Lopadostoma and Whalleya
(Daranagama et al. 2018) and accepted as a family in Xylariales according to the evolutionary
based ranking system (Hyde et al. 2017a). Antimicrobial azaphilone pigments discovered from
Creosphaeria sassafras are known metabolites discovered in the family (Quang et al. 2005).
Ecological and economic significance of Lopadostomataceae
Lopadostomataceae members play a major role as saprobes on corticated and decorticated
wood (Daranagama et al. 2018). Sassafrin type pigments were identified from this family, however,
investigation on potential bioactive compounds in this family is needed.
Genera included in Lopadostomataceae
Creosphaeria Theiss., Beih. bot. Zbl., Abt. 2 27: 396 (1910)
Index Fungorum number: IF1288; 2 morphological species (Daranagama et al. 2018, Species
Fungorum 2020), 1 species with sequence data.
Type species – Creosphaeria sassafras (Schwein.) Y.M. Ju, F. San Martín & J.D. Rogers
Notes – Ju et al. (1993) resurrected Creosphaeria based on the morphological differences
with Hypoxylon. In several phylogenetic studies, Creosphaeria is sister to Lopadostoma (Jaklitsch
et al. 2014, Daranagama et al. 2018). The genus is characterised by stromata with KOH- extractable
pigments, subglobose to broadly cylindrical ascomata, cylindrical, long-stipitate asci with discoid,
J+, apical rings and unicellular, ellipsoid, slightly inequilateral to oblong or cylindrical, brown
ascospores, with straight germ slit and libertella-like coelomycetous asexual morphs (Daranagama
et al. 2018).
Jumillera J.D. Rogers, Y.M. Ju & F. San Martín, Mycotaxon 64: 41 (1997)
Index Fungorum number: IF27739; 8 morphological species (Species Fungorum 2020).
Type species – Jumillera mexicana J.D. Rogers, Y.M. Ju & F. San Martín
Notes – Jumillera was introduced by Rogers et al. (1997) based on generally small
ascospores as compare to Biscogniauxia and libertella-like asexual morphs with geniculosporium689
like conidial state as synanamorphs including culture characteristics. At present the genus is placed
in Lopadostomataceae. However, the taxonomic placement of the genus needs to be revised with
phylogenetic studies (Daranagama et al. 2018, Wendt et al. 2018).
Lopadostoma (Nitschke) Traverso, Fl. ital. crypt. 1(2): 169 (1906)
Index Fungorum number: IF2925; 24 morphological species (Species Fungorum 2020), 11
species with sequence data.
Type species – Lopadostoma turgidum (Pers.) Traverso
Notes – Lopadostoma comprises 12 species from Europe which are saprobes producing
pseudostromatic sexual morphs and libertella-like asexual morphs (Fig. 152, Jaklitsch et al. 2014).
So far species delimitation of Lopadostoma is based only on ITS and LSU sequence data with low
resolution, while protein coding regions, especially rpb2, are essential for further species
delimitation (Daranagama et al. 2016, Jaklitsch et al. 2014). The Lopadostoma gastrinum sexual
morph and L. turgidum asexual morph are illustrated in this entry (Fig. 151, 152).
Figure 151 – Lopadostoma gastrinum (Material examined – ITALY, Province of Forlì-Cesena
[FC], Monte Mirabello – Predappio, on dead fallen branch of Quercus sp. (Fagaceae), 12 April
2017, Erio Camporesi IT3269C, MFLU 17-0941; ibid. HKAS 102310). a, b Stromata on the host. c
Cross section of stroma. d Peridium. e, f Paraphyses. g J+, apical ring in Melzer’s reagent. h-l
Ascospores (l-white arrows show germ slit). m-o Asci. Scale bars: d, m-o = 20 μm, g-l = 10 μm, e,
f = 5 μm.
Whalleya J.D. Rogers, Y.M. Ju & F. San Martín, Mycotaxon 64: 48 (1997)
690
Index Fungorum number: IF27738; 2 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Whalleya microplaca (Berk. & M.A. Curtis) J.D. Rogers, Y.M. Ju & F. San
Martín
Notes – Whalleya species have conspicuous, bipartite, stromata and asci with a J+, apical
ring, and dark, ellipsoidal ascospores and has scolecosporous conidia, similar to lopadostomaceous
species (Rogers et al. 1997). Based on both xylariaceous and lopadostomaceous morphological
affinities, Maharachchikumbura et al. (2016b) placed the genus in Xylariales incertae sedis.
However, based on recent phylogenetic studies, the genus is accommodated in Lopadostomataceae
(Daranagama et al. 2018, Wendt et al. (2018).
Figure 152 – Lopadostoma turgidum (ex-epitype culture CBS 133207). a, b Conidiophores (b.
showing phialide bearing conidium). c Conidia. Scale bars: 10 μm (redrawn from Jaklitsch et al.
2014).
Lulworthiaceae Kohlm., Spatafora & Volkm.-Kohlm., Mycologia 92(3): 456 (2000)
Index Fungorum number: IF82091; Facesoffungi number: FoF01295; 47 species.
Saprobic, on wood or growing on sea grasses, marsh plants and calcareous animal shells,
endoskeletons and feathers, parasitic on algae and endophytic in sea grasses. Sexual morph:
Ascomata pale to dark brown to black, occasionally purple, subglobose to cylindrical, immersed or
superficial, coriaceous, ostiolate, papillate, sometimes with a long neck. Peridium mostly two
layered, composed of an outer layer of cells of textura angularis and an inner layer of elongate
cells. Paraphyses lacking; centrum initially filled with a hyaline pseudoparenchyma, dissolving at
maturity. Asci 8-spored, unitunicate, cylindrical to fusiform, short pedicellate, deliquescent.
Ascospores fasciculate, hyaline, filiform, septate or aseptate, conical or semi-globose chambers at
the ends, with or without mucus. Asexual morph: hyphomycetous, hyphae pale brown, septate,
branched. Conidiophores micronematous or semi-micronematous. Conidia septate or aseptate,
some coiled (adapted from Maharachchikumbura et al. 2016b).
Type genus – Lulworthia G.K. Sutherl.
Notes – Lulworthia species have been recorded from various marine habitats and this is one
of the largest genera among the marine fungi (Kohlmeyer et al. 2000). Sutherland (1916)
introduced Lulworthia to accommodate the species L. fucicola, found on the brown seaweed,
bladder wrack at Lulworth Cove on the Dorset coast, UK, but no type material was ever deposited
for this taxon. Campbell et al. (2005) designated a neotype for the type species of Lulworthia (L.
691
fucicola) from a collection from Chile. Lulworthia has been revisited over many years (Kohlmeyer
1972, Kohlmeyer & Kohlmeyer 1979, Koch & Jones 1984, Schaumann et al. 1986). Johnson &
Sparrow (1961) listed 12 Lulworthia species, which were synonymized under L. medusa by
Cavaliere & Johnson (1966). Three species were described by Kohlmeyer (1972) and Koch &
Jones (1984) recognized six species. Eleven accepted species were included in Lulworthia with a
number of other taxa by Kohlmeyer et al. (2000). However, their morphological differences are not
sufficient to distinguish the species (Kohlmeyer et al. 2000). As a result of sequence analysis of
LSU and SSU sequences a number of transfers have been recommended, such as Lulworthia crassa
to Kohlmeyeriella, Lulworthia lignoarenaria to Lulwoidea and Lulworthia uniseptata to Lulwoana
by Campbell et al. (2005). Jones et al. (2015) showed that only L. fucicola can be accepted in
Lulworthia sensu stricto, while 11 species are referred to Lulworthia sensu lato until further taxa
are collected and sequenced. Abdel-Wahab et al. (2017) introduced a novel genus Sammeyersia to
accommodate L. grandispora based on molecular data. Azevedo et al. (2017) introduced a novel
species Lulworthia atlantica to this genus in the Lulworthia sensu stricto clade.
The genera Lulworthia and Lindra were previously placed in Halosphaeriales (Spatafora et al.
1998). Subsequently, phylogenetic analysis of several species of Lulworthia and Lindra with
filamentous ascospores led to the inclusion of their parent genera to a new family Lulworthiaceae
in a new order Lulworthiales (Kohlmeyer et al. 2000). Jones et al. (2008) showed that the asexual
genera Cumulospora and Orbimyces grouped in Lulworthiaceae (Lulworthiales) with statistical
support. Subsequently, in a molecular analysis of SSU and LSU sequence data, Abdel-Wahab et al.
(2010) demonstrated that the genera Cumulospora, and Cirrenalia were polyphyletic and
introduced the new genera Halazoon, Hydea, Matsusporium and Moheirospora for species
previously referred to Cirrenalia, and Moromyces to accommodate Cumulospora varia. They also
introduced the genera Moleospora and Glomerulispora in Lulworthiaceae. Currently this family
comprises the genera, Cumulospora, Halazoon, Haloguignardia, Hydea, Kohlmeyeriella, Lindra,
Lulwoana, Lulwoidea, Lulworthia, Matsusporium, Moleospora, Moromyces, Orbimyces,
Rostrupiella, and Sammeyersia (Jones et al. 2015, Maharachchikumbura et al. 2016b, AbdelWahab et al. 2017, Wijayawardene et al. 2017a, 2018).
Ecological and economic significance of Lulworthiaceae
Lulworthiaceae species are mostly saprobes occurring on wood or growing on sea grasses and
marsh plants (Kohlmeyer et al. 2000, Jones et al. 2015, Azevedo et al. 2017). Calado et al. (2015)
reported that Lulworthia sp. (in Castro Marim) occurrs mostly in the more frequently flooded plant
portions (basal portions) associated with stems and/or leaf sheaths of salt marsh plant Spartina.
Lindra thalassiae has been reported in the degradation of turtle grass which is an ecologically
important marine flowering plant (Orpurt et al. 1964). Haloguignardia sp. and Lulworthia kniepii
have been reported as parasites on red and brown algae (Kohlmeyer 1974, Alongi et al. 1999). Gall
formation in species of the genera Cystoseira and Sargassum is generally caused by
Haloguignardia spp. (Apt 1988, Alongi et al. 1999). The occurrence of Lulwoana has also been
reported in extreme environments such as saline soils (Hujslova et al. 2010). It is also believed that
Lulwoana sp. in Posidonia oceanica roots help the host to capture mineral nutrients, through lytic
activity by living as an endophyte (Torta et al. 2015). Nambiar & Raveendran (2015) reported
Matsusporium tropicalis was commonly found on four substrates including calcareous animal
shells, ‘chicken bone’, endoskeleton of Seibia and feathers.
Genera included in Lulworthiaceae
We do not provide notes on each genus as these can be found at marinefungi.org (Jones et al.
2019). A monograph on genera of Lulworthiaceae is also planned by Dayarathne et al. (2020).
Cumulospora I. Schmidt, Mycotaxon 24: 420 (1985)
Index Fungorum number: IF25713; 1 species with sequence data.
Type species – Cumulospora marina I. Schmidt
692
Halazoon Abdel-Aziz, Abdel-Wahab & Nagah., Mycol. Progr. 9(4): 545 (2010)
Index Fungorum number: IF543113; 2 species with sequence data.
Type species – Halazoon melhae Abdel-Aziz, Abdel-Wahab & Nagah.
Haloguignardia Cribb & J.W. Cribb, Pap. Dept. Bot. (formerly Biol.) Univ. Qd. 3: 97 (1956)
Index Fungorum number: IF2205; 6 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Haloguignardia deciduas Cribb & J.W. Cribb
Hydea K.L. Pang & E.B.G. Jones, Mycol. Progr. 9(4): 549 (2010)
Index Fungorum number: IF543118; 1 species with sequence data.
Type species – Hydea pygmea (Kohlm.) K.L. Pang & E.B.G. Jones
Notes – Abdel-Wahab et al. (2010) transferred Cirrenalia pygmea to Hydea based on
phylogenetic analyses of combined LSU and SSU sequence data. Hydea pygmea is commonly
found on mangrove wood, especially growing on Rhizophora species (Abdel-Wahab et al. 2010).
Hydea pygmea is different from all other cirrenalia-like species by the dark-brown to black hooked
nature of the conidia and by the internal proliferation of the conidia (Kohlmeyer & Kohlmeyer
1979).
Kohlmeyeriella E.B.G. Jones, R.G. Johnson & S.T. Moss, J. Linn. Soc., Bot. 87: 208 (1983)
Index Fungorum number: IF25816; 2 species with sequence data.
Type species – Kohlmeyeriella tubulata (Kohlm.) E.B.G. Jones, R.G. Johnson & S.T. Moss
Lindra I.M. Wilson, Trans. Br. mycol. Soc. 39(4): 411 (1956)
Index Fungorum number: IF2877; 5 morphological species (Species Fungorum 2020), 3
species with sequence data.
Type species – Lindra inflata I.M. Wilson
Lulwoana Kohlm., Volkm.-Kohlm., J. Campb., Spatafora & Gräfenhan, Mycol. Res. 109(5): 562
(2005)
Index Fungorum number: IF28935; 1 species with sequence data.
Type species – Lulwoana uniseptata (Nakagiri) Kohlm., Volkm.-Kohlm., J. Campb.,
Spatafora & Gräfenhan
Lulwoidea Kohlm., Volkm.-Kohlm., J. Campb., Spatafora & Gräfenhan, Mycol. Res. 109(5): 564
(2005)
Index Fungorum number: IF28936; 1 species with sequence data.
Type species – Lulwoidea lignoarenaria (Jørg. Koch & E.B.G. Jones) Kohlm., Volkm.Kohlm., J. Campb., Spatafora & Gräfenhan
Lulworthia G.K. Sutherl., Trans. Br. mycol. Soc. 5(2): 259 (1916)
Index Fungorum number: IF2953; 22 morphological species (Species Fungorum 2020), 3
species with sequence data.
Type species – Lulworthia fucicola G.K. Sutherl.
Matsusporium E.B.G. Jones & K.L. Pang, Mycol. Progr. 9(4): 550 (2010)
Index Fungorum number: IF543120; 1 species with sequence data.
Type species – Matsusporium tropicale (Kohlm.) E.B.G. Jones & K.L. Pang
Moleospora Abdel-Wahab, Abdel-Aziz & Nagah., Mycol. Progr. 9(4): 547 (2010)
Index Fungorum number: IF543116; 1 species with sequence data.
Type species – Moleospora maritima Abdel-Wahab, Abdel-Aziz & Nagah.
693
Figure 153 – Hydea pygmea (Material examined – JAPAN, Shiira Riv., Iriomote, Bruguiera
gymnrrhiza., on fallen leaf, July 8 1997, I. okane, IFO-H 12229). a Herbarium label. b, c
Appearance of conidia and conidiophores on CMSWA. d-f Conidia attached to the conidiophores.
g-i Helicoid conidia. Scale bars: d-f = 20 μm, g-i = 10 μm, j-n = 5 μm.
Moromyces Abdel-Wahab, K.L. Pang, Nagah., Abdel-Aziz & E.B.G. Jones, Mycol. Progr. 9(4):
555 (2010)
Index Fungorum number: IF543127; 1 species with sequence data.
Type species – Moromyces varius (Chatmala & Somrith.) Abdel-Wahab, K.L. Pang, Nagah.,
Abdel-Aziz & E.B.G. Jones
694
Orbimyces Linder, Farlowia 1: 404 (1944)
Index Fungorum number: IF9170; 1 species with sequence data.
Type species – Orbimyces spectabilis Linder
Rostrupiella Jørg. Koch, K.L. Pang & E.B.G. Jones, Bot. Mar. 50(5-6): 295 (2007)
Index Fungorum number: IF29175; 1 species with sequence data.
Type species – Rostrupiella danica Jørg. Koch, K.L. Pang & E.B.G. Jones
Sammeyersia S.Y. Guo, E.B.G. Jones & K.L. Pang, Botanica Marina 15 (2017)
Index Fungorum number: IF820458; 1 species with sequence data.
Type species – Sammeyersia grandispora (Meyers) S.Y. Guo, E.B.G. Jones & K.L. Pang
Notes – Sammeyersia was introduced by Abdel-Wahab et al. (2017) to accommodate
Sammeyersia grandispora which was earlier Lulworthia grandispora. Sammeyersia grandispora is
identified by the length of the ascospores over 400 μm (Abdel-Wahab et al. 2017). Our new isolate
groups with the S. grandispora isolates deposited in GenBank (Fig. 16). Therefore, based on both
morphological and phylogenetic data we identified our new isolate as S. grandispora. Sammeyersia
grandispora is illustred herein (Fig. 154).
Macrohilaceae Crous, IMA Fungus 6(1): 180 (2015)
Index Fungorum number: IF812795; Facesoffungi number: FoF06269; 1 species.
Endophytic or saprobic on leaves of Myrtaceae (dicotyledons). Sexual morph: Undetermined.
Asexual morph: Coelomycetous. Conidiomata pycnidial, immersed, becoming erumpent, medium
brown, globose to irregular. Conidiomata wall composed of pale brown-walled cells of textura
globulosa. Conidiophores reduced to conidiogenous cells. Conidiogenous cells pale brown,
cylindrical, percurrently proliferating at the apex with annellations. Conidia brown to dark brown,
1-septate, slightly constricted at septum, apex rounded, base flattened with a protruding hilum
(adapted from Maharachchikumbura et al. 2016b).
Type genus – Macrohilum H.J. Swart
Notes – The type species of Macrohilum, M. eucalypti formed a distinct phylogenetic lineage
in Diaporthales, thus Crous et al. (2015a) introduced Macrohilaceae to accommodate it. Subsequent
publications by Maharachchikumbura et al. (2015, 2016b), Wijayawardene et al. (2016b, 2018a)
and Senanayake et al. (2017a, 2018) accepted Macrohilaceae as a well-established family within
Diaporthales and basal to Diaporthaceae. Currently, Macrohilaceae comprises only one genus.
Ecological and economic significance of Macrohilaceae
Swart (1988a) reported Macrohilum eucalypti from dead Eucalyptus leaves, while Crous et
al. (2015a) isolated M. eucalypti from living leaves. However, neither Swart (1988a) nor Crous et
al. (2015a) reported whether it caused leaf spots or any other symptoms. But, both of these
publications did not carry out Koch's postulates to determine the pathogenicity of M. eucalypti. It
would be appropriate to carry out more sampling from diseased Eucalyptus leaves and perform
Koch’s postulates to determine whether Macrohilum species are pathogens as have been done for
Colletotrichum species on Artocarpus heterophyllus (Bhunjun et al. 2019).
Genus included in Macrohilaceae
Macrohilum H.J. Swart, Trans. Br. mycol. Soc. 90(2): 288 (1988)
Index Fungorum number: IF11123; 1 species with sequence data.
Type species – Macrohilum eucalypti H.J. Swart
Notes – The monotypic genus Macrohilum was introduced by Swart (1988a) with M.
eucalypti, the type species. Macrohilum resembles coniothyrium-like taxa but is phylogenetically
distinct as it has a protruding hilum (Crous et al. 2015a, Wijayawardene et al. 2016b). Furthermore,
the four base pairs differences observed in their ITS sequences suggest that the New Zealand
isolates may probably represent a novel Macrohilum species.
695
Figure 154 – Sammeyersia grandispora (Material examined – THAILAND, Phetchaburi Province,
Hat Chao Samran, 47° 72506’ E, 40° 25038’ N, 0 m asl., on intertidal decayed wood of Rhizopora
sp. at a mangrove stand, 28 August 2015, M. Dayarathne CHAM005, MFLU 16-1172). a Ascomata
on wood surface. b Section through ascoma in wood. c Peridium composed of thick-walled angular
cells. d, f Asci. g Filamentous ascospore. h Polar end chamber of ascospore. Scale bars: b, d, f, g =
100 μm, a = 50 μm, c, h = 10 μm.
696
Magnaporthaceae P.F. Cannon, Syst. Ascom. 13(1): 26 (1994)
Index Fungorum number: IF81963; Facesoffungi number: FoF01101; 135 species.
Pathogenic on monocotyledons or saprobic on plant material. Sexual morph: Ascomata
perithecial, solitary or scattered, black, superficial or immersed in plant tissue, globose to
subglobose, with cylindrical, black, periphysate neck. Peridium comprising a few to several layers
of cells of textura epidermoidea. Paraphyses hyaline, septate, intermingled among asci. Asci 8spored, unitunicate, subcylindrical, short-pedicellate, with an apical ring. Ascospores biseriate,
hyaline to olivaceous, filiform or fusoid, curved to sigmoid, with or without transverse septate ends,
bluntly rounded, lacking sheaths. Asexual morph: Hyphomycetous, at times formed from sclerotia.
Conidiophores unbranched or branched. Conidiogenous cells integrated, pigmented, phialidic with
collarettes, or denticulate. Conidia variable in shape, hyaline to pale brown, straight or curved, with
or without septa (adapted from Maharachchikumbura et al. 2016b).
Type genus – Nakataea Hara
Notes – Magnaporthaceae was introduced by Cannon (1994). The placement of taxa of
Magnaporthaceae has long been problematic due to the lack of convincing morphological and
inconclusive molecular data (Thongkantha et al. 2009). Based on DNA phylogenies,
Magnaporthaceae was placed as a family in the Sordariomycetes (Kirk et al. 2001, Lumbsch &
Huhndorf 2007, Maharachchikumbura et al. 2016b). Thongkantha et al. (2009) established a new
order, Magnaporthales to accommodate Magnaporthaceae based on characters and phylogenetic
analysis. The family was originally described with six genera (Cannon 1994). Kirk et al. (2001)
accepted nine genera while Kirk et al. (2008) accepted 13 genera. Luo et al. (2014) established
Pseudophialophoda in the family. Maharachchikumbura et al. (2016b) provided an updated
account with 22 genera and a key to species. A new genus, Bifusisporella was also introduced by
Silva et al. (2009). In this study we accept 23 genera in Magnaporthaceae based on publications and
taxonomy.
Ecological and economic significance of Magnaporthaceae
Most genera of Magnaporthaceae are necrotrophic and hemibiotrophic plant pathogens
infecting root and shoots of Poaceae and Cyperaceae (Zhang et al. 2011, Luo & Zhang 2013).
Endophytic or apparently saprotrophic taxa on non-gramineous hosts were added by Tibpromma et
al. (2018). Pyricularia oryzae (= Magnaporthe oryzae), a pathogen causing rice blast disease, was
accommodated in Pyriculariaceae, while Magnaporthe salvinii which causes stem rot in rice was
synonymized under Nakataea oryzae in Magnaporthaceae (Klaubauf et al. 2014). Other notable
pathogens of Magnaporthaceae include Gaeumannomyces graminis, Magnaporthiopsis poae and
M. rhizophila. Some members of Magnaporthaceae (e.g. Ophioceras and Ceratosphaeria) also
occur in aquatic habitats, or on dead plant material, such as wood (Shearer et al. 1999, Réblová
2006, Huhndorf et al. 2008, Thongkantha et al. 2009).
Genera included in Magnaporthaceae
Bifusisporella R.M.F. Silva, R.J.V. Oliveira, J.D.P. Bezerra, Souza-Motta & G.A. Silva, Mycol.
Progr. 18: 852 (2019)
Index Fungorum number: IF828222; 1 species with sequence data.
Type species – Bifusisporella sorghi R.M.F. Silva, R.J.V. Oliveira, J.D.P. Bezerra, SouzaMotta & G.A. Silva
Notes – Bifusisporella was introduced as an endophyte from healthy leaves of Sorghum
bicolor in Brazil based on morphology and phylogenetic analysis (Silva et al. 2019). It forms
phialides in culture consisting of curved, elongate and cylindrical-clavate conidiogenous cells, and
both macroconidia and microconidia (Silva et al. 2019). The sexual morph of Bifusisporella has not
been reported.
Budhanggurabania P. Wong, Khemmuk & R.G. Shivas, Persoonia, Mol. Phyl. Evol. Fungi 34:
241 (2015)
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Index Fungorum number: IF811696; 1 species with sequence data.
Type species – Budhanggurabania cynodonticola P. Wong, Khemmuk & R.G. Shivas
Notes – Budhanggurabania cynodonticola was introduced from diseased roots and stolons of
Cynodon in New South Wales, Queensland and the Northern Territory, Australia (Crous et al.
2015c, Wong et al. 2015). Sexual morphs of Budhanggurabania are characterized by ellipsoidal
ascospores, 3-dark brown septa, brown in central cells, hyaline to pale brown in apical cells, with
oblique striations in lateral view (Crous et al. 2015c). Hyphomycetous asexual morphs are
characterized by hyaline conidiophores, phialidic conidiogenous cells, straight to slightly curved,
and conidia that are hyaline, cylindrical or slightly curved, aseptate and aggregated in slimy heads
(Crous et al. 2015c).
Figure 155 – Macrohilum eucalypti (a. DAR 59000, holotype; b-g. CPC 19421, epitype). a
Conidioma. b Developing conidium attach to conidiogenous cell. c Developed conidia attach to
conidiogenous cells. d-g Conidia. Scale bars: a = 100 μm, b-f = 5 μm (a is extracted from
Wijayawardene et al. 2016b, which was redrawn from Swart 1988a).
Buergenerula Syd., Annls mycol. 34(4/5): 392 (1936)
Index Fungorum number: IF679; 5 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Buergenerula biseptata (Rostr.) Syd.
Notes – Buergenerula biseptata (≡ Metasphaeria biseptata) was found on Caricis vesicaria
in Norway. The genus is characterized by cylindrical to subfusiform asci, and clavate, hyaline
ascospores with 3–4-septa. The asexual morph has not been reported.
Bussabanomyces Klaubauf, M.-H. Lebrun & Crous, Stud. Mycol. 79: 99 (2014)
Index Fungorum number: IF810195; 1 species with sequence data.
Type species – Bussabanomyces longisporus (Bussaban) Klaubauf, M.-H. Lebrun & Crous
Notes – Bussabanomyces longispora (≡ Pyricularia longispora) was isolated as an endophyte
from leaves of Amomum siamense. The genus is characterized by macronematous conidiophores,
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cylindrical and denticulate conidiogenous cells, obclavate hyaline to pale brown, 4(–5)-septate
conidia. The sexual morph has not been reported (Klaubauf et al. 2014). Bussabanomyces is
morphologically similar to Pyricularia, however, it is distinct in its unbranched conidiophores and
terminal conidiogenous cells giving rise to 4(–5)-septate, pale brown conidia (Bussaban et al. 2003,
Klaubauf et al. 2014).
Figure 156 – Budhanggurabania cynodonticola (Material examined – AUSTRALIA, New South
Wales, from rotted roots of Cynodon dactylon, April 2012, Wong P.T.W., BRIP 59305 a, holotype)
a Herbarium packet. b, c Colonies on PDA (b-from above, c-from below). d, e Colonies and
mycelium on PDA. f-i Conidia with slimy heads (from slides). j Ascomata. k Asci with ascospores.
l Ascospores. Scale bars: f-i = 20 µm, j = 100 µm, k, l = 10 µm.
Ceratosphaerella Huhndorf, Greif, Mugambi & A.N. Mill., Mycologia 100(6): 941 (2008)
Index Fungorum number: IF508749; 2 species with sequence data.
Type species – Ceratosphaerella castillensis (C.L. Sm.) Huhndorf, Greif, Mugambi & A.N.
Mill.
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Figure 157 – Buergenerula biseptata (Material examined – NORWAY, Carex vesicaria, 1885,
Werenskiold F., O-F72662 (holotype of Metasphaeria biseptata) Sydow, H. 1936. Mycotheca
Germanica Fasc. LVII-LX no. 2801-3000). a-c Herbarium packet. d, e Fruiting bodies on substrate.
f Ascomata. g-l Asci. m Asci amoung with paraphyses. n Apical ring. o-r Ascospores. Scale bars: f
= 200 µm, g-m = 50 µm, n = 5 µm, o = 30 µm, p-r = 10 µm.
Notes – Huhndorf et al. (2008) introduced Ceratosphaerella castillensis (≡ Ceratosphaeria
castillensis) from bark of Castillo viejo and described a new species, C. rhizomorpha from
decaying wood in Kenya. The sexual morph of genus is characterized by clavate asci, fusiform and
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hyaline ascospores with 3-septa. The asexual morph has synnemata and ellipsoid to cylindrical
conidia, that are pale to darker brown, and 1–3-septate.
Clasterosporium Schwein., Trans. Am. phil. Soc., New Series 4(2): 300 (1832)
Index Fungorum number: IF7685; 41 morphological species (Species Fungorum 2020), 7
species with sequence data.
Type species – Clasterosporium caricinum Schwein.
Notes –The asexual morph of Clasterosporium is characterized by dark hypha, brownish
conidiophores, cylindrical or spindle-shaped, 3(+)-celled conidia. The sexual morph of
Clasterosporium has reported based on a two species in Clasterosphaeria, and later
Clasterosphaeria was synonymised under Clasterosporium (Zhang et al. 2016). The genus sexual
characters is superficial, globose, dark brown to black, ostiolate ascomata, later collapsing, 8spored asci and hyaline to subhyaline, fusiform, 3-septate ascospores (Sivanesan 1984, Hosagoudar
2004). However, these genera have to be recollected and sequenced to confirm their placement.
Clavatisporella K.D. Hyde, Mycotaxon 55: 276 (1995)
Index Fungorum number: IF27472; 1 morphological species.
Type species – Clavatisporella musicola K.D. Hyde, Mycotaxon 55: 276 (1995)
Notes – The monophyletic genus, Clavatisporella was introduced for a saprobe on senescent
leaves of Musa in Indonesia. The genus is characterized by immersed, globose to subglobose
ascomata, with broad cylindrical or clavate asci, with a subapical, J-, refractive ring and hyaline
ascospores that are fusiform to clavate, 1–2 septate, with a swollen base, and a mucilaginous padlike appendages at an apical (Hyde 1995e).
Falciphora J. Luo & N. Zhang, Mycologia 107(3): 643 (2015)
Index Fungorum number: IF810801; 1 species with sequence data.
Type species – Falciphora oryzae Z.L. Yuan, Chu L. Zhang & F.C. Lin
Notes – This monotypic genus was established based on F. oryzae (≡ Harpophora oryzae)
isolated from root tissues of Oryza granulata seedlings in Yunnan (Luo et al. 2015). The genus has
densely branched conidiophores, phialidic and straight conidiogenous cells and sickle-shaped and
strongly curved aseptate, hyaline conidia (Luo et al. 2015).
Falciphoriella Hern.-Restr. & Crous, Stud. Mycol. 83: 9 (2016)
Index Fungorum number: IF816902; 1 species with sequence data.
Type species – Falciphoriella solaniterrestris Hern.-Restr. & Crous
Notes – Falciphoriella solaniterrestris was isolated from soil in a potato field in the
Netherlands. The genus is characterized by phialidic, subcylindrical to elongated and fusoidellipsoid conidiogenous cells and fusiform to obovoid slightly curved conidia. The sexual morph
has not been reported.
Gaeumannomycella Hern.-Restr. & Crous, Stud. Mycol. 83: 9 (2016)
Index Fungorum number: IF816904; 2 species with sequence data.
Type species – Gaeumannomycella caricis Hern.-Restr. & Crous
Notes – Gaeumannomycella caricis was found on Carex rostrata in the UK. The genus
produces phialidic conidiogenous cells that are straight or curved, ampulliform to lageniform or
conical shaped, with lunate or cylindrical and hyaline conidia, and hyaline hyphopodia that become
brown when mature (Hernández-Restrepo et al. 2016b). Crous et al. (2019a) introduced a second
species, G. caricicola, found on a dead leaf of Carex remota in Germany.
Gaeumannomyces von Arx & D.L. Olivier, Trans. Br. mycol. Soc. 35: 32 (1952)
Index Fungorum number: IF2026; 20 morphological species (Species Fungorum 2020), 12
species with sequence data.
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Type species – Gaeumannomyces graminis (Sacc.) von Arx & D.L. Olivier
Notes –The sexual morph of this genus is characterized by globose or pyriform and immersed
ascomata with a conical to cylindrical neck, and fusiform, hyaline, multiseptate ascospores.
Asexual morphs are characterised by phialidic conidiogenous cells with refractive collarettes and
lunate or phialophora-like conidia.
Herbampulla Scheuer & Nograsek, Mycotaxon 47: 419 (1993)
Index Fungorum number: IF22427; 1 morphological species.
Type species – Herbampulla crassirostris Scheuer & Nograsek
Notes –Herbampulla crassirostris was found on dead and rotten leaves of Carex firma in
Austria. The genus produces cylindrical asci and fusiform 5-septate and yellowish brown
ascospores. The asexual morph has not been reported.
Kohlmeyeriopsis Klaubauf, M.-H. Lebrun & Crous, Stud. Mycol. 79: 101 (2014)
Index Fungorum number: IF810197; 1 species with sequence data.
Type species – Kohlmeyeriopsis medullaris (Kohlm., Volkm.-Kohlm. & O.E. Erikss.)
Klaubauf, M.-H. Lebrun & Crous
Notes – Kohlmeyeriopsis medullaris (≡ Gaeumannomyces medullaris) was reported on
senescent culms of Juncus roemerianus in North Carolina. The genus has trichocladium-like
asexual morphs (Klaubauf et al. 2014). The sexual morph has ellipsoid ascomata, fusoid to
cylindrical asci with a large apical ring staining in Meltzer’s reagent, and filamentous, hyaline
ascospores, which produce appressoria when germinating (Klaubauf et al. 2014).
Magnaporthiopsis J. Luo & N. Zhang, Mycologia 105: 1021 (2013)
Index Fungorum number: IF802972; 7 species with sequence data.
Type species – Magnaporthiopsis poae (Landsch. & N. Jacks.) J. Luo & N. Zhang
Notes – Magnaporthiopsis was introduced as a phialophora-like asexual genus. It was
identified to Magnaporthaceae based on phylogenetic analyses of multi-genes (SSU, ITS, LSU,
MCM7, rpb1, tef1) (Luo & Zhuang 2012). The sexual morph characters are globose ascomata with
a cylindrical neck and black, clavate asci with a refractive ring and fusiform to fusoid, septate,
hyaline or yellow brown ascospores (Luo & Zhuang 2012).
Muraeriata Huhndorf, Greif, Mugambi & A.N. Mill., Mycologia 100: 948 (2008)
Index Fungorum number: IF508755; 2 species with sequence data.
Type species – Muraeriata collapsa Huhndorf, Greif, Mugambi & A.N. Mill.
Notes – Muraeriata was introduced to accommodate two species found on wood, M. africana
(from Kenya) and M. collapsa (from Costa Rica). The genus is characterised by the vacuolate
middle ascomal wall layer and by phylogenetic analysis of LSU and SSU (Huhndorf et al. 2008).
Mycoleptodiscus Ostaz., Mycologia 59(6): 970 (1968)
Index Fungorum number: IF9029; 19 morphological species (Species Fungorum 2020), 5
species with sequence data.
Type species – Mycoleptodiscus terrestris (Gerd.) Ostaz.
Notes – The genus has superficial, sporodochial conidiomata, one cell thick, mostly thickwalled, dark brown conidiogenous cells with a prominent circular aperture in the upper wall,
hyaline conidia with cylindrical to fusiform shaped, and 0–2-septa, sometimes bearing an apical
and/or basal cellular unbranched filiform appendage (Katsuhiko 1996). Mycoleptodiscus
endophyticus was described as an endophyte in leaves of Freycinetia sp. (Tibpromma et al. 2018).
Nakataea Hara, Diseases Rice Plant, Edn 2: 185 (1939)
Index Fungorum number: IF565608; 6 morphological species (Species Fungorum 2020), 1
species with sequence data.
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Type species – Nakataea oryzae (Catt.) J. Luo & N. Zhang
Notes – Nakataea causes stem rot of rice (Ou 1985). The genus was introduced based on N.
oryzae (= N. sigmoidea) from Oryza sativa in Italy. Krause & Webster (1972) showed Nakataea
and Magnaporthe are congeneric with their type species, N. oryzae and M. salvinii belonging to the
same species. Therefore, Luo & Zhang (2013) synonymized M. oryzae under N. oryzae based on
the name Nakataea (1939), which is older than Magnaporthe (1972) This resulted in Nakataea
becoming the correct name for the type of the family (Murata et al. 2014, Klaubauf et al. 2014, Luo
et al. 2105, Maharachchikumbura et al. 2015b, 2016b). Khemmuk et al. (2016) confirmed the two
genera as congeneric, based on phylogenetic analysis of multi-genes (ITS, LSU, tef1, rpb1).Only
Nakataea oryzae has molecular data.
Neogaeumannomyces D.Q. Dai & K.D. Hyde, Fungal Divers. 72: 34 (2015)
Index Fungorum number: IF550936; 1 species with sequence data.
Type species – Neogaeumannomyces bambusicola D.Q. Dai & K.D. Hyde
Notes – The genus is monotypic with Neogaeumannomyces bambusicola found dead culm of
bamboo in Thailand (Liu et al. 2015). The genus is similar to Gaeumannomyces in perithecial
ascomata with long necks, cylindrical asci and filiform ascospores. However, Neogaeumannomyces
lacks hyphopodia whereas Gaeumannomyces species produce hyphopodiate mycelia (Walker 1972,
Liu et al. 2015).
Omnidemptus P.F. Cannon & Alcorn, Mycotaxon 51: 483 (1994)
Index Fungorum number: IF27275; 3 morphological species (Species Fungorum 2020), 2
species with sequence data.
Type species – Omnidemptus affinis P.F. Cannon & Alcorn
Notes – Omnidemptus affinis was found on leaves of Panicum effusum in Queensland
(Cannon & Alcorn 1994, Thongkantha et al. 2009). The sexual morph is characterized by perithecia
ascomata, cylindrical to clavate, short pedicellate asci with an obtuse apex and an apical pore that
stains dark blue with Melzer’s reagent and fusiform, 1–3-septate, hyaline ascospores (Cannon &
Alcorn 1994). The asexual morph is sporodochium-like, with cylindrical, clavate or ellipsoid
conidiogenous cells, and falcate, 2-septate conidia, the base of conidia sometimes with an abrupt
change in contour to short pedicel-like extension (Hernández-Restrepo et al. 2019).
Pseudophialophora J. Luo & N. Zhang, Mycologia 106(3): 581 (2014)
Index Fungorum number: IF807080; 9 species with sequence data.
Type species – Pseudophialophora eragrostis J. Luo & N. Zhang
Notes –Luo et al. (2014) confirmed the placement of Pseudophialophora in Magnaporthaceae
by phylogenetic analysis. The type species was isolated from roots of Eragrostis in New Jersey.
The genus is phialophora-like, producing dark septate hyphae, simple and short conidiophores,
typical phialides and unicellular conidia (Luo et al. 2014). (Crous et al. 2017) provided a
phylogenetic update to nine species..
Pyriculariopsis M.B. Ellis, Dematiaceous Hyphomycetes (Kew): 206 (1971)
Index Fungorum number: IF9671; 9 morphological species (Soares et al. 2011, Species
Fungorum 2020), 2 species with sequence data.
Type species – Pyriculariopsis parasitica (Sacc. & Berl.) M.B. Ellis
Notes – Pyriculariopsis parasitica (= Pyricularia musae) was recognized as differing from
Pyricularia by the absence of a septum to separate its denticles from the conidiophore (Ellis 1971).
In a molecular study, Bussaban et al. (2005) found Pyriculariopsis was outside a clade of
Magnaporthaceae. However, they concluded that Pyriculariopsis should be maintained based
mainly in its straight or curved, obclavate and rostrate conidia. Soares et al. (2011) suggested an
additional feature to delimit Pyriculariopsis, which was consistently observed or illustrated for
most species, as pointed out by Whitton et al. (2001). Pyriculariopsis contains two distinct
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ecological groups, one is composed of saprobes occurring on foliar and stem debris and the other
includes plant pathogens on monocot hosts in Zingiberales (Soares et al. 2011, Uchida & Kadooka
1994).
Slopeiomyces Klaubauf, M.-H. Lebrun & Crous, Stud. Mycol. 79: 102 (2014)
Index Fungorum number: IF810199; 1 species with sequence data.
Type species – Slopeiomyces cylindrosporus (D. Hornby, Slope, Gutter. & Sivan.) Klaubauf,
M.-H. Lebrun & Crous
Notes – Klaubauf et al. (2014) described the genus based on morphology of its
hyphomycetous asexual morph and phylogenetic analysis placement in Magnaporthaceae.
Malaysiascaceae Tibpromma & K.D. Hyde, Fungal Divers. 93: 88 (2018)
Index Fungorum number: IF554753; Facesoffungi number: FoF04611; 1 species.
Saprobic on dead or decaying leaves of plants in terrestrial habitats. Sexual morph: Ascomata
perithecial, immersed at the base, obpyriform, dark brown, with setae. Peridium composed of 3–5
layers of pale brown cells of textura prismatica. Paraphyses branched, septate. Asci 8-spored,
unitunicate, cylindrical-clavate, short pedicellate, apex truncate with shallow ring. Ascospores biseriate, hyaline, ellipsoid to oblong, apiculate at ends with minute mucoid cap, smooth-walled,
guttulate, becoming 1-septate and pale brown after discharge. Asexual morph: Conidiophores
subcylindrical, macronematous, unbranched, erect, flexuous, thick-walled, rounded at apex,
guttulate, multi-septate, pale brown to bark brown, smooth-walled. Conidiogenous cells
enteroblastic, phialidic, subcylindrical, subhyaline, terminal, integrated. Conidia solitary,
cylindrical-ellipsoid to clavate, rounded at apex, aseptate, hyaline, smooth-walled, guttulate, dry in
mass (adapted from Tibpromma et al. 2018).
Type genus – Malaysiasca Crous & M.J. Wingf.
Notes – Malaysiascaceae was introduced by Tibpromma et al. (2018) to accommodate
Malaysiasca which belong in Glomerellales. The family is similar to Australiascaceae, but the
latter differs in having phialidic conidiogenesis with hyaline 0(–3)-septate conidia aggregated in
slime or chains (Réblová et al. 2011).
Ecological and economic significance of Malaysiascaceae
Malaysiascaceae species are saprobic on dead or decaying leaves or wood in terrestrial
habitats (Crous et al. 2016b, Tibpromma et al. 2018) involved in nutrient cycling.
Genus included in Malaysiascaceae
Malaysiasca Crous & M.J. Wingf., Persoonia 36: 373 (2016)
Index Fungorum number: IF817044; 1 species with sequence data.
Type species – Malaysiasca phaii Crous & M.J. Wingf.
Notes – Malaysiasca was established by Crous et al. (2016b) to accommodate Malaysiasca
phaii, based on LSU gene analysis, and is related to members of Glomerellales. Tibpromma et al.
(2018) found that Malaysiasca phaii belongs in Glomerellales forming a well-supported clade with
other families and introduced a new family as Malaysiascaceae. This family is monotypic
comprising only Malaysiasca phaii, which was found on leaves of Phaius reflexipetalus (Crous et
al. 2016b) and Freycinetia javanica (Tibpromma et al. 2018).
Melanconidaceae G. Winter, Rabenh. Krypt.-Fl., Edn 2 (Leipzig) 1.2: 764 (1886)
Index Fungorum number: IF80988; Facesoffungi number: FoF01395; 36 species.
Saprobic or pathogenic on plants and wood, in terrestrial and aquatic habitats. Sexual morph:
Pseudostromata well-developed, obvious, erumpent. Ectostromatic disc surrounded by bark or not,
yellowish-white. Ostiolar canal opening around the disc. Ascomata arranged as circles around the
ectostromatic disc, oblique or horizontal, globose to subglobose, coriaceous and black with long,
periphysate and lateral ostiolar canals. Peridium with outer, thick-walled, brown cells of textura
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globosa to textura angularis and inner, thick-walled, flat, hyaline cells of textura angularis.
Paraphyses wide, hypha-like, deliquescent at maturity. Asci 8-spored, unitunicate, oblong to
fusiform, short pedicellate, with distinct, J-, apical ring. Ascospores overlapping uniseriate to
biseriate, hyaline, ellipsoid, 1-septate. Asexual morph: Coelomycetous. melanconium-like.
Conidiomata acervular, scattered, solitary, superficial, black, coriaceous. Conidiophores branched
at the base, septate, Conidiogenous cells annellidic, cylindrical. Conidia hyaline to brown, ellipsoid
or subglobose, smooth-walled, thick-walled (adapted from Senanayake et al. 2017a).
Figure 158 – Malaysiasca phaii (Material examined – THAILAND, Krabi Province, Khlong Thom
District, on dead leaves of Freycinetia javanica Blume., 15 December 2015, S. Tibpromma KB032,
MFLU 16-1910). a Colony on natural substrate. b Conidiogenous cells with conidiophores and
conidia. c Conidiophores. d-g Conidiogenous cells with conidia. h, i Conidia. Scale bars: a = 200
μm, b = 100 μm, c-g = 10 μm, h, i = 5 μm.
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Type genus – Melanconis Tul. & C. Tul.
Notes – Melanconidaceae includes species with yellowish-white ectostromatic discs,
surrounding the ascomata which are arranged in a circle. Most of the family are saprobes and
pathogens which cause diseases on economically important trees. Maharachchikumbura et al.
(2016b) included 24 genera in the family. Sequence data is available for Dicarpella, Melanconiella,
Melanconis, Melanconium, and Prosthecium. Voglmayr & Jaklitsch (2014) synonymized
Prosthecium under Stilbospora and included the genus in Stilbosporaceae. Crous et al. (2012d)
included more than half of known wuestneia-like species in Harknessia in Harknessiaceae.
Morphological and molecular evidence showed that the family was monogeneric represented by
Melanconis and its asexual morph Melanconium (Castlebury et al. 2002; Rossman et al. 2007).
Melanconium was synonymized under Melanconis by Rossman et al. (2015). Phylogenetic analysis
of Senanayake et al. (2017a) showed that Dicarpella and Melanconiella clustered away from the
family. As a result, Senanayake et al. (2017a) excluded all the genera listed in
Maharachchikumbura et al. (2016b) from Melanconidaceae, except Melanconis.
Figure 159 – Melanconis betulae (Material examined – CHINA, Gansu Province: Gannan Tibetan
Autonomous Prefecture, Zhouqu County, Qiban Forestry Centre, 33°56′35.36″N, 104°07′13.03″E,
3221 m asl, on twigs and branches of Betula albosinensis Burkill, 20 August 2014, Y.M. Liang,
BJFC-S1319, holotype). a-c Appearance on host. d Sections through conidiomata. f Conidiophores
and alpha conidia. g, h Alpha conidia. i, j Beta conidia. Scale bars: b-e = 500 μm, f-h =10 μm.
Ecological and economic significance of Melanconidaceae
Most sexual and asexual morphs of Melanconis cause cankers and shoot blights (Fan et al.
2016, Senanayake et al. 2018) with various disease symptoms (Fan et al. 2016). Melanconis
stilbostoma causes birch dieback (Fan et al. 2016) causing branches in the crown to die off. Some
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taxa produce antifungal compounds. Thermozymocidin (myriocin) and flavovirin were isolated
from Melanconis flavovirens (Sailer et al. 1989) and have shown antifungal activity against yeasts
and a few filamentous fungi.
Genus included in Melanconidaceae
Melanconis Tul. & C. Tul., Select. fung. carpol. (Paris) 2: 115 (1863)
Index Fungorum number: IF3060; 36 morphological species (Species Fungorum 2020), 8
species with sequence data.
Type species – Melanconis stilbostoma (Fr.) Tul. & C. Tul.
Notes – The genus comprises saprobic, pathogenic (Senanayake et al. 2018) and endophytic
(Barengo et al. 2000) species associated with the hardwood of plants in Betulaceae (Fan et al.
2016). The genus is characterised by ascomata with 8-spored asci, ellipsoid, 1-septate ascospores,
and acervular conidiomata with hyaline to brown, ellipsoid or subglobose conidia. Most of the
species are known in North America and Europe (Senanayake et al. 2017a). In this entry we
illustrate Melanconis betulae.
Melanconiellaceae Senan., Maharachch. & K.D. Hyde, Stud. Mycol. 86: 275 (2017)
Index Fungorum number: IF821561; Facesoffungi number: FoF03495; 41 species.
Phytopathogenic or saprobic. Sexual morph: Stromata present or absent. If present;
Pseudostromata inconspicuous, erumpent, pale or dark coloured ectostromatic disc or pulvillus
causing a more or less pustulate bark surface. Ectostromatic disc convex, flat to concave which can
be surrounded by bark or not. Central column beneath the disc is more or less conical, having
hyaline or pigmented hyphae. The hyphae are mixed with a pigmented powdery amorphous
substance in cream, yellow, olive, brownish or grey. Ascomata perithecial, epiphyllous without
stromatic tissues and immersed in host substrate, inconspicuous or appearing as rounded bumps
beneath the bark that surrounds the ectostromatic disc, oblique or horizontal, scattered or often
arranged in a circle around the central column, with long lateral ostioles that converge at the margin
of the central column. Ostioles emerging in various positions in the ectostromatic disc. Peridium
bears dark, thick-walled cells of textura angularis. Paraphyses broad, hyaline or lacking. Asci 2–8spored, unitunicate, cylindrical-clavate, oblong or fusoid, with a distinct, J-, apical ring, tapering
below to a short, narrow pedicel. Ascospores 1–2- overlapping seriate, hyaline, yellowish or brown,
oblong, fusoid or ellipsoid, 0–1-septate, septa central or slightly eccentric, slightly constricted or
not, smooth-walled, with or without short, blunt appendages and sometimes with a narrow
gelatinous sheath. Asexual morph: Coelomycetous. Conidiomata acervular or pycnidia, punctiform,
subcuticular, immersed or erumpent, sometimes with a central, well-developed, pale brown,
pseudoparenchymatous layer becoming thinner or absent at the margin of the conidiomata,
multiloculate, sometimes papillate, sometimes with pale coloured, ectostromatic disc and central
column or with radiate scutella. Scutella convex, membranous, brown, somewhat translucent, with
a central hyaline or pale disc, giving rise to radiating hyphae, thick-walled cells radiating from a
central point, rounded to pointed at the tips. Conidiophores often reduced to conidiogenous cells or
branched, sometimes septate only at the base, few-celled, smooth, hyaline to pale brown,
sometimes short, forming under the developing scutellum. Pseudoparaphyses filiform.
Conidiogenous cells annellidic or phialidic. Conidia initially hyaline becoming brown, ellipsoid,
obovoid, subglobose, ovoid or oblong, thick-walled, smooth to finely verrucose, with or without
distinct hyaline sheath, each with a truncate base and obtuse to bluntly pointed apex, sometimes
somewhat granular, sometimes with inconspicuous to conspicuous basal hilum, with or without
distinct hyaline sheath or frill (adapted from Senanayake et al. 2017a).
Type genus – Melanconiella Sacc.
Notes – Senanayake et al. (2017a) revised the family to accommodate the genera Dicarpella
(previously included in Melanconidaceae), Greeneria, Melanconiella, and Sphaeronaemella
fragariae (was in Microascales incertae sedis), Tubakia (previously Diaporthales incertae sedis),
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and a novel genus Microascospora. These taxa formed a distinct clade with moderate support in the
phylogenetic analysis and a new family was proposed.
Dicarpella was introduced based on D. bina and the asexual morph of this genus was
reported as Tubakia (Belisario 1991). Tubakia is typified by T. japonica. It was found that the type
species of these two genera are not linked. However, molecular data linked Tubakia and Diplacella
coupled with a few Diplacella species having Tubakia asexual morphs (Sogonov et al. 2008).
Tubakia is more commonly encountered than Dicarpella and is also a more widely used name than
Dicarpella. Senanayake et al. (2017a) showed a plausible relationship of Dicarpella dryina and
Tubakia seoraksanensis as a holomorphic genus. Senanayake et al. (2017a) mentioned without
analysing sequence data of the type species that it is hard to confirm that Dicarpella and Tubakia
are congeneric. Therefore, Senanayake et al. (2017a) maintained Dicarpella and Tubakia as two
separate genera until further sequence data becomes available. Senanayake et al. (2017a)
introduced a new genus Microascospora to this family with M. rubi as the type species.
Sphaeronaemella fragariae did not cluster in this analysis with other Sphaeronaemella species and
clustered with Microascospora rubi. Therefore, Sphaeronaemella fragariae was excluded from
Sphaeronaemella and placed in Microascospora as M. fragariae. Senanayake et al. (2017a)
therefore included Dicarpella, Greeneria, Melanconiella, Microascospora and Tubakia in
Melanconiellaceae. Braun et al. (2018) revised Tubakia using morphological and molecular data.
They sequenced the type species of Tubakia (T. dryina) and introduced a new family Tubakiaceae
to accommodate Tubakia. Braun et al. (2018) considered Tubakia suttoniana and Dicarpella dryina
as synonyms..
Senanayake et al. (2018) included Massariovalsa in Melanconiellaceae based on
morphology, but Massariovalsa was not included in the phylogenetic analysis due to the lack of
molecular data. Senanayake et al. (2018) therefore placed Greeneria, Massariovalsa,
Melanconiella and Microascospora in Melanconiellaceae. In the phylogenetic analysis of Fan et al.
(2018), Melanconiella cornuta formed a distinct clade basal to Melanconiella in the family. Based
on morphology and various host affinities (Cornus and Juglans vs. Betulaceae), M. cornuta was
excluded from Melanconiella by Fan et al. (2018) and placed in a new genus Sheathospora.
Phookamsak et al. (2019) included Septomelanconiella as a new genus in Melanconiellaceae based
on Septomelanconiella thailandica. Therefore, Melanconiellaceae presently accommodates the
genera Dicarpella, Greeneria, Massariovalsa, Melanconiella, Microascospora, Septomelanconiella
and Sheathospora.
Ecological and economic significance of Melanconiellaceae
Some species of the family are phytopathogens. Greeneria uvicola is a common pathogen
responsible for various diseases in grapes, such as grapevine dead-arm dieback, grapevine trunk
disease (Navarrete et al. 2009) and bitter-rot of grapes (Farr et al. 2001). The taxon occurs on stem
lesions, mummified berries, leaves, and tendrils. Greeneria uvicola infects several species of Vitis
(grapes), including V. aestivalis, V. labrusca, V. rotundifolia, and V. vinifera. (Navarrete et al.
2009). Melanconiella species are mainly restricted to overwintered plants and are able to cause
mild cankers on the hosts (Voglmayr et al. 2012).
Genera included in Melanconiellaceae
Dicarpella Syd. & P. Syd., Annls mycol. 18(4/6): 181 (1921)
Index Fungorum number: IF1512; 5 morphological species (Species Fungorum 2020).
Type species – Dicarpella bina (Harkn.) Syd. & P. Syd.
Notes – Dicarpella has seven species epithets in Index Fungorum (2020). Dicarpella
georgiana was transferred to Mastigosporella as M. georgiana (Barr 1978). Wuestniopsis was
proposed to accommodate Dicarpella georgiana and D. quercifolia (Reid & Dowsett 1990), while
Dicarpella dryina has been linked to Tubakia seoraksanensis (Senanayake et al. 2017a, Braun et al.
2018). Therefore, currently Dicarpella comprises only D. bina, D. liquidambaris-styracifluae and
D. rubicola. Sequence data for the type species are unavailable. The asexual morph of Dicarpella
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was reported as Tubakia (Belisario 1991) and a few Tubakia species have diplacella-like sexual
morphs (Sogonov et al. 2008, Senanayake et al. 2017a, 2018, Braun et al. 2018). Due to the
unavailability of sequence data for the type species in both genera, it cannot be confirmed that
Dicarpella and Tubakia are congeneric. Hence, Senanayake et al. (2017a) placed this genus in
Melanconiellaceae, which we follow here until molecular data is available.
Greeneria Scribn. & Viala, C. r. hebd. Séanc. Acad. Sci., Paris 105: 473 (1887)
Index Fungorum number: IF8399; 1 species with sequence data.
Type species – Greeneria uvicola (Berk. & M.A. Curtis) Punith.
Notes – Greeneria was introduced based on G. fuliginea (Scribner & Viala 1887) and
synonymized under Melanconium (Cavara 1889) as M. fuligineum. van derAa (1973) placed the
genus in Phyllosticta as P. ampelicida. Punithalingam (1974) renamed the taxon as Greeneria
uvicola with descriptions and illustrations. LSU rDNA sequence analysis (Farr et al. 2001) showed
that G. uvicola belonged in Diaporthales. Senanayake et al. (2017a) reported that G. uvicola and G.
saprophytica should be accommodated in Melanconiellaceae. Greeneria saprophytica did not have
a close affinity to G. uvicola and this needs further resolution. Greeneria does not have a sexual
morph (Zhang & Blackwell 2001). Based on Senanayake et al. (2018), Greeneria now has a single
species, G. uvicola.
Massariovalsa Sacc., Michelia 2 (no. 8): 569 (1882)
Index Fungorum number: IF3020; 13 morphological species (Senanayake et al. 2018).
Type species – Massariovalsa sudans (Berk. & M.A. Curtis) Sacc.
Notes – Massariovalsa is a saprobic genus found on woody bark. Massariovalsa was a
subgenus of Melanconis (Wehmeyer 1941). Petrak (1952b) and Muller & von Arx (1962) separated
the genus using the pycnidial asexual morph and the mucous epispore on the ascospores.
Massariovalsa sudans was considered to be the sexual morph of Melanconiopsis inquinans (type
species of Melanconiopsis) (Wehmeyer 1939, Rossman et al. 2015). Massariovalsa has integrated,
enteroblastic, phialidic conidiogenous cells. The stromatic disc is not present in Massariovalsa but
the conidiogenesis is similar to Melanconiella. Sequence data are not available for Massariovalsa
in GenBank. Therefore, Senanayake et al. (2018) accommodated Massariovalsa in
Melanconiellaceae based on similar morphology.
Melanconiella Sacc., Syll. fung. (Abellini) 1: 740 (1882)
Index Fungorum number: IF3059; 19 morphological species (Species Fungorum 2020), 16
species with sequence data.
Type species – Melanconiella spodiaea (Tul.) Sacc.
Notes – Melanconiella is restricted to recently dead corticated branches. The genus is hostspecific to the plant family Betulaceae as saprobes (Senanayake et al. 2018). Voglmayr et al. (2012)
revised Melanconiella using herbarium material and recently collected specimens. The
dissimilarities in morphology and phylogeny of Melanconiella and Melanconis were discussed by
Voglmayr et al. (2012). See Crous et al. (2016b) and Senanayake et al. (2018) for further details.
Melanconiella spodiaea is illustrated in this entry.
Microascospora Senan. & K.D. Hyde, Stud Mycol 86:217–296 (2017)
Index Fungorum number: IF821562; 1 species with sequence data.
Type species – Microascospora rubi Senan., Camporesi & K.D. Hyde
Notes – The sexual, saprobic genus Microascospora is phylogenetically distinct from other
genera in Melanconiellaceae. Species have comparatively small ascospores (< 20 µm long) with
wavy, filiform, long appendages. Sphaeronaemella fragariae (previously placed in Microascales
incertae sedis) clustered with Microascospora rubi and not with other Sphaeronaemella species
(incertae sedis in Microascales, Senanayake et al. 2017a). Therefore, S. fragariae was excluded
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from Sphaeronaemella and accommodated in Microascospora as M. fragariae (Senanayake et al.
2017a).
Figure 160 – Melanconiella spodiaea (Material examined – GERMANY, Berlin, Jungfernheide,
on bark of Carpinus sp. (Betulaceae) P. Sydow s. n., NY 02932468; SWITZERLAND, Kaferberg
und Uetliberg bei Zurich P. Sydow s. n., NY 02932469). a Herbarium packages. b, c Specimen. d
Vertical cross section of ascoma. d Peridium. e-i Asci. j Vertical cross section of conidioma. k-l
Conidiogenous cells and conidia. m Conidia. Scale bars: c = 1 mm, d-i =20 µm, J = 500 µm, k, l =
10 µm, m = 5 µm.
Septomelanconiella Samarak. & K.D. Hyde, Fungal Divers. Notes 95: 1–273 (2019)
Index Fungorum number: IF555301; 1 species with sequence data.
Type species – Septomelanconiella thailandica Samarak. & K.D. Hyde
Notes – Septomelanconiella is a monotypic genus distinguished from other species in
Melanconiellaceae by its 1-euseptate and luminate conidia. The genus is similar to Melanconiella
in its finely verrucose, brown mature conidia. Septomelanconiella thailandica forms an
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independent lineage, phylogenetically distint from other genera in Melanconiellaceae and this is in
agreement with the phylogeneties of Senanayake et al. (2017).
Sheathospora X.L. Fan, MycoKeys 42: 118 (2018)
Index Fungorum number: IF828429; 1 morphological species.
Type species – Sheathospora cornuta (C.M. Tian & Z. Du) X.L. Fan
Notes – Sheathospora was established to accommodate Melanconiella cornuta previously
included in the Melanconiella clade (Voglmayr et al. 2012, Du et al. 2017). The genus can be
differentiated from the other genera in Melanconiellaceae by its pycnidial conidiomata and conidia
with a distinct, hyaline sheath.
Meliolaceae W. Martin, Mycol. Pap. 15: 23(1946)
Index Fungorum number: IF80993; Facesoffungi number: FoF00741; 2371 species.
Colonies epiphyllous or hypophyllous. Hyphae superficial, brown, straight to flexuous,
septate, branched, reticulate, with hyphopodia and phialides. Hyphopodia alternate to unilateral or
opposite, 2-celled, brown. Setae straight to hooked, dark brown, forming on hyphae or ascomata.
Sexual morph: Ascomata superficial, globose to subglobose, or flattened. Peridium comprising
hyaline inner stratum, and dark brown outer stratum of textura angularis. Paraphyses evanescent.
Asci 2–4-spored, unitunicate, asci wall attenuated or broken when mature, without a fixed shape.
Ascospores ellipsoidal to oblong, 3–4-septate, constricted at the septum, hyaline when young,
becoming brown when mature. Asexual morph: Phialides ampulliform or flask-shaped on hyphae.
Conidiogenous cells formed directly from vegetative hyphae. Conidia unicellular, small and
hyaline (adapted from Cannon & Kirk 2007).
Type genus – Meliola Fr.
Figure 161 – Drawing of asexual morph of Meliolaceae.
Notes – Meliolaceae was established by Martin (1941) without a Latin diagnosis and later
validated by Hansford (1946). Roger (1953) placed the Meliolaceae with nine other families in
Hypocreales. This family has also been placed in Dothideales, Erysiphales, Meliolales,
Myriangiales and Hypocreales at various times (Martin 1941, Luttrell 1951, 1989, Roger 1953,
Ainsworth et al. 1971, Müller & von Arx 1973, Yarwood 1973, Barr 1976a, Eriksson 1981,
Hawksworth et al. 1983). Ainsworth et al. (1971) and Eriksson & Hawksworth (1993) listed 50
genera in the family; later studies listed 25 genera (Hawksworth et al. 1995), 22 genera (Kirk et al.
2008), and 26 genera (Lumbsch & Huhndorf 2010) and seven genera (Hongsanan et al. 2015). A
checklist provided by Zeng et al. (2017) included the number of species in each genus of
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Meliolaceae that are accepted in Index Fungorum and excluded some erroneous records. Species
with valid information were accepted, and records with only a name available was noted in that
study. Previous identifications of Meliolaceae were mainly based on host association and
morphology. A formula proposed by Beeli (1920) was applied and modified for rapid identification
among thousands of species in Meliolaceae. It used a numerical code to summarize key diagnostic
characters, such as ascospores, ascomata, setae and appressoria. Mueller et al. (1991) stated that the
basal cytoplasm in phialides have a conidiogenous function and conidia-like spores was observed
being extruded from terminal openings of phialides. The reason for lack of studies on phialides is
that they function only in some species, certain stages, or specific environmental conditions
(Mueller et al. 1991). Therefore, phialides are considered as the asexual morph of Meliolaceae.
Although members of Meliolaceae are unculturable, sequence data can be obtained as described in
Zeng et al. (2018). There are only 20 confirmed species of Meliolaceae with sequence data in
GenBank and it has not been determined if species are host-specific. In the future, analysis of
sequence data can replace the Beeli formula as a more reliable method to identify such groups, as
well as establishing host-specificity and the asexual morphs of Meliolaceae.
Ecological and economic significance of Meliolaceae
Species of Meliolales can cause significant effects on crops, such as reducing yield and
quality of fruits of Citrus (e.g. Meliola butleri, M. camelliae and M. citricola) (Rao 1969). They
produce hyphopodia that penetrate the leaf surface to gain nutrients from host plants which results
in a reduction of chlorophyll, starch, sugars, proteins and amino acids, without causing pathogenic
damage (Hosagoudar et al. 1997, Old et al. 2003, Rodríguez Justavino & Piepenbring 2007). They
could reduce photosynthesis by covering the host surface and increase the temperature and
respiration in those areas (Hosagoudar et al. 1997, Hosagoudar & Raju 2013, Hongsanan et al.
2015).
Genera included in Meliolaceae
Amazonia Theiss., Annls mycol. 11(6): 499 (1913)
Index Fungorum number: IF153; 65 morphological species (Species Fungorum 2020, Zeng et
al. 2017), 1 species with sequence data.
Type species – Amazonia psychotriae (Henn.) Theiss.
Notes – Amazonia shares similarities with genera in Meliolaceae based on its brown hyphae
with phialides, and brown, 4-septate ascospores, but differs in its flattened ascomata. It is also
similar with taxa of Microthyriaceae in the structure of ascomata, but differs in the hyphae and
ascospores.
Appendiculella Höhn., Sber. Akad. Wiss. Wien, Math.-naturw. Kl., Abt. 1 128(7-8): 556 (1919)
Index Fungorum number: IF289; 74 morphological species (Species Fungorum 2020, Zeng et
al. 2017), 1 species with sequence data.
Type species – Appendiculella calostroma (Desm.) Höhn.
Notes – Appendiculella is characterized by its conical appendages with transversely striate
walls that develop on the surface of the ascomata. Rodríguez Justavino et al. (2014) provided the
first sequence for this genus, and its phylogenetic placement is related with Asteridiella.
Asteridiella McAlpine, Proc. Linn. Soc. N.S.W. 22(1): 38 (1897)
Index Fungorum number: IF406; 365 morphological species (Species Fungorum 2020, Zeng
et al. 2017), 3 species with sequence data.
Type species – Asteridiella solani McAlpine
Notes – There are no setae or appendages on either the ascomata or hyphae in species of
Asteridiella. Asteridiella and Appendiculella are similar in morphology and related in phylogeny.
There are few sequences available for Asteridiella and Appendiculella and thus it is necessary to
verify their phylogenetic placement with more sequence data.
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Cryptomeliola S. Hughes & Piroz., Mycol. Pap. 174: 14 (1997)
Index Fungorum number: IF27826; 3 morphological species (Species Fungorum 2020, Zeng
et al. 2017).
Type species – Cryptomeliola orbicularis (Berk. & M.A. Curtis) S. Hughes & Piroz.
Notes – Fascicular hyphal setae with bulbous tips are the typical character to distinguish
Cryptomeliola species from other genera of Meliolaceae. Colonies are comprise densely reticulate
hyphae, which is similar with Asteridiella. Hosagoudar (2003) suggested not to include this genus
in Meliolales, due to lack of hyphopodia. However, Hongsanan et al. (2015) described the structure
of hyphopodia and placed it in Meliolales.
Endomeliola S. Hughes & Piroz., N.Z. Jl Bot. 32(1): 53 (1994)
Index Fungorum number: IF27386; 1 species with sequence data.
Type species – Endomeliola dingleyae S. Hughes & Piroz.
Notes – Endomeliola is the only genus lacking superficial hyphae in Meliolaceae. It contains
a single species with an unverified sequence in GenBank. The brown, 4-septate ascospores places
Endomeliola as a member of Meliolaceae. However, its placement needs to be confirmed with
reliable sequence data.
Irenopsis F. Stevens, Annls mycol. 25(5/6): 411 (1927)
Index Fungorum number: IF2505; 152 morphological species (Species Fungorum 2020, Zeng
et al. 2017), 7 species with sequence data.
Type species – Irenopsis tortuosa (G. Winter) F. Stevens
Notes – Irenopsis is typified by true setae on the ascomata and lack of hyphal setae. The
phylogeny supports the separation of Irenopsis, which forms a distinct and monophyletic clade in
Meliolaceae.
Meliola Fr., Syst. orb. veg. (Lundae) 1: 111(1825)
Index Fungorum number: IF3100; 1694 morphological species (Species Fungorum 2020,
Zeng et al. 2017), 20 species with sequence data.
Type species – Meliola nidulans (Schwein.) Cooke
Notes – Meliola is the largest genus in Meliolaceae, which comprises more than 70% of
species in the family. Members of this genus typically have hyphal setae and globose ascomata.
Meliola telosmae is illustrated in this entry (Fig. 162). Meliola telosmae differs from M.
clerodendricola in having longer appressoria and setae, and thinner, light brown ascospores.
Therefore, we assign this new collection as a reference specimen of M. telosmae with sequence data
provided.
Setameliola D.R. Reynolds, Gdns' Bull., Singapore 61(2): 424 (2010)
Index Fungorum number: IF540193; 17 morphological species (Species Fungorum 2020,
Zeng et al. 2017).
Type species – Setameliola argentina (Speg.) D.R. Reynolds
Notes – This combination of characters could be uninformative, and needs phylogenetic
confirmation.
Melogrammataceae G. Winter, Rabenh. Krypt. -Fl., Edn 2 (Leipzig) 1.2: 797 (1886)
Index Fungorum number: IF80994; Facesoffungi number: FoF00840; 17 species.
Saprobic on bark of woody plants. Sexual morph: Stromata superficial, erumpent, pulvinate
or discoid, reddish-brown, subglobose, soft-textured, with pseudoparenchymatous cells, with black
ostiole. Ascomata perithecial, dark brown to black, immersed, globose, ostiole periphysate.
Peridium comprising rows of cells, externally brown, internally hyaline. Paraphyses hypha-like,
septate. Asci 8-spored, unitunicate, clavate, or fusoid, short pedicellate, with a shallow, J-, apical
ring. Ascospores 1-3-seriate, hyaline or brown, filiform, aseptate or 1–3 septate, cylindrical or
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falcate. Asexual morph: Coelomycetous. Mycelium hyphae to yellow-brown, septate, branched.
Conidiophores long, cylindrical, stiffly upright, septate, verticillate and whorled or not.
Conidiogenous cells holoblastic, proliferating sympodially, hyaline, thin-walled. Conidia elongate
falcate or filiform (adapted from Maharachchikumbura et al. 2016b).
Type genus – Melogramma Fr.
Notes – Jaklitsch & Voglmayr (2012) showed that Melogrammataceae clustered in Xylariales
rather than Diaporthales based on analysis of ITS and LSU sequence data. Maharachchikumbura et
al. (2015, 2016b) confirmed the placement of Melogramma in Melogrammataceae in Xylariales
based on the analyses of combined LSU, SSU, tef1 and rpb2 sequence data. However, Senanayake
et al. (2015) showed it should be placed in Amphisphaeriales. Hongsanan et al. (2017) confirmed
the status in Amphisphaeriales based on phylogenetic and MCC trees.
Figure 162 – Meliola telosmae (Material examined – THAILAND, Nang Lae, Chiang Rai, on the
living leaves of Telosma cordata (Apocynaceae), 4 January 2014, Saranyaphat Boonmee, MFLU
14-0003, reference specimen designated here). a The host plant. b Ascomata on leaf surface. c
Cross section of ascoma. d, e Hyphae with appressoria. f Phialides. g Setae. h-k Asci from young to
mature state. l Peridium. m-s Ascospores. Scale bars: b = 100 µm, c-e = 50 µm, f-s = 20 µm.
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Ecological and economic significance of Melogrammataceae
Species of this family are saprobes on bark of woody plants and thus involved in nutrient
cycling.
Genus included in Melogrammataceae
Melogramma Fr., Summa veg. Scand., Section Post. (Stockholm): 386 (1849)
Index Fungorum number: IF3117; 17 morphological species (Species Fungorum 2020), 3
species with sequence data.
Type species – Melogramma campylosporum Fr.
Notes – Melogramma is characterized by reddish-brown stroma, dark brown to black,
globose, ostiolate and perithecial ascoma, unitunicate, straight, clavate, fusoid or sigmoid, short
pedicellate, 8-spored asci with a J-, apical ring and brown, 3-septate, falcate, often strongly curved
ascospores with subacute ends and guttulate when mature (Maharachchikumbura et al. 2016b). In
this entry we illustrate Melogramma campylosporum.
Microascaceae Luttr. ex Malloch, Mycologia 62(4): 734 (1970)
Index Fungorum number: IF81001; Facesoffungi number: FoF01798; 263 species.
Saprobic and pathogenic on various plants. Sexual morph: Ascomata black, globose
cleistothecial or perithecial, superficial to immersed, ostiolate, ampulliform, aggregated to
scattered, glabrous or covered with scattered hairs. Ostiole often comprises a neck with variable
length and shape, with ostiolar hairs. Peridium thick-walled, dark brown to black, composed of
cells of textura angularis or textura intricata. Hamathecium without interascal tissues. Asci 8spored, globose to barrel-shaped, evanescent, thin-walled, obovate, unitunicate, sessile to short
pedicellate, arranged in basipetal rows. Ascospores crowded, aseptate, seriate, pale to dark reddishbrown, asymmetrical, reniform, triangular or quadrangular, dextrinoid when immature, germ pores
present, or with germ slits, smooth-walled. Asexual morph: Conidiophores smooth-walled,
elongate, macronematous, mononematous or sporodochial to synnematous, with or without
rhizoids. Conidiogenous cells short simple or little branched, cylindrical annellides, borne singly or
in groups of 2–5 on the vegetative hyphae, ampulliform or lageniform, subhyaline to dark when
mature, smooth- to rough-walled. Conidia aseptate, pale yellow to dark brown, obovate or clavate,
base truncate and rounded to pointed at the apex, globose to subglobose, smooth thin-walled to
finely roughened thick-walled, produced singly or in basipetal dry chains. Presence of solitary
conidia in some species, sessile or on short stalks from the vegetative hyphae (adapted from
Sandoval-Denis et al. 2016a, Maharachchikumbura et al. 2016b).
Type genus – Microascus Zukal.
Notes – Microascaceae was circumscribed by Luttrell (1951) in Microascales and validated
by Malloch (1970) (Sandoval-Denis et al. 2016a, Maharachchikumbura et al. 2016b). It comprises
several closely related genera of saprobic, plant and opportunistic human pathogenic fungi (de
Hoog et al. 2011, Sandoval-Denis et al. 2013, 2016b, Maharachchikumbura et al. 2016b).
Sandoval-Denis et al. (2016a, b) revised and proposed several new taxa and combinations under
Microascaceae. Unlike the studies by Issakainen et al. (2003), Sandoval-Denis et al. (2016a) used
combined gene (LSU, ITS, tef1 and tub2) phylogenetic analysis and confirmed the polyphyletic
nature of the two groups, Microascus and Scopulariopsis. As a result, they reported several
uncertain genera which were later resolved in Sandoval-Denis et al. (2016b). For example,
Acaulium and Fairmania were circumscribed as a new lineage in Microascaceae (Sandoval-Denis
et al. 2016a). In our current revision of Microascaceae, we accommodate 23 genera. Yunnania,
Fairmania, Acaulium, Gamsia, and Rhinocladium are the latest additions in the family (SandovalDenis et al. 2016a, b, Wijayawardene et al. 2017a, 2018a).
Ecological and economic significance of Microascaceae
The species are mainly saprobic, coprophilous and are also clinically important.
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Figure 163 – 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, p-r redrawn from Laflamme 1976). a Stroma on the host.
b Cross section of stroma. c Vertical section of stroma. d Ascoma. e Periphyses. f Peridium. g
Paraphyses. h-k Asci. l-o Ascospores. p Pycnidia. q Conidiophores with conidia. r Conidia. Scale
bars: a = 500 µm, b-d = 200 µm, e-k, p = 30 µm, l-o = 15 µm, q, r = 5 µm.
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Genera included in Microascaceae
Acaulium Sopp, Skr. VidenskSelsk. Christiania, Kl. I, Math. -Natur. (11): 42 (1912)
Index Fungorum number: IF7011; 4 morphological species (Species Fungorum 2020), 3
species with sequence data.
Type species – Acaulium albonigrescens Sopp
Notes – The asexual morph is hyphomycetous and has been treated as a synonym of
Scopulariopsis by Seifert et al. (2011). Species occur in terrestrial habitats worldwide and are
characterized by annellidic conidiogenesis with mycelium forming hyphal fascicles and guttulate
conidia. Taxonomic revisions were carried out by Kirk et al. (2013). Sandoval-Denis et al. (2016a)
accepted, Acaulium, in Microascaceae by segregating it from Microascus and Scopulariopsis based
on molecular phylogenetic study. The placement of the genus was also accepted in
Maharachchikumbura et al. (2015, 2016b) and Wijayawardene et al. (2017a, 2018a).
Brachyconidiellopsis Decock, R.F. Castañeda & Adhikari, Cryptog. Mycol. 25(2): 140 (2004)
Index Fungorum number: IF28798; 1 species with sequence data.
Type species – Brachyconidiellopsis fimicola Decock
Notes – The asexual morph of Brachyconidiellopsis is hyphomycetous and species are
saprobes. The genus is characterized by sporodochial to synnematal conidiomata, with a narrow,
dark brown to black stipe, with a slight fasciculate fertile apex. It was described as coprophilous on
dung of an unidentified Cervidae in Nepal (Decock et al. 2004). The genus was accepted in
Microascaceae by Sandoval-Denis et al. (2016a), Maharachchikumbura et al. (2015, 2016b) and
Wijayawardene et al. (2017a, 2018a).
Canariomyces Arx, Persoonia 12(2): 185 (1984)
Index Fungorum number: IF25789; 6 morphological species (Species Fungorum 2020), 5
species with sequence data.
Type species – Canariomyces notabilis von Arx
Notes – Sexual morphs are saprobes and the mature ascospores are similar to some
Chaetomium species. The genus was accepted in Microascaceae by Sandoval-Denis et al. (2016a),
Maharachchikumbura et al. (2015, 2016b) and Wijayawardene et al. (2017a, 2018a). The asexual
morph of this genus is undetermined.
Cephalotrichum Link, Mag. Gesell. naturf. Freunde, Berlin 3(1–2): 20 (1809)
Index Fungorum number: IF7527; 35 morphological species (Species Fungorum 2020), 27
species with sequence data.
Type species – Cephalotrichum stemonitis (Pers.) Nees
Notes – The genus has a similar conidiogenous apparatus to Microascus and Scopulariopsis.
However, the conidiophores in Cephalotrichum arise from synnemata and produce conidia in dry
basipetal chains. Asexual morphs are hyphomycetous and saprobic on soil and dung with a
worldwide distribution. de Beer et al. (2013a) and Sandoval-Denis et al. (2016a) added new
species. The genus was accepted in Microascaceae by Sandoval-Denis et al. (2016a),
Maharachchikumbura et al. (2015, 2016b) and Wijayawardene et al. (2017a, 2018a). The sexual
morph of this genus is undetermined.
Doratomyces Corda, Deutschl. Fl., 3 Abt. (Pilze Deutschl.) 2: 65 (1829)
Index Fungorum number: IF8093; 3 morphological species (Species Fungorum 2020).
Type species – Doratomyces neesii Corda
Notes – Species of Doratomyces are similar to Cephalotrichum (Sandoval-Denis et al. 2016a)
and are common in soil, dung and decaying plant materials. The genus was accepted in
Microascaceae by Sandoval-Denis et al. (2016a), Maharachchikumbura et al. (2015, 2016b) and
Wijayawardene et al. (2017a, 2018a). The sexual morph of this genus is undetermined.
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Echinobotryum Corda, Deutschl. Fl., 3 Abt. (Pilze Deutschl.) 3(12): 51 (1831)
Index Fungorum number: IF8120; 2 morphological species (Species Fungorum 2020).
Type species – Echinobotryum atrum Corda
Notes – Echinobotryum species are found in terrestrial habitats as saprobes with a worldwide
distribution. The genus is characterized by spores clustered on the conidiophore apices and warts
on the spore wall and are similar to Wardomyces (Corda 1831). However, they can be distinguished
based on the germ slits on the ornamented spores in Echinobotryum. The genus was accepted in
Microascaceae by Kirk et al. (2013), Maharachchikumbura et al. (2015, 2016b) and Wijayawardene
et al. (2017a, 2018a). The sexual morph of this genus is undetermined.
Enterocarpus Locq.-Lin., Revue Mycol., Paris 41(4): 510 (1977)
Index Fungorum number: IF1826; 2 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Enterocarpus uniporus Locq. -Lin.
Notes – The genus is characterized by ascomata lacking ostioles with an apical tuft of hairs
and a hyphal ‘capillitium’ surrounding the ascospores. Species are saprobic and coprophilous in
terrestrial habitats and are distributed in Sahara Desert and Italy. The genus was accepted in
Microascaceae by Kirk et al. (2013), Maharachchikumbura et al. (2015, 2016b) and Wijayawardene
et al. (2017a, 2018a). The asexual morph of this genus is undetermined.
Fairmania Sacc., Annls mycol. 4(3): 276 (1906)
Index Fungorum number: IF1972; 1 morphological species.
Type species – Fairmania singularis Sacc.
Notes – Malloch & Cain (1971) placed F. singularis in Microascus as M. singularis as they
were similar and this was accepted by von Arx et al. (1988). However, due to the presence of
longitudinal striations on the conidial wall and erect and thick-walled annellides of the species and
phylogenetic differences, Sandoval-Denis et al. (2016), reinstated Fairmania in Microascaceae.
Fairmania species are wood-decay saprobes and have been reported from Canada, Japan and the
USA.
Gamsia M. Morelet, Ann. Soc. Sci. Nat. Arch. Toulon et du Var 21: 105 (1969)
Index Fungorum number: IF8306; 4 species with sequence data.
Type species – Gamsia dimera (W. Gams) M. Morelet
Notes – This genus was separated from Wardomyces on the basis of well-differentiated
conidiophores, and the conidial arrangement with large apical clusters. This separation is also
supported by the phylogenetic studies. The genus was accepted in Microascaceae by Kirk et al.
(2013), Sandoval-Denis et al. (2016a), Maharachchikumbura et al. (2015, 2016b) and
Wijayawardene et al. (2017a, 2018a). The sexual morph of the genus is undetermined.
Kernia Nieuwl., Am. Midl. Nat. 4: 379 (1916)
Index Fungorum number: IF2563; 14 morphological species (Species Fungorum 2020), 7
species with sequence data.
Type species – Kernia nitida (Sacc.) Nieuwl.
Notes – The genus is characterized by hairy ascomata and ellipsoidal reddish-brown to brown
ascospores with two germ pores and graphium- or scopulariopsis-like asexual morphs. In
phylogenetic studies the genus forms well-supported clades related to Scedosporium and allied
genera. The genus was accepted in Microascaceae by Kirk et al. (2013), Sandoval-Denis et al.
(2016a), Maharachchikumbura et al. (2015, 2016b) and Wijayawardene et al. (2017a, 2018a). The
sexual morph of this genus is undetermined.
718
Lomentospora Hennebert & B.G. Desai, Mycotaxon 1(1): 45 (1974)
Index Fungorum number: IF 8790; 1 morphological species.
Type species – Lomentospora prolificans Hennebert & B.G. Desai
Notes – Lomentospora prolificans was isolated from greenhouse soil in Belgium. However,
the name was thought to be invalid and was placed under Beauveria. Malloch and Salkin (1984)
synonymized the species as Scedosporium inflatum. Later the species was renamed as
Scedosporium prolificans by Geuho and de Hoog (1991). However, based on its phylogenetic
distance and distinguishable morphology from other Scedosporium species, the species was
accepted as Lomentospora prolificans (Lackner et al. 2014). Distribution of the genus is limited to
Australia, European regions and the USA. Lomentospora species have been reported to cause
fungal infections in immunocompromised patients, those with malignancy, and organ and stem cell
transplant recipients.
Lophotrichus R.K. Benj., Mycologia 41(3): 347 (1949)
Index Fungorum number: IF2946; 7 morphological species (Species Fungorum 2020), 6
species with sequence data.
Type species – Lophotrichus ampullus R.K. Benj.
Notes – Some species of the genus are coprophilous while others are cellulolytic, and that
their appearance on dung is more or less incidental. The genus was accepted in Microascaceae by
Kirk et al. (2013), Sandoval-Denis et al. (2016a), Maharachchikumbura et al. (2015, 2016b) and
Wijayawardene et al. (2017a, 2018a). The sexual morph of this genus is undetermined.
Microascus Zukal, Verh. Zool-Bot. Ges. Wien 35:339 (1886)
Index Fungorum number: IF3153; 54 morphological species (Species Fungorum 2020), 39
species with sequence data.
Type species – Microascus longirostris Zukal
Notes – Phylogenetic study by Issakainen et al. (2003) indicated that subdivision of the genus
was necessary. Sandoval-Denis et al. (2016b) confirmed that there were several distinct lineages
which were supported by morphology, such as germ slits, synnemata or conspicuously hairy
ascomata (Abbott 2000, Issakainen et al. 2003, Jagielski et al. 2016, Sandoval-Denis et al. 2016b).
Sandoval-Denis et al. (2016a, b) resolved the taxonomic complexity of the genus and introduced
seven new species, with new combinations in the Microascus lineage (Sandoval-Denis et al.
2016a). However, several taxa (e.g. M. longirostris and M. pseudolongirostris) have uncertain
position in a phylogenetic tree , suggesting further study is needed to resolve the taxonomic
placements of problematic taxa (Woudenberg et al. 2017). Microascus longirostris has a wide host
range, mostly from dung of mammals, soil, wood, seeds, air, and clinical samples (Barron et al.
1961, Sandoval-Denis et al. 2016a, b, Woudenberg et al. 2017). Neither a type culture nor
herbarium specimen is available (Barron et al. 1961, Sandoval-Denis et al. 2016a, Wijayawardene
et al. 2017a). A neotype of M. longirostris was designated by Sandoval-Denis et al. (2016a), which
was corresponds to the descriptions by Zukal (1885) and cultural characteristics descriptions
correspond to Barron et al. (1961), Morton & Smith (1963) and von Arx et al. (1988).
Parascedosporium Gilgado, Gené, Cano & Guarro, Int. J. Syst. Evol. Microbiol. 57(9): 2176
(2007)
Index Fungorum number: IF506554; 2 species with sequence data.
Type species – Parascedosporium tectonae (C. Booth) Gilgado, Gené, Cano & Guarro
Notes – The genus is characterized by denticulate conidiogenous cells with sympodial
conidia. Gilgado et al. (2007) proposed a new combination for Graphium tectonae as
Parascedosporium tectonae. The genus was accepted in Microascaceae by Kirk et al. (2013),
Sandoval-Denis et al. (2016a), Maharachchikumbura et al. (2015, 2016b) and Wijayawardene et al.
(2017a, 2018a).
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Figure 164 – Microascus longirostris (Material examined – USA, Maine, Kittery Point, from a
wasp’s nest, 1961, R. Thaxter, MUCL 9057, neotype). a-c Herbarium material of Microascus
longirostris (dried culture and micro slide). d, e Ascomata. f Peridium. g Asci. h, i Ascospores.
Scale bars: d = 50 μm, f = 20 μm, g = 10 μm, h, i = 5 μm.
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Petriella Curzi, Boll. R. Staz. Patalog. Veget. Roma 10: 384 (1930)
Index Fungorum number: IF3852; 7 morphological species (Species Fungorum 2020), 6
species with sequence data.
Type species – Petriella asymmetrica Curzi
Notes – Petriella resembles Microascus but was segregated from Microascus by its hairy
ascomata. The ascospores in Petriella are also larger than most other genera in Microascales. The
genus was accepted in Microascaceae by Kirk et al. (2013), Sandoval-Denis et al. (2016a),
Maharachchikumbura et al. (2015, 2016b) and Wijayawardene et al. (2017a, 2018a).
Pseudallescheria Negr. & I. Fisch., Revista Inst. Bacteriol. Dr. Carlos G. Malbrán’ 12 (201): 5–9
(1944)
Index Fungorum number: IF4403; 8 morphological species (Species Fungorum 2020), 3
species with sequence data.
Type species – Pseudallescheria shearii Negroni & I. Fisch.
Note – Pseudallescheria species are saprobes worldwide and have been isolated from soil,
sewage, contaminated water, and the manure of farm animals. Species are also opportunistic
pathogens that can cause various infections in humans. The genus was accepted in Microascaceae
by Kirk et al. (2013), Sandoval-Denis et al. (2016a), Maharachchikumbura et al. (2015, 2016b) and
Wijayawardene et al. (2017a, 2018a). The asexual morphs of Pseudallescheria are undetermined,
however, scedosporium-like and graphium-like taxa are probably linked.
Pseudoscopulariopsis Sand.-Den., Gené & Guarro, Persoonia, Mol. Phyl. Evol. Fungi 36: 24
(2015)
Index Fungorum number: IF809215; 2 species with sequence data.
Type species – Pseudoscopulariopsis schumacheri (E.C. Hansen) Sandoval-Denis Gené &
Guarro
Notes –Sandoval-Denis et al. (2016b) found that species of Pseudoscopulariopsis formed a
distinct clade basal to the Scopulariopsis clade. Pseudoscopulariopsis species are characterised by
hyaline to subhyaline conidiogenous cells, which usually comprise annellides arising from swollen
conidiophores (Sandoval-Denis et al. 2016b). The sexual morphs are undetermined. The genus was
accepted in Microascaceae by Kirk et al. (2013), Sandoval-Denis et al. (2016a,b),
Maharachchikumbura et al. (2015, 2016b) and Wijayawardene et al. (2017a, 2018a).
Rhinocladium Sacc. & Marchal, Bull. Soc. R. Bot. Belg. 24(1): 65 (1885)
Index Fungorum number: IF9722; 11 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Rhinocladium coprogenum Sacc. & Marchal
Notes – The genus is hyphomycetous with polyblastic conidiogenous cells integrated within
the conidiophores. Rhinocladium species are saprobic and occur in terrestrial habitats and have a
worldwide distribution. The genus was accepted in Microascaceae by Kirk et al. (2013), SandovalDenis et al. (2016a), Maharachchikumbura et al. (2015, 2016b) and Wijayawardene et al. (2017a,
2018a).
Scedosporium Sacc. ex Castell. & Chalm., Manual of tropical medicine (London): 1122 (1919)
Index Fungorum number: IF9794; 11 morphological species (Species Fungorum 2020), 8
species with sequence data.
Type species – Scedosporium apiospermum Sacc. ex Castell. & Chalm.
Notes – Scedosporium is a ubiquitous filamentous genus with a wide host range and
worldwide distribution (Wijayawardene et al. 2017a). The genus is characterized by hyaline,
cylindrical conidiogenous cells arising from undifferentiated hyphae, with obovoidal, hyaline,
sticky conidia. Graphium-like synnemata are formed with large, erect bundles of hyphae
721
terminating in a dense aggregate of conidiogenous cells. The asexual morphs produce conidia from
a short extension of the conidiogenous cells with annellidic development (Guého E. & De Hoog
1991, Lackner et al. 2014, Ramirez-Garcia et al. 2018). The genus was accepted in Microascaceae
by Sandoval-Denis et al. (2016a), Maharachchikumbura et al. (2015, 2016b) and Wijayawardene et
al. (2017a, 2018a).
Scopulariopsis Bainier, Bull. Soc. Mycol. Fr. 23: 98 (1907)
Index Fungorum number: IF9854; 71 morphological species (Species Fungorum 2020), 21
species with sequence data.
Type species – Scopulariopsis brevicaulis (Sacc.) Bainier
Notes – The asexual morphs are hyphomycetous and saprobic. Species of Scopulariopsis
have been recovered from environmental samples (soil, air, mouldy indoor environments), food
(cheese and butter), as well as from human clinical samples. The sexual morph is undetermined.
Scopulariopsis species are found in terrestrial habitats with worldwide distribution. The genus was
accepted in Microascaceae by Kirk et al. (2013), Sandoval-Denis et al. (2016a),
Maharachchikumbura et al. (2015, 2016b) and Wijayawardene et al. (2017a, 2018a).
Wardomyces F.T. Brooks & Hansf., Trans. Br. mycol. Soc. 8(3): 137 (1923)
Index Fungorum number: IF10433; 8 morphological species (Species Fungorum 2020), 7
species with sequence data.
Type species – Wardomyces anomalus F.T. Brooks & Hansf.
Notes – The asexual morph is hyphomycetous and saprobic. Sexual morphs can be identified
by significantly larger ascomata and dark ascospores with two germ pores (Sandoval-Denis et al.
2016a). The genus was accepted in Microascaceae by Ariyawansa et al. (2015), Seifert et al.
(2011), Maharachchikumbura et al. (2015, 2016b), Sandoval-Denis et al. (2016a) and
Wijayawardene et al. (2012, 2017a, 2018a).
Wardomycopsis Udagawa & Furuya, Mycotaxon 7(1): 92 (1978)
Index Fungorum number: IF10434; 8 morphological species (Species Fungorum 2020), 7
species with sequence data.
Type species – Wardomycopsis inopinata Udagawa & Furuya
Notes – The asexual morphs are hyphomycetous saprobes. Conidiation at the early stages of
Wardomycopsis resembles those of Wardomyces. However, the sexual morph of Wardomycopsis
differ from those of Wardomyces, in terms of having straw-coloured, reniform to triangular
ascospores with a single germ pore. Wardomycopsis species are distributed in Asia, Europe, and
North America and are mainly soil-borne. The genus was accepted in Microascaceae by
Ariyawansa et al. (2015), Seifert et al. (2011), Maharachchikumbura et al. (2015, 2016b),
Sandoval-Denis et al. (2016a) and Wijayawardene et al. (2012, 2017a, 2018a).
Yunnania H.Z. Kong, Mycotaxon 69: 320 (1998)
Index Fungorum number: IF27939; 3 species with sequence data.
Type species – Yunnania penicillata H.Z. Kong
Notes – Jagielski et al. (2016) stated that Scopulariopsis carbonaria does not fit in
Scopulariopsis and proposed a new genus, Fuscoannellis. Woudenber et al. (2017) gave priority to
the already described genus Yunnania (Kong 1998) and synonymized Fuscoannellis with
Yunnania. The genus was accepted in Microascaceae by Wijayawardene et al. (2017a).
Microdochiaceae Hern.-Restr., Crous & J.Z. Groenew., Persoonia 36: 64 (2016)
Index Fungorum number: IF811871; Facesoffungi number: FoF01907; 74 species.
Saprobic, endophytic or pathogenic on leaves, seeds and soil. Sexual morph: Stromata
present or absent. Ascomata perithecial. Asci 8-spored, cylindrical, oblong, clavate, with a J+,
722
funnel-shaped, apical ring. Ascospores biseriate or uniseriate, hyaline to pale brown, fusoid,
ellipsoid or oblong. Asexual morph: Coelomycetous. Conidiomata if present, sporodochial.
Conidiophores solitary or aggregated, mono- or biverticillate. Conidiogenous cells solitary or in
whorls, polyblastic, sympodial, denticulate, cylindrical often ampulliform, lageniform with
elongated necks and minute annellides from percurrent proliferations, hyaline to pale brown.
Conidia hyaline, lunate, oblong, fusiform or cylindrical, straight or curved, flattened at base.
Chlamydospores if present, brown (adapted from Hernández-Restrepo et al. 2016b).
Type genus – Microdochium Syd. & P. Syd.
Notes – Microdochiaceae was introduced by Hernández-Restrepo et al. (2016b) to
accommodate the genera Microdochium, Idriella and Selenodriella based on LSU sequence data. In
the phylogenetic analysis, the type of Microdochium (= Monographella) formed a distinct clade in
Xylariales. Species belonging to Microdochiaceae are characterized by monographella-like sexual
morphs and asexual morphs with polyblastic, sympodial or annellidic conidiogenous cells, and
hyaline conidia without appendages (Hernández-Restrepo et al. 2016b). Combined multi-gene and
divergence estimates have revealed that the family diverged around 79 MYA (Hyde et al. 2017a).
Ecological and economic significance of Microdochiaceae
Microdochiaceae includes many species which are of economic importance as
phytopathogens and saprobes. They have been isolated from living, fallen, rotten and decaying
leaves, stems, kernals, roots, oospores, soil and air (Kwaśna & Bateman 2007, Hernández-Restrepo
et al. 2016b, Zhang et al. 2017c, Farr & Rossman 2018). Microdochium nivale and M. majus are
both serious soil and seed borne pathogens causing microdochium patch, also known as pink snow
mould/fusarium patch, seedling blight on cereals and forage grasses (Glynn et al. 2005, Glynn &
Edwards 2009, Vogelgsang et al. 2013). Control of this disease has been investigated using
bacterial seed treatments and botanicals (Vogelgsang et al. 2013, Johansson et al. 2003). Some
Microdochiaceae species may act as biocontrol agents, e.g. Microdochium bolleyi as a biocontrol
agent of cereal root and stem base pathogens (Douglas & Deacon 1994). Endophytic Microdochium
species are also reported to have applied significance. Zhang et al. (2008) investigated effective
anti-fungal, anti-bacterial and anti-algal compounds against Bacillus megaterium, Chlorella fusca
Escherichia coli and Microbotryum violaceum using Microdochium bolleyi, an endophyte from
Fagonia cretica.
Genera included in Microdochiaceae
Idriella P.E. Nelson & S. Wilh., Mycologia 48: 550. (1956)
Index Fungorum number: IF8625; 30 morphological species (Hernández-Restrepo et al.
2016b), 4 species with sequence data.
Type species – Idriella lunata P.E. Nelson & S. Wilh.
Notes – Idriella comprises soil inhabiting hyphomycetes and terrestrial species worldwide.
The genus is characterised by brown, aseptate conidiophores and polyblastic conidiogenous cells
with hyaline, unicellular, smooth, lunate, curved conidia in heads (Hernández-Restrepo et al.
2016b).
Microdochium Syd. & P. Syd., Annls mycol. 22(3/6): 267 (1924)
Index Fungorum number: IF8926; 37 morphological species (Species Fungorum 2020), 20
species with sequence data.
Type species – Microdochium phragmitis Syd. & P. Syd.
Notes – Microdochium (asexual morph, Monographella - sexual morph) is the type genus of
Microdochiaceae and was established by Sydow (1924) with M. phragmitis as the type species,
found on living leaves of Phragmites australis in Germany. Since Microdochium has more species
and is more commonly encountered, and the name is more frequently used in literature, HernándezRestrepo et al. (2016b) suggested to use Microdochium over Monographella. Microdochium
species are economically important plant pathogens on grasses and cereals worldwide (Hernández723
Restrepo et al. 2016b, Wijayawardene et al. 2017a, Farr & Rossman 2018). However, species
identification mainly depends on multi-gene phylogenetic analyses with the prior the protein coding
region tub2. Thirty-seven species have been accepted under Microdochium (Hernández-Restrepo et
al. 2016b, Zhang et al. 2017c, Crous et al. 2018c, 2019b).
Microdochium phragmitis, the type species of Microdochium was isolated from leaves of
Phragmites communis. Hernández-Restrepo et al. (2016b) epitypified the species and provided an
ex-epitype with molecular data. However, variations among the strains and polymorphisms make it
difficult to delimit species in Microdochium. This entry is illustrated with Microdochium
phragmitis (Fig. 165).
Figure 165 – Microdochium phragmitis (Material examined – GERMANY, Brandenburg, Berlin,
on dead leaves of Phragmites communis (Poaceae), 8 November 1919, H. Sydow, K-IMI 193888,
holotype). a Herbarium details. b Host. c, d Sporodochia. e-g Conidiogenous cells and attached
conidia. h-l Conidia (d, e-g, k, l in Congo red). Scale bars: 10 µm.
Selenodriella R.F. Castañeda & W.B. Kendr., Univ. Waterloo Biol. Ser. 33: 34 (1990)
Index Fungorum number: IF11398; 7 morphological species (Hernández-Restrepo et al.
2016), 2 species with sequence data.
Type species – Selenodriella fertilis (Piroz. & Hodges) R.F. Castañeda & W.B. Kendr.
Notes – Selenodriella species are hyphomycetes which produce setiform conidiophores, and
conidiogenous cells and branches are disposed in whorls along the main axis of setiform
conidiophores. Hernández-Restrepo et al. (2016b) introduced S. cubensis and confirmed the
phylogenetic placement in Microdochiaceae.
Myelospermataceae K.D. Hyde & S.W. Wong, Mycol. Res. 44(1): 349 (1999)
Index Fungorum number: IF82090; Facesoffungi number: FoF06193; 5 species.
Pathogenic or saprobic on various plants. Sexual morph: Pseudostromata weakly raised,
visible as blackened ostiolar dots, immersed, multiloculate, clustered around a common central
pore. Ascomata in vertical section subglobose or ellipsoidal, brown, necks mostly eccentric
leading to the common central pore. Peridium comprising mostly brown-walled, elongate cells,
inwardly hyaline and becoming textura intricata and fusing with the host tissue at the outside.
Paraphyses numerous, hypha-like, filamentous, septate, unbranched, tapering distally. Asci 8spored, unitunicate, cylindrical, long-pedicellate, apically rounded, with a J-, refractive, subapical,
discoid ring. Ascospores overlapping uniseriate, hyaline, irregularly ellipsoidal, curved or straight,
some narrower at the center, some lunate, unicellular, and surrounded by a distinct mucilaginous
sheath. Asexual morph: Undetermined (adapted from Maharachchikumbura et al. 2016b).
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Type genus – Myelosperma Syd.
Notes – Myelospermataceae was introduced by Hyde & Wong (1999) to accommodate
Myelosperma. The family has been poorly studied. Four species were reported as saprobic and one
species as parasitic (M. parasitica). Most of the species occur on palms (Arecaceae), except M.
gigasporum, which was found on unidentified wood. Hyde & Wong (1999) accommodated
Myelospermataceae in Diaporthales based on lack of stroma (only pseudostroma present), the
valsoid nature of the ascomata, deliquescing paraphyses, and ascus morphology. Subsequently,
Kirk et al. (2001) placed the genus in Xylariales, which seemed controversial due to lack of
reference taxa and molecular analysis (Eriksson 1999, Kang et al. 2002). Maharachchikumbura et
al. (2016b) and Wijayawardene et al. (2017a, 2018a) followed the taxonomy of Kirk et al. (2001)
and accepted the family in Xylariales. However, it is still unclear if the family belongs in
Xylariales. Due to lack of molecular data, the different morphology from other xylariaceous genera
with morphological affinities to Diaporthales, herein we place Myelospermataceae in
Xylariomycetidae families, incertae sedis, until further studies can better resolve the placement.
Ecological and economic significance of Myelospermataceae
Species of Myelospermataceae are generally saprobes, although M. parasitica causes leaf
spots on living leaves of Phoenix reclinata (Hyde et al. 1996). The species are likely to be
endophytes which become saprobes on host senescence.
Genus included in Myelospermataceae
Myelosperma Syd. & P. Syd., Ann. Mycol. 13: 38 (1915)
Index Fungorum number: IF3351; 5 morphological species (Species Fungorum 2020).
Type species – Myelosperma tumidum Syd. & P. Syd.
Notes – Myelosperma was established by Sydow & Sydow (1915) to accommodate
Myelosperma tumidum which occurs on the dead rachis of Cocos nucifera. Four species were
introduced from different hosts, substrates and countries (Ahmad 1971, Hyde et al. 1996, Aptroot et
al. 1998, Hyde & Wong 1999).
Myrmecridiaceae Crous, Persoonia 34: 219 (2015)
Index Fungorum number: IF812461; Facesoffungi number: FoF06703; 16 species.
Saprobic on decaying wood submerged in freshwater, leaf litter, on stem or leaves of
herbaceous plants, occurring on soil and in house dust (Crous et al. 2015b, c, 2016b, 2018b, c,
Peintner et al. 2016, Réblová et al. 2016a, Tibpromma et al. 2017b). Sexual morph: Ascomata
immersed, solitary or aggregated, with subglobose to conical venter and ostiolate papilla as well as
dark brown clypeus, glabrous. Ostiole periphysate. Peridium leathery, two-layered with hyaline to
pale brown outer layer and subhyaline inner layer consisting of cells of textura angularis to textura
prismatica. Paraphyses cylindrical, septate. Asci 8-spored, unitunicate, cylindrical, with a slender
long pedicel, apex rounded. Ascospores uniseriate or overlapping uniseriate, hyaline, ellipsoidal, 3septate, slightly verrucose, with granular content (Réblová et al. 2016a). Asexual morph: In vitro,
Mycelium submerged, hyaline, thin-walled. Conidiophores erect from creeping hyphae,
unbranched, straight or flexuous, brown at base and paler towards the apex, subcylindrical, smooth,
septate. Conidiogenous cells differentiating from the apical region of conidiophore, integrated,
polyblastic, cylindrical, subhyaline to brown, with conidium-bearing denticles along the
conidiogenous axis. Conidia hyaline to pale brown, ellipsoid to obovoid to fusoid, solitary, smooth
or verrucose, aseptate or septate, some species present a wing-like sheath (adapted from Crous et al.
2015c, 2016a, 2018b, c, Peintner et al. 2016, Tibpromma et al. 2017b).
Type genus – Myrmecridium Arzanlou, W. Gams & Crous
Notes – Myrmecridiales and Myrmecridiaceae were established by Crous et al. (2015c) to
accommodate Myrmecridium species based on the analyses of LSU sequence data.
Myrmecridiaceae is the type family of Myrmecridiales (Sordariomycetes). Myrmecridiales is a
monotypic order in the subclass Diaporthomycetidae (Crous et al. 2015c) and has phylogenetic
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affinities with Distoseptisporaceae and Magnaporthales in a multi-gene phylogenetic tree based on
LSU, SSU, tef1 and rpb2 sequence data (Hongsanan et al. 2017). Myrmecridiaceae comprises two
genera Myrmecridium (Arzanlou et al. 2007) and Neomyrmecridium (Crous et al. 2018b).
Figure 166 – Myelosperma tumidum (Material examined – SRI LANKA, Peradeniya, on the dead
rachis of Cocos nucifera L. (Arecaceae), 1912, T. Petch, S (Swedish Museum of Natural History),
F9909, F9911, Petch 3369, holotype). a Herbarium package. b Host. c-e Pseudostromata. f
Horizontal section of ascomata. g, h Vertical section through clustered multiloculate ascomata. i
Peridium. j Paraphyses. k-m Asci. n Apex of ascus with the refractive subapical ring. o, p
Ascospores with sheath. q, r Ascospores. Scale bars: b = 1 cm, c, d = 1000 µm, e = 200 µm, f = 500
µm, j-m = 100 µm, n-r = 20 µm.
Ecological and economic significance of Myrmecridiaceae
As a member of soil fungal community, Myrmecridium species are positively related with the
death rate of notoginseng, which is a valuable herbal medicine (Dong et al. 2016). Myrmecridium
726
species also occur in the sugarcane rhizosphere, and might have a potential as biocontrol agents
against pathogens (Romão-Dumaresq et al. 2016). Myrmecridium schulzeri is an excellent
decomposer of wood of oak and pine. It was observed in the third stage of fungi succession during
wood decomposition (Wrzesień et al. 2016). Some Myrmecridium species displayed relatively high
antibacterial activity (Zhang et al. 2012a) and glyphosate-tolerance (Schlatter et al. 2017b).
Myrmecridium taxa showed high abundance in live and dying glyphosate-killed wheat roots and
may play an under-recognized role in green bridge dynamics of glyphosate degradation (Schlatter
et al. 2017a).
Genera included in Myrmecridiaceae
Myrmecridium Arzanlou, W. Gams & Crous, Stud. Mycol.58: 84 (2007)
Index Fungorum number: IF504559; 13 species with sequence data.
Type species – Myrmecridium schulzeri (Sacc.) Arzanlou, W. Gams & Crous
Notes – Myrmecridium accommodates ramichloridium-like taxa with hyaline mycelium and
pale brown to hyaline, conidiogenous cells with pimple-like denticles. Among the 15 epithets listed
in Index Fungorum (2020), M. montsegurinum is reported as a sexual morph in Myrmecridium
(Réblová et al. 2016a) and the remaining 14 taxa are based on their asexual morphs. Because
Myrmecridium sorbicola has septate conidia and it phylogenetically groups with species of
Neomyrmecridium, it was transferred to the latter genus (Crous et al. 2018b).
Myrmecridium montsegurinum, which is the only known sexual morph in Myrmecridium,
occurs on decaying wood of various deciduous trees submerged in freshwater (Réblová et al.
2016a). Myrmecridium montsegurinum shares a close phylogenetic affinity to M. schulzeri and M.
flexuosum, but statistical support for this alliance is not significant (Réblová et al. 2016a).
Neomyrmecridium Crous, Persoonia 41: 287 (2018)
Index Fungorum number: IF828190; 3 species with sequence data.
Type species – Neomyrmecridium septatum Crous
Notes – Neomyrmecridium was erected by Crous et al. (2018c) to accommodate N. asiaticum
Crous, N. septatum and N. sorbicola. This genus is phylogenetically sister to Myrmecridium,
however, Neomyrmecridium can be distinguished by its septate conidia.
Myrotheciomycetaceae Crous, Persoonia 40: 351 (2018)
Index Fungorum number: IF825408; Facesoffungi number: FoF06875; 45 species.
Associated with leaves. Sexual morph: Ascomata sphaerical to slightly irregular, transparent,
globose, covered with sparse to abundant, but undifferentiated hyphal attachments, occasionally
with conidiophores on the surface, lacking ostioles. Peridium relatively wide, composed of angular
cells forming a textura angularis in surface view and rather flattened in side view. Asci crowded, 8spored, uniformly distributed in the centrum, clavate or globose. Ascospores hyaline to pale yellow
or light brown, ellipsoidal, unicellular, smooth-walled, with or without pores or slits, with or
without wing-like appendages and gelatinous layers. Asexual morph: Conidiomata solitary to
sporodochial, with crystalline to white or orange conidial mass; with or without basal stroma.
Conidiophores, hyaline, smooth to warty, unbranched or branched, subcylindrical with terminal
and lateral conidiogenous cells, some progressively shortened with the formation of each conidium.
Conidiogenous cells hyaline, smooth, phialidic or with retrogressive conidiogenesis. Conidia
aggregated in slimy mass, with or without septa, hyaline or pale-yellow or pale-brown to brown,
smooth-walled, ellipsoid or fusoid-ellipsoid or clavate (adapted from Crous et al. 2018d).
Type genus – Myrotheciomyces Crous
Notes – Myrotheciomycetaceae was introduced by Crous et al. (2018d) in Hypocreales, to
accommodate a genus Myrotheciomyces based on characters and phylogenetic analyses. Presently,
the family includes Emericellopsis, Leucosphaerina, Myrotheciomyces and Trichothecium (Crous
et al. 2018d).
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Figure 167 – Myrmecridium montsegurinum (Material examined – FRANCE, Midi-Pyrénées:
Ariège, Montségur, Le Lasset stream along D9 road, 880−890 m a.s.l., on submerged wood of
Fraxinus excelsior, 1 October 2013, J. Fournier J.F. 13180, PRM 934684, holotype). a Specimen
label and host. b, c Appearance of stromata on substrate. d-f Vertical section of ascostroma. g
Ostiolar canal. h Peridium. i, j Asci and paraphyses. k-o Asci with ascospores (k = stained with
lugo's iodine). p-s Ascospores. Scale bars: d = 150 µm, e = 100 µm, g, i, k = 50 µm, j, l-o = 30 µm,
h = 25 µm, p-s = 10 µm
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Figure 168 – Myrmecridium schulzeri (CBS 325.74) (a, b, c, e), M. flexuosum (CBS 398.76) (d),
M. sorbicola (CBS 143433) (f), M. iridis Crous (CBS 139917) (g). a-d Conidiophores. e-g Conidia
(Redrawn from Arzanlou et al. 2007, Crous et al. 2015c, 2018c). Scale bars: a = 100 µm, b-g = 10
µm.
Ecological and economic significance of Myrotheciomycetaceae
Members of Myrotheciomycetaceae are pathogens on fruits and vegetables and also saprobes
on dead material, decaying vegetables, dung, agricultural and forest soils, peat, rhizomes, prairies
and freshwater-, estuarine- and marine-mud sediments, (Domsch et al. 1980, Pitt & Hocking 1999,
Malloch 1989, Giraldo et al. 2012). Trichothecium roseum causes pink rot disease on various fruits
and vegetables worldwide (Batt & Tortorello 2014, Domsch et al. 1980, Pitt & Hocking 1999). The
species produces a wide variety of secondary metabolites, such as roseotoxins and trichothecenes,
which can infect and spoil a variety of fruit crops (Batt & Tortorello 2014).
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Genera included in Myrotheciomycetaceae
Emericellopsis J.F.H. Beyma, Antonie van Leeuwenhoek 6: 264 (1940)
Index Fungorum number: IF1773; 15 morphological species (Species Fungorum 2020), 14
species with sequence data.
Type species – Emericellopsis terricola J.F.H. Beyma
Notes – Emericellopsis species are primarily soil-derived fungi. Species of Emericellopsis
have been isolated from various environments worldwide (viz. agricultural and forest soils, peat,
rhizomes, prairies and freshwater-, estuarine- and marine-mud sediments, (Domsch et al. 1980).
Globose asci contain 8 ascospores with light brown, ellipsoidal and wing-like appendages (Zuccaro
et al. 2004). The asexual morphs have conidiophores arising from substratum or aerial hyphae, and
hyaline, 1–2-guttulate, elongate-ellipsoidal conidia (Beyma 1940).
Figure 169 – Emericellopsis terricola (redrawn from Beyma 1940). a Perithecium. b Ascus. c
Ascospores. d Ascospores (front view). e Hypha with conidiophores. f Conidiophores. g Conidia.
Scale bars: a = 50 µm, b-d, g = 5 µm.
Leucosphaerina Arx, Persoonia 13(3): 294 (1987)
Index Fungorum number: IF26159; 2 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Leucosphaerina indica (Arx, Mukerji & N. Singh) Arx
Notes – Leucosphaerina was introduced by von Arx (1987) based on L. indica from dung of
Boselaphus tragocamelus in Delh, but lacks sequence data. Malloch (1989) introduced L. arxii
fungi from dung of Equus in North Carolina. Summerbell et al. (2011) provided sequence data for
L. arxii. The genus is characterized by clavate asci, ellipsoidal, hyaline ascospores with sheaths,
and an acremonium-like asexual morphs with solitary or paired tapering conidiogenous cells,
septate at the base, bearing small apical clusters of conidia, conidia are ellipsoidal to obovate,
hyaline, 1-celled and produced in wet masses (von Arx 1987, Malloch 1989).
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Figure 170 – Leucosphaerina arxii (redrawn from Malloch 1989). a Ascomatal initials. b
Ascomata (surface view at left, cross section at right). c Developing asci. d Mature asci. e
Ascospores. f Conidiogenous cells. g Conidia. Scale bars: b = 40 µm, a, c-g = 10 µm.
Myrotheciomyces Crous, Persoonia 40: 351 (2018)
Index Fungorum number: IF825409; 1 species with sequence data.
Type species – Myrotheciomyces corymbiae Crous
Notes – The genus was introduced from Corymbia based on morphology and an ITS
megablast search (Crous et al. 2018d).
Figure 171 – Myrotheciomyces corymbiae (redrawn from Crous et al. 2018d). a Conidiomata
sporulating on pine needle agar. b, c Conidiogenous cells. d Conidia. Scale bars: a = 200 μm, b-d =
10 μm.
Trichothecium Link, Mag. Gesell. naturf. Freunde, Berlin 3(1-2): 18 (1809)
Index Fungorum number: IF10303; 27 morphological species (Species Fungorum 2020), 5
species with sequence data.
Type species – Trichothecium roseum (Pers.) Link
731
Notes – Species of Trichothecium grow on various fruits and vegetables, as well as cereal
grains (Pitt & Hocking 1999) and on decaying vegetation and in soil. Species are characterized by
simple hyphae conidiophores and septate, 2-celled, clavate conidia, with the apical cell being larger
than the curved basal cell. The sexual morph is undetermined.
Figure 172 – Trichothecium roseum (redrawn from Halstead 1876 in Popular Science Monthly Vol
9). a-c Conidiophores with conidia. Scale bars: a, b = 20 µm, c = 10 µm.
Nectriaceae Tul. & C. Tul., Select. fung. carpol. (Paris) 3: 3 (1865)
Index Fungorum number: IF81059; Facesoffungi number: FoF01396; 2081 species.
Endophytic, foliicolous or saprobic on woody plant hosts, some are entomogenous, a few
species are human pathogens, in terrestrial and aquatic habitats. Sexual morph: Ascomata stromatic
or astromatic, white, red, dark red, reddish-brown, orange, orange-red, orange-brown, yellow, pale
yellow, brown, greyish yellow-green, dark bluish, bluish purple, bluish black or black, solitary or
aggregated in groups, perithecial, globose to subglobose, ovoid, elongate-ovoid, obpyriform,
obovoid or pyriform, KOH +/-, surface smooth to papillate, striate, warted, verrucose or scaly, with
or without setae, ostiolar region sometimes papillate, periphysate. Paraphyses present or absent.
Asci 4–8-spored, unitunicate, clavate to narrowly clavate, cylindrical or ellipsoidal, with or without
apical ring, with pointed or pedicellate base. Ascospores uniseriate to biseriate or overlapping,
hyaline to yellow, yellow-brown, golden-brown, pale-brown or green, fusiform, long-fusiform,
ellipsoidal, oblong, biconic, pyriform, reniform or allantoid, aseptate to multi-septate or muriform,
constricted at the septum or not, smooth-walled, spinulose, verruculose or striate. Asexual morph:
Mainly hyphomycetous, less commonly coelomycetous. Conidiomata synnematous, sporodochial
or pycnidial. Conidiophores branched or unbranched, penicillate, verticillate. Conidiogenous cells
monophialidic to polyphialidic, ampulliform to lageniform, cylindrical, elongate-ampulliform or
subcylindrical, hyaline, smooth-walled. Conidia globose, ovate, ellipsoidal, cylindrical to
subcylindrical, fusiform, long-fusiform, filiform, allantoid or falcate, straight to slightly or strongly
curved, hyaline, aseptate to multi-septate, constricted at septum or not, with or without visible
abscission scars, sometimes guttulate, smooth-walled. Chlamydospores present or absent (adapted
from Maharachchikumbura et al. 2016b).
Type genus – Nectria (Fr.) Fr.
Notes – Nectriaceae species occur worldwide, have higher diversity in warm temperate and
tropical regions (Rossman et al. 1999, Rossman 2000, Chaverri et al. 2011, Schroers et al. 2011,
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Hyde et al. 2014, Lombard et al. 2015). They are commonly found on bark of recently dead woody
substrates, especially in tropical regions. Some are plant endophytes or pathogens, other fungi or
insects (Rossman et al. 1999, Lombard et al. 2015).
Seaver (1909) divided Hypocreales into two families: Nectriaceae and Hypocreaceae, based
on stromatic and perithecial characters. Petch (1938) also accepted Nectriaceae as a family in
Hypocreales, while Munk (1957) and Dennis (1960) placed it in Sphaeriales. However, Kreisel
(1969) and Rossman et al. (1999) accepted Nectriaceae and Hypocreaceae as two separate families
in Hypocreales. Lumbsch & Huhndorf (2010) listed 26 genera in Nectriaceae.
Maharachchikumbura et al. (2016b) and Lombard et al. (2015) accepted 47 genera in the family
based on molecular data. The asexual morphs of Nectriaceae species are mainly hyphomycetous,
except Thyronectria, which forms pycnidia (Wijayawardene et al. 2017b). Corinectria was
introduced to the family by González & Chaverri (2017). Another genus Neothyronectria was
added to Nectriaceae by Yang et al. (2019). Aiello et al. (2017) introduced the new genus
Pleiocarpon, based on morphology and molecular data. Xenocalonectria was placed in synonymy
with Xenocylindrocladium by Rossman et al. (2016). Rossman et al. (2016) synonymised
Antipodium under Ophionectria. Pleonectria was considered as an synonym of Thyronectria by
Wijayawardene et al. (2017b). Stachybotryna, previously listed in Nectriaceae by
Maharachchikumbura et al. (2016b), was maintained in Ascomycota genera incertae sedis by
Wijayawardene et al. (2018a). Wijayawardene et al. (2018a) accepted 63 genera in Nectriaceae
including Baipadisphaeria, Varicosporella, Xenocylindrocladium and Xenoleptographium. A new
name was proposed for the monotypic genus Curvicladium by Crous et al. (2016a), as the generic
name “Curvicladium” was already occupied for a moss species.
Ecological and economic significance of Nectriaceae
The members of the Nectriaceae are facultative, sometimes virulent plant pathogens, causing
serious problems on crop plants, often encountered as asexual morphs eg. Albonectria rigidiuscula,
causes green point gall of cocoa (Ploetz 2007, Pérez et al. 2012, Yang et al. 2018c), while
Fusarium sambucinum causes hop canker, potato storage rot, and root rot of many crops (Ray &
Hammerschmidt 1998, Wharton et al. 2006). Fusarium oxysporum causes root rots, foot rots, and
wilt diseases of numerous crop plants (Hyde et al. 2014, Chittem et al. 2015, McGovern 2015,
Rossman et al. 1999). They also cause cankers, root rots, and other diseases on hardwood and
coniferous trees, e.g. Abies and Acer cankers caused by Neonectria castaneicola; beech (Fagus
spp.) bark disease caused by Nectria coccinea and N. ditissima, black foot disease of grapevines
(Vitis spp.) caused by Neonectria liriodendri; ‘coral spot Nectria canker’ caused by N. cinnabarina;
root rots caused by N. radicicola; and cankers caused by N. rugulosa, among others (Hirooka et al.
2005, 2012, Kobayashi et al. 2005, Castlebury et al. 2006, Halleen et al. 2006, Chaverri et al.
2011). Some of the species of Nectriaceae are important in agricultural industry as they can be used
for production of plant hormones eg. gibberellic acid production by Fusarium moniliforme
(Panchal & Desai 2016, Puyam et al. 2017).
Genera included in Nectriaceae
In this paper we only list the genera in Nectriaceae. Notes will be provided in a monograph.
Albonectria Rossman & Samuels, Stud. Mycol. 42: 105 (1999)
Index Fungorum number: IF27953; 1 morphological species.
Type species – Albonectria rigidiuscula (Berk. & Broome) Rossman & Samuels
Allantonectria Earle, Plant. Bak. 2(1): 11 (1901)
Index Fungorum number: IF128; 2 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Allantonectria miltina (Durieu & Mont.) Weese
733
Allonectella Petr., Sydowia 4(1–6): 345 (1950)
Index Fungorum number: IF140; 2 morphological species (Species Fungorum 2020).
Type species – Allonectella rubescens Petr.
Aphanocladium W. Gams, Cephalosporium-artige Schimmelpilze (Stuttgart): 196 (1971)
Index Fungorum number: IF7184; 3 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Aphanocladium album (Preuss) W. Gams
Aquanectria L. Lombard & Crous, Stud. Mycol. 80: 207 (2015)
Index Fungorum number: IF810949; 7 species with sequence data.
Type species – Aquanectria penicillioides (Ingold) L. Lombard & Crous
Atractium Link, Mag. Gesell. naturf. Freunde, Berlin 3(1–2): 10 (1809)
Index Fungorum number: IF7291; 5 morphological species (Species Fungorum 2020), 2
species with sequence data.
Type species – Atractium stilbaster Link, Mag. Gesell.
Baipadisphaeria Pinruan, Mycosphere 1: 58 (2011)
Index Fungorum number: IF518245; 1 species with sequence data.
Type species – Baipadisphaeria spathulospora Pinruan
Bisifusarium L. Lombard, Crous & W. Gams, Stud. Mycol. 80: 223 (2015)
Index Fungorum number: IF810226; 7 species with sequence data.
Type species – Bisifusarium dimerum (Penz.) L. Lombard & Crous
Calonectria De Not., Comm. Soc. crittog. Ital. 2(3): 477 (1867)
Index Fungorum number: IF746; 203 morphological species (Species Fungorum 2020), 157
species with sequence data.
Type species – Calonectria pyrochroa (Desm.) Sacc.
Calostilbe Sacc. & P. Syd., Syll. fung. (Abellini) 16: 591(1902)
Index Fungorum number: IF758; 3 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Calostilbe striispora (Ellis & Everh.) Seaver
Campylocarpon Halleen, Schroers & Crous, Stud. Mycol. 50(2): 448 (2004)
Index Fungorum number: IF28858; 3 species with sequence data.
Type species – Campylocarpon fasciculare Schroers, Halleen & Crous
Chaetonectrioides Matsush., Matsush. Mycol. Mem. 9: 5 (1996)
Index Fungorum number: IF27663; 1 morphological species.
Type species – Chaetonectrioides malaysiana Matsush.
Chaetopsina Rambelli, Atti Accad. Sci. Ist. Bologna, Cl. Sci. Fis. Rendiconti 3: 5 (1956)
Index Fungorum number: IF7584; 18 morphological species (Species Fungorum 2020), 7
species with sequence data
Type species – Chaetopsina fulva Rambelli
Coccinonectria L.Lombard & Crous, Stud. Mycol. 80: 218 (2015)
Index Fungorum number: IF810176; 2 species with sequence data.
Type species – Coccinonectria pachysandricola (B.O. Dodge) L. Lombard & Crous
734
Corallomycetella Henn., Hedwigia 43: 245 (1904)
Index Fungorum number: IF1237; 3 morphological species (Species Fungorum 2020), 2
species with sequence data.
Type species – Corallomycetella heinsenii Henn.
Corallonectria C. Herrera & P. Chaverri, Mycosystema 32(3): 539 (2013)
Index Fungorum number: IF803108; 1 species with sequence data.
Type species – Corallonectria jatrophae (Möller) C.Herrera & P. Chaverri
Corinectria C. González & P. Chaverri, Mycol. Progr. 16(11-12): 1021 (2017)
Index Fungorum number: IF822856; 1 species with sequence data.
Type species – Corinectria fuckeliana (C. Booth) C. González & P. Chaverri
Cosmospora Rabenh., Hedwigia 2: 59 (1862)
Index Fungorum number: IF1273; 48 morphological species (Species Fungorum 2020), 27
species with sequence data.
Type species – Cosmospora coccinea Rabenh.
Cosmosporella S.K. Huang, R. Jeewon & K.D. Hyde, Cryptog. Mycol. 39(2): 179 (2018)
Index Fungorum number: IF554371; 1 species with sequence data.
Type species – Cosmosporella olivacea S.K. Huang, R. Jeewon & K.D. Hyde
Curvicladiella Decock & Crous, Stud. Mycol. 55: 225(2006)
Index Fungorum number: IF500866; 1 species with sequence data.
Type species – Curvicladiella cignea (Decock & Crous) Decock & Crous
Cyanochyta Höhn., Sber. Akad. Wiss. Wien, Math.-naturw. Kl., Abt. 1 124: 92 (1915)
Index Fungorum number: IF7853; 1 morphological species.
Type species – Cyanochyta cyanogena (Speg.) Höhn.
Cyanonectria Samuels & P. Chaverri, Mycol. Progr. 8(1): 56 (2009)
Index Fungorum number: IF537057; 1 species with sequence data.
Type species – Cyanonectria cyanostoma (Sacc. & Flageolet) Samuels & P. Chaverri
Cyanophomella Höhn., Hedwigia 60: 156 (1918)
Index Fungorum number: IF25077; 1 morphological species.
Type species – Cyanophomella acervalis (Sacc.) Höhn.
Cylindrocladiella Boesew., Can. J. Bot. 60(11): 2289 (1982)
Index Fungorum number: IF7869; 45 morphological species (Species Fungorum 2020), 43
species with sequence data.
Type species – Cylindrocladiella parva (P.J. Anderson) Boesew.
Cylindrodendrum Bonord., Handb. Allgem. mykol. (Stuttgart): 98 (1851)
Index Fungorum number: IF7873; 42 morphological species (Species Fungorum 2020), 40
species with sequence data.
Type species – Cylindrodendrum album Bonord.
Dacryoma Samuels, Brittonia 40(3): 328 (1988)
Index Fungorum number: IF25250; 2 morphological species (Species Fungorum 2020).
Type species – Dacryoma album Samuels
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Dactylonectria L. Lombard & Crous, Phytopathol. Medit. 53: 348 (2014)
Index Fungorum number: IF810142; 13 morphological species (Species Fungorum 2020), 12
species with sequence data.
Type species – Dactylonectria macrodidyma (Halleen et al.) L. Lombard & Crous
Dematiocladium Allegr., Aramb., Cazau & Crous, Mycol. Res. 109(7): 836 (2005)
Index Fungorum number: IF28939; 2 species with sequence data.
Type species – Dematiocladium celtidis Allegr., Aramb., Cazau & Crous
Fusarium Link, Mag. Gesell. naturf. Freunde, Berlin 3(1–2): 10 (1809)
Index Fungorum number: IF8284; 298 morphological species (Species Fungorum 2020, Xia
et al. 2019), 189 species with sequence data.
Type species – Fusarium roseum Link
Fusicolla Bonord., Handb. Allgem.mykol. (Stuttgart): 150 (1851)
Index Fungorum number: IF8294; 10 morphological species (Species Fungorum 2020), 9
species with sequence data.
Type species – Fusicolla betae Bonord.
Geejayessia Schroers, Gräfenhan & Seifert, Stud.Mycol. 68(1): 124 (2011)
Index Fungorum number: IF519479; 6 morphological species (Species Fungorum 2020), 2
species with sequence data.
Type species – Geejayessia cicatricum (Berk.) Schroers
Gliocephalotrichum J.J. Ellis & Hesselt., Bull. Torrey bot. Club 89: 21 (1962)
Index Fungorum number: IF8340; 13 species with sequence data.
Type species – Gliocephalotrichum bulbilium J.J. Ellis & Hesselt.
Gliocladiopsis S.B. Saksena, Mycologia 46: 662 (1954)
Index Fungorum number: IF8341; 14 species with sequence data.
Type species – Gliocladiopsis sagariensis S.B. Saksena
Notes – A new isolate MFLUCC 19-0309 grouped with the ex-type of Gliocladiopsis tenuis
(IMI 68205) and other G. tenuis isolates (data not shown). Our collection is also similar to G. tenuis
in conidiophore and conidial dimensions (Crous & Wingfield 1993, Crous & Peerally 1996). Based
on morphological similarities and phylogenetic analysis we identified our fungus as Gliocladiopsis
tenuis, which is illustrated here (Fig. 15).
Ilyonectria P. Chaverri & C. Salgado, Stud. Mycol. 68(1): 69 (2011)
Index Fungorum number: IF518558; 23 species with sequence data.
Type species – Ilyonectria radicicola (Gerlach & L. Nilsson) P. Chaverri & C. Salgado
Macroconia (Wollenw.) Gräfenhan, Seifert & Schroers, Stud. Mycol. 68(1): 101 (2011)
Index Fungorum number: IF519441; 4 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Macroconia leptosphaeriae (Niessl) Gräfenhan & Schroers
Mariannaea G. Arnaud, Bull. trimest. Soc. mycol. Fr. 68:196 (1952)
Index Fungorum number: IF529380; 18 morphological species (Species Fungorum 2020),
16 species with sequence data.
Type species – Mariannaea elegans G. Arnaud
Microcera Desm., Annls Sci. Nat., Bot., sér. 3 10: 359 (1848)
736
Index Fungorum number: IF8920; 4 species with sequence data.
Type species – Microcera coccophila Desm.
Figure 173 – Gliocladiopsis tenuis (Material examined – THAILAND, Chiang Rai Province, Mae
Fah Luang University, on decaying fruit of a palm tree, 18 August 2017, R.H. Perera CR-F 12,
MFLU 19-0956; living culture, MFLUCC 19-0309). a, b Conidiomata on PDA. c-e Conidiophores,
conidiogenous cells with attached conidia. f, g Conidia. h Mycelium with chlamydospores. i
Chlamydospores. Scale bars: a, b = 1 mm, c-e, g = 50 µm, f, g = 20 µm, h, i = 50 µm.
737
Murinectria M. Niranjan & V.V. Sarma, gen. nov.
Index Fungorum number: IF556843; Facesoffungi number: FoF06587; 4 species with
sequence data.
Etymology – The generic name refers to Nectria spp. having muriform ascospores.
Sexual morph: Stromata immersed to erumpent, subiculate, solitary to aggregated, bright red.
Ascomata perithecial, superficial, globose, gregarious, immersed in the stroma, KOH+ve pigments,
often associated with synnemata of the asexual state. Peridium consists of cells of textura
globulosa and textura angularis, walls pigmented. Asci unitunicate, 8-spored, cylindric-clavate,
with or without an inconspicuous ring at apex, early deliquescent. Ascospores overlapping
biseriate, hyaline to pale brown, yellow in Lugol's solution, ellipsoidal to fusiform, muriform, often
constricted at each septum, straight or slightly curved. Asexual morph: Synnematous on natural
substrata, laterally oriented near perithecial ascomata, usually erumpent through epidermis, solitary
to gregarious around ascomata, erect or nodding, red-brown at base, becoming dark brown to black
with age, Conidiogenous cells phialidic. Conidia hyaline, obovoid, slightly flat to rounded apex,
narrow towards base, acute or obtuse ends.
Type species – Murinectria murispora M. Niranjan & V.V. Sarma.
Notes – Murinectria is characterized by muriform ascospores, which are similar to some
muriform species in Nectria. In our phylogenetic analysis, M. murispora branched separately from
other species of Nectria but clustered closely with Nectria species with muriform ascospores.
Hence, we establish a new genus Murinectria based on muriform ascospores in the sexual morph
and a synnematous asexual morph with phialospores.
Murinectria murispora M. Niranjan & V.V. Sarma, sp. nov.
Figs 175, 176
Index Fungorum number: IF556613; Facesoffungi number: FoF06267
Etymology – The specific epithet “murispora” refers to the fungus having muriform
ascospores.
Holotype – AMH-10077.
Saprobic on decaying climber. Sexual morph: Stromata up to 1 mm high and 3.2 mm diam.,
erumpent through epidermis, subiculate, pseudoparenchymatous cells forming textura prismatica
cell layers, intergrading with peridium. Ascomata 550–580 × 415–500 μm, perithecial, globose,
superficial, gregarious, KOH+ dark red, LA+ yellow, often associated with synnemata of the
asexual state, depressed apical region, periphysate, apical region without stroma. Peridium up to 43
μm thick, wall consists of textura globulosa and textura angularis cells, walls pigmented. Asci 64–
92 × 14.5–18 μm, 8-spored, unitunicate, cylindric-clavate, with an inconspicuous ring at apex, early
deliquescent. Ascospores (25–) 26–36.6 (–38) × 10 – 16.5 (–18.5) μm ( x = 32.8 × 14.9, n = 28),
overlapping biseriate, hyaline, yellow in Lugol's solution, ellipsoidal to fusiform, muriform, with
5–9 transverse septa and 1–5 longitudinal septa, often constricted at each septum, straight,
sometimes slightly curved, smooth-walled. Asexual morph: Synnematous on natural substrata, 892
μm length × 144–164 μm diameter, laterally oriented near perithecial ascomata, usually erumpent
through epidermis, solitary to gregarious around ascomata, erect or nodding, unbranched,
narrowing towards apex, red-brown at base, becoming dark brown to black with age, ovoid heads
consisting of pools of conidia, individual conidiophores septate. Conidiogenous cells phialidic.
Conidia 4.7–6.3 × 2.6–3.4 μm ( x = 5.5 × 3, n = 28), hyaline, obovoid, slightly flat to rounded apex,
narrow towards base, acute ends, smooth-walled.
Culture characteristics – White cottony colonies on malt extract agar, becoming gray-brown
at maturity, filamentous, radial, background pale yellow colour, 42 mm diameter in one-week old
culture grown at 28 °C.
Material examined – INDIA, Andaman and Nicobar Islands, South Andaman, Pongibalu,
Manjery (11˚52’25.7”N 92˚64’89.9”E), on decaying twig, 10 December, 2017. M. Niranjan PUFNI
17634 (AMH-10077, holotype), extype-living culture, NFCCI-4515.
GenBank numbers – ITS: MK860769, LSU: MK860767.
738
Notes – Murinectria murispora has similar characters to three species of Nectria in having
muriform ascospores (Table 1) (Hirooka et al. 2012). In comparison to Murinectria murispora,
Nectria polythalama stromata are shorter and narrower and have smaller ascomata and ascospores.
The synnemata (asexual state) on natural substrata are scattered or grow around the stromata of
Murinectria murispora, while in Nectria polythalama, the synnemata are frequently found in the
middle of the stromata. Nectria pseudotrichia also resembles Murinectria murispora in having
similar ascomata and muriform ascospores, but is distinct in having smaller ascomata and
ascospores. Nectria antarctica also has muriform ascospores. The ascostromata and ascospores of
N. antarctica, however, are smaller and have longer asci when compared to Murinectria murispora.
Murinectria murispora is quite similar to Nectria polythalama and N. pseudotrichia in terms of
septal constrictions, unlike N. antarctica. Murinectria murispora is distinct from all three species in
often having 5 longitudinal septa. Hence, based on the above mentioned differences and DNA
sequence analysis, a new species, M. murispora is introduced.
Murinectria antarctica (Speg.) M. Niranjan & V.V. Sarma, comb. nov.
Index Fungorum number: IF557053
Basionym – Nectria antarctica (Speg.) Rossman., Mem. New York Bot. Gard. 49: 25 (1989)
Murinectria pseudotrichia (Schwein) M. Niranjan & V.V. Sarma, comb. nov.
Index Fungorum number: IF557054
Basionym – Nectria pseudonectria (Schewein.) Berk. & M.A. Curtis, Journal of the Academy
of Natural Sciences, Phiadelphia 2: 289 (1853)
Murinectria polythalama (Berk) M. Niranjan & V.V. Sarma, comb. nov.
Index Fungorum number: IF557055
Basionym – Nectria polythalama Berk., The botany of the Antarctic Voyage II, Flora NovaeZealandiae 2: 203, t. 106:15 (1855)
Notes – Nectria was originally described with light to bright, soft-textured, superficial,
uniloculate perithecia, unitunicate asci and phialidic asexual morphs (Hirooka et al. 2012). For a
long time, the concept of Nectria was very broad with more than 1000 species described under
Nectria sensu lato. Many species of Nectria sensu lato have been placed in segregate genera in
Bionectriaceae and Nectriaceae (Rossman et al. 1999, Schoch et al. 2000, Luo & Zhuang 2010,
Chaverri et al. 2011). Saccardo (1883c) also separated genera based on single characters e.g.
species with medium length ascospores having more than one horizontal septum were placed in
Calonectria, while those with very long ascospores and multiple septa were placed in
Ophionectria.
Nectria and Thyronectria have some species with muriform ascospores (Jaklitsch &
Voglmayr 2014). Thyronectria species have immersed ascomata with a bright yellow scurf,
muriform ascospores that produce small ascoconidia in the asci and pycnidial asexual morphs. On
the contrary, Nectria is characterized by immersed to erumpent ascomata without a bright yellow
scurf and one to many horizontally septate to muriform ascospores, but lacks ascoconida and has
synnematous or sporodochial asexual morphs (Hirooka et al. 2012, Jaklitsch & Voglmayr 2014).
Rossman et al. (1999) accepted only 28 species in Nectria. Only N. antarctica, N. polythalama, and
N. pseudotrichia have muriform ascospores. A new taxon collected on decaying twigs in Andaman
Islands, India has similarities to Nectria and has muriform ascospores. In our molecular analysis
based on ITS and LSU sequence data (Fig. 174) it clustered with other muriform species of Nectria
(Fig. 15). These three species and the new taxon branched separately in our phylogram and were
distantly placed from other species of Nectria, such as N. balansiae. Hence, based on the
characteristics (muriform ascospores) and the molecular sequence analysis, a new genus
Murinectria is introduced with M. murispora as the type species. Nectria antarctica, N.
739
pseudotrichia and N. polythalama. which produce muriform ascospores are herein transferred to
Murinectria as M. antarctica, M. pseudotrichia and M. polythalama.
Figure 174 – Maximum parsimony phylogeny constructed using the ITS and LSU sequences from
strains of Nectriaceae with the new taxon Murinectria murispora.
740
Figure 175 – Murinectria murispora (AMH-10077, holotype). a, b Ascomata on host. c, d, g
Vertical section of ascomata. e Peridium. f Neck. h, i Culture on MEA plate. l Asci. j, k, m-o
Ascospores. Scale bars: d = 100 μm, d, f, g = 50 μm, j-o = 10 μm.
741
Figure 176 – Murinectria murispora (AMH 10077). a, b Synnemata on natural substrate. c-e
Synnemata. f, g Synnemata head. h, i Conidiophores and conidia. j, k Conidiogenous cells. l
Conidia. Scale bars: f = 200 μm, c, d, g, h = 50 μm, e, j, k 20 μm, i, l = 10 μm.
Nalanthamala Subram., J. Indian bot. Soc. 35: 478 (1956)
Index Fungorum number: IF9076; 6 morphological species (Species Fungorum 2020), 5
species with sequence data.
Type species – Nalanthamala madreeya Subram.
742
Table 1 List of the Murinectria species having muriform ascospores.
Fungi
Murinectria
antarctica
Ascomata
315–548 μm high
× 270–520 μm
diam
Asci
95–125 × 15–
25 μm
Ascospores
(19.4–)23.0–
30.4(–35.1) ×
(6.8–)8.1–10.9(–
13.6) μm
M. polythalama
300–435 μm high
× 290–345 μm
diam
70–96 × 15.7–
17.9 μm
(17.9–)21.8–
29.0(–35.4) ×
(6.1–)7.3–10.1(–
12.3) μm,
M. pseudotrichia
333–548 μm high
× 296–534 μm
diam
65–125 × 13–
32 μm
M. murispora
550–580 μm high
× 415–500 μm
diam
70–96 × 14.5–
18 μm
(14.8–)21.0–
28.8(–41.3) ×
(4.6–)7.5–11.4(–
15.0) μm
(25.0–)26.1–
36.6(–38.3) ×
(9.7–)10.2–16.3
(–18.6) μm,
Septation
5–8 transverse septa
and 1–2
longitudinal septum,
not constricted at
each septum
5–8 transverse
septate and 1–2
longitudinal septate,
often constricted at
each septum
5–8 transverse septa
and 1-2 longitudinal
septum, often
constricted
5–9 transverse
septate and 1–2 (–5)
longitudinal septate,
often constricted at
each septum
Nectria (Fr.) Fr., Summa veg. Scand., Sectio Post. (Stockholm): 387 (1849)
Index Fungorum number: IF3431; 931 morphological species (Species Fungorum 2020), 49
species with sequence data.
Type species – Nectria cinnabarina (Tode) Fr.
Notes – Nectria was recognized as Hypocrea sect. Nectria and raised to generic level by Fries
(1849) without designating a type species. Seaver (1909) designated N. peziza as the type, but it
was considered as illegitimate (Rossman et al. 1999). Clements & Shear (1931) made the first
legitimate typification of the genus by designating N. cinnabarina as the lectotype (Rossman et al.
1999). There were around 800 epithets proposed for Nectria sensu lato with 200 estimated species
(Booth 1959, Rossman 1996). Species in Nectria sensu stricto have been listed with a key to 28
species by Rossman (1989). Rossman et al. (1999) restricted Nectria sensu stricto to species
considered congeneric with the type species Nectria cinnabarina and accepted 27 species in the
genus. Based on morphology (Hirooka et al. 2010, Rossman et al. 1999) and LSU sequence data
(Rehner & Samuels 1995), Nectria sensu lato was separated into a number of genera.
Neocosmospora E.F. Sm., U.S.D.A. Div. Veg. Pathol. Bull.17: 45 (1899)
Index Fungorum number: IF3447; 70 morphological species (Species Fungorum 2020), 8
species with sequence data.
Type species – Neocosmospora vasinfecta E.F. Sm.
Neonectria Wollenw., Annls mycol. 15(1/2): 52 (1917)
Index Fungorum number: IF3469; 25 morphological species (Species Fungorum 2020), 18
species with sequence data.
Type species – Neonectria candida (Ehrenb.) Rossman, L. Lombard & Crous
Neothyronectria Crous & Thangavel, Persoonia 37: 329 (2016)
Index Fungorum number: IF819079; 2 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Neothyronectria sophorae Crous & Thangavel
743
Ophionectria Sacc., Michelia 1(no. 3): 323 (1878)
Index Fungorum number: IF3608; 11 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Ophionectria trichospora (Berk. & Broome) Sacc.
Pandanaceomyces Tibpromma & K.D. Hyde, Fungal Divers. 92: 107 (2018)
Index Fungorum number: IF554537; 1 species with sequence data.
Type species – Pandanaceomyces krabiensis Tibpromma & K.D. Hyde
Paracremonium L. Lombard & Crous, Stud. Mycol. 80: 233 (2015)
Index Fungorum number: IF810267; 5 species with sequence data.
Type species – Paracremonium inflatum L. Lombard & Crous
Payosphaeria W.F. Leong, Bot. Mar. 33: 511 (1990)
Index Fungorum number: IF25521; 1 morphological species.
Type species – Payosphaeria minuta H.Y.M. Leung
Penicillifer Emden, Acta bot. neerl. 17: 54 (1968)
Index Fungorum number: IF9256; 7 morphological species (Species Fungorum 2020), 5
species with sequence data.
Type species – Penicillifer pulcher Emden
Persiciospora P.F. Cannon & D. Hawksw., J. Linn. Soc.,Bot. 84: 133 (1982)
Index Fungorum number: IF3839; 4 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Persiciospora moreaui P.F. Cannon & D.Hawksw.
Pleiocarpon L. Lombard & D. Aiello, IMA Fungus 8(1): 73 (2017)
Index Fungorum number: IF820028; 3 species with sequence data.
Type species – Pleiocarpon strelitziae L. Lombard & D. Aiello
Pleogibberella Sacc., Syll.fung., Addit. I-IV (Abellini): 217 (1886)
Index Fungorum number: IF4211; 3 morphological species (Species Fungorum 2020).
Type species – Pleogibberella calamia (Cooke) Berl. & Voglino
Pleurocolla Petr., Annls mycol. 22(1/2): 15 (1924)
Index Fungorum number: IF9458; 1 morphological species.
Type species – Pleurocolla tiliae Petr.
Pseudoachroiostachys Tibpromma & K.D. Hyde, Fungal Divers. 92: 107 (2018)
Index Fungorum number: IF554547; 1 species with sequence data.
Type species – Pseudoachroiostachys krabiense Tibpromma & K.D. Hyde
Pseudocosmospora C. Herrera & P. Chaverri, Mycologia 105: 1291 (2013)
Index Fungorum number: IF802432; 13 morphological species (Species Fungorum 2020), 6
species with sequence data.
Type species – Pseudocosmospora eutypellae C. Herrera & P. Chaverri
Pseudonectria Seaver, Mycologia 1(2): 48 (1909)
Index Fungorum number: IF4460; 17 morphological species (Species Fungorum 2020), 2
species with sequence data.
Type species – Pseudonectria buxi DC.) Seifert, Gräfenhan & Schroers
744
Rectifusarium L. Lombard, Crous & W. Gams, Stud. Mycol. 80: 229 (2015)
Index Fungorum number: IF810252; 2 species with sequence data.
Type species – Rectifusarium ventricosum (Appel & Wollenw.) L. Lombard & Crous
Rugonectria P. Chaverri & Samuels, Stud. Mycol. 68(1): 73 (2011)
Index Fungorum number: IF518563; 5 species with sequence data.
Type species – Rugonectria rugulosa (Pat. & Gaillard) Samuels, P. Chaverri & Salgado
Sarcopodium Ehrenb., Synop. Pl. Crypt. 2: 101. (1824)
Index Fungorum number: IF9788; 15 morphological species (Species Fungorum 2020), 6
species with sequence data.
Type species – Sarcopodium circinatum Ehrenb.
Stylonectria Höhn., Sber. Akad. Wiss. Wien, Math.-naturw. Kl., Abt. 1 124: 52 (1915)
Index Fungorum number: IF5301; 5 species with sequence data.
Type species – Stylonectria applanata Höhn.
Thelonectria P. Chaverri & C. Salgado, Stud. Mycol. 68(1): 76 (2011)
Index Fungorum number: IF518567; 41 morphological species (Species Fungorum 2020),
29 species with sequence data.
Type species – Thelonectria discophora (Mont.) P. Chaverri & C. Salgado
Thyronectria Sacc., Grevillea 4(no. 29): 21 (1875)
Index Fungorum number: IF5469; 41 morphological species (Species Fungorum 2020), 33
species with sequence data.
Type species – Thyronectria rhodochlora (Mont.) Seeler
Varicosporella Lechat & J. Fourn., Ascomycete.org 7(1): 2 (2015)
Index Fungorum number: IF810690; 1 species with sequence data.
Type species – Varicosporella aquatica Lechat & J. Fourn.
Varicosporellopsis Lechat & J. Fourn., Ascomycete.org 8(3): 96 (2016)
Index Fungorum number: IF815311; 1 species with sequence data
Type species – Varicosporellopsis aquatilis Lechat & J. Fourn.
Volutella Tode, Fung. mecklenb. sel. (Lüneburg) 1: 28 (1790)
Index Fungorum number: IF39165; 32 morphological species (Species Fungorum 2020), 9
species with sequence data.
Type species – Volutella ciliata (Alb. & Schwein.) Fr.
Xenoacremonium L. Lombard& Crous, Stud. Mycol. 80: 234 (2015)
Index Fungorum number: IF810270; 2 species with sequence data.
Type species – Xenoacremonium recifei (Leão & Lõbo) L. Lombard & Crous
Xenocylindrocladium Decock, Hennebert & Crous, Mycol. Res. 101(7): 788 (1997)
Index Fungorum number: IF27788; 3 species with sequence data.
Type species – Xenocylindrocladium serpens Decock, Hennebert & Crous
Xenogliocladiopsis Crous & W.B. Kendr., Canad. J. Bot. 72: 63 (1994)
Index Fungorum number: IF27282; 2 species with sequence data.
Type species – Xenogliocladiopsis eucalyptorum Crous & W.B. Kendr.
745
Xenoleptographium Marinc., T.A. Duong, Z.W. de Beer & M.J. Wingf., Persoonia 35: 319 (2015)
Index Fungorum number: IF812683; 1 species with sequence data.
Type species – Xenoleptographium phialoconidium Marinc.
Xenonectriella Weese, Sber. Akad. Wiss. Wien, Math.-naturw. Kl., Abt. 1 128: 749 (1919)
Index Fungorum number: IF5822; 15 morphological species (Species Fungorum 2020).
Type species – Xenonectriella lutescens (Arnold) Weese
Neomelanconiellaceae Crous, Persoonia 41: 267 (2018)
Index Fungorum number: IF828247; Facesoffungi number: FoF07079; 1 species.
Endophytic forming leaf spots. Sexual morph: Undetermined. Asexual morph: Conidiomata
solitary to aggregated, pycnidial, brown with central ostiole. Conidiomatal wall comprising several
layers of medium brown cells of textura angularis. Conidiophores lining the inner cavity, septate,
hyaline, smooth, subcylindrical with slight apical taper, branched or not. Conidiogenous cells
hyaline, smooth, ampulliform, terminal and intercalary, phialidic. Conidia solitary, aseptate,
hyaline, smooth, guttulate, subcylindrical to narrowly ellipsoid, apex obtuse, tapering to truncate
hilum (adapted from Crous et al. 2018b).
Type – Neomelanconiella Crous
Notes – Neomelanconiellaceae was introduced by Crous et al. (2018b) based on single
collection of Neomelanconiella combreti. Phylogenetically, this family clusters between
Melanconiellaceae and Harknessiaceae.
Ecological and economic significance of Neomelanconiellaceae
Neomelanconiella combreti forms spots on leaves of Combretum sp., which is a native plant
to southern Africa, Madagascar, tropical Asia and tropical America. This plant is used in African or
Indian herbal medicine.
Genus included in Neomelanconiellaceae
Neomelanconiella Crous, Persoonia 41: 267 (2018)
Index Fungorum number: IF828246; 1 species with sequence data.
Type species – Neomelanconiella combreti Crous
Notes – The monotypic genus Neomelanconiella was introduced by Crous et al. (2018b)
based on N. combreti. The sexual morph is undetermined and the asexual morph is similar to
Melanconiella (Crous et al. 2018b). However, most of Melanconiella species occur on recently
dead twigs and branches of Betulaceae in the north temperate zone (Voglmayr et al. 2012), while
Neomelanconiella combreti occurs on leaves of Combretaceae in the Southern Hemisphere.
Neomelanconiella differs from Melanconiella in having pycnidial conidiomata without
ectostromatic discs and central columns and ellipsoid conidia without two large and numerous
small guttules. Phylogenetically Neomelanconiella combreti clustered with Cryptodiaporthe vepris
forming a distinct clade between Melanconiellaceae and Harknessiaceae (Crous et al. 2018b).
Niessliaceae Kirschst., Annls mycol. 37(1/2): 89 (1939)
Index Fungorum number: IF81070; Facesoffungi number: FoF01126; 13 species.
Saprobic or parasitic on wood, leaves and stems in freshwater and terrestrial habitats, some
possibly fungicolous and lichenicolous. Sexual morph: Ascomata perithecial or cleistothecial,
yellow, brown, dark brown to black, solitary or gregarious, superficial, erumpent to immersed in a
subiculum or crustose stroma, cupulate, globose to subglobose, membranaceous, tuberculate, with
or without circinately coiled, lightly pigmented, brown, septate, apical setae, collabent, or
collapsing laterally, or not collapsing, papilla present or lacking, when present with periphysate
ostiole. Peridium thick, membranaceous, outer layer composed of yellow, light brown or brown
cells of textura angularis; inner layer composed of hyaline cells of textura prismatica. Paraphyses-
746
Figure 177 – Neomelanconiella combreti (redrawn from Crous et al. 2018b). a Symptomatic leaf. b
Conidiogenous cell with conidia. c-d Conidia. Scale bars: b-c = 10 µm, d = 5 µm.
-filiform, hyaline, septate, or absent. Asci 8-spored, unitunicate, oblong to clavate, long or short
pedicellate, with narrowly discoid, J-, apical ring or ring absent, evanescent at maturity. Ascospores
1–3-seriate, hyaline to brown, ellipsoid to fusiform, slightly curved, aseptate to 1-septate, striate or
verrucose, smooth-walled. Asexual morph: Hyphomycetous. Mycelium white to light brown,
superficial, effused, thick subiculum. Conidiophores thin-walled, hyaline, unbranched, aseptate,
swollen, with a short and distinct collarette, terminating in a single conidium. Phialides straight,
smooth. Conidia oval to ellipsoid, aseptate to 1-septate, hyaline, smooth. Coelomycetous,
Conidiomata subimmersed, globose to subglobose, dark brown. Peridium thick-walled, wall
composed of brown to hyaline cells of textura angularis, becoming hyaline towards inner
conidiogenous region. Conidiophores hyaline, subcylindrical, branched apically, 1–2-septate.
Conidiogenous cells phialidic, hyaline, fusiform to ellipsoid, straight to curved, tapering towards a
subtruncate apex. Conidia oval to ellipsoid, aseptate to 2-septate, hyaline, with guttulate, smoothwalled (adapted from Maharachchikumbura et al. 2016b).
Type genus – Niesslia Auersw.
Notes – Niessliaceae was introduced by Kirschstein (1939) based on superficial, dark, setose
perithecial or cleistothecial ascomata, and is typified by Niesslia. Barr (1990b) proposed that
members of this family be accepted in a narrow sense and transferred genera from Sphaeriaceae
(Müller & von Arx 1962, 1973) and Trichosphaeriaceae, such as Melchioria, Pseudorhynchia and
Valetoniella (Barr 1983, Hawksworth et al. 1983, Eriksson & Hawksworth 1990). Samuels & Barr
(1997) referred the family to Hypocreales (including Atronectria, Circinoniesslia, Cryptoniesslia,
Malmeomyces, Melanopsamma, Miyakeomyces, Myrmaeciella, Niesslia, Paraniesslia,
Pseudorhynchia, Trichosphaerella, Taiwanascus, Valetoniella, Valetoniellopsis), based on the
short ostiolar periphyses and phialidic conidiogenous cells (Starbäck 1899, Hara 1913, Eriksson &
Hawksworth 1991, 1993, Hawksworth et al. 1995, Rossman et al. 1999, Cai & Hyde 2007b, Etayo
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& Sancho 2008). The placement of Niessliaceae in the Hypocreales is supported by molecular data
and includes Eucasphaeria, Hyaloseta, Myrtacremonium, Niesslia, Rosasphaeria, Trichosphaerella
and Valetoniellopsis (Tang et al. 2007, Jaklitsch & Voglmayr 2012, Lombard et al. 2015, Crous et
al. 2016a, 2017b). Many genera included in Niessliaceae lack sequence data and several are
monotypic and thus recollection and sequence data is needed.
Ecological and economic significance of Niessliaceae
Members of Niessliaceae are usually saprobic or/and parasitic on wood or lichens or other
fungi, such as Niesslia echinoides on Erioderma barbellatum and Pseudorhynchia mauritiana on
Cordemoya integrifolia (Müller & von Arx 1962, 1973, Barr 1990b, Samuels & Barr 1997,
Dulymamode et al. 2001, Tang et al. 2007, Jaklitsch & Voglmayr 2012, Etayo et al. 2013, Lombard
et al. 2015, Crous et al. 2016a, 2017b).
Genera included in Niessliaceae
Atronectria Etayo, Biblthca Lichenol. 98: 52 (2008)
Index Fungorum number: IF532866; 1 morphological species.
Type species – Atronectria magellanica Etayo
Notes – Atronectria was established based on Atronectria magellanica which has clavate asci
with an outer layer of textura epidermoidea and oblong, 1-septate ascospores (Etayo & Sancho
2008). Atronectria is similar to Pseudonectriella in having hyaline, oblong ascospores and it is the
only species with ascoma having an outer layer of textura epidermoidea in Niessliaceae (Etayo &
Sancho 2008).
Circinoniesslia Samuels & M.E. Barr, Can. J. Bot. 75(12): 2166 (1998)
Index Fungorum number: IF27818; 1 morphological species.
Type species – Circinoniesslia nectriae Samuels & M.E. Barr
Notes – The monotypic Circinoniesslia was established by C. nectriae and has ascomata
surrounded by circinate setae, cylindrical asci and oblong to ellipsoid ascospores with striate
appendages (Samuels & Barr 1997). The ascospores are similar to Valetoniella and it was separated
from other members of Niessliaceae based on the special circinate setae on the ascomata (Samuels
& Barr 1997).
Cryptoniesslia Scheuer, Mycol. Res. 97(5): 543 (1993)
Index Fungorum number: IF22420; 1 morphological species.
Type species – Cryptoniesslia setulosa Scheuer
Notes – Cryptoniesslia setulosa, the only species of Cryptoniesslia, has ascomata surrounded
by short, branched, apical setae, clavate asci and elongate ascospores (Scheuer 1993). The setae and
ascospores are similar to Niesslia erysipheoides and should be given generic status in Niessliaceae
based on different characters of immersed ascomata and clavate asci (Samuels & Barr 1997).
Eucasphaeria Crous, Fungal Divers. 25: 21 (2007)
Index Fungorum number: IF501093; 2 species with sequence data.
Type species – Eucasphaeria capensis Crous
Notes – Eucasphaeria capensis is characterized by brown ascomata, clavate asci with J+,
apical ring and fusoid-ellipsoidal ascospores and falcate conidia (Crous et al. 2007b). The second
species, Eucasphaeria rustici, was introduced by Crous et al. (2016a) based on phialidic
conidiogenous cells and falcate conidia. This genus is related to Rosasphaeria and Niesslia in
Niessliaceae (Crous et al. 2016a).
Hyaloseta A.W. Ramaley, Mycotaxon 79: 269 (2001)
Index Fungorum number: IF28517; 1 species with sequence data.
Type species – Hyaloseta nolinae A.W. Ramaley
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Notes – The monotypic Hyaloseta has ascomata surrounded by hyaline setae with swollen
apices, cylindrical asci and ellipsoid, 1-septate ascospores (Ramaley 2001). The genus is closely
related to Trichosphaerella based on multi-gene analysis (Lombard et al. 2015; Fig. 15) and has a
monocillium-like asexual morph.
Malmeomyces Starbäck, Bih. K. svenska VetenskAkad. Handl., Afd. 3 25(no. 1): 32 (1899)
Index Fungorum number: IF2989; 1 morphological species.
Type species – Malmeomyces pulchellus Starbäck
Notes – Malmeomyces pulchellus as the only member in Malmeomyces with cupulate
ascomata with septate setae, clavate asci and cylindrical ascospores (Starbäck 1899, Lumbsch &
Huhndorf 2007).
Melchioria Penz. & Sacc., Malpighia 11(9-10): 399 (1897)
Index Fungorum number: IF3099; 6 morphological species (Species Fungorum 2020).
Type species – Melchioria leucomelaena Penz. & Sacc.
Notes – Melchioria leucomelaena has glabrous ascomata, clavate asci and cylindrical
ascospores (Penzig & Saccardo 1897a).
Miyakeomyces Hara, Bot. Mag., Tokyo 27(no. 317): (248) (1913)
Index Fungorum number: IF3226; 1 morphological species.
Type species – Miyakeomyces bambusae Hara
Notes – The monotypic Miyakeomyces has cushion-like ascomata, cylindrical asci and
oblong-fusiform ascospores (Hara 1913).
Myrmaeciella Lindau, Nat. Pflanzenfam., Teil. I (Leipzig) 1(1): 478 (1897)
Index Fungorum number: IF3384; 2 morphological species (Species Fungorum 2020).
Type species – Myrmaeciella endoleuca (Sacc.) Lindau
Notes – Myrmaeciella endoleuca has black, turbinate stromata and clavate asci with ellipsoid,
verruculose ascospores (Saccardo 1880b; Rossman et al. 1999, Jaklitsch & Voglmayr 2011).
Rossman et al. (1999) suggested that Myrmaeciella is allied with members of Niessliaceae.
Myrtacremonium Crous, Persoonia 38: 259 (2017)
Index Fungorum number: IF820937; 1 species with sequence data.
Type species – Myrtacremonium eucalypti Crous
Notes – The asexual genus Myrtacremonium has solitary conidiophores and subcylindrical
straight conidia. The genus is closely related to Valetoniellopsis in Niessliaceae (Fig. 15).
Neoeucasphaeria Crous, Persoonia 41: 293 (2018)
Index Fungorum number: IF828249; 1 species with sequence data.
Type species – Neoeucasphaeria eucalypti Crous
Notes – Crous et al. (2018b) found Neoeucasphaeria eucalypti on Eucalyptus sp. in Australia.
The genus is related to Eucasphaeria, but has aseptate conidia that are different from Eucasphaeria
(Crous et al. 2018b).
Niesslia Auersw., Myc. Europ. Pyren. 5-6: 30 (1869)
Index Fungorum number: IF3506; 98 morphological species (Species Fungorum 2020), 11
species with sequence data.
Type species – Niesslia chaetomium (Ces. & De Not.) Auersw.
Notes – Niesslia is characterized by tuberculate perithecia, surrounded by brown, septate
setae, clavate asci and filiform ascospores (Gonnermann & Rabenhorst 1869, Gams et al. 2019).
The members of Niessliaceae nest in this family based on multi-gene analysis (Fig. 15).
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Paraniesslia C.K.M. Tsui, K.D. Hyde & Hodgkiss, Mycologia 93(5): 1002 (2001)
Index Fungorum number: IF28568; 2 morphological species (Species Fungorum 2020).
Type species – Paraniesslia tuberculata C.K.M. Tsui, K.D. Hyde & Hodgkiss
Notes – Paraniesslia tuberculata has pyriform to subglobose ascomata and clavate asci with
ellipsoid, verruculose ascospores (Saccardo 1880b, Rossman et al. 1999, Tsui et al. 2001b,
Jaklitsch & Voglmayr 2011). The second species Paraniesslia aquatica, a freshwater taxon, was
introduced by Cai & Hyde (2007b).
Pseudonectriella Petr., Sydowia 13(1-6): 127 (1959)
Index Fungorum number: IF4461; 1 morphological species.
Type species – Pseudonectriella ahmadii Petr.
Notes – Pseudonectriella ahmadii has cupulate, verruculose ascomata and cylindrical asci
with ellipsoid to oval ascospores (Petrak 1959).
Pseudohyaloseta Tibpromma & K.D. Hyde, Fungal Divers. 93: 113 (2018)
Index Fungorum number: IF 554542; 1 species with sequence data.
Type species – Pseudohyaloseta pandanicola Tibpromma & K.D. Hyde
Notes – Pseudohyaloseta is similar to Hyaloseta in having globose to subglobose ascomata
and obclavate asci with cylindrical and 1-septate ascospores, but they are well-separated in
phylogenetic analyses (Tibpromma et al. 2018).
Pseudorhynchia Höhn., Sber. Akad. Wiss. Wien, Math.-naturw. Kl., Abt. 1 118: 1206 (1909)
Index Fungorum number: IF4483; 2 morphological species (Species Fungorum 2020).
Type species – Pseudorhynchia polyrrhyncha (Penz. & Sacc.) Höhn.
Notes – Pseudonectriella was established by Pseudorhynchia polyrrhyncha which found on
Elettaria sp. in Java (Höhnel 1909b). The genus is characterized by subglobose ascomata with setae
surrounded by ostiole and cylindrical to clavate asci with ellipsoid ascospores. Dulymamode et al.
(2001) introduced the second species, Pseudorhynchia mauritiana, found on fallen leaves of
Cordemoya integrifolia in Mauritius.
Rosasphaeria Jaklitsch & Voglmayr, Fungal Divers. 52(1): 93 (2012)
Index Fungorum number: IF561139; 1 species with sequence data.
Type species – Rosasphaeria moravica (Petr.) Jaklitsch & Voglmayr
Notes – Rosasphaeria moravica has immersed, subglobose ascomata and cylindrical asci
with 1-septate, fusiform to cylindrical ascospores. The genus is closely related to Eucasphaeria in
Niessliaceae (Jaklitsch & Voglmayr 2012; Fig. 15).
Taiwanascus Sivan. & H.S. Chang, Mycol. Res. 101(2): 176 (1997)
Index Fungorum number: IF27752; 2 morphological species (Species Fungorum 2020).
Type species – Taiwanascus tetrasporus Sivan. & H.S. Chang
Notes – Taiwanascus tetrasporus has cleistothecial ascomata with branched setae and broadly
cylindrical asci with fusiform to ellipsoid ascospores (Sivanesan & Chang 1997). The second
species Taiwanascus samuelsii was collected in India (Rajeshkumar & Rossman 2013).
Trichosphaerella E. Bommer, M. Rousseau & Sacc., Bull. Soc. R. Bot. Belg. 29(no. 1): 261 (1890)
Index Fungorum number: IF5587; 7 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Trichosphaerella decipiens E. Bommer, M. Rousseau & Sacc.
Notes – Trichosphaerella decipiens has tuberculate ascomata with setae and cylindrical to
clavate asci with ellipsoid ascospores (Saccardo 1891, Döbbeler et al. 2015). Sequence data of T.
ceratophora were provided by Lombard et al. (2015) and reveals that Trichosphaerella is closely
related to Hyaloseta in Niessliaceae.
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Valetoniella Höhn., Sber. Akad. Wiss. Wien, Math.-naturw. Kl., Abt. 1 118: 1499 (1909)
Index Fungorum number: IF5700; 3 morphological species (Species Fungorum 2020).
Type species – Valetoniella crucipila Höhn.
Notes – Valetoniella was established by Valetoniella crucipila and has globose ascomata with
branched setae, clavate asci with ellipsoid ascospores (Höhnel 1909a). The other two species, V.
claviornata and V. pauciornata, were introduced by Samuels & Barr (1997). Valetoniella crucipila
is illustrated in this entry (Fig. 175).
Figure 178 – Valetoniella crucipila (Material examined – NEW ZEALAND, Auckland, Titirangi
beach reserve; on ascomata of Nectria haematococca and N. mammoidea, 18 September 1980,
Samuels GJ & Johnston PR, PDD no. 41743). a, b Material label. c Ascomata. d Ascoma cross
section. e Peridium. f Setae. g Asci stained in Congo red reagent. h Ascospores (d-f, h. from
permanent slides). Scale bars: c = 200 µm, d = 100 µm, e-g = 20 µm, h = 10 µm.
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Valetoniellopsis Samuels & M.E. Barr, Can. J. Bot. 75(12): 2175 (1998)
Index Fungorum number: IF27817; 1 species with sequence data.
Type species – Valetoniellopsis laxa Samuels & M.E. Barr
Notes – The monotypic Valetoniellopsis with V. laxa has globose ascomata with setae and
clavate asci with ellipsoid to cylindrical ascospores (Samuels & Barr 1997). The genus is closely
related to Myrtacremonium based on multi-gene analysis (Fig. 15).
Nitschkiaceae Nannf., Nova Acta R. Soc. Scient. upsal., Ser. 4 8(no. 2): 56 (1932)
Index Fungorum number: IF81071; Facesoffungi number: FoF01120; 110 species.
Saprobic or parasitic on wood, leaves or lichens, in freshwater, marine and terrestrial
habitats. Sexual morph: Ascomata perithecial, dark brown to black, gregarious, superficial to
erumpent, sitting on or in a subiculum or subiculum absent, cupuliform, globose to subglobose,
carbonaceous, coriaceous to membranaceous, rough, tuberculate, smooth or with short spines, with
or ostiolar opening indistinct, the apex collapsing when dry or shallowly cupulate when moist.
Subiculum thin or thick-walled, numerous or scanty, brown to dark brown, septate, branched
hyphae, with spiny termination. Peridium thick, composed of three layers, outer layer comprising
dark tissues, thin, carbonaceous; middle layer comprising dark brown to brown cells of textura
angularis, thick, membranaceous, while the inner layer has hyaline cells of textura prismatica, thin,
and membranaceous, munk pores present or absent. Paraphyes when present, filamentous, septate,
simple or branched. Asci 4- to multi-spored, unitunicate, clavate to cylindrical, long or short
pedicellate or sessile, ends blunt. Ascospores 2–3-seriate or irregularly arranged, hyaline or brown,
fusiform, allantoid or subcylindrical, slightly curved, 0–4-aseptate, concolourous, smooth-walled,
sheath and appendages present or absent, with or without guttules. Asexual morph: Undetermined
(adapted from Maharachchikumbura et al. 2016b).
Type genus – Nitschkia G.H. Otth
Notes – Nitschkiaceae was introduced by Nannfeldt (1932) based on erumpent, black, rough
ascomata and clavate asci with hyaline allantoid ascospores. Nannfeldt (1975) concluded that
Nitschkiaceae comprised three separate groups, but that the differences between them were not
sufficient to place them in separate families. He accepted five genera in this family and provided
additional taxonomic details with descriptions. Mugambi & Huhndorf (2010) accepted 12 genera
based on a multi-gene dataset and morphology (Lumbsch & Huhndorf 2010). The monotypic
genera Neochaetosphaerella and Tortulomyces were introduced based on morpholy (Vasilyeva et al.
2012, 2013). In this study, 14 genera are included in this family. Most taxa assigned to the family
need to be re-evaluated with sequence data analysis.
Ecological and economic significance of Nitschkiaceae
Members of Nitschkiaceae are usually saprobic or/and parasitic on wood, lichens or other
fungi and are distributed worldwide, such as Nitschkia parasitans on stromata of Nectria and
Rhagadostoma corrugatum on Peltigera sp. (Hawksworth 1980, Gonzalez & Martin 1996,
Vasilyeva et al. 2010).
Genera included in Nitschkiaceae
Acanthonitschkea Speg., Anal. Mus. nac. B. Aires, Ser. 3 17(10): 116 (1908)
Index Fungorum number: IF12; 9 morphological species (Species Fungorum 2020), 2
species with sequence data.
Type species – Acanthonitschkea argentinensis Speg.
Notes – Acanthonitschkea argentinensis has spinate ascomata and clavate asci with reniform
ascospores (Spegazzini 1908). It was considered as a member of Coronophoraceae (Müller & von
Arx 1973, von Arx 1981a, Subramanian & Sekar 1990). Mugambi & Huhndorf (2010) transferred
the genus to Nitschkiaceae based on multi-gene analysis. The genus is closely related to Nitschkia
in Fig. 15.
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Biciliosporina Subram. & Sekar, Kavaka 18(1-2): 69 (1993)
Index Fungorum number: IF21072; 1 morphological species.
Type species – Biciliosporina karwarensis Subram. & Sekar
Notes – Biciliosporina karwarensis was placed in Coronophoraceae and is characterized by
cylindrical asci and oval to reniform ascospores with filiform appendages at each end
(Subramanian & Sekar 1990). Maharachchikumbura et al. (2015, 2016b) and Wijayawardene et al.
(2018b) accepted it as member of Nitschkiaceae.
Botryola Bat. & J.L. Bezerra, Publicações Inst. Micol. Recife 431: 11 (1964)
Index Fungorum number: IF634; 1 morphological species.
Type species – Botryola tetrasperma Bat. & J.L. Bezerra
Notes – The monotypic Botryola has clavate asci and ellipsoid, 4-spored ascospores with 1
septa (Batista et al. 1964).
Fracchiaea Sacc., Atti Soc. Veneto-Trent. Sci. Nat. 2(1): 163 (1873)
Index Fungorum number: IF2008; 22 morphological species (Species Fungorum 2020), 2
species with sequence data.
Type species – Fracchiaea heterogenea Sacc.
Notes – Fracchiaea heterogenea has clavate, polysporous asci and oblong ascospores
(Saccardo 1873). The genus was placed in Coronophoraceae (Müller & von Arx 1973,
Subramanian & Sekar 1990) and Mugambi & Huhndorf (2010) transferred it to Nitschkiaceae
based on multi-gene analysis.
Groenhiella Jørg. Koch, E.B.G. Jones & S.T. Moss, Bot. Mar. 26(6): 265 (1983)
Index Fungorum number: IF25806; 1 morphological species.
Type species – Groenhiella bivestia Jørg. Koch, E.B.G. Jones & S.T. Moss
Notes – Groenhiella has clavate asci and subglobose ascospores with appendages formed by
fragmentation of a sheath (Koch et al. 1983). Groenhiella, a marine genus, was initially placed in
Coronophoraceae (Koch et al. 1983), but assigned to Nitschkiaceae (Eriksson & Hawksworth
1987).
Janannfeldtia Subram. & Sekar, Kavaka 18(1-2): 69 (1993)
Index Fungorum number: IF27492; 1 morphological species.
Type species – Janannfeldtia karwarensis Subram. & Sekar
Notes – The monotypic Janannfeldtia has clavate asci and reniform ascospores with filiform
appendages at each end (Subramanian & Sekar 1990).
Lasiosphaeriopsis D. Hawksw. & Sivan., Trans. Br. mycol. Soc. 74(2): 371 (1980)
Index Fungorum number: IF2658; 7 morphological species (Species Fungorum 2020).
Type species – Lasiosphaeriopsis salisburyi D. Hawksw. & Sivan.
Notes – Lasiosphaeriopsis salisburyi has 4-spored asci and broadly fusiform ascospores with
3–4 septa (Hawksworth 1980). The genus was collected on Peltigera rufescens associated with
Rhagadostoma lichenicola (Hawksworth 1980).
Loranitschkia Lar.N. Vassiljeva, Mikol. Fitopatol. 24(3): 207 (1990)
Index Fungorum number: IF25456; 1 morphological species.
Type species – Loranitschkia viticola Lar.N. Vassiljeva
Notes – Loranitschkia viticola is characterised by clavate asci and clavate ascospores with a
tail-like appendage and was isolated from Vitis sp. in Russia (Vasilyeva 1990, Vasilyeva et al.
2010).
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Neochaetosphaerella Lar.N. Vassiljeva, S.L. Stephenson & Chernyshev, Fungal Divers. 52(1): 192
(2012)
Index Fungorum number: IF519185; 4 morphological species (Species Fungorum 2020).
Type species – Neochaetosphaerella thaxteriospora Lar.N. Vassiljeva, S.L. Stephenson &
Chernyshev
Notes – The monotypic Neochaetosphaerella has clavate asci and broadly allantoid
ascospores with 3 septa (Vasilyeva et al. 2012).
Neotrotteria Sacc., Bulletino dell'orto Botanico della R. Universitá di Napoli 6: 45 (1921)
Index Fungorum number: IF3487; 1 morphological species.
Type species – Neotrotteria pulchella Sacc.
Notes – Neotrotteria has multi-spored asci with ellipsoid ascospores (Saccardo 1918) and
was treated as synonym of Acanthonitschkea (Nannfeldt 1975, Nannfeldt & Santesson 1975).
Vasilyeva et al. (2010) accepted the genus as synonym of Fracchiaea based on similar morphs such
as polysporous asci and hyaline ascospores. Maharachchikumbura et al. (2016b) considered
Neotrotteria as an independent genus in Nitschkiaceae based on a Quellkörper in the peridium and
its difference from Fracchiaea.
Nitschkia G.H. Otth, Bidr. Känn. Finl. Nat. Folk 23: 13 (1873)
Index Fungorum number: IF3515; 52 morphological species (Species Fungorum 2020), 5
species with sequence data.
Type species – Nitschkia fuckelii Nitschke
Notes – The taxonomy of Nitschkia introduced by Karsten (1873) is confused and currently
includes 65 species (Index Fungorum 2020). The name Nitschkia fuckelii was first introduced by
Nitschke (1870), but Saccardo (1873) suggested Coelosphaeria to replace Nitschkia, because the
spelling of “Nitschkia” was similar to the generic name for a diatom. Fitzpatrick (1923) reviewed
the nomenclature of Coelosphaeria and the name was rejected. Confusion as to the type species
arises because Fuckel (1870) introduced the type species N. fuckelii based on Sphaeria cupularis.
Neither taxon has been sequenced, and phylogenetic analysis is based on Nitschkia calyculus, N.
grevillei, N. meniscoidea and N. tetraspora (Mugambi & Huhndorf 2010).
Rhagadostoma Körb., Parerga lichenol. (Breslau) 5: 472 (1865)
Index Fungorum number: IF4689; 8 morphological species (Species Fungorum 2020).
Type species – Rhagadostoma corrugatum Körb.
Notes – Rhagadostoma was established by Rhagadostoma corrugatum and has clavate, 2–4spored asci and cylindrical ascospores with 1 septum (Körber 1865). The lichenicolous genus,
Rhagadostoma, was considered as a member of Sordariales (Navarro-Rosinés & Hladun 1994,
Navarro-Rosinés et al. 1999) and Huhndorf et al. (2004a) proposed a possible relationship with
Bertia. Maharachchikumbura et al. (2016b) accepted this genus in Nitschkiaceae.
Rhagadostomella Etayo, Biblthca Lichenol. 84: 109 (2002)
Index Fungorum number: IF28689; 1 morphological species.
Type species – Rhagadostomella gregaria Etayo
Notes – Rhagadostomella was established by Rhagadostomella gregaria and has clavate asci
and cylindrical ascospores with 1 septa (Etayo 2002, Diederich 2003). Diederich (2003) proposed
this genus to be included in Nitschkiaceae.
Tortulomyces Lar.N. Vassiljeva, S.L. Stephenson, Chernyshev & K.D. Hyde, Mycoscience 54(1):
111 (2013)
Index Fungorum number: IF564276; 1 morphological species.
Type species – Tortulomyces thailandicus Lar.N. Vassiljeva, S.L. Stephenson, Chernyshev &
K.D. Hyde
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Notes – The monotypic Tortulomyces was established for T. thailandicus which has
cylindrical asci and subglobose, 1-septate ascospores (Vasilyeva et al. 2013).
Figure 179 – Neotrotteria pulchella (Material examined – Taiwan, Kukuan; on stem of Lycium
chinense, Chen CY, 17 July 1990, IMI no. 354793). a Material label. b Ascostroma on the host. c
Collapsed ascoma. d Section through perithecium with quellkörper. e Ascoma cross section. f
Setea. g Branched hyphe of subiculum. h-i Asci. j Asocspores. Scale bars: b = 2 mm, c = 500 µm,
d-e = 200 µm, f-g = 50 µm, h-i = 20 µm, j = 10 µm.
Obryzaceae Körb., Syst. lich. germ. (Breslau): 427 (1855)
Index Fungorum number: IF81075; Facesoffungi number: FoF05205; 3 species.
Mainly lichenicolous with green algae on rocks or bark and seldom on mosses. Sexual
morph: Ascomata perithecial, pyriform, astromatic, immersed. Ostiole periphysate. Peridium
hyaline with ostiolar region appearing light brown at times. Asci 4-8-spored, unitunicate, clavate,
closely ellipsoid to ovoid, short-pedicellate, with barely visible apical ring, deliquescent at the base.
Ascospores overlapping 1–2-seriate, hyaline, aseptate, smooth-walled, fusiform to limoniform or
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ellipsoid, lacking any mucilaginous sheath or appendages. Asexual morph: Undetermined (adapted
from Cannon & Kirk 2007, Maharachchikumbura et al. 2016b).
Type genus – Obryzum Wallr.
Notes – Obryzaceae is an insufficiently studied family which has been closely associated with
plant pathogenic taxa in Gnomoniaceae, although it was mainly mentioned in lichenological studies
(McNeill et al. 2006). Gnomoniaceae was subsequently recommended for conservation by
Hawksworth & Eriksson (1988) with rejection of Obryzaceae and this suggestion was
acknowledged by McNeill et al. (2006). However, members in Gnomoniaceae can be distinguished
as the latter have ascomata with minimally developed stroma and most often, the ascospores are
small, typically less than 25 µm and aseptate or one-septate either at the median or excentric
position (Sogonov et al. 2008). Furthermore, the host substrates for Gnomoniaceae range from
leaves, twigs and stems from herbaceous plants to bark or wood (Sogonov et al. 2008). Lumbsch &
Huhndorf (2010) listed Obryzaceae in Sordariomycetes, families incertae sedis, which was
followed by Maharachchikumbura et al. (2015). Later, it was excluded from Sordariomycetes,
families incertae sedis (Maharachchikumbura et al. 2016b). Furthermore, based on the ICN Article
36.1, the name Obryzaceae is invalid since Körber (1855) established this family name “ad int.”. Its
validation should be attributed to Eriksson (1981.
Ecological and economic significance of Obryzaceae
Species of this family are lichenicolous.
Genus included in Obryzaceae
Obryzum Wallr., Naturgesch. Flecht. 1: 253 (1825)
Index Fungorum number: IF3537; 3 morphological species (Species Fungorum 2020).
Type species – Obryzum corniculatum (Hoffm.) Wallr.
Notes – Species of this genus have been reported to be derived mainly from temperate areas
and observed to be occasionally parasitic on cyanobacterial lichens of Leptogium genus. At first,
Obryzum corniculatum was Collema corniculatum. However, after it had been considered improper
to give the lichenicolous fungus similar name as its host, Collema corniculatum, the generic name
was changed to Obryzum (Lücking et al. 2017). Given the fact that Obryzum was at first mistakenly
introduced as generic name for a presumed pyrenocarpous gelatinous lichen (Wallroth 1825),
neither C. corniculatum nor Obryzum can be lectotypified on the lichenicolous fungus (Lücking et
al. 2017). In this case, a conservation proposal is essential if O. corniculatum and Obryzum are to
be maintained in their currently used sense (Lücking et al. 2017). Obryzum is now maintained in
the Pezizomycotina, genera incertae sedis and is still a poorly studied genus with insufficient
molecular data (Lücking et al. 2017). Obryzum corniculatum is illustrated in this entry.
Ophioceraceae Klaubauf, E.G. LeBrun & Crous, Studies in Mycology 79: 85–120 (2014)
Index Fungorum number: IF810201; Facesoffungi number: FoF01254; 23 species.
Saprobic on wood and other plant material, commonly isolated in aquatic habitats. Sexual
morph: Ascomata perithecial, solitary or in small groups, immersed or semi-immersed, dark
brown to black, globose to subglobose, coriaceous, ostiolate. Ostiole a long neck, lined with
periphyses. Paraphyses numerous, septate, tapering, hypha-like, thin-walled, filamentous,
branched, intermingled among asci. Peridium thick, 3-layered, blackened. Asci 8-spored,
unitunicate, subcylindrical to narrowly fusoid, apedicellate, with thimble-shaped, J-, apical ring.
Ascospores fasciculate, hyaline, yellowish in mass, filiform, septate. Asexual morph:
Undetermined (adapted from Maharachchikumbura et al. 2016b).
Type genus – Ophioceras Sacc.
Notes – The monotypic family Ophioceraceae was introduced by Klaubauf et al. (2014) in
Magnaporthales to accommodate Ophioceras. Ophiocera, contains species that mostly occur on
wood submerged in freshwater. Previously, Ophioceras was placed in Magnaporthaceae following-
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Figure 180 – Obryzum corniculatum (Material examined – UK, March 1876 W. Joshua W, M0042649, neotype). a-c Herbarium material. d Appearance of ascomata on host substrate. e
Transverse section through ascoma. f Transverse section through ostiole. g Peridium. h Paraphyses
i-j Asci. k Lack of apical ring. l, m Ascospores. Scale bars: d = 300 µm, e = 50 µm, f-g, j = 30 µm,
h, l = 10 µm, i = 20 µm.
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-the phylogenetic analysis of Chen et al. (1999) and Inderbitzin & Berbee (2001). However, based
on analysis of combined LSU and rpb1, Ophioceras clusters separately from Magnaporthaceae in
Magnaporthales (Thongkantha et al. 2009) and hence a new monotypic family was introduced
(Klaubauf et al. 2014).
Ecological and economic significance of Ophioceraceae
Saprobic taxa such as Ophioceras species have the ability to decompose lignocellulosic
matter in woody litter, resulting in softening of the wood and releasing nutrients in the form of
simple molecules that go back into the soil and can be reused by plants and all other organisms
(Yuen et al. 1998, Bucher et al. 2004). Thus, they play an important role in nutrient and carbon
cycling, biological diversity and ecosystem functioning (Palmer et al. 1997, Wong et al. 1998a).
Reátegui et al. (2005) isolated four new tetrahydropyran derivatives called ophiocerins A-D and a
new africane sesquiterpenoid (ophioceric acid) from cultures of the aquatic fungus venezuelense,
provided structures and relative stereochemistry of these compounds. Dong et al. (2010) isolated a
novel neolignan with an unprecedented dibenzo-1, 6-dioxacyclodecane carbon skeleton, ophiocerol
from cultural filtrates of the freshwater fungus Ophioceras dolichostomum. Ophioceras is therefore
a source of novel biologically active secondary metabolites.
Genus included in Ophioceraceae
Ophioceras Sacc., Syll. fung. (Abellini) 2: 358 (1883)
Index Fungorum number: IF3595; 23 morphological species (Species Fungorum 2020), 11
species with sequence data.
Type species – Ophioceras dolichostomum (Berk. & M.A. Curtis) Sacc.
Notes – Saccardo (1883c) introduced Ophioceras based on O. dolichostomum. The genus is
characterized by black ascomata with long necks, cylindrical asci with small, refractive, apical
rings and filiform ascospores (Teng 1934, Conway & Barr 1977, Shearer et al.1999, Tsui et al.
2001c, Thongkantha et al. 2009, Klaubauf et al. 2014). Ophioceras species are commonly
encountered on decaying woody substrates in freshwater habitats all over the world (Hyde 1992c,
Hyde & Goh 1998, Shearer et al. 1999, Tsui et al. 2001c, Thongkantha et al. 2009, Hu et al. 2012).
In this entry, Ophioceras commune is illustrated (Fig. 181).
Ophiocordycipitaceae G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, Stud. Mycol. 57: 35
(2007)
Index Fungorum number: IF504190; Facesoffungi number: FoF01315; 469 species.
Parasitic on arthropods, protozoans, rotifers, nematodes, humans, animals and fungi
worldwide. Sexual morph: Stromata or subiculum darkly pigmented or brightly coloured, tough,
fibrous, pliant to wiry, rarely fleshy, often with aperithecial apices or lateral pads. Ascomata
superficial to completely immersed, ordinal or oblique in arrangement. Asci unitunicate,
cylindrical, rarely fusoid to ellipsoid, usually with thickened and almost bulbous ascus apex.
Ascospores fasciculate, hyaline, usually filiform, multi-septate, disarticulating into part-spores or
non-disarticulating. Asexual morph: see notes (adapted from Maharachchikumbura et al. 2016b).
Type genus – Ophiocordyceps Petch
Notes – Ophiocordycipitaceae was introduced based on phylogenetic analyses. Quandt et al.
(2014) included Ophiocordyceps, Tolypocladium, Polycephalomyces, Purpureocillium, Drechmeria
and Harposporium in Ophiocordycipitaceae based on morphology and phylogenetic analyses.
Spatafora et al. (2015) confirmed this classification and introduced species combinations ito
Drechmeria, Harposporium, Ophiocordyceps and Purpureocillium. Perennicordyceps was
introduced by Matočec et al. (2014) based on morphology and phylogenetic analyses.
Later,Simmons et al. (2015) provided 23 species of Hirsutella with sequence data for 23 species of
Hirsutella. Most species in this family produce dark pigmented, tough to pliant stromata, that often
have aperithecial apices (Sung et al. 2007). Asexual generic names associated with-
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Figure 181 – Ophioceras commune (Material examined – CHINA, Yunnan Province, saprobic on
decaying wood submerged in Erhai lake, March 2015, X.Y. Liu, S-578, HKAS 92587; March
2015, H.Y. Su, S-536, HKAS 92569; June 2015, S.M. Tang, S-338, HKAS 92640). a, b Neck. c, d
Longitudinal section through ascomata. e Longitudinal section of peridium. f Paraphyses. g-j Asci.
k-o Ascospores. Scale bars: c, d = 200 μm, f-j = 50 μm, e, k-o = 30 μm.
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-Ophiocordyceps include Sorosporella (Sorokin 1888), Hirsutella (Doassans & Patouillard 1892,
Gams & Zare 2003), Hymenostilbe (Petch 1931a), Stilbella (Seifert 1985), Syngliocladium (Petch
1932), and Paraisaria (Samson & Brady 1983). Based on morphology and phylogenetic analyses,
Ophiocordyceps was protected and other names suppressed in accordance with the “one fungus one
name” (Quandt et al. 2014). Paraisaria was also resurrected as a genus under this family
(Mongkolsamrit et al. 2019).
Tolypocladium was proposed for protection over Elaphocordyceps and Chaunopycnis based
on molecular analysis (Quandt et al. 2014). There is a wide ranges of asexual lineages associated
with this family, some of which are restricted in their phylogenetic distribution, while others are
often found in distant lineages (Quandt et al. 2014). For example, verticillium-like conidiophores, a
common asexual morph of many species in several hypocrealean families, including
Ophiocordycipitaceae, Cordycipitaceae and Clavicipitaceae (Zare et al. 2000, Gams & Zare 2001,
Sung et al. 2001, 2007). Thus the asexual genera links are unresolved and need further research.
Ecological and economic significance of Ophiocordycipitaceae
Many species in Ophiocordycipitaceae are economically important fungi and Ophiocordyceps
sinensis is collected throughout the Himalaya’s with great value to the local populations.
Tolypocladium ophioglossoides is also reported as a Traditional Chinese Medicine (Quandt et al.
2015, Huang et al. 2017). Tolypocladium sp. is well-known as a producer of cyclosporine, which is
an important immunosuppressant drug and also increases the competitiveness of the fungus in the
environment (Bushley et al. 2013, Yang et al. 2018). The Zombie ant fungi have important roles in
the ecosystem by controlling the behaviour of ants (Evans et al. 2011, Araújo et al. 2018, de Bekker
et al. 2018).
Genera included in Ophiocordycipitaceae
Drechmeria W. Gams & H.-B. Jansson, Mycotaxon 22(1): 36 (1985)
Index Fungorum number: IF105294; 12 morphological species (Species Fungorum 2020), 5
species with sequence data
Type species – Drechmeria coniospora (Drechsler) W. Gams & H.B. Jansson
Notes – The genus was introduced to accommodate two hyphomycetes species (Gams &
Jansson 1985). Quandt et al. (2014) placed ten species (most of them pathogens of protozoans) in
the genus based on phylogenetic analyses, including species of Haptocillium and Cordyceps gunnii.
Species of the genus infect protozoa and have crescent-shaped, helical conidia (Gams & Jansson
1985, Quandt et al. 2014).
Harposporium Lohde, Tagbl. Versamml. Ges. Deutsch. Naturf. 47: 206 (1874)
Index Fungorum number: IF8454; 36 morphological species (Species Fungorum 2020), 9
species with sequence data.
Type species – Harposporium guillulae Lohde
Notes – Most species in this genus were reported as pathogens of nematodes and sexual
morphs of this genus are podocrella-like (Quandt et al. 2014, Spatafora et al. 2015). Species of the
genus infect nematodes and rotifers and have ascospores breaking into two lanceolate multi-septate
part-spores, sphaerical or hirsutella-like conidiogenous cells, and crescent-shaped or helicoid
conidia (Quandt et al. 2014, Spatafora et al. 2015).
Hirsutella Pat., Revue mycol., Toulouse 14(no. 54): 67 (1892)
Index Fungorum number: IF8538; 83 morphological species (Species Fungorum 2020), 39
species with sequence data
Type species – Hirsutella entomophila Pat.
Notes – The genus was introduced for hyphomycetes pathogenic on insects, mites and
nematodes (Simmons et al. 2015). The sexual morph of this genus was linked to Ophiocordyceps
(Simmons et al. 2015). The genus is characterised by synnematous or mononematous
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conidiophores, fusiform, conidia, with a distinct chromophilic slime layer or covered by a finely
warted epispore (Doassans & Patouillard 1892, Quandt et al. 2014).
Hymenostilbe Petch, Naturalist (Hull), ser. 3: 101 (1931)
Index Fungorum number: IF8598; 12 morphological species (Species Fungorum 2020), 1
species with sequence data
Type species – Hymenostilbe muscaria Petch
Notes – This genus was considered as pathogenic on arthropods (Samson & Evans 1975).
The sexual morph was linked to Ophiocordyceps (White et al. 2003). The genus is characterised by
polyblastic, cylindrical to clavate conidiogenous cells, bearing multiple apical or subapical
denticles (Samson & Evans 1975).
Ophiocordyceps Petch, Trans. Br. mycol. Soc. 16(1): 73 (1931)
Index Fungorum number: IF3598; 242 morphological species (Species Fungorum 2020), 125
species with sequence data.
Type species – Ophiocordyceps blattae (Petch) Petch
Notes – This genus is the type genus of Ophiocordycipitaceae (Sung et al. 2007). Quandt et
al. (2014) emended this genus based on morphology and phylogenetic analyses. Based on
phylogenetic analyses, the oldest name in the Ophiocordycipitaceae clade is Sorosporella, a
synonym of Syngliocladium. Sorosporella was suppressed (Quandt et al. 2014), while
Syngliocladium was previously treated with respect to the second oldest name Hirsutella (Evans &
Samson 1982). Ophiocordyceps should be protected over the other names. Most species of the
genus are considered as entomopathogenic and have darkly pigmented, tough, fibrous, pliant to
wiry, stipitate or sessile stromata, superficial to completely immersed perithecia and ascospores
disarticulating into part-spores or non-disarticulating (Sung et al. 2007). The asexual morphs of this
genus are members of Hirsutella, Hymenostible, Paraisaria and Syngliocladium (Sung et al. 2007).
Paraisaria Samson & B.L. Brady, Trans. Br. mycol. Soc. 81(2): 285 (1983)
Index Fungorum number: IF25834; 11 species with sequence data.
Type species – Paraisaria gracilis (Grev.) Luangsa-ard, Mongkolsamrit & Samson
Notes – The genus was resurrected from Ophiocordyceps to accommodate three new species
and eight combinations based on morphology and phylogenetic analyses (Mongkolsamrit et al.
2019). The genus is characterised by robust solitary stroma with globose or ovoid, fertile heads, and
fleshy and cream stipes (Mongkolsamrit et al. 2019). Asexual morphs are developed as white, loose
synnemata, with verticillately branched conidiophores and narrowly cylindrical to fusiform conidia
(Mongkolsamrit et al. 2019).
Perennicordyceps Matočec & I. Kušan, Ascomycete.org 6(5): 129 (2014)
Index Fungorum number: IF810775; 4 species with sequence data.
Type species – Perennicordyceps prolifica (Kobayasi) Matočec & I. Kušan
Notes – This genus was introduced to accommodate 4 species, which have superficial
perithecia and hirsutella-like or acremonium-like asexual morphs.
Polycephalomyces Kobayasi, Sci. Rep. Tokyo Bunrika Daig., Sect. B 5: 245 (1941)
Index Fungorum number: IF9494; 17 morphological species (Species Fungorum 2020), 13
species with sequence data.
Type species – Polycephalomyces formosus Kobayasi
Notes – Polycephalomyces species are found in subtropical regions and are parasitic or
hyperparasitic on insects (Xiao et al. 2018). This genus was considered as pathogenic on insects or
mycoparasites on Ophiocordyceps species and has stipitate, bright stromata, immersed perithecia,
ascospores which break into part spores, lanceolate to narrowly lageniform phialides and one or
two types of conidia (Matočec et al. 2014, Xiao et al. 2018).
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Figure 182 – Ophiocordyceps xuefengensis (Material examined – CHINA, Hunan Province, on
Phassus nodus in root of living Clerodendrum cyrtophyllum, 20 October 2012, Ru-Cai Zhu, GZUH
2012HN14, holotype). a Habitat of Ophiocordyceps xuefengensis. b Overview of stromata and the
host. c Adult of the host. d Brown, superficial ascomata on stroma. e-g Cross sections showing the
complete stromata and perithecia. h-j Asci. k Part of ascospore. Scale bars: Scale bars: e-g = 200
µm, h-k= 50 µm.
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Figure 183 – Ophiocordyceps xuefengensis (Material examined – CHINA, Hunan Province, on
Phassus nodus in root of living Clerodendrum cyrtophyllum, 20 October 2012, Ru-Cai Zhu,
GZUCCHN131, ex-paratype living culture). a Overview of stromata and the host. b Colonies on
PDA after 1 week. c Back of colonies on PDA after 1 week. d Colonies on PDA after 3 weeks. e
Back of colonies on PDA after 1 week. f Solitary conidiogenous cells from vegetative hyphae. g
Opposite conidiogenous cells. h Conidiogenous cells in whorls. i Conidia. Scale bars: b-e = 2 cm, fh = 10 µm, i= 5 µm.
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Purpureocillium (Thom) Luangsa-ard, Houbraken, Hywel-Jones & Samson, FEMS Microbiol.
Lett. 321(2): 144 (2011)
Index Fungorum number: IF519529; 5 species with sequence data.
Type species – Purpureocillium lilacinum (Thom) Luangsa-ard, Houbraken, Hywel-Jones &
Samson
Notes – The genus was described for one emended medical species, Paecilomyces lilacinum,
based on morphology and phylogenetic analyses (Luangsa-ard et al. 2011). Two new species were
added by Perdomo et al. (2013a) and Hyde et al. (2016b) and 2 new combinations were introduced
by Ban et al. (2015) and Spatafora et al. (2015). No sexual morph has been reported for this genus.
The genus is characterised by ovate to cylindric phialides with distinct necks or erect and densely
grouped phialides, which are purple in mass (Matočec et al. 2014, Xiao et al. 2018).
Tolypocladium W. Gams, Persoonia 6(2): 185 (1971)
Index Fungorum number: IF10242; 47 morphological species (Species Fungorum 2020), 24
species with sequence data.
Type species – Tolypocladiumin inflatum W. Gams
Notes – This genus was was protected over Elaphocordyceps and Chaunopycnis by Quandt et
al. (2014). It contains parasites of insects, rotifers and other fungi or is a soil-inhabitant (Quandt et
al. 2014). The asexual morph is characterised by solitary or branched, cylindrical to conical
phialides and cylindrical to globose conidia, the sexual morph is characterised by darkly
pigmented, stipitate stromata, half-immersed to immersed perithecia, a yellow to brown pigmented
peridium and ascospores breaking into part-spores (Gams 1971, Bissett 1983, Gazis et al. 2014).
Ophiostomataceae Nannf., Nova Acta R. Soc. Scient. upsal., Ser. 4 8(no. 2): 30 (1932)
Index Fungorum number: IF91154; Facesoffungi number: FoF01807; 353 species.
Saprobic or parasitic on woody plants, occasionally on herbaceous plants, symbionts of or
associated with beetles and mites. Sexual morph: Ascomata superficial, perithecial or cleistothecial,
globose to subglobose, with external hairs, pale brown, dark brown or black, with long or short
ostiolar necks, or sometimes lacking ostioles. Ostiolar neck short or long, slender, straight or
flexuous, brown to dark brown at the base, pale brown to hyaline at the apex, containing parallel
hyphae with pigmented cell walls around the ostiole. Ostiolar hyphae present or absent; when
present, convergent or divergent, filamentous, septate, pale brown to hyaline, with some genera
containing ostiolar beak (e.g. Subbaromyces). Peridium pseudoparenchymatous, with outer wall
layers composed of angular cells with thickened, pigmented walls, and lined by thin-walled,
hyaline cells of textura angularis. Asci 8-spored, unitunicate, evanescent, fusiform, oblong to
clavate, some genera globose to subglobose (e.g. Fragosphaeria), with thin, deliquescent walls.
Ascospores bi- to multi-seriate, variable in shape, frequently asymmetrical, oval, allantoid,
fusiform, cylindrical, ellipsoidal, reniform, or falcate, aseptate or 1-septate, hyaline or appearing
yellowish in mass, with or without a sheath. Asexual morph: Hyphomycetous. Conidiophores
mononematous or synnematous, simple or branched, with some bearing a brush-like apical
branching structure (e.g. Leptographium), erect, hyaline or dark brown to black. Conidiogenous
cells holoblastic, polyblasic, sympodial, denticulate, phiailidic, with some proliferating at or
somewhat below the apex and giving rise to another series of denticles or conidiogenous loci (e.g.
Ophiostoma, Sporothrix). Conidia solitary or aggregated, fusiform, obovate to oblong, cylindrical,
aseptate, hyaline, some bearing conidia in a viscoid pale reddish-brown head (e.g. pesotum-like)
(adapted from Maharachchikumbura et al. 2016b).
Type genus – Ophiostoma Syd. & P. Syd.
Notes – Ophiostomataceae was originally described with three genera (Ophiostoma,
Ceratostomella, Endoconidiophora) (Nannfeldt 1932). de Beer et al. (2013a) accepted six genera
(Ceratocystiopsis, Ophiostoma, Fragosphaeria, Leptographium, Raffaelea, Graphium) based on
phylogenetic analyses. Maharachchikumbura et al. (2015, 2016b) provided an updated outline of
Ophiostomataceae with 12 genera (Ceratocystiopsis, Fragosphaeria, Hyalobelemnospora,
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Hyalorhinocladiella, Klasterskya, Leptographium, Ophiostoma, Pesotum, Phialographium,
Raffaelea, Spumatoria, Subbaromyces), which included sexual and asexual morphs, based on a
phylogenetic analysis. Pesotum and Hyalorhinocladiella appear to be synonyms of Ophiostoma,
but this may need further study. de Beer et al. (2016a) accepted Sporothrix in Ophiostomatales
based on DNA sequence data and morphology of both morphs. Bateman et al. (2016) introduced
Afroraffaelea. We accept 13 genera based on previous research (Afroraffaelea, Aureovirgo,
Ceratocystiopsis, Fragosphaeria, Graphilbum, Hawksworthiomyces, Klasterskya, Leptographium,
Ophiostoma, Raffaelea, Sporothrix, Spumatoria, and Subbaromyces) (Bateman et al. 2016, de Beer
et al. 2013a, 2016a, Maharachchikumbura et al. 2015b, 2016b).
Ecological and economic significance of Ophiostomataceae
Members of this family are mostly saprobes on sap-wood. Species are specialized with sticky
spores to facilitate insect dispersal. Many bark beetles act as vectors of ophiostomatoid fungi, e.g.
Ophiostoma, Leptographium, Ceratocytiopsis species (Upadhyay 1981, Wingfield et al. 1993,
Jacobs & Wingfield 2001, Zipfel et al. 2006). Most species cause sap stain or blue stain of freshly
cut logs and affect timber quality (Seifert 1993), and several species are important pathogens on
species of Protea, Pinus, Larix, Tsuga and Pistacia and other tree species (Wingfield et al. 1988,
Jacobs & Wingfield 2001, Roets et al. 2013, Zhou et al. 2013).
Genera included in Ophiostomataceae
Afroraffaelea C.C. Bateman, Y.T. Huang & D.R. Simmons, Fungal Ecology 25: 46 (2016)
Index Fungorum number: IF816236; 1 species with sequence data.
Type species – Afroraffaelea ambrosiae C.C. Bateman, Y.T. Huang & D.R. Simmons
Notes – This monotypic genus was identified as symbionts of Premnobius cavipennis from
pharyngeal mycangia, which were collected in the USA. The fungus was introduced based on
culture characteristic on potato dextrose agar (PDA), sequencing data, and microtome sectioning
wtih micro-CT scanning of mycangia (Bateman et al. 2016).
Aureovirgo J.A. van der Linde, Z.W. de Beer & Jol. Roux, Antonie van Leeuwenhoek 109: 593
(2016)
Index Fungorum number: IF813870; 1 species with sequence data.
Type species – Aureovirgo volantis J.A. van der Linde, Z.W. de Beer & Jol. Roux
Notes – Aureovirgo volantis was isolated from galleries of Cyrtogenius africus, on diseased
Euphorbia ingens in Limpopo Province, South Africa (van der Linde et al. 2016). The sexual
morph has globose ascomata with ostiolar hyphae and contains lunate ascospores with sheaths
attenuated around each end. The asexual morph is leptographium-like with holoblastic, oblongelliptical and oval conidia (van der Linde et al. 2016).
Ceratocystiopsis H.P. Upadhyay & W.B. Kendr., Mycologia 67(4): 799 (1975)
Index Fungorum number: IF889; 5 morphological species (Species Fungorum 2020), 4
species with sequence data.
Type species – Ceratocystiopsis minuta (Siemaszko) H.P. Upadhyay& W.B. Kendr.
Notes – Ceratocystiopsis has globose to subglobose ascomata with long necks and hyaline
ascospores with or without gelatinous sheaths; the asexual morph has sympodially proliferating
conidiogenous cells, with holoblastic, hyaline, aseptate conidia.
Fragosphaeria Shear, Mycologia 15(3): 124 (1923)
Index Fungorum number: IF2011; 2 species with sequence data.
Type species – Fragosphaeria purpurea Shear
Notes – Fragosphaeria is characterised by globose ascomata lacking ostioles and globose
ascospores produced in yellowish-brown masses from subglobose asci. The asexual morph has
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micronematous, unbranched or sparingly branched, sympodial and holoblastic conidia (Shear
1923).
Graphilbum H.P. Upadhyay & W.B. Kendr., Mycologia 67(4): 800 (1975)
Index Fungorum number: IF8393; 13 morphological species (Species Fungorum 2020), 10
species with sequence data.
Type species – Graphilbum sparsum H.P. Upadhyay& W.B. Kendr.
Notes – Graphilbum sparsum was collected from Picea glauca in Alaska. The genus is
characterised by allantoid to reniform ascospores with sheaths and the asexual morph has
synnematous/sporothrix-like conidiophores with solitary and holoblastic reniform to obovate
conidia.
Hawksworthiomyces Z.W. de Beer, Marinc. & M.J. Wingf., Fungal Biology 120(11): 1329 (2016)
Index Fungorum number: IF815685; 4 species with sequence data.
Type species – Hawksworthiomyces lignivorus (De Mey., Z.W. de Beer & M.J. Wingf.) Z.W.
de Beer, Marinc. & M.J. Wingf.
Notes – The genus was introduced to accommodate the type species, Hawksworthiomyces
lignivorus, based on morphology and phylogenetic analyses (de Beer et al. 2016b). The asexual
morphs have micronematous, unbranched or sparingly branched conidiophores and holoblastic,
ellipsoidal to cylindrical conidia (de Beer et al. 2016b). The sexual morph is undetermined. Three
species were detected by using environmental nucleic acid sequences (ENAS) to assess Fungal
Diversity. (de Beer et al. 2016b).
Klasterskya Petr., Annls mycol. 38(2/4): 225 (1940)
Index Fungorum number: IF2569; 3 morphological species (Species Fungorum 2020).
Type species – Klasterskya acuum (Mouton) Petr.
Notes – Klasterskya was introduced to accommodate Lentomita acuum found on dry leaves
of Pinus sylvestris. The genus is characterised by subglobose to obpyriform ascomata having
ostioles with fimbriate hyphae and 1-septate, elliptical to cylindrical ascospores, surrounded by a
hyaline gelatinous sheath, discharged in gummy masses (Petrak 1940, Valldosera & Guarro 1989).
The asexual morph is undetermined.
Leptographium Lagerb. & Melin, Svensk SkogsvårdsföreningTidskr. 25: 257 (1927)
Index Fungorum number: IF8749; 74 morphological species (Species Fungorum 2020), 68
species with sequence data.
Type species – Leptographium lundbergii Lagerb. &Melin
Notes – Leptographium lundbergii was introduced from wood of Coniferae in Sweden. The
strain was introduced as the asexual morph of Grosmannia (Lagerberg et al. 1927). de Beer et al.
(2013b) synonymized Grosmannia under Leptographium. The genus causes sap-stain of mainly
conifer timber, and less often of hardwoods (Harrington & Cobb 1988, Wingfield et al. 1993,
Jacobs & Wingfield 2001). A few species are saprobes found in the soil or decaying plant material,
and some are important tree pathogens (Harrington & Cobb 1988). Like most ophiostomatoid taxa,
Leptographium spp. are best known for their association with bark beetles (Harrington & Cobb
1988, Kirisits 2004). They are adapted to be carried by these insects, having erect conidiophores or
ascomata with long necks and conidia and ascospores produced in slimy masses at the apices of
these structures (Six 2003, Kirisits 2004, Cardoza et al. 2008).
Ophiostoma Syd. & P. Syd., Annls mycol. 17(1): 43 (1919)
Index Fungorum number: IF3614; 134 morphological species (de Beer et al. 2013), 62
species with sequence data.
Type species – Ophiostoma piliferum (Fr.) Sydow & P. Sydow
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Notes – de Beer et al. (2013a) accepted 134 species in Ophiostoma and some species are
important tree pathogens, such as O. ulmi and O. novo-ulmi, the causal agents of Dutch elm
disease, transmitted by elm bark beetles (Webber & Gibbs 1989, Brasier 1991). The sexual morph
is characterized by globose to subglobose ascomata, surrounded by external hairs, with ostiolar
necks and some species containing divergent, ostiolar hyphae. The asexual morph has sporothrixlike or pesotum-like conidiogenous cells with ellipsoidal to cylindrical obovate, globose and
aseptate conidia. The asexual morph of Ophiostoma piliferum is illustrated in this entry (Figs. 184,
185).
Figure 184 – Ophiostoma piliferum (redrawn from Osorio 1985). a Ascoma. b Ostiolar region with
ostiolar hyphae. c Ascospores. Scale bars: a-c = 10 µm, b = 5 µm.
Raffaelea Arx & Hennebert, Mycopath. Mycol. appl. 25: 310 (1965)
Index Fungorum number: IF9685; 33 morphological species (Species Fungorum 2020), 30
species with sequence data.
Type species – Raffaelea ambrosiae Arx & Hennebert
Notes – Raffaelea ambrosiae is associated with Platypus cylindrus on Quercus in the UK.
Raffaelea is a genus associated with ambrosia beetles worldwide (Harrington et al. 2010, de Beer et
al. 2013a, Musvuugwa et al. 2015, Simmons et al. 2016). These fungi commonly occur in
symbioses with wood-boring ambrosia beetles, Coleoptera, Curculionidae, Scolytinae and
Platypodina and cause sap stain or blue stain of freshly cut logs and affect timber quality. The
genus is characterised by hyphae bearing typical phialides and conidiogenous cells with
globose/ovoid to elongated conidia, often truncate at the base, with conidia budding yeast-like
daughter cells. The sexual morph is undetermined.
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Figure 185 – Ophiostoma piliferum. a Conidiophores and conidia with sporothrix-like
conidiogenous cells. b Conidiophores and conidia with raffaelea-like conidiogenous cell structure
(redrawn from Osorio 1985). c Conidial apparatus. d Conidial apparatus from various living
cultures with conidiogenous cells. e Conidia (redrawn from de Hoog 1974). Scale bars: a-e = 10
µm.
Sporothrix Hektoen & C.F. Perkins, J. Exp. Med. 5: 80 (1901)
Index Fungorum number: IF10046; 80 morphological species (Species Fungorum 2020), 60
species with sequence data
Type species – Sporothrix schenckii Hektoen & C.F. Perkins
Notes –“Sporotrichosis”, a human disease caused by Sporothrix schenckii was reported by
Benjamin Schenck in 1898. The disease agent was isolated from a specimen taken from cutaneous
lesions of a patient (Hektoen & Perkins 1900). Sporothrix schenckii lives in soil, plants and many
mammals (Barros et al. 2011, Chakrabarti et al. 2015). The S. schenckii species complex occurs
worldwide, being most commonly found in Africa, America and Asia (Sizar & Talati 2019). de
Beer et al. (2013b) reported Sporothrix schenckii/Ophiostoma stenoceras is a species complex in
Ophiostomasensu lato, representing a distinct genus in Ophiostomatales. Based on phylogenetic
analyses of four gene regions (LSU, ITS, tub2, calM), de Beer et al. (2016b) concluded that
Sporothrix was distinct from Ophiostoma sensu stricto, and synonymised Sporotrichopsis and
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Dolichoascus under this genus. The sexual morph is characterized by globose ascomata with long
necks and brown to black bases, 8-spored, evanescent globose to broadly clavate asci and hyaline,
aseptate, lunate, allantoid, reniform ascospores. The asexual morph is dimorphic with a mycelial
and a yeast phase (de Hoog 1974).
Spumatoria Massee & E.S. Salmon, Ann. Bot., Lond. 15: 350 (1901)
Index Fungorum number: IF5175; 1 species with sequence data.
Type species – Spumatoria longicollis Massee & E.S. Salmon
Notes –Spumatoria longicollis was found on dung of Equus caballus in the UK. The genus is
characterised by globose perithecia, with one or seldom two necks, with ostiolar hyphae and
hyaline, 2(−3)-celled, clavate, coarsely guttulate ascospores. The asexual morph is sporothrix-like,
with dacrioid conidia and blastoconidia growing directly from undifferentiated hyphae (Giraldo et
al. 2017a).
Subbaromyces Hesselt., Bull. Torrey bot. Club 80: 511 (1953)
Index Fungorum number: IF5303; 2 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Subbaromyces splendens Hesselt.
Notes – Subbaromyces splendens was repeatedly found growing on trickling filter beds in
New York. Hesseltine (1953) reported both the sexual and asexual morphs. The genus is
characterised by ascomata with ostiolar necks and 1-septate ascospores surrounded by a sheath. The
asexual morph has micronematous conidiophores, with non-apically branching structures and
solitary, holoblastic, oblong-elliptical to obovate conidia.
Oxydothidaceae S. Konta & K.D. Hyde, Mycosphere 7(9): 1431 (2016)
Index Fungorum number: IF552561; Facesoffungi number: FoF02700; 79 species.
Saprobic, endophytic or parasitic on monocotyledons. Sexual morph: Ascomata solitary or
irregularly scattered, ellipsoidal or subglobose, immersed or semi-immersed, slightly raised from
host surface with light or darkened discs, with most taxa lying horizontal to the host surface.
Peridium thick-walled, brown to dark brown, and/or peridium cells merging with the host tissue.
Paraphyses hypha-like, filamentous, irregular, septate, persisting between asci, but often
fragmenting in dried material. Asci 8-spored, unitunicate, cylindrical, pedicellate, with a J+ (rarely
J-), subapical ring. Ascospores fasciculate, hyaline, sometimes appearing yellow in mass, fusiform
or filiform, 1-septate, tapering from the center to spine-like, pointed or rounded ends. Appressoria
produced by germinating ascospores in some species, solitary, hyaline or greenish, or pale brown to
brown, irregular in shape, thick-walled. Asexual morph: Hyphomycetous. Selenosporella sp.
Conidiophores mononematous, (1–)2–3-septate, unbranched or 1-branced, brown olivaceous, thickwalled at below, thin-walled and colourless above, unicellular, colourless, distinct obviously
differentiated apex or base (adapted from Samuels & Rossman 1987, Konta et al. 2016).
Type genus – Oxydothis Penz. & Sacc.
Notes – Oxydothidaceae (Konta et al. 2016) was established to accommodate the monotypic
genus Oxydothis, and it is phylogenetically related to Vialaeaceae and Iodosphaeriaceae in
Xylariales. Hyde et al. (2017a) provided a molecular clock estimates to evaluate the ranking of taxa
of Sordariomycetes, and indicated that the status of Oxydothidaceae is supported with a divergence
of 114 MYA (stem age), which is in the recommended divergence range for families (50–130
MYA).
Ecological and economic significance of Oxydothidaceae
Oxydothis is one of the most common genera encountered in monocotyledons, such as palms
(Arecaceae), Pandanus (Pandanaceae) and bamboo (Gramineae) (Hyde 1993b, c, 1994a, Wang &
Hyde 1999, Wong & Hyde 2001, Fröhlich & Hyde 2000, Taylor & Hyde 2003, Shenoy et al. 2005,
Hidayat et al. 2006, Tibpromma et al. 2018) and are saprobes important in nutrient cycling. The
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taxa are also pathogens (Fröhlich & Hyde 1994) and endophytes (Hyde 1994a). It is important to
identify species and study their biotrophic nature, which may provide resources for biological
control and protect plants during planting.
Genus included in Oxydothidaceae
Oxydothis Penz. & Sacc., Malpighia 11(11-12): 505 (1897)
Index Fungorum number: IF3661; 79 morphological species (Species Fungorum 2020), 12
species with sequence data.
Type species – Oxydothis grisea Penz. & Sacc.
Notes – Oxydothis was described from Cibodas, Java, Indonesia by Penzig & Saccardo
(1897b) with three species O. grisea, O. nigricans and O. maculosa, and placed in
Amphisphaeriaceae (sensu Eriksson & Hawksworth 1991). The genus is characterized by two types
of ascomata, one is developed singly or in clusters, in darkened, ellipsoidal raised areas on the host
surface, and have distinctive eccentric ostioles, whereas another are those developed below a raised
sheet of host epidermis, and usually not darkened (Fröhlich & Hyde 2000). Hyde reviewed the taxa
of Oxydothis and emphasized that ascus and ascospore morphology in this genus is consistent and
reliable for distinguishing species, and also discussed the morphology of closely related genera,
Ceriospora, Frondispora, Lasiobertia and Leiosphaerella (Hyde 1993b, 1994a). However, the
familial classification of this genus was uncertain and it has been placed in Hyponectriaceae (Hyde
1993b) and Clypeosphaeriaceae (Kang et al. 1999b) based on morphology. Jeewon et al. (2003b)
analysed DNA sequence datas and mentioned that Oxydothis was closely related to Leiosphaerella
(Xylariales, genera incertae sedis), and Konta et al. (2016) placed the genus in Oxydothidaceae
(Xylariales). Konta et al. (2016) also suggested that Oxydothis species may be associated with
healthy plants as endophytes and become saprobes, based on their observations of appressoria. In
this study, we introduce a new species, Oxydothis phoenicis, from mangrove habitats.
Oxydothis phoenicis S.N. Zhang, K.D. Hyde & J.K. Liu, sp. nov.
Fig. 186
Index Fungorum number: IF555516; Facesoffungi number: FoF05092
Saprobic on rachis of Phoenix paludosa. Sexual morph: Ascomata 230–600 μm diam.,
solitary or clustered in groups, forming under slightly raised, blackened, ellipsoidal regions on the
host surface. Peridium 20–25 μm wide, composed of 2–3 layers of pale brown to hyaline cells with
textura prismatica. Paraphyses seems to dissolve during maturation. Asci 142–159 μm × 14–21 μm
( x = 148.9 × 17 μm, n = 20), 8-spored, cylindrical, mostly straight, pedicellate, unitunicate, with a
wedge-shaped, J+, subapical ring, 3.5–5 μm high, 3.5–4.5 μm diam. Ascospores 60–78 μm× 6–7.5
μm ( x = 72 × 6.7 μm, n = 30), fusiform, hyaline, obliquely 1–2-seriate, tapering gradually from a
constricted central or seldom upper central septum to short, pointed processes. Asexual morph:
Undetermined.
Material examined – THAILAND, Chang Wat Chanthaburi, Amphoe Khlung, Tambon Wan
Yao, on rachis of Phoenix paludosa (Arecaceae Roxb.), intertidal, 25 April 2017, S.N. Zhang,
SNT134 (MFLU 18-1074 holotype); ibid., (HKAS 97485, isotype); ex-type living culture
MFLUCC 18-0269 (= MFLUCC 18-0270).
GenBank numbers – Sequence data used in this study: ITS: MK088065 (MFLUCC 18-0269),
LSU: MK088061 (MFLUCC 18-0269), SSU: MK088063 (MFLUCC 18-0269), ITS: MK088066
(MFLUCC 18-0270), LSU: MK088062 (MFLUCC 18-0270), SSU: MK088064 (MFLUCC 180270). Sequence data not used in this study but submitted in GenBank: tef1: MK087667 (MFLUCC
18-0269), tef1: MK087668 (MFLUCC 18-0270).
Notes – Oxydothis phoenicis is similar to O. asiatica, O. mauritiae, O. uniseriata and O.
rattanica, but differs in ascospore and apical ring size (Fröhlich & Hyde 2000). Oxydothis
phoenicis also shares similar ascospores and ascal apical rings with O. calami, O licualae (and O.
rubella, but is distinct in ascomata morphology (Hyde 1993c). The phylogeny based on multi-gene
analysis also supports O. phoenicis as an independent lineage and indicates it is distinct from other
species with sequence data.
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Figure 186 – Oxydothis phoenicis (MFLU 18-1074, holotype). a Appearance of fruiting bodies on
host substrate. b Section of ascoma. c Peridium. d-g Asci. g Reaction of apical ring in Melzer’s
reagent. h-m Ascospores. n Germinating ascospores. o Appressoria. p Colonies on PDA. q Colonies
on MEA. Scale bars: a = 500 µm, b, d-g, n = 50 µm, c, h-m, o = 20 µm.
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Papulosaceae Winka & O.E. Erikss., Mycoscience 41 (2):102 (2000)
Index Fungorum number: IF82096; Facesoffungi number: FoF01284; 9 species.
Saprobic on wood in freshwater or marine habitats, and pathogenic on roots in terrestrial
habitats. Sexual morph: Ascomata perithecial, solitary, immersed or semi-immersed, black to dark
brown, globose to ellipsoidal, coriaceous, ostiolate. Necks cylindrical, periphysate. Peridium
composed of cells of textura angularis or brown-walled compressed cells. Paraphyses tapering,
simple or branched, septate. Asci 8-spored, unitunicate, cylindrical, short pedicellate, with a J-, or
J+, refractive, bipartite or discoid, apical ring. Ascospores uniseriate to biseriate, hyaline or brown,
ellipsoidal or fusiform, unicellular to 3-septate, smooth or verruculose, with or without cup-like,
bipolar appendages. Asexual morph: Undetermined (adapted from Réblová et al. 2013).
Type genus – Papulosa Kohlm. & Volkm.-Kohlm.
Notes – Winka & Eriksson (2000) introduced Papulosaceae and concluded that the exact
phylogenetic position of Papulosa could not be established due to the small number of SSU
sequences available for unitunicate pyrenomycetes at that time. Papulosa shared similarities with
Diaporthales, Ophiostomatales, Phyllachorales and Sordariales based on molecular data, thus the
genus was accommodated in the subclass Sordariomycetidae. Maharachchikumbura et al. (2015)
placed Papulosaceae as a sister clade to Coniochaetaceae and Cordanaceae. Abdel-Wahab et al.
(2011) analyzed LSU sequence data and found that Brunneosporella and Fluminicola clustered
with Papulosa amerospora and Cataractispora receptaculorum. A similar relationship was seen in
phylogenetic analyses carried out by Réblová et al. (2013) using LSU, SSU, and rpb2 sequence
data. Both Brunneosporella and Fluminicola appear in the same clade as that of Papulosa and
Pleurophragmium parvisporum and were named as Papulosaceae (Réblová et al. 2013). LSU and
SSU sequence analyses of Boonyuen et al. (2012), showed that Brunneosporella (as
Ascobrunneispora aquatica) clustered with Papulosa. Maharachchikumbura et al. (2015, 2016b)
placed Brunneosporella and Fluminicola in Papulosaceae. Platytrachelon which was introduced by
Réblová et al. (2013) has a close phylogenetic relationship with the Papulosaceae. Réblová et al.
(2013) included Platytrachelon in Sordariomycetes incertae sedis while Wijayawardene et al.
(2018a) listed it under Papulosaceae. However, the phylogenetic placement of Platytrachelon
should be verified with more taxon sampling. Khemmuk et al. (2016) introduced a novel genus,
Wongia to Papulosaceae based on phylogenetic analyses of combined ITS, LSU, rpb1 and tef1
sequence data. Hence, the family comprises four genera at present.
Ecological and economic significance of Papulosaceae
Most members in this family are found on submerged wood in freshwater or marine habitats
as saprobes (Ranghoo et al. 2001).
Genera included in Papulosaceae
Brunneosporella Ranghoo & K.D. Hyde, Mycol. Res. 105(5): 625 (2001)
Index Fungorum number: IF28492; 1 species with sequence data.
Type species – Brunneosporella aquatica Ranghoo & K.D. Hyde
Notes – Brunneosporella was introduced by Ranghoo et al. (2001) which comprises 1-septate
and concolorous ascospores and asci with a discoid apical ring. Brunneosporella aquatica is the
only species assigned to this genus.
Fluminicola S.W. Wong, K.D. Hyde & E.B.G. Jones, Fungal Divers. Res. Ser. 2: 190 (1999)
Index Fungorum number: IF28031; 4 morphological species (Species Fungorum 2020), 3
species with sequence data.
Type species – Fluminicola bipolaris S.W. Wong, K.D. Hyde & E.B.G. Jones
Notes – Wong et al. (1999) introduced Fluminicola for the freshwater species F. bipolaris.
The genus is characterized by cylindrical asci with a relatively massive, refractive apical ring and
fusiform and hyaline ascospores with irregular bifurcate or cup-like bipolar appendages (Wong et
al. 1999, Zhang et al. 2017a). Five species are accommodated in the genus, i.e. F. aquatica, F.
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bipolaris, F. coronata, F. saprophytica and F. thailandensis (Zhang et al. 2017a, Index Fungorum
2020). Fluminicola saprophytica and F. thailandensis are illustrated below; for descriptions see
Zhang et al. (2017a).
Papulosa Kohlm. & Volkm.-Kohlm., Syst. Ascom. 11(2): 96 (1993)
Index Fungorum number: IF26426; 1 species with sequence data
Type species – Papulosa amerospora Kohlm. & Volkm.-Kohlm.
Notes – Kohlmeyer & Volkmann-Kohlmeyer (1993) established the monotypic genus
Papulosa, but were unable to place this genus with certainty into any order or family.
Wongia Khemmuk, Geering & R.G. Shivas, IMA fungus 7(2): 247–252 (2016)
Index Fungorum number: IF817529; 3 species with sequence data.
Type species – Wongia garrettii (P. Wong & M.L. Dickinson) Khemmuk, Geering & R.G.
Shivas
Notes – This genus was introduced to accommodate two root infecting fungi from Australia
Magnaporthe garrettii and M. griffinii, which formed a well-supported separate clade in
Papulosaceae and clustered outside Magnaporthales (Khemmuket al. 2016). Wongia species have J, apical rings in the asci and 3-septate ascospores that have dark brown middle cells and pale brown
to subhyaline shorter distal cells (Khemmuket al. 2016).
Pararamichloridiaceae Crous, Persoonia 39: 357 (2017)
Index Fungorum number: IF823463; Facesoffungi number: FoF05314; 3 species.
Pathogenic on plant leaves. Sexual morph: Undetermined. Asexual morph: Mycelium
consisting of hyaline, smooth, septate, branched, hyphae. Conidiophores erect, solitary, straight
to flexuous, septate, branched at apex or not, subcylindrical, subhyaline to medium brown,
smooth. Conidiogenous cells terminal and intercalary, subcylindrical, subhyaline to medium
brown, smooth, polyblastic, denticulate. Conidia solitary, hyaline, smooth, aseptate, thin-walled,
clavate to ellipsoid (adapted from Crous et al. 2017a).
Type genus – Pararamichloridium Crous
Notes – Crous et al. (2017a) introduced Pararamichloridiaceae and accepted
Pararamichloridium and Woswasia. However, Woswasia, along with Xylochrysis and
Cyanoannulus, was placed in Diaporthomycetidae families incertae sedis by Zhang et al. (2017a)
based on morphology, phylogenetic analysis and molecular dating. In the multi-loci ITS, LSU, SSU
and rpb2 phylogenetic tree, Pararamichloridium livistonae (CBS 143166) and P. verrucosum (CBS
128.86) grouped together and formed a separate clade with 99% MP bootstrap support, 92% ML
bootstrap support and 100% Bayesian posterior probabilities (data not shown). Woswasia
atropurpurea (CBS 133167), Xylochrysis lucida (CBS 135996) and Cyanoannulus petersenii
(R044a and R044b) formed a separate branch which is distant from Pararamichloridium clade
(data not shown). In this study, we exclude Woswasia from Pararamichloridiaceae based on its
close phylogenetic affinity with Xylochrysis and Cyanoannulus in Woswasiaceae.
Ecological and economic significance of Pararamichloridiaceae
Pararamichloridium livistonae was collected from the leaves of Livistona australis in
Australia and it causes leaf spots of the host plant (Crous et al. 2017a).
Genus included in Pararamichloridiaceae
Pararamichloridium Crous, Persoonia 39: 357 (2017)
Index Fungorum number: IF823369; 3 species with sequence data.
Type species – Pararamichloridium livistonae Crous
Notes – Currently, three species, Pararamichloridium caricicola, P. livistonae and P.
verrucosum are accepted in this genus. The genus is characterised by branched or unbranched,
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medium brown, septate conidiophores, medium brown, denticulate conidiogenous cells and
solitary, hyaline, smooth, granular, aseptate, and clavate conidia (Crous et al. 2017a).
Figure 187 – Fluminicola saprophytica (Material examined – THAILAND, Prachuap Khiri Khan,
on submerged bamboo in a small river, 30 July 2015, W. Dong 04A, MFLU 15-2694, holotype; extype culture MFLUCC 15-0976) and Fluminicola thailandensis (Material examined – THAILAND,
Prachuap Khiri Khan, on submerged wood in a small river, 30 July 2015, W. Dong 22A (MFLU
15-2704, holotype). a, b Appearance of necks on host. c, d Vertical sections of ascomata. e, f
Peridium. g, h Asci. i, j Apex of asci. k, l Ascospores. Scale bars: c, h = 100 µm, d = 50 µm, e, f =
10 µm, g, k, l = 20 µm, i, j = 5 µm.
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Figure 188 – Pararamichloridium livistonae (Redrawn from Crous et al. 2017a).
Parasympodiellaceae Hern.-Restr., Gené, Guarro & Crous, Stud. Mycol. 86: 87 (2017)
Index Fungorum number: IF820298; Facesoffungi number: FoF05187; 9 species.
Saprobic on leaves and twigs. Sexual morph: Undetermined. Asexual morph: Colonies
effuse, cottony, partly superficial, greyish. Conidiophores mononematous, erect, conspicuous and
properly differentiated from vegetative hyphae, flexuous, smooth, regularly septate, branched,
brown. Conidiogenous cells smooth, pale brown or hyaline, holoblastic, multiplying sympodially,
undergoing basipetal secession to produce conidia. Conidia thallic-arthric, aseptate or septate,
hyaline, produced in unbranched pseudo-chains. Synasexual morph: often stylaspergillus-like
(adapted from Hernández-Restrepo et al. 2017).
Type genus – Parasympodiella Ponnappa
Notes – Parasympodiellaceae, as established by Hernández-Restrepo et al. (2017), is
accommodated in Parasympodiellales and it includes the monotypic genus Parasympodiella
(Hernández-Restrepo et al. 2017). Species of this family most commonly grow as saprobes on plant
litter, conifers and dicotyledons (Subramanian & Vittal 1973, Crous et al. 1995, Cheewangkoon et
al. 2009, Seifert et al. 2011, Hernández-Restrepo et al. 2017). Parasympodiellales along with
Parasympodiellaceae have been introduced to represent the clade which includes four of the nine
Parasympodiella species, namely, P. lauri, P. elongata, P. eucalypti and P. laxa (HernándezRestrepo et al. 2017).
Ecological and economic significance of Parasympodiellaceae
Saprotrophic fungi are prominently important decomposers of plant litter through secretion of
lignocellulolytic enzymes. In so doing, they regulate cycling of nutrients as well as carbon in the
terrestrial ecosystem (Baldrian & Valášková 2008, Crowther et al. 2012). Additionally,
Parasympodiella species may cause commercial damage to economically important plants since
they often colonize coniferous and dicotyledonous plants (Crous et al. 1995, Cheewangkoon et al.
2009, Seifert et al. 2011).
Genus included in Parasympodiellaceae
Parasympodiella Ponnappa, Trans. Br. mycol. Soc. 64(2): 344 (1975).
Index Fungorum number: IF820298; 9 morphological species (Species Fungorum 2020), 4
species with sequence data.
Type species – Parasympodiella laxa (Subram. & Vittal) Ponnappa
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Figure 189 – Parasympodiella laxa (Material examined – USA, shoots of Juniperus virginiana,
April 1968, C.S Hodges, IMI 134194; o, p redrawn from Cheewangkoon et al. 2009). a-c
Herbarium material. d, e Appearance of colonies on host substrate. f Mount of colonies. g
Conidiophore. h-j Conidiophores, conidiogenous cells and conidia. k Chain of conidia. l-p Conidia.
Scale bars: c = 1 mm, f-i, k = 100 µm, j = 50 µm, l, n = 20 µm, m = 30 µm, o, p = 10 µm.
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Notes – Parasympodiella was introduced by Ponnappa (1975), with P. laxa as the type
species. The latter species was earlier placed in Sympodiella (Ponnappa 1975). However, whereas
Sympodiella species have small conidiophores, up to 280 μm along with terminal or subterminal
conidiogenous cells which give rise to conidial chains reaching up to six conidia, Parasympodiella
species have larger conidiophores, reaching up to 700 μm, together with conidiogenous cells
residing along the conidiophore stipe at irregular distances and conidia being produced in ever
extending loose chains (Kendrick 1958). Parasympodiella includes distinct and solitary
conidiophores with conidiogenous cells which extend sympodially and produce catenate,
cylindrical, thallic conidia when the fertile hyaline branches disarticulate, characters which
distinguish this genus from similar genera such as Polyscytalum (Riess 1853), Sympodiella
(Kendrick 1958), Bahusakala (Subramanian 1958) and Neoscytalidium (Crous et al. 2006a).
Parasympodiella laxa is illustrated in this entry.
Phaeoappendicosporaceae Crous & M.J. Wingf., Fungal Systematics and Evolution. 3: 96 (2019).
Index Fungorum number: IF829458; Facesoffungi number: FoF06297; 2 species.
Saprobic on dead woody bark. Sexual morph: Pseudostroma immersed, becoming erumpent;
ectostroma pale brown to grey, containing periphyses; ostioles cylindrical. Perithecia globose to
lenticular, dark brown, wall of cells of textura angularis. Paraphyses septate, unbranched, hyphalike. Asci 8-spored, ellipsoid to fusoid, without a refractive canal at apex. Ascospores brown,
ellipsoid-fusoid, 1-euseptate, with gelatinous appendage at each truncate end. Asexual morph:
Coelomycetous. Conidiomata pycnidial, multilocular, forming a long neck. Paraphyses hyaline,
cylindrical, septate, unbranched, hypha-like. Conidiophores subcylindrical, hyaline to pale brown,
septate, unbranched. Conidiogenous cells cylindrical, hyaline to pale brown, proliferating
percurrently at apex. Conidia ellipsoid to oblong, straight to slightly curved, thick-walled,
transversely euseptate with oblique septa.
Type genus – Phaeoappendicospora Senan., Q.R. Li & K.D. Hyde
Notes – Phaeoappendicosporaceae was introduced by Crous et al. (2019b) based on
Phaeoappendicospora as the type genus and Neophaeoappendicospora as the second genus. Both
genera have similar characters, but Neophaeoappendicospora is distinct from
Phaeoappendicospora by its larger ascospores and presence of asexual morph.
Ecological and economic significance of Phaeoappendicosporaceae
All members in this family are saprobes. Neophaeoappendicospora leucaenae was isolated
from Leucaena leucocephala, a commercially and environmental important Fabaceae used as
fodder, charcoal, organic manure and in the paper industry. However, members in both genera are
involved in nutrient recycling through litter decomposition.
Genera included in Phaeoappendicosporaceae
Neophaeoappendicospora Crous & M.J. Wingf., Fungal Systematics and Evolution 3: 96 (2019)
Index Fungorum number: IF829323; 1 species with sequence data.
Type species – Neophaeoappendicospora leucaenae Crous & M.J. Wingf.
Notes – Neophaeoappendicospora was isolated from Leucaena leucocephala from France,
which is a source of quality animal feed, firewood or used in charcoal production, green manure
and biomass, food for humans and pulpwood for paper industry.
Phaeoappendicospora Senan., Q.R. Li & K.D. Hyde, Stud. Mycol. 86: 217–296 (2017)
Index Fungorum number: IF821572; 1 species with sequence data.
Type species – Phaeoappendicospora thailandensis Q.R. Li, Senan. & K.D. Hyde
Notes – Phaeoappendicospora was introduced in Diaporthales genera incertae sedis. The
genus has been introduced based on Phaeoappendicospora thailandensis, obtained from a dead
branch of Quercus sp. The species resembles Hapalocystis berkeleyi (Sydowiellaceae). It did not,
however, cluster with any Hapalocystis taxa nor with any family in the Diaporthales, which
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resulted to its being classified as genera incertae sedis (Senanayake et al. 2017b). Crous et al.
(2019b) introduced Phaeoappendicosporaceae for this genus with Neophaeoappendicospora. The
genus is characterised by brown and small ascospores. Phaeoappendicospora thailandensis is
illustrated in this entry.
Figure 190 – Neophaeoappendicospora leucaenae (CPC 27240 redrawn from Crous et al. 2019b).
a Asci. b Paraphyses. c Ascospores. d Conidiogeneous cells giving rise to conidia. e Conidia. Scale
bars: 10 µm.
Phaeochoraceae K.D. Hyde, P.F. Cannon & M.E. Barr. Syst. Ascom. 15(1–2):118 (1997)
Index Fungorum number: IF81916; Facesoffungi number: FoF01910; 17 species.
Saprobic or biotrophic on palms leaves. Sexual morph: Stromata as blackened regions
scattered on the host, usually raising above the substrate surface, rarely inconspicuous, one or
multi-loculate. Ascomata ellipsoidal to subglobose, dark brown, clustered and fusing to form multiostiolate cavity or solitary with one ostiole. Ostiole conspicuous or inconspicuous. Peridium several
layered, composed of flattened, brown to dark brown-walled cells, thinner at the base. Paraphyses
wide, cylindrical hypha-like, septate, thin-walled, evanescent early. Asci 6–8-spored, unitunicate,
saccate or fusiform, with long or short pedicellate, very thin-walled, usually without apical
structures, deliquescing early. Ascospores uniseriate to biseriate, various shades of brown,
ellipsoidal, fusiform or cylindrical, aseptate, thick-walled, sometimes with one flattened face,
sometimes delicately striated, with or without appendages. Asexual morph: Coelomycetous.
Conidiogenous cells narrowly conical, usually percurrently proliferating, but occasionally
sympodially proliferating, with periclinal thickening, without a collarette. Conidia aseptate,
narrowly fusiform to bacillary, hyaline, thin-walled (adapted from Cannon 1992, Hyde et al. 1997a,
Taylor & Hyde 2003, Maharachchikumbura et al. 2016b).
Type genus – Phaeochora Höhn.
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Figure 191 – Phaeoappendicospora thailandensis (Material examined – THAILAND, Chiang Rai,
near Khun Korn water fall, on branch of Quercus sp. (Fagaceae), 25 December 2012, Q.R. Li,
TL19, MFLU 12–2131, holotype). a Appearance of fungus on host substrate. b Horizontal cross
section of stroma. c Vertical cross section of stroma. d, e Asci. f-i Ascospores. Scale bars: a = 1
mm, b = 500 μm, c = 100 μm, d-i = 10 μm.
Notes – Phaeochoraceae was introduced by Hyde et al. (1997a) to accommodate the genera
Cocoicola, Phaeochora and Serenomyces. Phaeochoropsis was subsequently included to
Phaeochoraceae. Species of Phaeochoraceae are biotrophic and saprobic on palms (Arecaceae), and
characterized by a well-developed stroma and usually form tar-spots. Phaeochoraceae was
previously assigned to Phyllachorales mainly because of the stromatic characteristics. Mardones et
al. (2017) showed the phylogenetic placement of Phaeochoraceae in Phyllachorales.
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Ecological and economic significance of Phaeochoraceae
Phaeochoraceae accommodates species that visible as tar spot or lesions on plant leaves,
which may cause diseases of host plant. Species of Cocoicola are pathogens associated with petiole
blight. Serenomyces invades palms and causes rachis blight or petiole blight (Elliott & Des Jardin
2014). The diseased palms may be significantly affected in terms of yield and quality including
incidence of fruit rots.
Genera included in Phaeochoraceae
Cocoicola K.D. Hyde, Nova Hedwigia 60(3–4): 600 (1995a)
Index Fungorum number: IF27575; 5 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Cocoicola cylindrospora (C. Booth & D.E. Shaw) K.D. Hyde
Notes – Cocoicola was introduced by Hyde (1995a) to accommodate C. cylindrospora (=
Anthostomella cylindrospora) and C. fusispora (= A. fusispora). Currently, Cocoicola contains five
species, which were found from palms. They are characterized by strongly flattened ascomata,
clustered around a central ostiole, or with a series of separate ostioles and surrounded by a stroma,
visible as ostiolar dots and present pale brown blister-like regions on the host surface. The
ascospores are cylindrical to fusiform, yellow to olivaceous, aseptate, with longitudinal striations.
Cocoicola is similar to Serenomyces by striated ascospores. However, they are distinct as the latter
has exclusively uniloculate ascomata with elongated necks. Serenomyces californica was
synonymized as Cocoicola californica because of its fusiform ascospores and multi-ostiolate
ascomata lacking necks (Hyde 1995a).
Phaeochora Höhn., Sber. Akad. Wiss. Wien, Math.-naturw. Kl., Abt. I, 118: 1513 (1909)
Index Fungorum number: IF3898; 4 morphological species (Species Fungorum 2020).
Type species – Phaeochora chamaeropis (Cooke) Höhn.
Notes – Phaeochora was introduced by Höhnel (1909a) to accommodate Auerswaldia
chamaeropis (= Dothidea chamaeropsis). Höhnel (1909a) treated A. chamaeropis as the type, and
also considered A. densa belongs to Phaeochora. However, Shear (1909) transferred A. densa to
Sphaerodothis found an earlier name S. steinheilii for the type species of Phaeochora, and
synonymized it as P. steinheilii. The two genera Phaeochora and Sphaerodothis were superficially
similar and were treated as synonyms by von Arx & Müller (1954) and Joly (1961). Species in
Phaeochora cause blackened regions on the host surface with stromata and ellipsoidal, fusiform or
cylindrical ascospores, but are different in ascomata and ascospores structure; while Sphaerodothis
has cylindrical asci (Hyde & Cannon 1999). Phaeochora steinheilii is characterized by ascospores
having one flattened face, with two, ±pulvinate extrusions of the epispore near the apices of the
flattened surface (Cannon 1992, Hyde et al. 1997a).
Phaeochoropsis K.D. Hyde & P.F. Cannon, Mycol. Pap.175: 30 (1999)
Index Fungorum number: IF27928; 4 morphological species (Species Fungorum 2020).
Type species – Phaeochoropsis neowashingtoniae (Shear) K.D. Hyde & P.F. Cannon
Notes – Phaeochoropsis was introduced by Hyde & Cannon (1999) based on P.
diplothemiifolii, P. mucosa, P. neowashingtoniae and P. palmicola. Species of Phaeochoropsis
occur on leaves and rachides of the host plant with lesions which become yellow and contain a
number of stromata.
Serenomyces Petr., Sydowia 6(1–4): 296 (1952b)
Index Fungorum number: IF5008; 4 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Serenomyces shearii Petr.
Notes – Serenomyces (Petrak 1952b) was described based on S. shearii from a dead palm leaf
of Serenoa serrulata collected in Florida and other Serenomyces species occur only in association
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with palms (Arecaceae). The genus causes of leaf disease and rachis/petiole blight (Elliott & Des
Jardin 2014). Serenomyces was placed in Ceratostomataceae by Petrak (1952b) because of
evanescent asci and elongated neck, in Melanosporaceae by von Arx & Müller (1954), and
tentatively assigned in Phyllachoraceae by Barr et al. (1989). Hyde et al. (1997a) included
Serenomyces in Phaeochoraceae, because of immersed ascomata in pseudostromatic tissues and
paraphyses. Barr et al. (1989) revised Serenomyces and accepted four species, and later included S.
virginiae (Barr et al. 1997). Serenomyces californica was transferred to Cocoicola californica
(Hyde & Cannon 1999). Serenomyces is presumed as biotrophic and it is difficult to culture strains
from palm tissues. Elliott & Des Jardin (2014) obtained cultures and provided molecular data for
three Serenomyces species, which were isolated from palm leaves. Serenomyces is characterized by
immersed ascomata or stromata with elongated neck, evanescent asci and ovoid or fusiform, pale
brown and aseptate ascospores (Hyde & Cannon 1999).
Figure 192 – Phaeochora steinheilii (redrawn from Cannon 1992, Hyde & Cannon 1999). a
Stromata visible as blackened spot on leaf surfaces. b, c Asci without apical structures, evanescent.
d, e Ascospores, golden to mid-brown, with two ± globose hyaline appendages near the apices of
the flattened surface. f Conidiogenous cells developing conidia. g Conidia. Scale bars: a = 1 mm, bc = 10 µm, d−g = 5 µm.
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Phaeochorellaceae Guterres, Galvão-Elias & Dianese, Mycologia 111: 660–675 (2019)
MycoBank number: IF825364; Facesoffungi number: FoF06856; 2 species.
Biotrophic on living leaves or saprotrophs on decaying plant materials. Sexual morph:
Pseudostromata dark brown to black, subepidermal on living leaves, also present in saprobic state.
Ascomata perithecial, globose to subglobose or pyriform, wall heavily pigmented, ostiolate. Ostiole
comprising many thin, filiform periphyses. Paraphyses filiform. Asci 8-spored, unitunicate, thinwalled, cylindrical to subclavate or ellipsoidal. Ascospores ellipsoidal to cylindrical, aseptate to 1septate, dark brown. Asexual morph: Undetermined (adapted from Guterres et al. 2019).
Type genus – Phaeochorella (Henn.) Theiss. & Syd.
Notes – Phaeochorellaceae has been introduced based on two genera, namely, Phaeochorella
and Phaeoappendicospora (Guterres et al. 2019). The former genus was initially accommodated in
Phyllachoraceae (Phyllachorales) only based on morphology since no species had molecular data.
The acquisition of DNA sequence data for Phaeochorella parinarii was instrumental since a multigene phylogeny confirmed the placement of the genus in Diaporthales, forming a sister clade with
Phaeoappendicospora thailandensis. Since the two genera formed a distinct clade, phylogenetically
distant from other sister clades, the new family Phaeochorellaceae was established (Guterres et al.
2019). Crous et al. (2019b) later introduced family Phaeoappendicosporaceae to accommodate
Phaeoappendicospora with a new genus Neophaeoappendicospora.
Ecological and economic significance of Phaeochorellaceae
Species of Phaeochorella are parasitic mostly on leaves of plants belonging to the
Chrysobalanaceae and Fabaceae families (Guterres et al. 2019).
Genus included in Phaeochorellaceae
Phaeochorella (Henn.) Theiss. & Syd., Ann Mycol 13:405. 1915.
Index Fungorum number: IF3899; 2 morphological species (Guterres et al. 2019), 1 species
with sequence data.
Type species – Phaeochorella parinarii (Henn.) Theiss. & Syd.
Notes – Phaeochorella, typified by P. parinarii, was initially accommodated in several
families, viz. Phacidiaceae, Phyllachoraceae and Polystigmataceae (Theissen & Sydow 1915,
Müller & von Arx 1973). It was then maintained in Phyllachoraceae, Phyllachorales (Cannon 1991)
until now, where it has finally been transferred to a new family, Phaeochorellaceae (Guterres et al.
2019). The type species is a synonym of the basionym, Cocconia parinarii, reported on the leaves
of Parinari curatellifolia (Hennings 1901) and its asexual morph was initially recognised as
Phomachorella parinarii, reported from Pa. capensis (Petrak 1947c). However, this was reported
as a misidentification since it was observed that P. parinarii produces both micro- and macroconidia while no macroconidia were seen in “Phomachorella parinarii” (Swart 1965). Conidia of
“Phomachorella parinarii” also originated from the basal layer of stroma and were surrounded by
slimy material, while conidia produced by P. parinarii were produced from all over the surface of
the stroma cavity and formed a powdery conidial mass. It was concluded that the conidia observed
by Petrak were from another fungus parasitizing the stroma of P. parinarii (Swart 1965).
Phaeochorella originally comprised five species, namely, P. parinarii, P. zonata, P.
artocarpi, P. ciliata, and P. machaerii which originated from tropical regions (Theissen & Sydow
1915, Petrak 1947c, Ramakrishnan & Ramakrishnan 1948, Batista & Peres 1960, Batista & Bezerra
1961). However, at present the genus comprises only P. parinarii and P. zonata (Guterres et al.
2019). Phaeochorella machaerii is now considered as synonym of Phyllachora puncta, while P.
ciliata and P. artocarpi comprise bitunicate asci and ascospores without any equatorial hyaline
band (Guterres et al. 2019). Auerswaldia clypeata was also placed in Phaeochorella and
synonymised to Phaeochorella clypeata, but it was subsequently excluded from the genus based on
ascospore morphology (Theissen & Sydow 1915, Petrak 1947c). Similarly, Phaeochorella
sphaerospora was not accepted as a species belonging to Phaeochorella but recognised as
synonym of Phyllachora conica since ascospore colour was considered insufficient to differentiate
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between the two species (Cannon 1991). The holotype of P. sphaerospora, was however, not
examined since it was apparently lost, and thus, only the colour of the ascospores was deemed as
insufficient character to differentiate between the two species (Cannon 1991).
Figure 193 – Phaeochorella parinarii (epitype UB Mycol. Coll. 23270, redrawn from Guterres et
al. 2019). a Pseudostromata on leaf of Parinari obtusifolia. b Immersed perithecial ascoma with a
periphysate ostiole. c Peridium comprising cells of textura prismatica. d Septate paraphyses. e
Ascus containing mature ascospores with typical subhyaline equatorial band. f, g Immature and
mature ascospores. Scale bars: a = 2 cm, b = 100 µm, c, d = 20 µm, e-g = 10 µm
Phlogicylindriaceae Senan. & K.D. Hyde, Fungal Divers. 73: 35 (2015)
Index Fungorum number: IF551190; Facesoffungi number: FoF00681; 10 species.
Saprobic on leaves, twigs and branches of dicotyledons especially on Eucalyptus
(Myrtaceae). Sexual morph: Pseudostroma solitary, black, scattered, immersed. Ascomata forming
under pseudostroma, solitary, brown to black, scattered, globose, coriaceous, ostiolate, papillate.
Papilla short, narrow, surrounding thickened. Peridium comprising outer, brown thick-walled cells
of textura angularis and inner, hyaline, thin-walled cells of textura angularis. Asci 8-spored,
unitunicate, cylindrical, short pedicellate, rounded at apex, with J+, discoid, subapical ring.
Ascospores overlapping uniseriate, hyaline, fusiform, rounded at ends, 1-septate, slightly
constricted at septa. Asexual morph: Coelomycetous. Conidiomata appearing as slimy, erect tufts
of hyaline conidial masses, eustromatic, unilocular or multilocular, sporodochial, synnematous,
indeterminate, turning brown with age. Conidiophores brown, smooth, branched. Conidiogenous
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cells annellidic, smooth, hyaline becoming light brown with maturity, subcylindrical, ampulliform
with elongated necks, often having percurrent proliferations. Conidia cylindrical to subcylindrical,
fusoid, hyaline, straight, smooth, rounded apex, rounded or subobtuse apex, truncate base, aseptate
to 1-septate, guttulate (adapted from Senanayake et al. 2015).
Type genus – Phlogicylindrium Crous, Summerb. & Summerell
Notes – Senanayake et al. (2015) introduced Phlogicylindriaceae to accommodate the genera
Phlogicylindrium and Ciferriascosea. The asexual genus produces slimy erect flame-like tufts of
hyaline conidial masses, and cylindrical, unicellular to 1-septate, hyaline conidia (Summerell et al.
2006). The sexual morph of Phlogicylindrium has not been reported. Ciferriascosea is a sexual
genus with no reported asexual morph, and was included in Phlogicylindriaceae due to its
phylogenetic affinity to Phlogicylindrium (Senanayake et al. 2015). The sexual genus Idriellomyces
was introduced in Phlogicylindriaceae by Crous et al. (2018c) based on its distinct morphology and
molecular data.
Ecological and economic significance of Phlogicylindriaceae
Phlogicylindrium eucalypti and P. uniforme were reported occurring in association with a
Mycosphaerella spp. causing lesions on living leaves of Eucalyptus sp. (Summerell et al. 2006,
Crous et al. 2011). Therefore, they possibly play a role as secondary invaders (Crous et al. 2011).
Phlogicylindrium tereticornis was found on leaves of Eucalyptus tereticornis infected with circular
to angular, brown leaf spots and red-purple margins (Crous et al. 2017a). All Phlogicylindrium
species have so far been reported on leaves from various Eucalyptus species in Australia.
Ciferriascosea species were isolated as saprobes on Spartium junceum from Italy (Senanayake et
al. 2015). All Phlogicylindrium species and Idriellomyces eucalypti have been reported on leaves
from various Eucalyptus species in Australia. Chitonospora and Ciferriascosea species have been
reported as saprobes from Africa and Europe.
Genera included in Phlogicylindriaceae
Ciferriascosea Senan., Bhat, Camporesi & K.D. Hyde, Fungal Divers. 73: 35 (2015)
Index Fungorum number: IF551191; 2 species with sequence data.
Type species – Ciferriascosea rectamurum Senan., Bhat, Camporesi & K.D. Hyde
Notes – Ciferriascosea was introduced to accommodate two sexual morph species
Ciferriascosea fluctamurum and C. rectamurum occurring on Sparticum junceum collected, from
Italy (Senanayake et al. 2015). They are characterized by having immersed, black pseudostromata,
ostiolate and papillate ascomata that produce asci with an indistinct J+, apical ring, and hyaline, 1septate, fusiform ascospores with narrowly rounded ends (Senanayake et al. 2015). Both species of
Ciferriascosea were introduced based on LSU and ITS sequence data.
Idriellomyces Crous, Persoonia 40: 369 (2018)
Index Fungorum number: IF825426; 1 species with sequence data.
Type species – Idriellomyces eucalypti Crous
Notes – Idriellomyces introduced by Crous et al. (2018d) is characterized by branched
conidiophores arranged as thick, erect synnemata, subcylindrical conidiogenous cells tapering
towards the apex to a rachis and fusoid, hyaline, aseptate conidia. It is similar to Idriella in
Microdochiaceae but distinct in having pigmented conidiophores often in aggregated synnemata
and in lacking chlamydospores (Crous et al. 2018d). The type species was found on leaves of
Eucalyptus obliqua from Australia (Crous et al. 2018d).
Phlogicylindrium Crous, Summerb. & Summerell, Fungal Divers. 23: 340 (2006)
Index Fungorum number: IF510013; 7 species with sequence data.
Type species – Phlogicylindrium eucalypti Crous, Summerb. & Summerell
Notes – All species of Phlogicylindrium have been reported from Australian Eucalyptus
(Summerell et al. 2006, Crous et al. 2011, 2016a, 2019b, Farr & Rossman 2019). Phlogicylindrium
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dunnii differs from other Phlogicylindrium species by eustromatic, multilocular conidiomata with
subcylindrical, 1-septate macroconidia and cylindrical aseptate microconidia in the same conidioma
(Crous et al. 2019b). Here, we provide an illustration of P. uniforme (Fig. 194). All species of
Phlogicylindrium are introduced based on LSU or ITS sequence data, while rpb2 and tef1 sequence
data are available for P. dunnii and P. tereticornis (Crous et al. 2017a, 2019b).
Figure 194 – Phlogicylindrium uniforme (redrawn from Crous et al. 2011). a-c Conidiophores with
conidia. d Conidia. Scale bars: a-c = 10 µm.
Phomatosporaceae von Arx, Antonie van Leeuwenhoek 17: 271 (1951)
Index Fungorum number: IF552312; Facesoffungi number: FoF02486; 74 species.
Saprobic on submerged wood or decaying twigs in both terrestrial and aquatic habitats.
Sexual morph: Ascomata perithecial, solitary, seldom gregarious, immersed or gradually becoming
erumpent with age, subglobose to globose, light brown to dark brown or black, coriaceous,
occasionally developing beneath a small blackened clypeus, ostiolate, with a short papilla or rarely
with a rostrate or cylindrical, hyaline neck, central or eccentric. Ostiole periphysate. Peridium
composed of small, brown pseudoparenchymatous cells. Paraphyses filamentous, aseptate or
septate, tapering. Asci 8-spored, unitunicate, cylindrical or oblong-fusiform, thin-walled, shortpedicellate or sessile, J- ring in the ascal apex. Ascospores overlapping uniseriate to rarely biseriate,
hyaline, ellipsoidal to fusiform, aseptate or septate, not constricted at the septum, at times biguttulate, with striations or appendages or a sticky mucilaginous sheath (Senanayake et al. 2016).
Asexual morph: sporothrix-like (adapted from Rappaz 1992, Fournier & Lechat 2010, Senanayake
et al. 2016).
Type genus – Phomatospora Sacc.
Notes – After von Arx (1951) invalidly established Phomatosporaceae, Senanayake et al.
(2016) formally introduced the family to accommodate Phomatospora, Lanspora and Tenuimurus.
Three genera, based on LSU, SSU and ITS phylogeny, formed a clade distantly related to other
existing orders in the Diaporthomycetidae, for which, the new order Phomatosporales, was
established (Senanayake et al. 2016).
Ecological and economic significance of Phomatosporaceae
Saprobic fungi secrete enzymes to degrade celluloses and lignocelluloses, thereby, giving rise
to soft-rot cavities in woody tissues, which lead to decay of dead plant materials both in the
terrestrial and freshwater ecosystems (Zare-Maivan & Shearer 1988a, b, Shearer 1993).
Genera included in Phomatosporaceae
Lanspora K.D. Hyde & E.B.G. Jones, Can. J. Bot. 64(8): 1581 (1986)
Index Fungorum number: IF25051; 1 species with sequence data.
Type species – Lanspora coronata K.D. Hyde & E.B.G. Jones
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Notes – The monotypic marine genus Lanspora was introduced by Hyde & Jones (1986)
based on Lanspora coronata, isolated from driftwood collected on rocky coasts. The species differs
from species of other marine genera in having ascospores with crown-like appendages on getting
into contact with water (Hyde & Jones 1986). The asexual morph of the type species is
undetermined.
Phomatospora Sacc., Nuovo G. bot. ital. 7: 306 (1875).
Index Fungorum number: IF4015; 72 morphological species (Species Fungorum 2020), 4
species with sequence data.
Type species – Phomatospora berkeleyi Sacc.
Notes – Phomatospora, typified by P. berkeleyi, originates from terrestrial, freshwater and
marine habitats (Hyde 1993a, Raja & Shearer 2008). Ascospores of aquatic species have adaptive
features such as appendages or slimy sheaths in order to facilitate entrapment of and/or adhesion to
substrates (Hyde 1993a, Raja & Shearer 2008). Phomatospora berkeleyi and P. arenaria were
initially observed to produce a Sporothrix asexual morph in culture, similar to some taxa in the
Xylariales, which led to Phomatospora being placed in Xylariales (Rappaz 1992). Molecular
analyses conducted by Lumbsch & Huhndorf (2007), however, did not support this placement and
subsequently, the genus was placed in Sordariomycetes genera incertae sedis. Réblová et al.
(2016a) as well as Senanayake et al. (2016) revealed through phylogenetic analyses that
Phomatospora groups together with Lanspora, for which Phomatosporaceae was introduced.
Phomatospora biseriata is excluded from the phylogenetic analysis conducted in the present study
since it does not cluster with other species of Phomatospora. Further studies should confirm the
definite identity of the species introduced as ‘Phomatospora biseriata’. Phomatospora berkeleyi is
illustrated herein (Fig. 195).
Tenuimurus Senan., Camporesi & K.D. Hyde, Mycosphere 7(5): 637 (2016)
Index Fungorum number: IF552315; 1 species with sequence data.
Type species – Tenuimurus clematidis Senan., Camporesi & K.D. Hyde
Notes – Tenuimurus typified by T. clematidis, which was introduced by Senanayake et al.
(2016) to accommodate species primarily having thin, delicate peridium as compared to other
species in Phomatosporaceae. The species is saprobic on overwintered plants, for example, on
Clematis vitalba (Ranunculaceae) (Senanayake et al. 2016).
Phyllachoraceae Theiss. & P. Syd., Annls mycol. 13(3/4): 168 (1915)
Index Fungorum number: IF81156; Facesoffungi number: FoF01329; 1125 species.
Parasitic on living leaves or saprobic on dead wood submerged in water. Sexual morph: Leaf
spots on host black, abundant, scattered, raised, mostly rounded to oblong or elongated, sometimes
parallel with leaf venation, surrounded by light brown necrotic region. Ascomata flattened, globose
to subglobose, with thin walls, lying in leaf tissues or in a pseudostroma or stroma and maturing in
living leaves, ostiolate. Ostiolar canal conical, wide, lacking periphyses. Peridium clypeate,
thickest adjacent to ostiolar canal, composed of a deeply melanized, brown-black, amorphous layer
of host cuticle and epidermal cells, often merging with a lighter pigmented region of brownish,
distorted parenchyma cells infiltrated with fungal hyphae, beneath the clypeus. Lower and lateral
peridium composed of two layers; an outer region comprising several layers of dark brown,
flattened, thin-walled fungal cells, which internally fuse with several layers of hyaline, flattened,
thin-walled fungal cells. Lateral peridium fuses outwardly with an irregular, narrow region of
distorted host parenchyma infiltrated by fungal cells. The basal peridium merges outwardly with
either a narrow zone of infiltrated and distorted host parenchyma and occasionally lower epidermal
cells, or integrates directly with a lower clypeus, similar in construction to that basal peridium.
Paraphyses numerous, persistent, filiform, branched or unbranched, septate, slightly longer than
asci. Asci 8-spored, persistent, cylindrical to fusiform, short pedicellate, with walls uniform in
thickness, not especially thickened at apex, apical ring often present, rarely absent, J-, or J+.
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Ascospores multi-seriate, fusiform to narrowly oval, usually hyaline, aseptate or rarely septate,
often with a mucilaginous sheath, with or without an attenuated base, sometimes with pad like
appendages. Asexual morph: Coelomycetous, with conidia in locules in a stroma, spermatial or
disseminative. Conidiomata similar to ascomata, immersed, subcuticular, strongly raising the host
surface, occasionally pycnidial, occupying the region between the cuticle and epidermis, ostiolate.
Conidiogenous cells arising from the basal and lateral walls, cylindrical, phialidic, aseptate,
hyaline, forming conidia singly at the apex. Conidia hyaline, filiform, aseptate (adapted from
Maharachchikumbura et al. 2016b).
Type genus – Phyllachora Nitschke
Notes – Theissen & Sydow (1915) introduced Phyllachoraceae, previously placed in several
orders, including Dothideales (Horst 1990), Sphaeriales (Nannfeldt 1932, Miller 1951, 1954,
Muller & von Arx 1962, Wehmeyer 1975), Xylariales (Luttrell 1951, Barr 1990a), Glomerellales
(Chadefaud 1960, Locquin 1984), Phyllachorales (Barr 1976a, b, 1983), Polystigmatales (Eriksson
1982b, Hawksworth et al. 1983), and Diaporthales (Cannon 1988). Currently, it is placed in order
Phyllachorales (Maharachchikumbura et al. 2015b, 2016b, Santos et al. 2016, Dayarathne et al.
2017). Phyllachoraceae is characterised by ascohymenial development with paraphyses, thinwalled asci, which may have an apical ring, that does not stain blue in iodine (J-) and ascospores
that are often hyaline and 1-celled (Cannon 1991). Asexual morphs are coelomycetes, spermatial or
disseminative (Hawksworth et al. 1995). Munk (1957) and Barr (1990a) had a different concept of
the family, including genera with J+, apical rings, in the ascus. However, in a study based on
available molecular data and literature Maharachchikumbura et al. (2015, 2016b) listed 58 genera
in Phyllachoraceae. Mardones et al. (2017) proposed a new family Telimenaceae with Telimena as
the type genus, resulting in three families in Phyllachorales and removal of Telimena from
Phyllachoraceae. Dayarathne et al. (2017) excluded Polystigma from the Phyllachorales based on
analysis of combined LSU, SSU and ITS sequence data. Hence, at present family Phyllachoraceae
comprises 54 genera. However, sequence data are available only for a few members of the genera
Ascovaginospora, Coccodiella and Phyllachora because of the difficulties in obtaining cultures.
The family contains a large number of species which are probably host-specific. Most genera
are monotypic and the majority lack sequence data. The family therefore needs recollecting with
sequence data. Many genera are poorly known and therefore the notes below are brief.
Ecological and economic significance of Phyllachoraceae
Members of family Phyllachoraceae are mostly obligate biotrophs known to be minor
pathogens, and facilitate pathways for secondary infection by other severe pathogenic organisms
(Dayarathne et al. 2017). They produce distinct leaf spots on forage grasses of Poaceae especially
members of Acer (Karami et al. 2014); species of Rosaceae (Cannon 1996); on Leguminosae spp.
(Cannon 1991), on Duranta spp. in the tropics (Hanlin & Tortolero 1991); small scabby leaf spots
or "lixa-pequena" on coconut palms in Brazil (Subileau et al. 1993); on plants of family
Asclepiadaceae (Pearce et al. 1999); Prunus species (Habibi et al. 2015); and on Myrcia spp.
(Santos et al. 2016). They are limited in the area of action and when they kill leaf tissues, they
increase lignin content and decreasing host protein content, acceptability and digestibility (Burton
& Wills 1981). Furthermore, leaf spots caused by phytopathogenic fungi including tar spot
pathogens on leaves decrease the rate of photosynthesis and alter the physiological and biochemical
aspects thereby reducing crop production (Martinez et al. 2010, Karami et al. 2014).
Genera included in Phyllachoraceae
Ascovaginospora Fallah, Shearer & W.D. Chen, Mycologia 89(5): 813 (1997)
Index Fungorum number: IF27832; 1 species with sequence data.
Type species – Ascovaginospora stellipala Fallah, Shearer & W.D. Chen
Notes – This monotypic genus was introduced by Fallah et al. (1997) to accommodate A.
stellipala, which is characterized by 1-celled, hyaline, rhomboid ascospores, surrounded by a
tetraradiate sheath, the arms of which elongate in water (Fallah et al. 1997).
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Figure 195 – Phomatospora berkeleyi (Material examined – ENGLAND, Gloucestershire, dead
stalks of Solanum sp., M.J. Berkeley, K(M): 49573, isotype). a-d Herbarium material. e
Appearance of ascomata on host substrate. f squash mount of ascoma. g Peridium. i Ascus (redrawn
from Fallah & Shearer 1998). h, j-n Ascospores. Scale bars: e, f = 100 µm, g = 20 µm, h, j-n = 3
µm, i = 10 µm.
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Figure 196 – Ascovaginospora stellipala (Material examined – USA. Wisconsin, on Carex limosa,
28 May 1996, Payam M. Fallah, ILLS 52319, holotype). a Herbarium material. b Ascomata on
Carex limosa leaf. c, d Transverse section through ascoma. e Peridium. f-h Asci. i Paraphyses. j, k
Ascospores with tetraradiate sheath. i Ascospores stained with Indian ink. Scale bars: b = 200 μm,
c, d = 50, f-i = 20 μm, e, j-l = 5 μm.
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Ascovaginospora stellipala occurs on submerged dead stems and leaves of Carex limosa
collected from two sphagnum bogs in northern Wisconsin (Fallah et al. 1997). Based on
morphology and nuclear ribosomal DNA sequence data, this species was first placed in
Lasiosphaeriaceae, Sordariales by Fallah et al. (1997), in Phyllachorales by Maharachchikumbura
et al. (2015, 2016b) and this is supported by Dayarathne et al. (2017). In our observation,
tetraradiate sheaths of ascospores arms did not extend in the water as described by Fallah et al.
(1997), perhaps due to age.
Brobdingnagia K.D. Hyde & P.F. Cannon, Mycol. Pap. 175: 47 (1999)
Index Fungorum number: IF27915; 2 morphological species (Species Fungorum 2020).
Type species – Brobdingnagia nigeriensis (Sivan. & Okpala) K.D. Hyde & P.F. Cannon
Notes – Brobdingnagia is characterised by large cylindrical to clavate ascospores inside a
gelatinous, evanescent, unitunicate ascus (Hyde & Cannon 1999, Sivanesan & Shivas 2002). They
can be saprobes or pathogens, while cultures and sequences are not available for any of the four
species (Hyde & Cannon 1999, Sivanesan & Shivas 2002, Wijayawardene et al. 2018a).
Camarotella Theiss. & Syd., Annls mycol. 13(3/4): 370 (1915)
Index Fungorum number: IF776; 8 morphological species (Species Fungorum 2020), 2
species with sequence data.
Type species – Camarotella astrocaryae (Rehm) Theiss. & Syd.
Notes – Theissen & Sydow (1915) described Camarotella as having thick-walled asci and
hyaline, 4-celled ascospores. Petrak (1940) revised this genus, while Clements & Shear (1931) and
Wehmeyer (1975) treated Camarotella, but ignored the changes introduced by Petrak (1940). Hyde
& Cannon (1999), who examined parasitic fungi causing tar spots and similar diseases on palms,
revaluated and accepted the genus.
Coccodiella Hara, Bot. Mag., Tokyo 25: 224 (1910)
Index Fungorum number: IF1139; 21 morphological species (Species Fungorum 2020), 5
species with sequence data.
Type species – Coccodiella arundinariae Hara
Notes – Hara (1911) introduced Coccodiella for a plant parasitic species characterized by a
stroma originating in the mesophyll, which then proliferates through the lower epidermis, forming a
sessile hypostroma attached to the host tissue. Members of Coccodiella are host-specific and
tropical in distribution (Mardones et al. 2017).
Cyclodomus Höhn., Sber. Akad. Wiss. Wien, Math.-naturw. Kl., Abt. 1 118: 1527 [67 repr.] (1909)
Index Fungorum number: IF7859; 2 morphological species (Species Fungorum 2020).
Type species – Cyclodomus umbellulariae Höhn.
Notes – Cyclodomus is a coelomycetous genus, which currently comprises five species
(Wijayawardene et al. 2018a) and they are saprobes that can be found in terrestrial environments
(Hyde et al. 1996).
Deshpandiella Kamat & Ullasa, Bull. Torrey bot. Club 100: 41 (1973)
Index Fungorum number: IF1479; 1 morphological species.
Type species – Deshpandiella jambolana (T.S. Ramakr., Sriniv. & Sundaram) Kamat &
Ullasa
Notes – The monotypic genus Deshpandiella is typified by D. jambolana, which causes leaf
spots and was collected from Coorg Forests, Mysore State, India parasitizing leaves of Eugenia
cuminii (Syzygium jambolanum) (Ullasa & Rao 1973). The genus is characterized by astromatic,
perithecial ascomata with eccentric necks and bent ostioles (Ullasa & Rao 1973). Asci are 8-spored,
unitunicate, with a J-, apical ring and pedicels gelatinize at maturity. Ascospores are brown and
equally 2-celled (Ullasa & Rao 1973).
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Figure 197 – Cyclodomus aequatoriensis (Material examined – ECUADOR, Oriente, Napo
Province, Rio Cuyabeno, Cuyabeno rainforest, on living leaves of young palm (probably
Astrocayum) in rainforest, August 1993, K.D. Hyde, BRIP 23240, holotype). a Herbarium material.
b Leaf spots on palm leaf. c, d Transverse section through conidioma. e, f Conidiogenous cells
attached to conidia. g Conidia. Scale bars: b = 1000 μm, c = 200 μm, d = 100 μm, e, f = 10 μm, f =
5 μm.
Diachora Müll. Arg., Jb. wiss. Bot. 25: 623 (1893)
Index Fungorum number: IF1488; 4 morphological species (Species Fungorum 2020).
Type species – Diachora onobrychidis (DC.) Jul. Müll.
Notes – Diachora comprises obligately biotrophic plant parasites, forming restricted or
expanded pseudostromata in leaves or stems, filling the epidermal host cells with dark walled cells,
while the fungal structures of the inner host tissues are colourless (Muller 1986). The asexual
morph was in Diachorella which are coelomycetous saprobes found in terrestrial habitats
(Ciccarone 1963, Sutton 1967).
Diatractium Syd. & P. Syd., Annls mycol. 18(4/6): 183 (1921)
Index Fungorum number: IF1502; 2 morphological species (Species Fungorum 2020); 1
species with sequence data.
Type species – Diatractium cordiae (F. Stevens) Syd. & P. Syd.
Notes – Diatractium have ascospores that are isthmoid (narrowed over a significant length of
the central portion of the spore, Cannon 1989). Species of Diatractium are parasitic on leaves
791
(Cannon 1989). Palmateer et al. (2008) provided sequence data for D. cordianum (Ellis & Kelsey)
Syd.
Erikssonia Penz. & Sacc., Malpighia 11(11-12): 526 (1898)
Index Fungorum number: IF1884; 5 morphological species (Species Fungorum 2020).
Type species – Erikssonia pulchella Penz. & Sacc.
Notes – Erikssonia was named by Penzig & Saccardo as a member of Hysteriaceae
(Stevenson 1943). Erikssonia can be distinguished by erumpent to superficial stromata, with
uniloculate epistromata, which sometimes are attached to the host tissue by a partly immersed
hypostroma (Penzig & Saccardo 1897a, b).
Fremitomyces P.F. Cannon & H.C. Evans, Mycol. Res. 103(5): 585 (1999)
Index Fungorum number: IF28323; 2 morphological species (Species Fungorum 2020).
Type species – Fremitomyces punctatus P.F. Cannon & H.C. Evans
Notes – Fremitomyces was introduced by Cannon & Evans (1999) for two species, F.
punctatus and F. mahe from East Africa and the Seychelles occurring on Erythroxylaceae with
initially brightly coloured stromata (Cannon & Evans 1999, Lumbsch & Huhndorf 2007).
Geminispora Pat., Bull. Soc. mycol. Fr. 9: 151 (1893)
Index Fungorum number: IF2041; 2 morphological species (Species Fungorum 2020).
Type species – Geminispora mimosae Pat.
Notes – Geminispora includes two terrestrial saprobes, G. derridis and G. mimosae (Cannon
1991, Wijayawardene et al. 2018a).
Gibellina Pass., Revue mycol., Toulouse 8: 177 (1886)
Index Fungorum number: IF2064; 2 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Gibellina cerealis (Pass.) Pass.
Notes – This genus includes two species, G. cerealis and G. rehmiana that are pathogens.
Imazekia Tak. Kobay. & Y. Kawabe, Japanese Journal of Tropical Agriculture 36(3): 201 (1992)
Index Fungorum number: IF27271; 1 morphological species.
Type species – Imazekia ryukyuensis Tak. Kobay. & Y. Kawabe
Notes – Imazekia ryukyuensis causes black leaf spots of Stephania living leaves. There are
two species in this genus, which are distinct from the other known genera of Phyllachorales by
having apiospores (Kobayashi & Kawabe 1992).
Isothea Fr., Summa veg. Scand., Section Post. (Stockholm): 421 (1849)
Index Fungorum number: IF2516; 4 morphological species (Species Fungorum 2020).
Type species – Isothea rhytismoides (Bab.) Fr.
Notes – Isothea was erected for the single species Sphaeria rhytismoides (Cannon 1996).
Five taxa have been transferred to Isothea by different authors (Cannon 1996).
Lichenochora Hafellner, Nova Hedwigia 48(3-4): 358 (1989)
Index Fungorum number: IF25318; 45 morphological species (Species Fungorum 2020).
Type species – Lichenochora thallina (Cooke) Hafellner
Notes – Lichenochora (Phyllachorales) was introduced by Hafellner (1989) to include
different lichenicolous taxa characterized by typical perithecia and unitunicate asci. However, some
of the species with bitunicate asci have been referred to other genera such as Didymella and
Stigmidium. Hafellner (1989) included six species of Lichenochora; L. constrictella, L.
inconspicua, L. galligena, L. polycoccoides, L. thallina (currently L. obscuroides) and L. weillii, in
Physciaceae. Subsequently, the number of Lichenochora taxa has increased with the description of792
Figure 198 – Fremitomyces punctatus (Material examined – KENYA, from living leaf of
Erythroxylum fischeri, IMI 355095, holotype). a, b Herbarium material of Fremitomyces
paunctatus. c Fruiting bodies on Erythroxylum fischeri leaf surface. d Section through ascoma. e
Immature asci. f Mature asci. g Asci with paraphyses. Scale bars: e = 10 μm, f, g = 20 μm, d = 50
μm.
793
-new species or the inclusion of some new combinations in various studies (Triebel et al. 1991,
Roux & Triebel 1994, Navarro-rosinés et al. 1994, Navrotskaya et al. 1996, Aptroot et al. 1997,
Hafellner et al. 2008).
Lindauella Rehm, Hedwigia 39: 82 (1900)
Index Fungorum number: IF2874; 1 morphological species.
Type species – Lindauella pyrenocarpoidea Rehm
Notes – This genus included two taxa L. amylospora and L. pyrenocarpoidea. Currently L.
amylospora is referred to Flaminia amylospora. Hence, this genus is currently monotypic.
Linochora Höhn., Sber. Akad. Wiss. Wien, Math.-naturw. Kl., Abt. 1 119: 638 (1910)
Index Fungorum number: IF8781; 25 morphological species (Species Fungorum 2020).
Type species – Linochora leptospermi Höhn.
Notes – A spermatial state of Phyllachora has been placed in the form genus Linochora by
Höhnel (1910), but currently maintained as a Phyllachora species (Species Fungorum 2020). A
genus with many species (circa 40) mostly causing leaf spots of grasses.
Lohwagia Petr., Bot. Arch. 43: 205 (1942)
Index Fungorum number: IF2917; 3 morphological species (Species Fungorum 2020).
Type species – Lohwagia intermedia (Speg.) Petr.
Notes – Lohwagia was erected by Petrak (1942) to accommodate Phyllachora intermedia. Its
erumpent stromata cause galls and hypertrophy to the host plant.
Maculatifrondes K.D. Hyde, Mycol. Res. 100(12): 1509 (1996)
Index Fungorum number: IF27719; 1 morphological species.
Type species – Maculatifrondes aequatoriensis K.D. Hyde
Notes – Hyde et al. (1996) introduced Maculatifrondes to accommodate M. aequatoriensis,
obtained from living leaves of an unidentified palm in Cuyabeno, Ecuador. Maculatifrondes
aequatoriensis has early deliquescing asci and a cyclodomus-like asexual morph (Hyde et al.
1996).
Malthomyces K.D. Hyde & P.F. Cannon, Mycol. Pap. 175: 69 (1999)
Index Fungorum number: IF27923; 2 morphological species (Species Fungorum 2020).
Type species – Malthomyces calamigena (Berk. & Broome) K.D. Hyde & P.F. Cannon
Notes – Malthomyces is typified by M. calamigena and currently comprises two species
including M. coimbatoricus (Species Fungorum 2020). Both these species cause tar spots on palms
(Hyde & Cannon 1999) and are saprobes found from India and Sri Lanka. The asexual morph of
Malthomyces is undetermined (Wijayawardene et al. 2018a).
Muelleromyces Kamat & Anahosur, Experientia 24: 849 (1968)
Index Fungorum number: IF3284; 1 morphological species.
Type species – Muelleromyces indicus Kamat & Anahosur
Notes – Muelleromyces is characterized by astromatic perithecia with a well-developed
clypeus, 8-spored, unitunicate, evanescent asci and with a thick apical canal, and brown, unequally
2-celled ascospores (Anahosur 1968). This genus comprises one species, which parasitizes leaves
of Eugenia jambolana. and is characterized by ascospores with a wide equatorial pallid band above
the septum on the large apical cell (Anahosur 1968; Kang et al. 1999a). A second species
Muelleromyces variisporus has been referred to Kamatella (Anahosur 1969).
Neoflageoletia J. Reid & C. Booth, Can. J. Bot.44: 450 (1966)
Index Fungorum number: IF3453; 1 morphological species.
Type species – Neoflageoletia bambusina (Syd.) J. Reid & C. Booth
794
Notes – Neoflageoletia is a monotypic genus that was introduced to accommodate
Flageoletia bambusina by Reid & Booth (1966).
Figure 199 – Malthomyces calamigena (Material examined – SRI LANKA, Peradeniya, on leaves
of Calamus rudentum Lour. (Arecaceae), January 1868, G.H.K. Thwaites (ex-herb. M.J. Berkeley),
K(M) 35077, holotype). a Leaf spots on Calamus rudentum leaf. b Stromata on host substrate. c
Transverse section through stroma. d Peridium. e Paraphyses-like hyphae. f, g Asci. h-j Immature
and mature ascospores. Scale bars: b = 500 μm, c = 100, e-g = 50 μm, d, h-j = 10 μm.
Neophyllachora Dayar. & K.D. Hyde, Mycosphere 8(10): 1613 (2017)
Index Fungorum number: IF553633; 4 species with sequence data.
Type species – Neophyllachora myrciae (Lév.) Dayar. & K.D. Hyde
Notes – Members of Neophyllachora have subepidermal, intra-epidermal stromata without a
deep invasion of the mesophyl, and clavate asci. Three species appear to specifically infect Myrcia
795
species, one on Myrciaria species, and one on different Psidium species (Dayarathne et al. 2017).
Neophyllachora myrciae has characteristically lunate ascospores, and two types of conidia
(ellipsoidal and falcate) formed in separate conidiomata, but both with phialidic conidiogenesis.
Neophyllachora trucatisporum, on Myrciaria camapuanensis has sublunate ascospores with both
ends showing characteristic wall thickenings; N. cerradensis on M. torta shows elliptical
ascospores with predominantly longer dimensions when compared to N. subcircinans, which is
found only on Psidium species; and P. myrciariae has ascospores similar in form to those of P.
cerradensis, but also forms fusoid-clavate phialidic conidia (Santos et al. 2016, Dayarathne et al.
2017).
Ophiodothis Sacc., Sylloge Fungorum 2: 652 (1883)
Index Fungorum number: IF3601; 8 morphological species (Species Fungorum 2020).
Type species – Ophiodothis vorax (Berk. & M.A. Curtis) Sacc.
Notes – There are around 33 species described in this genus (Index Fungorum 2020),
however, only six species currently belong to Phyllachoraceae, while others are referred to various
families, such as the Clavicipitaceae. Sequence data are not available for any of Ophiodothis
species.
Ophiodothella Henn., Hedwigia 43: 258 (1904)
Index Fungorum number: IF3600; 31 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Ophiodothella atromaculans (Henn.) Höhn.
Notes – This genus comprises saprobic, terrestrial taxa which have perithecial ascomata that
are characterized by the formation of ostiolate perithecia immersed in host leaves in which a
blackened clypeus typically occurs around the ostiole and beneath the perithecium opposite the
ostiole (Hanlin et al. 1992). Their unitunicate asci comprise a J+, apical ring and contain hyaline,
scolecosporous ascospores (Hanlin et al. 2002). Most species of Ophiodothella are leaf parasites.
Orphnodactylis Malloch & A. Mallik, Can. J. Bot.76(7): 1267 (1998)
Index Fungorum number: IF539031; 2 morphological species (Species Fungorum 2020).
Type species – Orphnodactylis kalmiae Malloch & A. Mallik
Notes – Two species, Orphnodactylis kalmiae and O. wittrockii were described in this genus
based on morphology (Malloch & Mallik 1998). Orphnodactylis kalmiae causes twig blight disease
of Kalmia angustifolia (Ericaceae).
Oxodeora K.D. Hyde & P.F. Cannon, Mycol. Pap. 175: 75 (1999)
Index Fungorum number: IF27926; 1 morphological species.
Type species – Oxodeora petrakii (Cif.) K.D. Hyde & P.F. Cannon
Notes – This is a monotypic genus containing Oxodeora petrakii which was described from
palms (Hyde & Cannon 1999). Oxodeora shows substantial ascospore ornamentation and
comprises erumpent, linear stromata and light brown ascospores, covered with wide, longitudinal,
irregular ridges (Hyde & Cannon 1999).
Parberya C.A. Pearce & K.D. Hyde, Fungal Divers. 6: 90 (2001)
Index Fungorum number: IF28524; 2 morphological species (Species Fungorum 2020).
Type species – Parberya kosciuskoa C.A. Pearce & K.D. Hyde
Notes – Parberya was introduced to accommodate phyllachoraceous species forming tar
spots on leaves, which contain golden-brown to brown, spinose ornamented ascospores (Pearce &
Hyde 2001). The distinctly ornamented ascospores of Parberya appear to be unique in
Phyllachoraceae (Pearce & Hyde 2001). This genus includes two taxa, Parberya arxii and P.
kosciuskoa.
796
Petrakiella Syd., Annls mycol. 22(1/2): 230 (1924)
Index Fungorum number: IF3849; 1 morphological species.
Type species – Petrakiella insignis Syd.
Notes – This is a monotypic genus, containing the single species Petrakiella insignis, which
is characterized by the markedly developed stroma having a perithecial layer on its peripheral part
and phragmosporous ascospores (Kobayashi et al. 1939). Sydow (1924) stated that the centrum
structure of perithecia in P. insignis is quite similar to the members of Diaporthaceae.
Phycomelaina Kohlm., Phytopath. Z. 63: 350 (1968)
Index Fungorum number: IF4044; 1 morphological species.
Type species – Phycomelaina laminariae (Rostr.) Kohlm.
Notes – Phycomelaina is a monotypic genus which is typified by P. laminariae that was
originally described by Rostrup (1894-95) in Dothidella. Phycomelaina laminariae is a parasitic on
marine algal kelps; Laminaria saccacharina and other Laminaria species (Schatz 1983 and is not a
common species.
Phyllachora Nitschke, Fungi rhenani exsic., suppl., fasc. 6(nos 2001-2100): no. 2056 (1867)
Index Fungorum number: IF4049; 866 morphological species (Species Fungorum 2020), 23
species with sequence data.
Type species – Phyllachora graminis (Pers.) Fuckel
Notes – Phyllachora, which is typified by P. graminis and is the type genus of
Phyllachoraceae with around 1513 species currently recognized (Maharachchikumbura et al.
2016b, Dayarathne et al. 2017, Species Fungorum 2020). Most species are named based on their
host association and this may not reflect the actual number of species (Cannon 1988). Phyllachora
comprises species that grow immersed in a clypeate pseudostroma in leaf tissues, varying from a
subcuticular or intra epidermal, to a generalized infection of the entire section of the mesophyll,
inducing characteristic black shiny superficial symptoms, commonly known as tar spots (Santos et
al. 2016). However, differences in depth of ascomata can be influenced by the consistency of the
host tissue (Cannon 1991), therefore, this is not a valid character to distinguish genera.
Phylleutypa Petr., Annls mycol. 32(5/6): 429 (1934)
Index Fungorum number: IF4052; 2 morphological species (Species Fungorum 2020).
Type species – Phylleutypa dioscoreae (Wakef.) Petr.
Notes – Phylleutypa, typified by P. dioscoreae, is a stromatic parasite on Dioscorea species
(Wakefield 1918, Petrak 1934). This genus currently comprises two additional taxa, P. kalmiae and
P. wittrockii.
Phyllocrea Höhn., Annls mycol. 16(1/2): 38 (1918)
Index Fungorum number: IF4063; 3 morphological species (Species Fungorum 2020).
Type species – Phyllocrea quitensis (Pat.) Höhn.
Notes – The plant parasitic species, Phyllocrea indica, P. paulliniae and P. quitensis
collectively represent Phyllocrea. These species are characterized by ascomata immersed in a
bright stroma and 2-celled ascospores (Ainsworth et al. 1973).
Pseudothiella Petr., Hedwigia 68(5): 257 (1928)
Index Fungorum number: IF25396; 1 morphological species.
Type species – Pseudothiella hirtellae (Henn.) Petr.
Notes – Pseudothiella is a monotypic genus typified by P. hirtellae which is parasitic on
Hirtella. It can be distinguished by ascomata immersed in a stroma or pseudostroma that is covered
by a stromatic clypeus and ascospores with a hyaline equatorial band (Ainsworth et al. 1973).
Pseudothiopsella Petr., Hedwigia 68(5): 259 (1928)
797
Index Fungorum number: IF9618; 1 morphological species.
Type species – Pseudothiopsella hirtellae Petr.
Notes – Pseudothiopsella, a coelomycete, comprises a single species P. hirtellae, which has
branched conidiophores that give rise to discrete, hyaline, cylindrical, indeterminate, smooth
conidiogenous cells with 1–3 enteroblastic-percurrent proliferations, resulting in clearly visible
collarettes (Furlanetto & Dianese1998).
Pterosporidium W.H. Ho & K.D. Hyde, Can. J. Bot. 74(11): 1826 (1996)
Index Fungorum number: IF27720; 2 morphological species (Species Fungorum 2020).
Type species – Pterosporidium rhizophorae (Vizioli) W.H. Ho & K.D. Hyde
Notes – Pterosporidium was introduced by Ho & Hyde (1996) to accommodate P.
rhizomorphae, and P. rhizophorae, associated with foliar diseases of mangrove leaves (Ho & Hyde
1996). Species are characterized by thin-walled, clavate and unicellular brown ascospores that are
covered by a thin paper like sheath (Ho & Hyde 1996). The genus differs from Anthostomella as
the ascospores lack a germ slit and are covered in a thin paper-like sheath.
Rehmiodothis Theiss. & Syd., Annls mycol. 12(2): 192 (1914)
Index Fungorum number: IF4666; 8 morphological species (Species Fungorum 2020).
Type species – Rehmiodothis osbeckiae (Berk. & Broome) Theiss. & Syd.
Notes – Rehmiodothis species are typical tar spot pathogens which have flattened domeshaped, glossy, black stromata, with ascomatal ostioles emerging through the leaf surface as
slightly protruding structures (Swart 1987). The stromata originate in or beneath the epidermis,
become heavily melanised and contain a few ascomata with oblique necks. The haustoria originate
from vegetative hyphae growing between palisade cells (Park et al. 2000). Rehmiodothis osbeckiae
causes black, shiny and tar-spots in living leaves of Melastoma malabathricum, with oblong to eggshaped, light-coloured, unequally 2-celled and ascospores that are constricted at septa (Narendra &
Rao 1976).
Retroa P.F. Cannon, Mycol. Pap. 163: 201 (1991)
Index Fungorum number: IF15437; 2 morphological species (Species Fungorum 2020).
Type species – Retroa dimorphandrae (F. Stevens) P.F. Cannon
Notes – This genus comprises R. centrolobii and R. dimorphandrae that parasitize the
angiosperm family Leguminosae (Cannon 1991).
Rhodosticta Woron., Izv. Imp. St.-Peterburgsk. Bot. Sada 11: 13 (1911)
Index Fungorum number: IF9739; 3 morphological species (Species Fungorum 2020).
Type species – Rhodosticta caraganae Woron.
Notes – Rhodosticta comprises plant parasitic taxa on a variety of hosts (Davay et al. 1965).
Rhodosticta caraganae has conidia formed on branched, short, almost doliiform conidiogenous
cells, supported by short septate conidiophores (Sutton 1980)
Rikatlia P.F. Cannon, Syst. Ascom. 11(2): 83 (1993)
Index Fungorum number: IF25550; 1 morphological species.
Type species – Rikatlia lungusaensis (Henn.) P.F. Cannon
Notes – Cannon (1992) introduced Rikatlia, which is unique in having banded ascospores.
This genus is monotypic and lacks molecular data to substantiate its classification.
Schizochora Syd. & P. Syd., Annls mycol. 11(3): 265 (1913)
Index Fungorum number: IF4894; 3 morphological species (Species Fungorum 2020).
Type species – Schizochora elmeri Syd. & P. Syd.
Notes – Currently, S. calocarpa, S. elmeri, S. pandani and S. stenosperm, are included in
Schizochora.
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Sphaerodothella C.A. Pearce & K.D. Hyde, Fungal Divers. 6: 85 (2001)
Index Fungorum number: IF28523; 1 morphological species.
Type species – Sphaerodothella danthoniae (McAlpine) C.A. Pearce & K.D. Hyde
Notes – Sphaerodothella was introduced by Pearce & Hyde (2001) to accommodate S.
danthoniae which produces black tar spots on leaves and culms of Danthonia spp. Sphaerodothella
danthoniae has a peridium composed of hyaline to brown, thin-walled, flattened fungal cells which
merge with host cells, rather than a distinct stroma of melanized textura globosa, and pale brown
ascospores which have a distinctive, dark brown, mucilaginous perisporium (Pearce & Hyde 2001).
Sphaerodothis (Sacc. & P. Syd.) Shear, Mycologia 1(4): 162 (1909)
Index Fungorum number: IF5100; 18 morphological species (Species Fungorum 2020).
Type species – Sphaerodothis arengae (Racib.) Shear
Notes – Sphaerodothis was introduced by Saccardo & Sydow (1902) for a subgenus of
Auerswaldia to include the single species Sphaerodothis arengae. The prominent character used in
separating this subgenus from Auerswaldia was the shape of the spores which are sphaerical or
subsphaerical (Shear 1909).
Stigmatula (Sacc.) Syd. & P. Syd., Bull. Herb. Boissier, 2 sér. 1: 77 (1901)
Index Fungorum number: IF5258; 9 morphological species (Species Fungorum 2020).
Type species – Stigmatula sutherlandiae (Kalchbr. & Cooke) Syd. & P. Syd.
Notes – Stigmatula is a genus confined to the two tribes Galegeae and Hedysareae of the
Leguminosae (Cannon 1994). Stigmatula has brightly coloured stromata which contain starch
granules (Müller 1986).
Stigmochora Theiss. & Syd., Annls mycol. 12(3): 272 (1914)
Index Fungorum number: IF5262; 9 morphological species (Species Fungorum 2020).
Type species – Stigmochora controversa (Starbäck) Theiss. & Syd.
Notes – Stigmochora species are characterized by hyaline, unequally septate ascospores (Barr
& Hodges 1987). All Stigmochora species recognized by Muller & von Arx (1962) are parasitic on
tropical legumes forming tar spots on the leaves.
Stromaster (Pat.) Höhn., Mitt. bot. Inst. tech. Hochsch. Wien 7(3): 93 (1930)
Index Fungorum number: IF5287; 1 morphological species.
Type species – Stromaster tuberculatus (Pat.) Höhn
Notes – Stromaster is a poorly known monotypic genus which lacks molecular data.
Tamsiniella S.W. Wong, K.D. Hyde, W.H. Ho & S.J. Stanley, Can. J. Bot. 76(2): 334 (1998)
Index Fungorum number: IF27935; 1 morphological species.
Type species – Tamsiniella labiosa S.W. Wong, K.D. Hyde, W.H. Ho & S.J. Stanley
Notes – Tamsiniella was introduced by Wong et al. (1998c) to accommodate the freshwater
fungus, T. labiosa (Wong et al. 1998c). The genus can be distinguished by dark brown, immersed
to semi-immersed, subglobose ascomata, with periphyses, papillate, thin-walled, pale brown
peridium, paraphyses, 8-spored, unitunicate, cylindrical asci, with an unusual J-, lip-like, refractive
apical ring, and hyaline, ellipsoidal-fusiform, aseptate ascospores, with narrow, roughed
mucilaginous sheath (Wong et al. 1998c, Phookamsak et al. 2019).
Telimenella Petr., Annln K. K. naturh. Hofmus. Wien 50: 468 (1940)
Index Fungorum number: IF5365; 2 morphological species (Species Fungorum 2020).
Type species – Telimenella persica Petr.
Notes – Petrak (1940) erected Telimenella which comprises saprobes or weak parasites
forming conspicuous, black stromata on grasses (Barr 1977, Makelii & Koponen1974).
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Telimenochora Sivan., Trans. Br. mycol. Soc. 88(4): 474 (1987)
Index Fungorum number: IF25143; 1 morphological species.
Type species – Telimenochora abortiva (F. Stevens) Sivan.
Notes – The monotypic genus, Telimenochora is characterized by 2-septate ascospores,
persistent periphysoids (apical downwardly growing free, colourless, filamentous hyphae), and
evanescent, filamentous, colourless, branched paraphyses (Sivanesan 1987).
Trabutia Sacc. & Roum., Revue mycol., Toulouse 3(no. 9): 27 (1881)
Index Fungorum number: IF5509; 1 morphological species.
Type species – Trabutia quercina (F. Rudolphi) Sacc. & Roum.
Notes – Trabutia quercina forms ascomata embedded in stromata with 8-spored, clavate or
saccate asci with short pedicels with apex obtuse acute and fusiform to ellipsoidal-fusiform,
hyaline, aseptate, smooth-walled ascospores, often curved and flattened on one side, without a
gelatinous sheath or appendage (Trapero & Sánchez 2003).
Tribulatia Joanne E. Taylor, K.D. Hyde & E.B.G. Jones, Fungal Divers. Res. Ser. 12: 189 (2003)
Index Fungorum number: IF28739; 1 morphological species.
Type species – Tribulatia appendicospora Joanne E. Taylor, K.D. Hyde & E.B.G. Jones
Notes – Tribulatia is a monotypic genus, containing T. appendicospora growing on the palm
Archontophoeniox alexandrae. The genus is characterized by dark, ostiolate ascomata forming
under a clypeus, cylindrical asci with discoid, J-, apical ring, and hyaline, fusiform, aseptate,
ascospores with a mucilaginous appendage (Taylor et al. 2003).
Uropolystigma Maubl., Bull. Soc. mycol. Fr. 36: 36 (1920)
Index Fungorum number: IF5688; 1 morphological species.
Type species – Uropolystigma atrotestaceum Maubl.
Notes – Uropolystigma is a poorly known monotypic genus and no molecular data are
available to confirm its phylogenetic affinities.
Vitreostroma P.F. Cannon, Mycol. Pap. 163: 206 (1991)
Index Fungorum number: IF25153; 1 morphological species.
Type species – Vitreostroma desmodii (Henn.) P.F. Cannon
Notes – This is also a poorly known monotypic genus with V. desmodii as the type species.
Zimmermanniella Henn., Hedwigia 41: 142 (1902)
Index Fungorum number: IF5870; 1 morphological species.
Type species – Zimmermanniella trispora Henn.
Notes – Zimmermanniella was introduced to accommodate Z. trispora, a leaf parasite of
mango in Malaysia.
Pisorisporiaceae Réblová & J. Fourn., Persoonia 34: 43 (2015)
Index Fungorum number: IF810339; Facesoffungi number: FoF01286; 4 species.
Saprobic on decaying wood in freshwater and terrestrial habitats. Sexual morph: Ascomata
astromatic, perithecial, solitary to small groups, immersed to superficial, subhyaline to pale brown
or dark brown to black, upright or obliquely lying, papillate or with a long cylindrical neck,
sometimes hairy, ostiolate, periphysate. Peridium soft-textured, leathery to fragile, two-layered,
inner layer of hyaline, elongated cells, outer layer partly carbonaceous. Paraphyses abundant,
persistent, cylindrical, septate, tapering, longer than asci. Asci 8-spored, unitunicate, cylindricalclavate, pedicellate, with a J+, or J-, apical ring. Ascospores uni- to biseriate, smooth-walled,
fusiform, hyaline, cylindrical to cymbiform, slightly falcate, multi-septate, without appendages or
mucilaginous sheath, with or without guttules. Asexual morph: Undetermined (adapted from
Réblová et al. 2010, 2015a).
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Type genus – Pisorisporium Réblová & J. Fourn.
Notes – Based on morpho-molecular data, Pisorisporiaceae was introduced by Réblová et al.
(2015a) in Pisorisporiales. The family predominantly includes aquatic fungi and few terrestrial taxa
in Achroceratosphaeria and Pisorisporium. In the phylogenetic analysis, the two genera cluster in a
sister clade. However, the genera differ in ascomata and presence of a J+, apical ring in the asci:
Achroceratosphaeria has hyaline to pale brown ascomata and a J-, apical ascus ring, while
Pisorisporium has carbonaceous ascomata with a J+, ascal apical ring (Réblová et al. 2015a,
2016c). The apical ring colouration reactions found with Pisorisporium spp. are commonly
encountered in genera with chitinoid apical rings (Réblová et al. 2015a). Distribution of the family
is limited to countries in Europe and Asia (Maharachchikumbura et al. 2016b).
Ecological and economic significance of Pisorisporiaceae
Decomposes wood with its saprobic activity.
Genera included in Pisorisporiaceae
Achroceratosphaeria Réblová, J. Fourn., K.D. Hyde & Ranghoo, Fungal Divers. 43: 79 (2010)
Index Fungorum number: IF515196; 2 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Achroceratosphaeria potamia Réblová, J. Fourn. & K.D. Hyde
Notes – Currently, two species are accepted in Achroceratosphaeria (Réblová et al. 2010,
Wijayawardene et al. 2017a, 2018a), although molecular data is only available for A. potamia. The
asexual morph is undetermined. Achroceratosphaeria differs from Pisorisporium in having hyaline
to pale brown ascomata and J-, ascus, apical ring.
Pisorisporium Réblová & J. Fourn., Persoonia 34: 45 (2015)
Index Fungorum number: IF810340; 2 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Pisorisporium cymbiforme Réblová & J. Fourn.
Notes – The genus was introduced by Réblová et al. (2015a), to accommodate taxa isolated
from decaying wood submerged in freshwater and is characterized by brown to black astromatic
ascomata and a J+, apical ring in the asci. The ascospores of Pisorisporium cymbiforme are slightly
constricted at the septa, while the paraphyses and ascospores have thick walls.
Plectosphaerellaceae W. Gams, Summerb. & Zare, Nova Hedwigia 85(3–4): 476 (2007)
Index Fungorum number: IF510694; Facesoffungi number: FoF01334; 164 species.
Saprobic on dead plant material or in soil and water, or pathogenic on plants or animal or
mycoparasitic on fungi. Sexual morph: Ascomata perithecial or cleistothecial, superficial, brown to
dark brown, subglobose to pyriform or globose, with elongate neck, with or without setae around
the base of the neck. Peridium multi-layered, composed of dark brown cells of textura angularis.
Paraphyses conspicuous in young ascomata or absent. Asci 8-spored, unitunicate, cylindrical,
clavate or saccate, without an apical ring. Ascospores irregularly arranged, hyaline or pale brown,
fusiform or ellipsoidal or ovoid, 1-celled or 2-celled, wall asperulate or smooth. Asexual morph:
Hyphomycetous or coelomycetous. Conidiomata synnematous, sporodochial or acervular when
present. Conidiophores simple or branched, hyaline or centrally pale brown or pale olivaceous.
Conidiophores cells blastic or phialidic, hyaline, some become orange-brown or olivaceous-brown,
partly melanized, verticillate or single, pigmented chlamydospores and microsclerotia formed by
some species. Conidia aggregated in slimy heads, hyaline or pigmented, shapes variable (ranging
from subglobose to oval or cylindrical, slightly curved, or fusiform with pointed ends), 1–2-celled,
smooth-walled (adapted from Maharachchikumbura et al. 2016b, Giraldo & Crous 2019)
Type genus – Plectosphaerella Kleb.
Notes – Plectosphaerellaceae was introduced by Zare et al. (2007) typified by
Plectosphaerella (Zare et al. 2007, Kirk et al. 2008). The family is referred to Glomerellales by
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Maharachchikumbura et al. (2016b). Based on Maharachchikumbura et al. (2016b), there are nine
genera accepted and listed in the family (Zare et al. 2007, Kirk et al. 2007, Réblová et al. 2011,
Cannon et al. 2012a, Wijayawardene et al. 2012, Grum-Grzhimaylo et al. 2013, 2016, Hirooka et
al. 2014, Maharachchikumbura et al. 2015b, 2016b). Brunneomyces was introduced and typified by
Brunneomyces brunnescens (Giraldo et al. 2017b). Based on phylogenetic analyses of a combined
LSU, SSU, ITS and tef1 sequence data and the difference in morphology, Longitudinalis (Hyde et
al. 2017b) and Acremoniisimulans (Tibpromma et al. 2018) were introduced in
Plectosphaerellaceae. Giraldo & Crous (2019) introduced another 12 genera in Plectosphaerellaceae
based on phylogenetic analyses with a combined LSU, SSU, ITS, tef1, tryptophan synthase (ts), act
and rpb2 dataset and their unique morphology. Currently, 24 genera are recognized in the
Plectosphaerellaceae
viz.
Acremoniisimulans,
Acrostalagmus,
Brunneomyces,
Brunneochlamydosporium, Chlamydosporiella, Chordomyces, Furcasterigmium, Fuscohypha,
Gibellulopsis, Lectera, Longitudinalis, Musicillium, Musidium, Nigrocephalum, Paragibellulopsis,
Paramusicillium, Phialoparvum, Plectosphaerella, Sayamraella, Stachylidium, Sodiomyces,
Summerbellia, Theobromium and Verticillium sensu stricto. The placement of Cephalosporium
serrae, Gliocladium cibotii and several Acremonium species remains unresolved, but are presently
included in Plectosphaerellaceae.
Figure 200 – Pisorisporium cymbiforme (redrawn from Réblová et al. 2015a). a Ascomata b
Peridium. c Paraphyses. d Asci. e Ascospores. Scale bars: a = 200 μm, b = 20 μm, c-e = 10 μm.
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Ecological and economic significance of Plectosphaerellaceae
Plectosphaerellaceae species play an important role in nature as saprobic and pathogenic taxa.
Many in genera such as Acrostalagmus, Lectera, Musicillium, Plectosphaerella and Verticillium are
pathogens of different plants worldwide (Cannon et al. 2012a, Carlucci et al. 2012, Garibaldi et al.
2012, Mersha et al. 2012, Masudi & Bonjar 2012, Kanakala & Singh 2013, Hyde et al. 2014,
O’Neal & Davis 2015, Zhang et al. 2015b, Carrieri et al. 2017, Giraldo et al. 2017b, Raimondo &
Carlucci 2018, Giraldo & Crous 2019). Brunneomyces hominis, and some Chordomyces species
have been isolated from human patients, while others occur in plant debris, or are saprobes of
plants. Gibellulopsis and Plectosphaerella have been reported as opportunistic pathogens of
animals (Batista & Da Silva Maia 1959, Domsch et al. 2007, Duc et al. 2009). Verticillium
nonalfalfae and some Acrostalagmus species are used as biological control agents. (Mohammadi &
Amini 2015, Kasson et al. 2019). Gibellulopsis nigrescens, Verticillium longisporum and many
plectosphaerellaceous Acremonium species are reported as soil-borne saprobes (Gams 1975,
Domsch et al. 2007, Zare et al. 2007, Fogelqvist et al. 2018). Moreover, some Sodiomyces members
have alkaliphilic or alkalitolerant properties (Okada et al. 1993, Grum-Grzhimaylo et al. 2013,
2016, 2018, Kozlova et al. 2019).
Genera included in Plectosphaerellaceae
Acremoniisimulans Tibpromma & K.D. Hyde, Fungal Divers. 93(1): 1–160 (2018)
Index Fungorum number: IF555329; 1 species with sequence data.
Type species – Acremoniisimulans thailandensis Tibpromma & K.D. Hyde
Notes – Acremoniisimulans was introduced by Tibpromma et al. (2018) based on
phylogenetic analysis of a combined LSU, SSU, ITS and tef1 dataset and its unique morphology.
Acremoniisimulans is characterized by septate, unbranched conidiophores, which are brown to pale
brown to hyaline at the apex, and have solitary, pale brown to brown, oval, aseptate conidia.
Acrostalagmus Corda, Icon. fung. (Prague) 2: 15 (1838)
Index Fungorum number: IF7044; 13 morphological species (Species Fungorum 2020), 2
species with sequence data.
Type species – Acrostalagmus luteoalbus (Link) Zare, W. Gams & Schroers
Notes – Acrostalagmus cinnabarinus turned out to be identical to Verticillium luteoalbum (≡
Sporotrichum luteoalbum), introduced by Link (1809). Thus, the new combination Acrostalagmus
luteoalbus was proposed as the type species of Acrostalagmus (Zare et al. 2004). Acrostalagmus
has mononematous or synnematous conidiophores, enteroblastic, monophialidic conidiogenous
cells, and hyaline or bright orange to reddish, oval, ellipsoidal to oblong-ellipsoidal conidia, held
together by slime (Corda 1838, Giraldo & Crous 2019). Based on morphology, substrate
relationships, and phylogenetic analyses, Zare et al. (2004) demonstrated the asexual-sexual
connection was erroneous for Acrostalagmus. Its sexual morph is undetermined.
Brunneochlamydosporium Giraldo López & Crous, Stud. Mycol. 92: 260 (2018)
Index Fungorum number: IF828053; 4 species with sequence data.
Type species – Brunneochlamydosporium nepalense (W. Gams) Giraldo López & Crous
Notes – Based on phylogenetic analyses of combined LSU, ITS, tef1 and rpb2 loci,
Brunneochlamydosporium was introduced by Giraldo & Crous (2019) to accommodate the new
combinations Brunneochlamydosporium nepalense, B. cibotii and the new species B.
macroclavatum, and B. terrestre. The genus is characterized by enteroblastic, monophialidic,
sometimes polyphialidic, terminal, lateral, (sub)cylindrical to subulate, hyaline, conidiogenous
cells, with conspicuous collarette and a periclinal thickening at the conidiogenous locus. Conidia
are ellipsoidal, 1-celled, hyaline, thin- and smooth-walled, containing two guttules, arranged in
slimy heads. Chlamydospores are lateral, terminal, intercalary, solitary, in pairs or short chains, 1–
2-celled, pale to dark brown, smooth and thick-walled (Giraldo & Crous 2019).
803
Brunneomyces Giraldo, Gene & Guarro, Mycol. Progr. 16(4): 357 (2017)
Index Fungorum number: IF811471; 3 species with sequence data.
Type species – Brunneomyces brunnescens (W. Gams) Giraldo, Gene & Guarro
Notes – Brunneomyces was introduced by Giraldo et al. (2017b) with B. brunnescens as the
type species. The genus is characterized by brown hyphae, sympodial conidiophores and chains of
ovoidal to ellipsoidal conidia. Three species, B. brunnescens, B. europaeus and B. hominis are
including in this genus.
Chlamydosporiella Giraldo López & Crous, Stud. Mycol. 92: 270 (2019)
Index Fungorum number: IF828069; 1 species with sequence data.
Type species – Chlamydosporiella restricta (J.F.H. Beyma) Giraldo López & Crous
Notes – Based on the research of Gams (1971), the study of original material for seven strains
and additional specimens and the phylogenetic analysis of the combined alignment of the LSU,
ITS, tef1 and rpb2 loci, Giraldo & Crous (2019) introduced Chlamydosporiella to accommodate
Acremonium restrictum (≡ Verticillium dahliae f. cerebriforme) (Beyma 1940, Gams 1971). The
genus is characterized by bent, unbranched or basitonously branched conidiophores and
conidiogenous cells with short collarettes and inconspicuous periclinal thickening at the
conidiogenous locus. Conidia are obovoid, widely ellipsoidal, 1-celled, hyaline, thick- and smoothwalled, arranged in slimy heads. Chlamydospores are terminal or intercalary, mostly chains, dark
olive green and thick-walled. The sexual morph is undetermined (Giraldo & Crous 2019).
Chordomyces Bilanenko, Georgieva & Grum-Grzhim., Fungal Divers. 71(1): 29 (2015)
Index Fungorum number: IF811265; 2 species with sequence data.
Type species – Chordomyces antarcticus Bilanenko, Georgieva & Grum-Grzhim.
Notes – Grum-Grzhimaylo et al. (2016) introduced Chordomyces based on the phylogenetic
analyses of a combine ITS and LSU dataset for 12 alkaliphilic isolates. Two species, Chordomyces
antarcticum and C. albus were introduced in the genus (Grum-Grzhimaylo et al. 2016, Giraldo et
al. 2017b). Chordomyces is characterized by erect conidiophores, solitary or forming in synnemata.
Conidiogenous cells are enteroblastic, mono- or polyphialidic, tapering towards the apex and often
proliferating sympodially. Conidia are subglobose, limoniform, ellipsoidal to cylindrical, rounded
at the apex, sometimes with protuberant hilum and1(‒2)-celled (Giraldo et al. 2017b).
Furcasterigmium Giraldo López & Crous, Stud. Mycol. 92: 251 (2019)
Index Fungorum number: IF828041; 1 species with sequence data.
Type species – Furcasterigmium furcatum (Moreau & V. Moreau) Giraldo López & Crous
Notes – Based on the phylogenetic analyses of combined LSU, ITS, tef1 and rpb2 sequence
data from 20 isolates of Acremonium furcatum, Giraldo & Crous (2019) introduced
Furcasterigmium for species of A. furcatum. This genus is characterized by erect conidiophores,
often proliferating sympodially, with conidiogenous cells as short lateral and cylindrical
asymmetrical projections. Conidiogenous cells are enteroblastic, mono- and polyphialidic, terminal,
lateral, subulate, hyaline, with conspicuous collarette and periclinal thickening at the conidiogenous
locus. Conidia are ellipsoidal, 1-celled, hyaline, smooth-walled and arranged in slimy heads
(Giraldo & Crous 2019).
Fuscohypha Giraldo López & Crous, Stud. Mycol. 92: 264 (2019)
Index Fungorum number: IF828058; 1 species with sequence data.
Type species – Fuscohypha expansa Giraldo López & Crous
Notes – Based on the analyses of combined LSU, ITS, tef1 and rpb2 sequence data and
morphological differences from Verticillium, Giraldo & Crous (2019) introduced Fuscohypha to
accommodate a single species F. expansa. The genus is characterized by simple or verticillate, erect
conidiophores. Conidiogenous cells are enteroblastic, monophialidic, terminal, lateral, cylindrical
to subulate, hyaline, with cylindrical collarette and a conspicuous periclinal thickening at the
804
conidiogenous locus. Conidia are subglobose or ellipsoidal, 1-celled and hyaline (Giraldo & Crous
2019).
Gibellulopsis Bat. & H. Maia, Anais Soc. Biol. Pernambuco 16(1): 153 (1959)
Index Fungorum number: IF8331; 7 species with sequence data.
Type species – Gibellulopsis piscis Bat. & H. Maia
Notes – Gibellulopsis was introduced by Batista & da Silva Maia (1959) and reused by Zare
et al. (2007) with G. piscis as its type species. Currently the genus comprises seven species (Zare et
al. 2007, Hirooka et al. 2014, Giraldo & Crous 2019, Index Fungorum 2020). This genus is
characterized by conidiophores arising from submerged or superficial hyphae, which are erect,
mostly terminal, usually 1–2 times branched, bearing one or two verticillate branches at a node.
Conidiogenous cells have inconspicuous collarettes and distinct periclinal thickenings at the
conidiogenous locus. Conidia are elongate ellipsoidal to cylindrical, 1- or 2-celled, hyaline,
smooth-walled and produced in slimy heads. Chlamydospores are lateral, terminal or intercalary,
pale to dark brown and form singly or in chains (Zare et al. 2007, Giraldo & Crous 2019).
Lectera P.F. Cannon, MycoKeys 3: 28 (2012)
Index Fungorum number: IF550041; 6 morphological species (Species Fungorum 2020), 4
species with sequence data.
Type species – Lectera colletotrichoides (J.E. Chilton) P.F. Cannon
Notes – Lectera established by Cannon et al. (2012a) for the plant pathogen Lectera
colletotrichoides and L. longa, based on the difference in ITS and gapdh sequences and the size of
conidia. Lectera capsici, L. humicola, L. nordwiniana and L. phaseoli were introduced in the genus
(Crous et al. 2017b, Giraldo & Crous 2019). This genus is characterized by sporodochial or
acervular conidiomata, erumpent through host tissues and without a clear upper wall, globose to
subglobose, pink or flesh coloured, with few marginal, erect setae. Setae are dark brown, septate,
and taper towards the apex. Conidiophores are often reduced to the conidiogenous cell. Conidia are
navicular or fusiform with pointed ends, and slightly curved (Cannon et al. 2012a).
Longitudinalis Tibpromma & K.D. Hyde, Fungal Divers. 87: 155 (2017)
Index Fungorum number: IF553099; 1 species with sequence data.
Type species – Longitudinalis nabanheensis Tibpromma & K.D. Hyde
Notes – Based on the phylogenetic analyses of a combined LSU, ITS and tef1 dataset in
Plectosphaerellaceae and the morphological difference from Synnemellisia and other
Plectosphaerellaceae genera, Longitudinalis was established by Tibpromma et al. (2018) for a taxon
L. nabanheensis (Giraldo & Crous 2019, Index Fungorum 2020, NCBI 2019). The genus is
characterized by synnemata with yellow-green to dark-brown stem and a dark-brown pin-like head
of conidiogenous cells. Conidia are aggregated on a pin-like head, fusoid-ellipsoid, tapering
towards both ends, with apex subobtuse to obtuse, at the beginning hyaline with dark brown at the
towards both ends and immature conidia yellow-green with brown longitudinal striations
(Tibpromma et al. 2018).
Musicillium Zare & W. Gams, Nova Hedwigia 85(3–4): 482 (2007)
Index Fungorum number: IF510696; 4 species with sequence data.
Type species – Musicillium theobromae (Turconi) Zare & W. Gams
Notes – Musicillium was introduced for the causal agent of cigar-end rot by Zare et al. (2007)
based on the morphological difference from Verticillium species. Musicillium elettariae, M.
pandanicola, M. theobromae and M. tropicale were included in this genus (Zare et al. 2007,
Tibpromma et al. 2018, Giraldo & Crous 2019, Index Fungorum 2020). The genus is characterized
by conidiophores that arise from submerged hyphae, and are erect, septate, repeatedly verticillate
towards the apex and distinctly brown throughout (slightly pale only near the tip). Chlamydospores
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are lacking, but moniliform hyphae are initially subhyaline, but turning brown (Zare et al. 2007,
Giraldo & Crous 2019).
Musidium Giraldo López & Crous, Stud. Mycol. 92: 253 (2019)
Index Fungorum number: IF828045; 1 species with sequence data.
Type species – Musidium stromaticum (W. Gams & R.H. Stover) Giraldo López & Crous
Notes – Musidium was established by Giraldo & Crous (2019) to accommodate the isolate
previously classified as Acremonium stromaticum. The genus is closely related to Sayamraella,
Summerbellia and Theobromium based on the analyses of a combine LSU, ITS, tef1 and rpb2
dataset, however, Musidium differs in its branched, dark olivaceous, incrusted or smooth and thick
walled stromatic hyphae (Giraldo & Crous 2019).
Nigrocephalum Giraldo López & Crous, Stud. Mycol. 92: 271 (2019)
Index Fungorum number: IF828071; 1 species with sequence data.
Type species – Nigrocephalum collariferum (Weisenb. & R. Kirschner) Giraldo López &
Crous
Notes – The monotypic genus Nigrocephalum was introduced by Giraldo & Crous (2019) to
accommodate two isolates formerly described as Acremonium collariferum, which were isolated
from the skin and nail lesions of humans. Nigrocephalum species can degrade keratin in vitro at 33
°C. The genus is characterized by erect, slightly bent, simple or basitonously branched
conidiophores. Conidiogenous cells have a conspicuous funnel-shaped collarette and periclinal
thickening at the conidiogenous locus. Conidia are ellipsoidal, concave in lateral view and
pigmented (Giraldo & Crous 2019).
Paragibellulopsis Giraldo López & Crous, Stud. Mycol. 92: 265 (2019)
Index Fungorum number: IF828060; 1 species with sequence data.
Type species – Paragibellulopsis chrysanthemi (Hirooka, Kawaradani & Toy. Sato) Giraldo
López & Crous
Notes – Paragibellulopsis is characterized by straight or slightly curved, 2-celled cylindrical
conidia, with tapering ends. Chlamydospores are grey brown, intercalary and single or in short in
chains (Giraldo & Crous 2019).
Paramusicillium Giraldo López & Crous, Stud. Mycol. 92: 269 (2019)
Index Fungorum number: IF828067; 1 species with sequence data.
Type species – Paramusicillium asperulatum Giraldo López & Crous
Notes – Giraldo & Crous (2019) introduced Paramusicillium to accommodate Musicillium
theobromae, based on the length and rough walls of conidiophores, and the phylogenetic analyses
of a combined LSU, ITS, tef1 and rpb2 dataset of the type species.
Phialoparvum Giraldo López & Crous, Stud. Mycol. 92: 265 (2019)
Index Fungorum number: IF828063; 1 species with sequence data.
Type species – Phialoparvum bifurcatum Giraldo López & Crous
Notes – Phialoparvum was established by Giraldo & Crous (2019) to accommodate the strain
CBS 299.70B, which was formerly classified as Acremonium furcatum. The genus was introduced
based on its difference in the apex of the conidiogenous cells and its conidia being longer than the
type of Acremonium and phylogenetically distant from the type strain as a single lineage.
Plectosphaerella Kleb., Phytopath. Z. 1: 43 (1929)
Index Fungorum number: IF4197; 20 morphological species (Species Fungorum 2020), 14
species with sequence data.
Type species – Plectosphaerella cucumerina (Lindf.) W. Gams
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Notes – Plectosphaerella was introduced by Klebahn in 1929 (Palm et al. 1995). The type
species is Plectosphaerella cucumerina, a holomorphic fungus (Zare et al. 2007, Carlucci et al.
2012). Plectosphaerella can be distinguished from other taxa by its asexual morph with the ratio of
septate conidia, conidial shape and dimensions and presence or absence of chlamydospores
(Carlucci et al. 2012).
Sayamraella Giraldo López & Crous, Stud. Mycol. 92: 255 (2019)
Index Fungorum number: IF828047; 1 species with sequence data.
Type species – Sayamraella subulata Giraldo López & Crous
Notes – Sayamraella was introduced as a monotypic genus for the isolates from soil based on
analysis of a LSU, ITS, tef1 and rpb2 dataset. The genus is characterized by conidiophores often
proliferating sympodially and conidiogenous cells as short lateral and cylindrical asymmetrical
projections. Conidiogenous cells have minute cylindrical collarettes and an inconspicuous
periclinal thickening at the conidiogenous locus.
Sodiomyces A.A. Grum-Grzhim., Debets & Bilanenko, Fungal Systematics and Evolution 3: 131
(2019)
Index Fungorum number: IF829354; 4 species with sequence data.
Type species – Sodiomyces alkalinus Grum-Grzhim., Debets & Bilanenko
Notes – Sodiomyces was introduced by Grum-Grzhimaylo et al. (2013) to accommodate the
holomorphic alkaliphilic species Heleococcum alkalinum based on the taxonomic reference and
analyses of a combined ITS, LSU, SSU, rpb2 and tef1 dataset. All Sodiomyces species were invalid
as the basionym lacked a valid type species (Giraldo & Crous 2019). Giraldo & Crous (2019) and
Crous et al. (2019b) redescribed, accepted and validated all Sodiomyces species. This genus is
characterized by cleistothecial ascomata, saccate asci, ellipsoidal or ovoid 2-celled and pale brown
ascospores and subglobose, cylindrical to oval or allantoid conidia (Grum-Grzhimaylo et al. 2013;
Giraldo & Crous 2019).
Stachylidium Link, Mag. Gesell. naturf. Freunde, Berlin 3(1–2): 15 (1809)
Index Fungorum number: IF10053; 7 morphological species (Species Fungorum 2020), 2
species with sequence data.
Type species – Stachylidium bicolor Link
Notes – Stachylidium was introduced by Link (1809) without assigning a type specimen.
Stachylidium bicolor was recognized as the type specimen (Hughes 1951b, Ellis 1971, Matsushima
1971, Barnett & Hunter 1972, von Arx 1981b, Dewi 2006). The lectotype of the S. bicolor was
designated by Hughes (1951b) and Gams (2017) based on the specimen under the name and
synonym of S. bicolor (Hughes 1958, Giraldo & Crous 2019). However, the lectotype designation
was controversial (Fries 1832, Giraldo & Crous 2019). Based on specimen examination from B and
L, Giraldo & Crous (2019) proposed the specimen with code B700016303 from B to be the most
suitable lectotype. The genus is characterized by septate, verticillate conidiophores which are pale
brown to brown at the base, sometimes pale to hyaline towards the apex, and roughened, singly or
in groups, sometimes forming lax synnemata. Conidiogenous cells are enteroblastic, monophialidic,
in whorls, terminal, lateral, cylindrical, ellipsoidal, hyaline or pale brown. Conidia are ellipsoidal or
cylindrical, 1-celled and pale brown to brown.
Summerbellia Giraldo López & Crous, Stud. Mycol. 92: 252 (2019)
Index Fungorum number: IF828043; 1 species with sequence data.
Type species – Summerbellia oligotrophica Giraldo López & Crous
Notes – Summerbellia was established by Giraldo & Crous (2019) with Summerbellia
oligotrophica as the type species. It was introduced as the monotypic genus for a group of isolates,
which clustered as a monophyletic lineage in the analyses of a combined LSU, ITS, tef1 and rpb2-
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Figure 201 – Plectosphaerella cucumerina (Material examined – AUSTRALIA, Queensland,
Tewantin, on the leaf of Lactuca sativa, 22 January 1991, Mayers, P.E., BRIP 17392, =
Monographella cucumerina). a Herbarium material. b Host. c Colony on the host. d Appearance of
colonies on leaf surface. e, f Conidiophores. g Conidiogenous cell. h, i Conidia. Scale bars: c = 100
µm, d = 1 mm, e, f = 25 µm, g = 15 µm, h-i = 5 µm.
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-dataset and previously identified as Gliocladium cibotii. This genus is characterized by erect or
(sub-)erect, unbranched or poorly branched conidiophores. Chlamydospores are terminal or
intercalary, mostly in chains, pale to dark brown, smooth- and thick-walled (Giraldo & Crous
2019).
Theobromium Giraldo López & Crous, Stud. Mycol. 92: 256 (2019)
Index Fungorum number: IF828049; 1 species with sequence data.
Type species – Theobromium fuscum Giraldo López & Crous
Notes – Theobromium was introduced by Giraldo & Crous (2019) to accommodate a strain
isolated from Theobroma sp. This genus is characterized by conidiophores often proliferating
sympodially and conidiogenous cells with short lateral and cylindrical asymmetrical projections
(Giraldo & Crous 2019).
Verticillium Nees, Syst. Pilze (Würzburg): 57 (1816)
Index Fungorum number: IF10400; 81 morphological species (Species Fungorum 2020), 20
species with sequence data.
Type species – Verticillium dahliae Kleb.
Notes – Species of this genus are plant pathogens (Domsch et al. 2007). This genus is
characterized by conidiophores arising from submerged hyphae or aerial mycelium, erect, mostly
verticillate, septate, hyaline to subhyaline, sometimes turning brown at the base. Conidiogenous
cells are enteroblastic, monophialidic, inserted in a mesotonous to acrotonous position, terminal,
lateral, flask-shaped or aculeate, hyaline, with inconspicuous collarettes. Resting structures include
pigmented resting mycelium, chlamydospores in short chains, and/or microsclerotia (Giraldo &
Crous 2019).
Pleurostomataceae Réblová, L. Mostert, W. Gams & Crous, Stud. Mycol. 50: 540 (2004)
Index Fungorum number: IF500153 Facesoffungi number: FoF01136; 5 species.
Saprobic on wood or soil in terrestrial habitats, or in sewage or pathogen of humans causing
subcutaneous phaeohyphomycosis. Sexual morph: Ascomata perithecial, black, gregarious or
scattered, superficial, stipitate, globose to subglobose, coriaceous, smooth, without setae, papillate;
papilla positioned laterally. Peridium composed of two to four layers, outer layer comprising brown
cells of textura intricata or epidermoidea, thick, coriaceous; inner layer comprising hyaline cells of
textura prismatica or angularis, thin, membranaceous; sometimes middle layers comprised of two
types of cells, forming thin-walled and brown cells of textura epidermoidea to in the outer part,
merging with thin-walled and dark brown cells of textura angularis in the inner part, thick,
coriaceous. Paraphyses numerous, hyaline, filamentous, early deliquescing. Asci polysporous,
unitunicate, reniform or oval, with short pedicel or sessile, apical ring lacking. Ascospores 2–3seriate, hyaline, oblong to allantoid, curved, aseptate, smooth-walled. Asexual morph: Mycelium
composed of branched, septate, hyaline or brown hyphae. Conidiophores in vitro arising from
aerial or submerged hyphae, hyaline, straight or flexuous, 0–2-septate, tuberculate or smoothwalled. Conidiogenous cells mono- or polyphialidic, cylindrical, hyaline, smooth-walled. Conidia
aggregated in slimy droplets, hyaline, aseptate, oblong to allantoid, curved or straight, smooth, with
or without guttules (adapted from Réblová et al. 2015b, Maharachchikumbura et al. 2015, 2016b).
Type genus – Pleurostoma Tul. & C. Tul.
Notes – Pleurostomataceae was introduced by Réblová et al. (2004). The genus was placed in
Calosphaeriales based on allantoid ascospores, ascogenous hyphae and other characteristics uniting
members, as well analyses of SSU and LSU sequence data (Réblová et al. 2004). Berlese (1900)
introduced Neoarcangelia with N. ootheca based on upright papillae on the ascomata. However,
Barr (1985) maintained there were insufficient features to separate the species from Pleurostoma
candollei and placed Neoarcangelia as the synonym of Pleurostoma (Höhnel 1918b, Barr 1985).
Shear (1937) also reexamined Sphaeria ootheca, the basionym of P. ootheca from Virginia, and
concluded the species was similar to P. candollei. Pleurostomophora was accepted as the asexual
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morph of Pleurostoma based on morphology and phylogenetic evidence (Réblová et al. 2004,
Vijaykrishna et al. 2004, Najwa et al. 2012). However, only the life history of Pleurostoma ootheca
has been experimentally verified and linked with Pleurostomophora ootheca. The asexual morph of
P. candollei is undetermined, except for an illustration in the protologue showing a sporodochial
conidiomata (Tulasne & Tulasne 1863). Réblová et al. (2015b) and Maharachchikumbura et al.
(2015) reported that Pleurostoma and Pleurostomophora are congeneric as both genera constitute a
strongly supported monophyletic clade in Pleurostomataceae in a multi-gene phylogenetic analysis.
Réblová et al. (2015b) proposed Pleurostoma as the correct name for the genus following the
principle of priority.
Ecological and economic significance of Pleurostomataceae
Several members of Pleurostomataceae are pathogenic on plants or/and humans. Pleurostoma
richardsiae has been isolated from wood, sewage and soil (Schol-Schwarz 1970) and in grafted
unions of nursery vines and diseased vines (Halleen et al. 2003, Halleen & Groenewald 2005,
Carlucci et al. 2015) and also infects humans (de Hoog et al. 2000). Pleurostoma repens was
associated with subcutaneous infections with granulomatous nodules in humans (Hironaga et al.
1989).
Genus included in Pleurostomataceae
Pleurostoma Tul. & C. Tul., Select. fung. carpol. (Paris) 2: 247 (1863)
Index Fungorum number: IF4247; 5 morphological species (Species Fungorum 2020), 4
species with sequence data.
Type species – Pleurostoma candollei Tul. & C. Tul.
Notes – Pleurostoma candollei is characterized by globose, black ascomata, with reniform to
oblong asci and allantoid, aseptate ascospores (Tulasne & Tulasne 1863, Réblová et al. 2015b,
Maharachchikumbura et al. 2016b).
Pleurotheciaceae Réblová & Seifert, Persoonia 37: 63 (2015)
Index Fungorum number: IF813229; Facesoffungi number: FoF05316; 85 species.
Saprobic on plant tissues of a variety of hosts. Sexual morph: Ascomata perithecial,
immersed, semi-immersed or superficial, papillate or with a central rarely eccentric neck. Ostiole
periphysate. Peridium leathery to fragile, carbonaceous, comprising two layers, outer layer
comprising brown to dark brown cells, inner layer comprising hyaline to pale brown cells.
Paraphyses abundant, sparsely branched, partially disintegrating, cylindrical. Asci 8-spored,
unitunicate, cylindrical or cylindrical-clavate, short or long pedicellate, with a pronounced J-, apical
ring. Ascospores overlapping or 1–3-seriate, hyaline or versicolorous with polar cells hyaline and
middle cells brown, ellipsoidal to fusiform, transversely multi-septate, lacking a mucilaginous
sheath or appendages. Asexual morph: Conidiomata present or absent, when present indeterminate
synnemata or loose fascicles. Conidiophores macronematous or semi-macronematous, sometimes
elongating percurrently. Conidiogenous cells holoblastic, conidial secession rhexolytic on short
denticles or rachis on sympodially extending polyblastic conidiogenous cells, or schizolytic on
monoblastic or solitary thallic conidiogenous cells. Conidia hyaline, sometimes with protracted
maturation of the middle cells, which turn brown, or brown or versicolorous, septate or aseptate.
(adapted from Réblová et al. 2016c).
Type genus – Pleurothecium Höhn.
Notes – Pleurotheciaceae was introduced by Réblová et al. (2016c) in Pleurotheciales. Ten
genera, i.e. Adelosphaeria, Brachysporiella, Helicoon, Melanotrigonum, Phaeoisaria,
Phragmocephala, Pleurotheciella, Pleurothecium, Sterigmatobotrys and Taeniolella, were included
in this family (Réblová et al. 2016c). Maharachchikumbura et al. (2016b) accepted eleven genera in
Pleurotheciaceae including Plagiascoma, which was placed in Fuscosporellales (Réblová et al.
2016c, Yang et al. 2016b, Wijayawardene et al. 2017a, 2018a). In addition, Monotosporella and
Helicoon were considered as members of Savoryellaceae (Savoryellales) in Maharachchikumbura
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et al. (2016b), while they were confirmed in Pleurotheciaceae by Réblová et al. (2012, 2016c).
Monotosporella was treated as a synonym of Brachysporiella by Ellis (1959). Hernández-Restrepo
et al. (2017) considered the genera Brachysporiella and Monotosporella distinct with support of
multi-locus phylogenetic analysis, and placed the genera in Kirschsteiniotheliales and
Pleurotheciales, respectively. Wijayawardene et al. (2017a) placed Brachysporiella in
Pleurotheciaceae and Monotosporella in Savoryellaceae, however, Wijayawardene et al. (2018a)
transferred Brachysporiella to Kirschsteiniotheliales genera incertae sedis and retained
Monotosporella in Savoryellaceae. Anapleurothecium was introduced by Hernández-Restrepo et al.
(2017) and placed in Pleurotheciaceae. Plagiascoma was accepted in Pleurotheciaceae by
Wijayawardene et al. (2017a, 2018a).
Figure 202 – Pleurostoma candollei (Material examined – FRANCE, Hauts-de-Seine, Chaville, on
dead wood of Quercus sp., 8 March 1859, PC 0167640). a Material. b Ascomata on host. c Ascoma
cross section. d Peridium. e, f Asci. g, h Ascospores. Scale bars: b = 200 µm, c = 100 µm, d = 20
µm, e-f = 5 µm, g-h = 1 µm.
Taeniolella, based on T. rudis, was included in Pleurotheciaceae by Réblová et al. (2016c) as
sister to Sterigmatobotrys. The fertile, penicillate sterigmatobotrys-like conidiophores developing
at the apex of the Taeniolella conidium were earlier reported by Réblová & Seifert (2011)
suggesting a close relationship between the two genera. Ertz et al. (2016) transferred Taeniolella
rudis to Sterigmatobotrys, based on the morphology of the penicillate synasexual morph and
molecular data. The type species, Taeniolella exilis, was placed in Kirschsteiniotheliaceae in
Dothideomycetes, and Taeniolella was recovered as strongly polyphyletic (Ertz et al. 2016).
Wijayawardene et al. (2017a) placed in Kirschsteiniotheliaceae (Kirschsteiniotheliales,
Dothideomycetes), however, Ekanayaka et al. (2017) transferred this genus to Mytilinidiaceae
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(Mytilinidiales, Dothideomycetes) and Wijayawardene et al. (2018a) accepted this treatment. In
this study, we accept the treatment of Ertz et al. (2016) and excluded Taeniolella from
Pleurotheciaceae.
It was shown that Helicoon is strongly polyphyletic. It was placed in three different classes,
viz. Leotiomycetes, Sordariomycetes and Dothideomycetes (Tsui & Berbee 2006). Helicoon
farinosum was included in Pleurotheciaceae by Réblová et al. (2016c) and Maharachchikumbura et
al. (2016b) based on the support of the multi-locus phylogenetic analyses. Wijayawardene et al.
(2017a) placed this genus in Orbiliaceae (Orbiliales, Orbiliomycetes), however, Wijayawardene et
al. (2018a) placed it in Savoryellaceae (Savoryellales, Sordariomycetes). Dayarathne et al. (2019a)
introduced Helicoascotaiwania to accommodate Ascotaiwania hughesii (asexual morph is Helicoon
farinosum) and placed this genus in Pleurotheciaceae.
Phragmocephala is polyphyletic (Su et al. 2015, Réblová et al. 2016b, Hernández-Restrepo et
al. 2017). Currently, there are nine species epithets in Index Fungorum (2020), however, only three
of them, P. garethjonesii, P. glanduliformis and P. stemphylioides, have molecular data available.
Su et al. (2015) introduced P. garethjonesii based on DNA sequence data and morphology and
placed this species in Melanommataceae in Dothideomycetes. Réblová et al. (2016c) placed P.
stemphylioides in Pleurotheciaceae in Sordariomycetes based on multi-locus phylogenetic analyses.
Phragmocephala glanduliformis was placed in Microthyriaceae in Dothideomycetes by HernándezRestrepo et al. (2017).
In this study, a multi-locus phylogenetic analyses based on a combined ITS, LSU, SSU and
rpb2 sequence data of Pleurotheciales is presented (Fig. 10). Bayesian inference, maximum
parsimony and maximum likelihood were used for phylogenetic analyses. The analyses provided
similar tree topologies, which are similar with those in Réblová et al. (2016c), Yang et al. (2016b),
Hernández-Restrepo et al. (2017), Hyde et al. (2017b, 2018b) and Luo et al. (2018). The
problematic genera and species and the newly introduced genus after Réblová et al. (2016c),
Phragmocephala stemphylioides (DAOM 673211), “Brachysporiella setosa” (HKUCC 3713)
(current name: Monotosporella setosa), Anapleurothecium botulisporum (FMR 11490),
“Taeniolella rudis” (DAOM 229838) (current name: Sterigmatobotrys rudis), Helicoon farinosum
(current name: Helicoascotaiwania hughesii) (ILLS 53605 and DAOM 241947), are grouped in a
robust clade Pleurotheciaceae (Fig. 10).
In conclusion, genera that accepted in the family in this study are: Adelosphaeria,
Anapleurothecium, Helicoascotaiwania, Melanotrigonum, Monotosporella, Phaeoisaria,
Phragmocephala, Pleurotheciella, Pleurothecium and Sterigmatobotrys.
Ecological and economic significance of Pleurotheciaceae
Some species of Pleurotheciaceae are human pathogens. A case of infectious keratitis caused
by presumed Carpoligna pleurothecii (current name: Pleurothecium recurvatum) was reported by
Chew et al. (2010). Taeniolella exilis was reported to infect humans and cause human subcutaneous
phaeohyphomycosis (Réblová et al. 2016c). Other species are likely to be aquatic and terrestrial
saprobes which are important in nutrient cycling.
Genera included in Pleurotheciaceae
Adelosphaeria Réblová, Persoonia 37: 63 (2016)
Index Fungorum number: IF813230; 1 species with sequence data.
Type species – Adelosphaeria catenata Réblová
Notes – This genus is similar to Pleurotheciella, however, its asexual morph is undetermined
(Réblová et al. 2016c). The genus is characterised by perithecial, astromatic, subglobose, dark
brown, papillate ascomata, periphysate ostioles, abundant, persistent, septate paraphyses,
unitunicate, cylindrical-clavate, pedicellate, 8-spored asci with a J-, apical ring, and ellipsoidal,
slightly curved, hyaline, transversely septate ascospores (Réblová et al. 2016c).
Anapleurothecium Hern.-Restr., R.F. Castañeda & Gené, Stud. Mycol. 86: 87 (2017)
812
Index Fungorum number: IF820300; 1 species with sequence data.
Type species – Anapleurothecium botulisporum Hern.-Restr., R.F. Castañeda & Gené
Notes – The genus is characterised by mononematous, unbranched, brown conidiophores,
sympodial, denticulate, brown conidiogenous cells and solitary, acropleurogenous, septate,
botuliform to cylindrical, brown conidia (Hernández-Restrepo et al. 2016a). It differs from other
genera by its polyblastic, sympodial, denticulate, brown conidiogenous cells that produce septate,
botuliform to cylindrical, brown conidia. Conidial secession is schizolytic.
Helicoascotaiwania Dayar., Maharachch. & K.D. Hyde, Frontiers in Microbiology 10 (no. 840): 22
(2019)
Index Fungorum number: IF555625; 1 species with sequence data.
Type species – Helicoascotaiwania hughesii (Fallah, J.L. Crane & Shearer) Dayar. & K.D.
Hyde
Notes – See the notes for Pleurotheciaceae. The genus is characterised by rarely branched,
hyaline conidiophores, determinate, denticles conidiogenous cells and helicosporous hyaline
conidia (Dayarathne et al. 2019a).
Melanotrigonum Réblová, Persoonia 37: 65 (2016)
Index Fungorum number: IF813232; 1 species with sequence data.
Type species – Melanotrigonum ovale Réblová
Notes – Melanotrigonum differs from Pleurotheciella by its triangular to ampulliform
conidiogenous cells which taper towards the apex with a single or rarely two denticles (Réblová et
al. 2016c).
Monotosporella S. Hughes, Can. J. Bot. 36: 786 (1958)
Index Fungorum number: IF8991; 4 morphological species (Species Fungorum 2020).
Type species – Monotosporella setosa (Berk. & M.A. Curtis) S. Hughes
Notes – See the notes for Pleurotheciaceae. The genus is characterised by unbranched, brown
conidiophores, monoblastic, percurrent, brown conidiogenous cells and septate, brown conidia
(Seifert et al. 2011).
Phaeoisaria Höhn., Math.-naturw. Kl., Abt. 1 118: 330 (1909)
Index Fungorum number: IF9305; 23 morphological species (Species Fungorum 2020), 11
species with sequence data.
Type species – Phaeoisaria bambusae Höhn.
Notes – Phaeoisaria is characterized by indeterminate synnemata with septate or aseptate
ellipsoidal, obovoidal, fusiform-cylindrical or falcate conidia.
Phragmocephala E.W. Mason & S. Hughes, Naturalist (Hull), ser. 3, 1951: 97 (1951)
Index Fungorum number: IF9367; 9 morphological species (Species Fungorum 2020), 4
species with sequence data.
Type species – Phragmocephala cookei E.W. Mason & S. Hughes
Notes – See the notes for Pleurotheciaceae. The genus is characterised by mononematous or
synnematous, unbranched, brown, percurrent conidiophores, monoblastic or solitary thallic, brown
conidiogenous cells and brown conidia with dark bands around septa (Seifert et al. 2011).
Pleurotheciella Réblová, Seifert & J. Fourn., Mycologia 104(6): 1304 (2012)
Index Fungorum number: IF564282; 11 species with sequence data.
Type species – Pleurotheciella rivularia Réblová, Seifert & J. Fourn.
Notes – Pleurotheciella was established by Réblová et al. (2012) with a dactylaria-like
asexual morph. The genus is characterised by unbranched to sparingly branched, septate
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conidiophores, integrated, denticles conidiogenous cells and ellipsoidal to clavate, hyaline, 0–
several-septate (Réblová et al. 2012). Pleurotheciella fusiformis is illustrated in this entry.
Figure 203 – Pleurotheciella fusiformis (Material examined – CHINA, Yunnan Province, saprobic
on decaying wood submerged in Erhai Lake, March 2015, H.Y. Su 1EHS A 17–5, MFLU 17–0908,
HKAS 92611; living culture, MFLUCC 17–0115, KUMCC 15–0196). a, b Ascomata on the natural
substratum. c, d Vertical sections of the peridium. e, f Asci. g-k Ascospores. Scale bars: c = 100
μm, d-f = 30 μm, g-k = 10 μm.
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Pleurothecium Höhn., Ber. dt. bot. Ges. 37: 154 (1919)
Index Fungorum number: IF9475; 11 morphological species (Species Fungorum 2020), 6
species with sequence data.
Type species – Pleurothecium recurvatum (Morgan) Höhn.
Notes – Pleurothecium was introduced by Höhnel (1919) to accommodate P. recurvatum
(Morgan) Höhn (Goos 1969). Fernández et al. (1999) introduced a sexual genus Carpoligna and
linked it to Pleurothecium. Réblová et al. (2016b) proposed using Pleurothecium rather than
Carpoligna, because of its priority and greater number of species. Pleurothecium floriforme is
illustrated in this entry.
Figure 204 – Pleurothecium floriforme (Material examined – THAILAND, Prachuap Khiri Khan
Province, on decaying wood submerged in a freshwater stream, 25 December 2014, Jaap van
Strien, Site 5-2-1, MFLU 15-1163, holotype, GZAAS 17-0013, isotype), ex-type living cultures
MFLUCC 15-0628, GZCC 15-0063). a Colonies on natural substrate. b Conidiogenous cells. c, d
Conidiogenous cells with conidia. e, f Conidiophores with conidia. g-j Conidia. Scale bars: a = 500
μm, b-d = 20 μm, e, f = 100 μm, g-j = 15 μm.
Sterigmatobotrys Oudem., Ned. kruidk. Archf, 2 sér. 4: 548 (1886)
Index Fungorum number: IF10090; 4 morphological species (Species Fungorum 2020), 3
species with sequence data.
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Type species – Sterigmatobotrys elatus (Sacc.) Oudem.
Notes – The genus is characterised by penicillately branched, brown conidiophores,
sympodial conidiogenous cells without scars or denticles and septate, hyaline or brown, slimy
conidia (Seifert et al. 2011). It differs from other genera in the family by its penicillately branched
conidiophores. Réblová & Seifert (2011) reported that the sexual morph of S. macrocarpa is similar
to species in Carpoligna and Chaetosphaeria.
Podosporaceae X. Wei Wang & Houbraken, Stud. Mycol. (2019)
Index Fungorum number: IF829841; Facesoffungi number: FoF06877; 92 species.
Saprobic on dung. Sexual morph: Ascomata superficial to immersed in medium, solitary or
loosely aggregated, ostiolate and ovoid to obpyriform, or lacking ostioles, globose to subglobose,
glabrous or having hypha-like to seta-like hairs. Peridium membranaceous to coriaceous, usually
opaque, in some species semi-translucent. Asci (2–)4- or 8- or multi-spored, unitunicate, cylindrical
to elongated clavate or fusiform, pedicellate, with or without an apical thickened ring, evanescent
or persistent until ascospores mature. Ascospores 1–2-seriate, 1-celled and pigmented, or 2-celled
and composed of a larger, pigmented upper cell and a smaller, pale or hyaline cell, with or without
appendage, usually smooth, in a few species ornamented. Asexual morph: Hyphomycetous. not
observed or cladorrhinum-like: Conidiophores micronematous, reduced to conidiogenous cells.
Conidiogenous cells intercalary or occasionally terminal, originating lateral or terminal peg-like
structure with a flaring collarette, with blastic conidia. Conidia single-celled, hyaline, smooth,
usually with a truncated base and a rounded apex (adapted from Wang et al. 2019a).
Type genus – Podospora Ces.
Notes – Podosporaceae was invalidly introduced by Hochberzanke (1930). The proposed
Podosporaceae introduced by Wang et al. (2019a) is based on Podospora, but they are delimited
based on morphology and DNA sequence data. Podosporaceae is sister to Chaetomiaceae in
Sordariales and accommodates Podospora, Trangularia and Cladorrhinum, which were positioned
in the polyphyletic family Lasiosphaeriaceae (Wang et al. 2019a). Based on the available sequences
of Apiosordaria, we consider this genus should be placed in Podosporaceae (Fig. 23). However,
Mouchacca & Gams (1993) indicated that several Apiosordaria species have asexual morph
connections with Cladorrhinum. Further work is required to determine the additional taxa in this
family.
Ecological and economic significance of Podosporaceae
Members of Podosporaceae mostly grow on dung of herbivore as saprobes and some are
endophytes (Cladorrhinum). Some Cladorrhinum species have activity against plant pathogenic
fungi and have potential for industrial and biotechnological applications (Martin et al. 2019).
Genera included in Podosporaceae
Cladorrhinum Sacc. & Marchal, Bull. Soc. Roy. Bot. Belgique 24: 64. (1885)
Index Fungorum number: IF7678; 11 morphological species (Species Fungorum 2020), 9
species with sequence data.
Type species – Cladorrhinum foecundissimum Sacc. & Marchal
Notes – Cladorrhinum was introduced for asexually reproducing species in 1885 by Saccardo
& Marchal, and the sexual morphs have been linked to Apiosordaria and Cercophora (Mouchacca
& Gams 1993). The species with Cladorrhinum asexual morphs, Apiosordaria verruculosa,
Cercophora samala, C. striata, and Podospora fimiseda, are phylogenetically closely related (Cai
et al. 2006b). Two species from Thielavia (Th. hyalocarpa and Th. intermedia) were placed in
Cladorrhinum based on their phylogenetic affinities with the type species of this genus. Therefore,
this genus is re-defined to accommodate sexually producing species. Phylogenetic evidence
indicated that the morphologically defined Cladorrhinum is polyphyletic (Carmarán et al. 2015).
Based on the phylogenetic analysis (Wang et al. 2019a), Cladorrhinum bulbillosum and Clad.
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phialophoroides, belong to the redefined genera Podospora and Triangularia, respectively. Further
work is needed to delimit the phylogenetic position of the Cladorrhinum species.
Cladorrhinum is found in soil as saprobes on dung or plant material (Lewis & Larkin 1998,
Madrid et al. 2011), or in roots as endophytes (Gasoni & Stegman de Gurfinkel 1997). Also, C.
foecundissimum and some species have activity against damping-off of the various crops caused by
Rhizoctonia solani and Pythium ultimum (Domsch & Gams 1968, Lewis & Larkin 1998, Gasoni &
Gurfinkel 2009).
Podospora Ces., Hedwigia 1(15): 103 (1856)
Index Fungorum number: IF4284; 72 morphological species, 2 species with sequence data
(Wang et al. 2019a).
Type species – Podospora fimiseda (Ces. & De Not.) Niessl
Notes – Podospora is one of the most common of coprophilous ascomycetes genera
worldwide, growing on herbivore dung (Doveri 2008b). Podospora is common in the temperate
regions but is rare in the tropics (Lundqvist 1972). This genus has over 100 species epithets. The
morphologically defined Podospora species are polyphyletic and distributed over at least 7 genericor higher-level clades based on rpb2 phylogeny in Wang et al. (2019a). Among the species in
Wang et al. (2019a), based on the phylogenetic analysis, only the type species and P. bulbillosum
are maintained in Podospora sensu stricto (Wang et al. 2019a). Further work is required to
determine the additional taxa in the genus.
Figure 205 – Podospora fimiseda (redrawn from Cesati 1856). a, b Ascomata. c Immature ascus
with paraphyses. d Asci. e Ascomal hairs. f Ascospores.
Triangularia Boedijn, Ann. Mycol. 32: 302 (1934)
Index Fungorum number: IF5534; 14 morphological species (Wang et al. 2019a), 9 species
with sequence data.
Type species – Triangularia bambusae (J.F.H. Beyma) Boedijn
Notes – Triangularia was introduced by Boedijn and defined by cylindrical or clavate asci
with a thickened apical ring and 2-celled smooth ascospores, with a larger, pigmented and conical
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to triangular upper cell and a smaller, paler or hyaline and triangular to hemisphaerical lower cell,
without gelatinous appendages (Guarro & Gene 1988). However, phylogenetic analysis in Wang et
al. (2019a) did not support the morphologically defined genus concept of Triangularia. Therefore,
this genus is re-defined which includes morphologically diverse species. Triangularia species are
usually isolated from soil and plant debris and occasional from aquatic sediments, generally from
warm places (Guarro & Cano 1988).
Polystigmataceae Höhn., Nova Acta R. Soc. Scient. upsal., Ser. 4 8(no. 2): 51 (1932)
Index Fungorum number: IF81595; Facesoffungi number: FoF03518; 20 species.
Parasitic on deciduous living leaves and shoots of Rosaceae. Stromata typically developing
in late spring and summer on living leaves, producing conidia in summer and autumn, and
ascospores from fallen overwintered leaves the following spring, irregular, bright red or orange,
surrounded apparently by healthy leaf tissue. Sexual morph: Ascostromata usually roughly circular
and covering a large area of the leaf, significantly raising the adaxial surface of the leaf, reddishbrown to black, the ostioles occasionally conspicuous. Ascomata sphaerical, immersed, distinct
walls, thin-walled. Paraphyses thinly dispersed, gradually tapering towards the apex, very thinwalled, strongly inflated between the septa. Asci 8-spored, unitunicate, clavate, long-pedicellate,
thin-walled in every stage, the apex obtuse, with an apical ring. Ascospores biseriate, cylindrical to
ellipsoidal or obovoid, occasionally slightly curved (fabiform), hyaline, aseptate, with or without
gelatinous sheath. Asexual morph: Conidial stromata irregular, yellowish-brown in very young
lesions, changing to orange to reddish-brown to black on maturity, occasionally causing small
creases on the leaves but without apparent significant hypertrophic lesion, composed of an upper
layer of epidermal cells filled with bright orange-brown material, a middle layer of hyaline fungal
cells which are angular to vertically elongated, and the lower layer resembling the upper one.
Conidiomata sphaerical, epigenous or hologenous, the ostiole unnoticeable and whose quantity
corresponds to the number of conidiomata, papillate or apapillate. Conidiomatal wall poorly
developed and not clearly distinguishable, composed of thin layer of small hyaline thick-walled
textura angularis. Conidiogenous cells nearly cylindrical, narrowing towards the upper region,
which appears somewhat irregular as a result of successive conidial scars, undergoing sympodial
proliferation, spreading the entire conidiomatal wall inner surface, emerging as lateral or terminal
branches from short and relatively undifferentiated conidiophores. Conidia hyaline, aseptate,
apparently smooth-walled, widest nearly at the base which is lanceolate to fusiform and truncate,
the upper part filiform, sigmoidally curved (adapted from Cannon 1996, Dayarathne et al. 2017).
Type genus – Polystigma DC.
Notes – Polystigmataceae was earlier considered a synonym of Phyllachoraceae and,
consequently, all taxa belonging to Phyllachoraceae were accommodated in Polystigmataceae
(Dennis 1968, Müller & von Arx 1973). Polystigmataceae was later raised to Polystigmatales (syn.
Phyllachorales) and this order accommodated Phyllachoraceae as the sole family (Hawksworth et
al. 1983, Mehrotra & Aneja 1990). This led to all Polystigma species then being placed in
Phyllachoraceae (Hawksworth et al. 1983, Mehrotra & Aneja 1990). Phylogenetic analyses
involving DNA sequence data for Polystigma amygdalinumis, however, showed that Polystigma
grouped in the subclass Xylariomycetidae, where it diverged from the Xylariales around 90 Million
years ago in the late Cretaceous (Habibi et al. 2015, Habibi & Banihashemi 2017). Therefore, it
cannot be accommodated in Phyllachorales (Sordariomycetidae) (Habibi et al. 2015, Habibi &
Banihashemi 2017). Phylogenetic analyses by Dayarathne et al. (2017) confirm that taxa of
Polystigma are phylogenetically distant from the Phyllachorales, and belong to Xylariales. They
have thus re-established Polystigmataceae in the subclass Xylariomycetidae (Dayarathne et al.
2017).
Ecological and economic significance of Polystigmataceae
Several species of Polystigma are foliicolous parasites on Prunus species (Suzuki et al. 2008).
Polystigma amygdalinum is often responsible for red blotch of almonds, while P. rubrum causes
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red leaf spot of plum (Habibi et al. 2015). Borovinova (2001) observed that, among the 16 plum
cultivars in the Institute of Agriculture in Kyustendil, Bulgaria (during the period 1997-1999), the
cultivars ‘Green gage’ and ‘Cacanska najbolja’ were most susceptible to infection from P. rubrum,
while cultivars ‘Lyubimtsa Hramova’, ‘Ashatan’, ‘Sofia-2’, ‘Gilej’, ‘Strinava’ and ‘Stanley plum’
were the least affected (Borovinova 2001).
Genus included in Polystigmataceae
Polystigma DC., Fl. franç., Edn 3 (Paris) 5/6: 164 (1815)
Index Fungorum number: IF4324; 20 morphological species (Species Fungorum 2020), 3
species with sequence data.
Type species – Polystigma rubrum (Pers.) DC.
Notes – Polystigma is the type genus of Polystigmataceae, established by Lamarck & de
Candolle (1815). It is a small genus, parasitic mainly to Prunus, with almost all of the species
reported with both asexual and sexual morphs (Cannon 1996). The generic name of Polystigma has
been recommended for use instead of the usually associated additional names, Polystigmina and
Rhodoseptoria, since it contains a greater number of species. Five taxa of Polystigmina are already
synonyms of Polystigma rubrum while the type species of Rhodoseptoria is synonym of
Polystigma rubrum (Réblová et al. 2016b). Suzuki et al. (2008) reported Polystigmina pallescens as
the asexual state of Polystigma fulvum.
Prosopidicolaceae Senan. & K.D. Hyde, Stud. Mycol. 86: 281 (2017)
Index Fungorum number: IF821565; Facesoffungi number: FoF03502; 2 species.
Pathogenic on species of Fabaceae. Sexual morph: Undetermined. Asexual morph:
Coelomycetous. Conidiomata pycnidial, rarely acervular, solitary or aggregated in a eustromatic
stroma with one to several ostioles or astromatic, grey to black, erumpent to immersed. Peridium
comprising grey-brown cells of textura angularis. Conidiophores reduced to conidiogenous cells or
lining the whole inner layer of the wall, subcylindrical, branched, septate, straight to irregularly
curved, base pale brown, becoming medium green-brown at apex. Conidiogenous cells mono- to
polyphialidic, tightly aggregated, hyaline, smooth, ampulliform, subcylindrical to lageniform,
prominent periclinal thickening, at times with percurrent proliferation. Conidia solitary, subhyaline
to grey-brown, smooth, guttulate, straight to variously curved, ellipsoid to fusoid-ellipsoid, apex
obtuse, base truncate to bluntly round (adapted from Senanayake et al. 2017a).
Type genus – Prosopidicola Crous & C.L. Lennox
Notes – Prosopidicolaceae was introduced by Senanayake et al. (2017a) for Prosopidicola
species. The sexual morph has not been reported for this family and it is distinct from other families
of Diaporthales in having pycnidial, rarely acervular, eustromatic conidiomata with one to several
ostioles, mono- to polyphialidic, ampulliform, subcylindrical to lageniform conidiogenous cells
with periclinal thickening and subhyaline to brown, ellipsoid to fusoid-ellipsoid conidia.
Ecological and economic significance of Prosopidicolaceae
Prosopidicola mexicana is the causative agent of pod disease of Prosopis glandulosa in
Mexico and the USA (Lennox et al. 2004). Prosopidicola albizziae causes stem cankers on
Albizzia falcataria and has been reported from leaves of Albizzia (Crous et al. 2016b).
Genus included in Prosopidicolaceae
Prosopidicola Crous & C.L. Lennox, Stud. Mycol. 50(1): 187 (2004)
Index Fungorum number: IF500048; 2 species with sequence data.
Type species – Prosopidicola mexicana Crous & C.L. Lennox
Notes – Prosopidicola albizziae is the second species of this genus (Lennox et al. 2004,
Crous et al. 2016b). Prosopidicola mexicana forms pod disease of Prosopis glandulosa and it has
been proposed as a biocontrol agent (Lennox et al. 2004).
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Figure 206 – Polystigma fulvum (Material examined – BELARUS, Minsk region, Maladzyechna
District, Sychevichi Village, near children's health camp “Brigantina”, deciduous forest on Prunus
padus (Rosaceae), 15 Oct. 2017, A.K. Khramtsov T-2083, MFLU 18-0261). a Swollen leaf spots on
Prunus padus L. b Close-up of ascomata. c Vertical section through ascoma. d Close-up of
inconspicuous ostiole. e Peridium of ascoma. f Asci. g Ascospore. h Faint sheath around ascospore.
Scale bars: a = 200 μm, c = 50 μm, d, f = 50 μm, e = 20 μm, g, h = 5 μm.
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Figure 207 – Polystigma rubrum (Material examined – RUSSIA, Rostov region, Shakhty City,
Cotton fabric microdistrict, torn shrubs near Grushevka river on Prunus stepposa Kotov., 9 October
2017, Timur S. Bulgakov, T-2093, MFLU 18-0271). a Swollen leaf spots on Prunus stepposa K. b
Close-up of leaf spots. c Vertical section through conidioma. d Conidiogenous cells and developing
conidia. e Conidiomatal wall. f-h Conidia. Scale bars: a = 200 μm, c = 50 μm, d-h = 10 μm.
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Figure 208 – Prosopidicola mexicana (redrawn from Lennox et al. 2004). a, b Disease symptoms
associated with Prosopidicola mexicana on pods of Prosopis glandulosa. c Cross section of
conidioma. d Conidiophores, conidiogenous cells and conidia. Scale bars: 10 µm.
Pseudodactylariaceae Crous, Persoonia 39: 421 (2017)
Index Fungorum number: IF823469; Facesoffungi number: FoF05318; 2 species.
Saprobic on plants. Sexual morph: Undetermined. Asexual morph: Conidiophores hyaline,
subcylindrical, straight to flexuous, unbranched or branched, thick-walled, septate.
Conidiogenous cells terminal, integrated, subcylindrical, denticulate. Conidia solitary,
aggregating in slimy mass, fusoid-ellipsoid, hyaline, smooth-walled, guttulate, septate,
surrounded by a thin mucilaginous sheath (adapted from Crous et al. 2017a).
Type genus – Pseudodactylaria Crous
Notes – A new order Pseudodactylariales and new family Pseudodactylariaceae were
introduced by Crous et al. (2017a) to accommodate a single genus Pseudodactylaria.
Pseudodactylariales and the new family Pseudodactylariaceae differs from other orders and
families as Pseudodactylaria species have 1-septate conidia encased in a mucoid sheath (Crous et
al. 2017a). Presently, one genus (Pseudodactylaria) with three species (P. brevis, P. hyalotunicata
and P. xanthorrhoeae) are accepted in Pseudodactylariaceae (Crous et al. 2017a, Lin et al. 2018).
Ecological and economic significance of Pseudodactylariaceae
All of the three accepted species are saprobic and thus are important in nutrient recycling.
Genus included in Pseudodactylariaceae
Pseudodactylaria Crous, Persoonia 39: 421 (2017)
Index Fungorum number: IF823411; 2 species with sequence data.
Type species – Pseudodactylaria xanthorrhoeae Crous
Notes – Pseudodactylaria was introduced by Crous et al. (2017a) for two dactylaria-like
species, P. hyalotunicata and P. xanthorrhoeae, in the subclass Sordariomycetidae and class
822
Sordariomycetes (Fig. 22). The genus is characterised by distinct, single, hyaline, unbranched and
septate conidiophores, terminal, integrated, polyblastic, denticulate conidiogenous cells and
solitary, fusoid-ellipsoid, hyaline conidia (Lin et al. 2018). Pseudodactylaria brevis is illustrated
below.
Figure 209 – Pseudodactylaria brevis (Material examined – THAILAND, Krabi, Wat Thum Sua,
on decaying wood, 15 December 2015, S. Tibpromma M 1-5, MFLU 18-1038; HKAS 102202,
paratype; living culture MFLUCC 16-0034). a Conidiophores on the host surface. b, c
Conidiophores, conidiogenous cells with denticles and conidia. d, e Conidiogenous cells with
denticles. f-i Conidia. Scale bars: b, c = 10 μm, d-i = 5 μm.
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Pseudohalonectriaceae Hongsanan & K.D. Hyde, Fungal Divers. 84:33 (2017)
Index Fungorum number: IF553215; Facesoffungi number: FoF03355; 14 species.
Saprobic on wood and other plant material, commonly isolated in marine and terrestrial or
freshwater habitats. Sexual morph: Ascomata erumpent to immerse with a protruding neck,
cylindrical, periphysate necks, greenish yellow, bright yellow to brown. Neck conical, composed of
parallel hyphae, outer hyphae outwardly directed, subglobose with enlarged ends, greenish yellow,
periphysate. Peridium multi-layered. Paraphyses tapering towards the apex, thin-walled, attached
to ascogenous hyphae. Asci 8-spored, unitunicate, cylindrical to clavate, with a J-, thimble-shaped,
refractive apical ring. Ascospores overlapping uniseriate to biseriate, hyaline to slightly coloured
and pale brown, pink/orange in mass in some species, cylindrical or ellipsoidal, straight to curved,
usually multi-septate, constricted or not-constricted at the septa, smooth-walled. Asexual morph:
Undetermined (adapted from Shearer & Zare-Maivan 1988, Hyde et al. 1999b).
Type genus – Pseudohalonectria Minoura & T. Muroi
Notes – Pseudohalonectriaceae was introduced in Magnaporthales by Hongsanan et al. (2017)
and comprises a single genus. Pseudohalonectria was previously placed in Lasiosphaeriaceae,
Sordariales supported by the preliminary phylogenetic analysis of SSU sequence data by Chen et
al. (1999). Shearer et al. (1999) transferred Pseudohalonectria to Magnaporthaceae. Klaubauf et al.
(2014) found that Pseudohalonectria clustered with species of Magnaporthaceae, Pyriculariaceae
and Ophioceraceae in the Magnaporthales in their phylogenetic study of Pyriculariaceae. Similar
results were obtained by Maharachchikumbura et al. (2015, 2016b), and they maintained
Pseudohalonectria in Magnaporthales genera, incertae sedis.
Ecological and economic significance of Pseudohalonectriaceae
Most species of Pseudohalonectria have antagonistic activity against other fungi and
bacteria. Asthana & Shearer (1990) tested paired cultures on agar against representatives of
Pseudohalonectria and showed that most tested species were strongly inhibited at a distance.
Pseudohalonectria adversaria exhibited potent nematicidal activity against pine wood nematodes
(Dong et al. 2004). Pseudohalonectria adversaria produce azaphilone compounds which are active
against several pests, weeds, nematodes, bacteria and fungi (Dong et al. 2006). The azaphilone
compounds can be developed as effective and alternative natural pest management products
(Foremska et al. 1992, Park et al. 2005, Dong et al. 2006). Pseudohalonectria species also degrade
lignocellulose and are involved in nutrient cycling.
Genus included in Pseudohalonectriaceae
Pseudohalonectria Minoura & T. Muroi, Trans. Mycol. Soc. Japan 19(2): 132 (1978)
Index Fungorum number: IF4437; 14 morphological species (Species Fungorum 2020), 5
species with sequence data.
Type species – Pseudohalonectria lignicola Minoura & T. Muroi
Notes – Pseudohalonectria was introduced to accommodate P. lignicola (Minoura & Muroi
1978), without being assigned to an order or family (Minoura & Muroi 1978). Hongsanan et al.
(2017) reviewed this genus and established a new family Pseudohalonectriaceae to accommodate
this genus in Magnaporthales. The genus is characterized by bright yellow to brown ascomata, with
long cylindrical, periphysate necks; unitunicate, cylindrical to clavate asci, with a J-, thimbleshaped, refractive apical rings and cylindrical, hyaline to slightly pigmented, and usually septate
ascospores (Shearer 1989). Members in Pseudohalonectria are found in marine and terrestrial or
freshwater habitats (Perera et al. 2016a). In this study, Pseudohalonectria lutea is illustrated.
Pseudomassariaceae Senan., Maharachch. & K.D. Hyde, Fungal Divers. 73: 132 (2015)
Index Fungorum number: IF551208; Facesoffungi number: FoF00843; 42 species.
Saprobic on recently dead twigs attached to the trees, immersed below rounded, slightly
elevated epidermis of the host. Sexual morph: Ascomata perithecial, scattered, solitary or-
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Figure 210 – Pseudohalonectria lutea (Material examined – CHINA, Tibet Autonomous Region,
on submerged decaying wood, 28 April 2015, Z.L. Luo, XZ A 2–2–1, S-1048, MFLU 18–1463;
living culture, MFLUCC 18–1297.). a, b Ascomata necks on decaying wood. c, d Sections of
ascomata. e Peridium. f-h Asci. i Paraphyses. j Ascal apical ring in side view. k-m Ascospores. n
Germinating ascospore. o, p Culture on PDA. Scale bars: c, d = 150 μm, e, i, j, n = 20 μm, f, g = 40
μm, i = 45 μm, k-m = 25 μm.
-aggregated, immersed, depressed globose to ellipsoid, coriaceous, black, ostiolate, papillate.
Ostiolar papilla short, cylindrical, wide at the apex, periphysate. Peridium comprising strongly
compressed, narrow, light to medium brown, thin-walled cells of textura angularis. Paraphyses
numerous, apically narrow, basally wide, hyaline. Asci 8-spored, unitunicate, easily disintegrated
when fresh, clavate to fusoid, short pedicellate, apically rounded, with J+, or J-, apical ring.
Ascospores biseriate or partially uniseriate, hyaline, broadly ellipsoid, oblong or narrowly clavate,
mostly apiosporous, with a rounded to subconical, small, lower cell or equally uniseptate, straight
or curved, thick-walled, smooth-walled. Asexual morph: Hyphomycetous. Setae erect, dark brown,
straight to flexuous, arising from superficial hyphae, branched at base, subcylindrical, tapering to
825
obtuse apex, 3–7-septate, basal cell slightly swollen. Conidiophores subcylindrical to setiform, with
radially lobed basal cells, pale brown, smooth, arising from superficial mycelium, straight to
flexuous, 1–10 septate. Conidiogenous cells terminal or lateral, polyblastic, subcylindrical to
somewhat clavate, pale brown, smooth, with 1–4 denticulate loci. Conidia lageniform, distal end
free, truncate, rostrate at proximal end, pale brown, with a subhyaline transverse band at equatorial
zone, smooth, guttulate, aseptate (adapted from Senanayake et al. 2015, Jaklitsch et al. 2016b).
Type genus – Pseudomassaria Jacz.
Notes – Jaklitsch & Voglmayr (2012) showed a separate taxonomic clade that included
Pseudomassaria and Leiosphaerella through separate analyses of ITS and LSU sequence data.
With emphasis on combined ITS-LSU analyses and morphological similarities the clade was
introduced as a family Pseudomassariaceae in Senanayake et al. (2015). Maharachchikumbura et al.
(2016b) accepted Pseudomassaria with apiosporous ascospores and Leiosphaerella with elongate,
bi-celled ascospores. Jaklitsch et al. (2016b) treated Leiosphaerella, Pseudomassaria,
Pseudapiospora and Pseudomassariella in Pseudomassariaceae. However, Pseudomassariella has
been treated in this family with no phylogenetic support. Pseudomassariaceae commonly appears
with Apiosporaceae, Beltraniaceae, Hyponectriaceae and Melogrammataceae clades in
Amphisphaeriales in various phylogenetic placements (Senanayake et al. 2015, Konta et al. 2016,
Maharachchikumbura et al. 2016b, Samarakoon et al. 2016, Hongsanan et al. 2017, Wijayawardene
et al. 2018a). In this study, we accept Pseudomassariaceae in Amphisphaeriales comprising four
genera as Leiosphaerella, Pseudomassaria, Pseudapiospora and Pseudomassariella. Only ITS and
LSU sequences are available and revisions are required.
Ecological and economic significance of Pseudomassariaceae
Pseudomassariaceae species are common as saprobes on recently dead twigs and leaves or
branches in terrestrial habitats (Barr 1964, Senanayake et al. 2015). The leaf-inhabiting species are
commonly recorded in overwintered leaves and rarely as parasites on dead spots on leaves i.e.
Pseudomassaria leucothoës has been reported in leaf spots of Leucothoë sp. (Barr 1964).
Genera included in Pseudomassariaceae
Leiosphaerella Höhn., Sber. Akad. Wiss. Wien, Math.-naturw. Kl., Abt. 1 128(7-8): 579 (1919)
Index Fungorum number: IF2717; 14 morphological species (Jaklitsch & Voglmayr 2012), 3
species with sequence data.
Type species – Leiosphaerella praeclara (Rehm) Höhn.
Notes – Leiosphaerella species are characterized by immersed perithecial ascomata with
apically free paraphyses, asci with J+, apical rings, and elongate, bicelled, hyaline ascospores. Barr
(1990b) treated the genus in Hyponectriaceae. Jaklitsch & Voglmayr (2012) revisited the genus and
placed it in Pseudomassariaceae.
Leiosphaerella praeclara is typified Leiosphaerella previously placed in Xylariales genera
incertae sedis and are associated with Vaccinium species (Ericaceae). Jaklitsch & Voglmayr (2012)
provided an epitype with living culture and sequence data. Combined LSU-ITS phylogenetic
analyses revealed that the Leiosphaerella belongs in Pseudomassariaceae (Senanayake et al. 2015,
Jaklitsch et al. 2016b). Hyde (1993d) described the similar genera to Leiosphaerella with the shape
of ascomata, asci, ascospores and J+/J- reactions of the apical ring. The genera Leiosphaerella,
Marinospora and Oxydothis share similar characters of cylindic-clavate asci, J+, apical ring and 2celled ascospores (Hyde 1993d). However, Marinospora is only known from marine habitats and
phylogenetically placed in Halosphaeriaceae (Sakayaroj et al. 2011), while Konta et al. (2016)
introduced Oxydothidaceae to accommodate Oxydothis. Leiosphaerella praeclara is illustrated in
this entry (Fig. 211).
Pseudapiospora Petr., Hedwigia 68: 233 (1928)
Index Fungorum number: IF4404; 3 morphological species (Jaklitsch & Voglmayr 2012), 1
species with sequence data.
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Type species – Pseudapiospora corni (Sowerby) Petr.
Notes – Senanayake et al. (2015) treated P. corni under Pseudomassaria corni. However,
Jaklitsch et al. (2016b) re-established Pseudapiospora corni, based on known host-specificity and
septum characterization.
Figure 211 – Leiosphaerella praeclara (redrawn from Jaklitsch & Voglmayr 2012). a Ostioles and
perithecial dots on bark surface. b Cross section of ascoma. c Peridium. d Apical ascus ring. e-g
Ascospores. h Ascus. Scale bars: a = 150 μm, b = 300 μm, c = 10 μm, e-h = 10 μm.
Pseudomassaria Jacz., Bull. Herb. Boissier 2: 663 (1894)
Index Fungorum number: IF4452; 24 morphological species (Species Fungorum 2020), 2
species with sequence data.
Type species – Pseudomassaria chondrospora (Ces.) Jacz.
Notes – Pseudomassaria was earlier accepted in Hyponectriaceae and Jaklitsch & Voglmayr
(2012) and Senanayake et al. (2015) showed the taxonomic placement of Pseudomassaria in
Pseudomassariaceae.
Pseudomassariella Petr., Sydowia 9(1-6): 602 (1955)
Index Fungorum number: IF4453; 1 species with sequence data.
Type species – Pseudomassariella vexata (Sacc.) Petr.
Notes – Petrak (1955b) erected Pseudomassariella as closely related to Pseudomassaria with
the type P. vexata. Müller & von Arx (1962) transferred the species to Leiosphaerella. Jaklitsch et
al. (2016b) provided taxonomic any phylogenetic revisions and provisionally accepted the genus in
Pseudomassariaceae.
Pseudomelanconidaceae C.M. Tian & X.L. Fan, Persoonia 40: 128 (2018)
Index Fungorum number: IF823991; Facesoffungi number: FoF04889; 2 species.
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Pathogenic forming cankers on trees. Sexual morph: Undetermined. Asexual morph:
Coelomycetous, melanconium-like. Conidiophores acervular, hyaline to pale brown, aseptate,
unbranched, cylindrical, smooth-walled. Conidiogenous cells annellidic, occasionally with distinct
annellations and collarettes. Conidia hyaline to greyish sepia to olivaceous as it matures, ellipsoid
to oblong, multiguttulate, aseptate, with distinct hyaline sheath, becoming inconspicuous when
mature. Conidial wall smooth (adapted from Fan et al. 2018).
Type genus – Pseudomelanconis C.M. Tian & X.L. Fan
Notes – Pseudomelanconidaceae shares similar characteristics to other families in
Diaporthales having acervular conidiomata covered by a pustulate conidial mass on the bark
surface, similar to Juglanconidaceae, Melanconiellaceae and Melanconidaceae (Fan et al. 2016,
Voglmayr et al. 2017). However DNA sequence data confirmed them to represent a distinct
phylogenetic lineage. Molecular investigations have revealed a remarkably high diversity of
corticolous melanconium-like fungi in Diaporthales (Fan et al. 2016, Voglmayr et al. 2012, 2017),
thus Fan et al. (2018) intrduced Pseudomelanconidaceae. In phylogenetic analysis of ITS, LSU,
rpb2 and tef1 sequence data in Fan et al. (2018), Pseudomelanconis forms a distinct clade sister to
Melanconiellaceae (Senanayake et al. 2018). Jiang et al. (2018) introduced a new genus
Neopseudomelanconis in Pseudomelanconidaceae from dead branches of Castanea mollissima in
China.
Ecological and economic significance of Pseudomelanconidaceae
Pseudomelanconis caryae causes branch canker and dieback of hickory trees, Carya
cathayensis, a popular nut food tree species mainly distributed in southeastern China (Wu et al.
2014a, Fan et al. 2018).
Genera included in Pseudomelanconidaceae
Neopseudomelanconis C.M. Tian & N. Jiang, Mycosphere 9: 1273 (2018)
Index Fungorum number: IF825183; 1 species with sequence data.
Type species – Neopseudomelanconis castaneae C.M. Tian & N. Jiang
Notes – The genus was introduced to accommodate Neopseudomelanconis castaneae, a
species on dead branches of Castanea mollissima in China (Jiang et al. 2018). The new genus was
based on morphology and phylogenetic evidence of ITS and LSU genes (Jiang et al. 2018).
Neopseudomelanconis was reported to have acervular conidiomata, cylindrical and hyaline
conidiophores, conidiogenous cells with apical collaretes and discreet annellations, and ellipsoid to
oblong, 2-celled, hyaline to brown mature conidia with a hyaline conidial sheath (Jiang et al. 2018).
Pseudomelanconis C.M. Tian & X.L. Fan, Persoonia 40: 128 (2018)
Index Fungorum number: IF823992; 1 species with sequence data.
Type species – Pseudomelanconis caryae C.M. Tian & X.L. Fan
Notes – The monospecific genus Pseudomelanconis has acervular conidiomata covered by
pustulate conidial masses on the bark surface. This genus is significantly different from other taxa
due to its distinct conidiogenous cells with apical collarettes showing discrete annellations. The
conidial sheath becomes inconspicuous when mature (Fan et al. 2018).
Pseudoplagiostomataceae Cheew., M.J. Wingf. & Crous, Fungal Divers. 44: 95 (2010)
Index Fungorum number: IF542097; Facesoffungi number: FoF01403; 7 species.
Pathogen on leaves, forming spots. Sexual morph: Ascomata solitary, immersed, scattered,
slanted to horizontal on host tissue, globose or ellipsoidal, black, coriaceous, papillate, ostiolate.
Papilla short, internally covered with hyaline, periphyses. Peridium composed of few layers of
brown, thick-walled cells of textura angularis. Paraphyses lacking. Asci 8-spored, unitunicate,
apedicellate, cylindrical, with J-, subapical ring. Ascospores overlapping uniseriate to biseriate,
hyaline, fusiform to ellipsoid, 1-septate, with terminal, elongate, hyaline appendages. Asexual
morph: Coelomycetous. Conidiomata brown, acervular or pycnidial. Peridium comprising small,
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brown cells of textura angularis. Conidiophores reduced to conidiogenous cells. Conidiogenous
cells enteroblastic, cylindrical to ampulliform, percurrently proliferating with periclinal
thickening and collarette. Conidia unicellular, ellipsoid, hyaline to brown, subglobose to broadlyallantoid, with obtuse apex and a flat protruding scar at the base (adapted from Cheewangkoon et
al. 2010).
Type genus – Pseudoplagiostoma Cheew. Cheew., M.J. Wingf. & Crous
Notes – Pseudoplagiostomataceae is distinct from other related families due to its unique
morphology with ascomata slanted horizontally to the host with an aparaphysate hamathecium and
ascospores with terminal, elongate, hyaline appendages. It has a distinct placement in Diaporthales
based on the maximum parsimony analysis of LSU sequence data (Cheewangkoon et al. 2010).
This placement was confirmed by the studies of Suwannarach et al. (2016), Du et al. (2017) and
Phookamsak et al. (2019).
Ecological and economic significance of Pseudoplagiostomataceae
This monotypic family comprises foliar pathogens recorded from Eucalyptus and Mangifera
and an endophyte on Dipterocarpus tuberculatus. This family produces leaf spot and shoot blight
diseases which can become a problem for eucalyptus plantations, especially in the case of P.
eucalypti (Lueangpraplut et al. 2013). Moreover, the species P. mangiferae is also associated with
leaf blight of Mangifera sp. (Phookamsak et al. 2019).
Genus included in Pseudoplagiostomataceae
Pseudoplagiostoma Cheew., M.J. Wingf. & Crous, Fungal Divers. 44: 96 (2010)
Index Fungorum number: IF516496; 7 species with sequence data.
Type species – Pseudoplagiostoma eucalypti Cheew., M.J. Wingf. & Crous
Notes – Pseudoplagiostoma is a foliar pathogen of Eucalyptus with astromatic, slanted to
horizontal, globose perithecia lacking paraphyses, with aseptate ascospores with terminal, elongate,
hyaline appendages and a cryptosporiopsis-like asexual morph. Cryptosporiopsis eucalypti falls in
Diaporthales in the phylogenetic analyses using combined gene analysis of ITS and tub2 gene
sequence data which resolved the species in Pseudoplagiostoma (Cheewangkoon et al. 2010). A
newly added species is P. dipterocarpi, an asexual endophyte isolated from the leaves of
Dipterocarpus tuberculatus in Chiang Mai, Thailand (Suwannarach et al. 2016). The asexual
morph of Pseudoplagiostoma mangiferae isolated from a living leaf of Mangifera in Yunnan,
China was introduced by Phookamsak et al. (2019) based on the morphology and phylogenetic
analyses of a combined ITS, LSU, tub2 and tef1 dataset.
Pseudoproboscisporaceae H. Zhang, K.D. Hyde & Maharachch., Fungal Divers. 85: 93 (2017)
Index Fungorum number: IF553760; Facesoffungi number: FoF03338; 5 species.
Saprobic on decaying wood and bamboo submerged in freshwater. Sexual morph: Ascomata
scattered or solitary, immersed, with long or short neck, dark brown to black, ostiolate. Neck central
or lateral, dark brown, erect or curve upwards. Peridium comprising several layers of compressed
cells. Paraphyses hypha-like, septate, tapering. Asci 8-spored, unitunicate, cylindrical, pedicellate,
with a relatively large, J-, apical ring. Ascospores uni- to tri-seriate, hyaline, fusiform, aseptate or
septate, smooth or rugose, thin-walled, with bipolar filamentous appendages. Appendages initially
coiled then unfurling to form long threads on release. Asexual morph: Undetermined (adapted from
Zhang et al. 2017a).
Type genus – Pseudoproboscispora Punith.
Notes – Pseudoproboscisporaceae was introduced in a newly established order
Atractosporales by Zhang et al. (2017a) based on phylogenetic and morphology and included two
genera Diluviicola and Pseudoproboscispora. Cateractispora also clustered in
Pseudoproboscisporaceae in the phylogenetic tree (Zhang et al. 2017a), however, it has not been
validly published (Ranghoo et al. 1999). Zhang et al. (2017a) observed the photo-plate of C.
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recepticuli from the PhD thesis of Ranghoo (1998) and thought that Cateractispora was likely to be
a synonym of Pseudoproboscispora.
Figure 212 – Pseudomelanconis caryae (Material examined – CHINA, Zhejiang Province,
Hangzhou City, Linan, Tianmu Mountain, N30°18'48.85" E119°26'36.99", 288 m asl, on twigs and
branches of Carya cathayensis, 21 April 2017, Q. Yang & Z. Du, CF 2017466, holotype). a
Conidiomata in host. b-d Locules. e Conidiophores and conidiogenous cells. f-i Conidia. Scale
bars: a = 1 mm, b-d = 200 µm, e-h = 10 µm.
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Figure 213 – Pseudoplagiostoma mangiferae (Material examined – CHINA, Yunnan Province,
Xishuangbanna, Jinghong, Nabanhe, associated with leaf blight symtom on living leaf of
Mangifera sp. (Anacardiaceae), 21 November 2015, R. Phookamsak, XB010, KUN-HKAS
102244, holotype). a Leaf blight symptom on living leaf of Mangifera sp. b Conidiomata on host
surface. c, d Sections through conidiomata. e Conidioma wall and the base. f, g Conidiogenous
cells. h Paraphyses. i-k Conidia. Scale bars: c-e = 50 μm, f-h = 20 μm, j = 25 μm, i, k = 10 μm.
Ecological and economic significance of Pseudoproboscisporaceae
Pseudoproboscisporaceae currently comprises three species, one of which is invalidly
published (Ranghoo et al. 1999, Zhang et al. 2017a), isolated from decaying bamboo and are
important decomposers of bamboo in freshwater and involved in nutrient recycling.
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Genera included in Pseudoproboscisporaceae
Diluviicola K.D. Hyde, S.W. Wong & E.B.G. Jones, Fungal Divers. Res. Ser. 1: 141 (1998)
Index Fungorum number: IF24138; 2 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Diluviicola capensis K.D. Hyde, S.W. Wong & E.B.G. Jones
Notes – Diluviicola was introduced by Hyde et al. (1998b) to accommodate D. capensis from
submerged wood in freshwater in Brunei. Diluviicola aquatica was introduced by Zhang et al.
(2017a) from submerged bamboo in a small river in Thailand and considered to differ from the type
species, D. capensis, in having subglobose or ellipsoidal ascomata with dark brown necks and
mostly biseriate ascospores. Sequence data for D. capensis are not available.
Pseudoproboscispora Punith., Kew Bull. 54(1): 234 (1999)
Index Fungorum number: IF28391; 3 morphological species (Species Fungorum 2020), 2
species with sequence data.
Type species – Pseudoproboscispora aquatica (S.W. Wong & K.D. Hyde) Punith.
Notes – Proboscispora was introduced by Wong & Hyde (1999) to accommodate P. aquatica
which was collected on submerged wood in freshwater. It was renamed Pseudoproboscispora by
Punithalingam (1999). Pseudoproboscispora was placed in Annulatascaceae by
Maharachchikumbura et al. (2016b) and later transferred to a new family Pseudoproboscisporaceae
based on phylogenetic analysis (Zhang et al. 2017a). However, Pseudoproboscispora caudae-suis
clustered in Annulatascaceae which is distant from Pseudoproboscisporaceae (Zhang et al. 2017a).
Pseudosporidesmiaceae Crous, Persoonia 39: 365 (2017)
Index Fungorum number: IF823464; Facesoffungi number: FoF05493; 2 species.
On leaves of living plants. Sexual morph: Undetermined. Asexual morph: Hyphomycetous.
Mycelium consisting of branched, septate, pale brown hyphae. Conidiophores solitary or in clusters,
flexuous or erect, arising from superficial hyphae, rejuvenating percurrently, with cylindrical stipe,
brown, smooth, thick-walled, mostly unbranched. Conidiogenous cells terminal, cylindrical, brown,
scars truncate, not thickened. Conidia solitary, obclavate, brown, subobtuse at apex, truncate at
base, euseptate, smooth-walled (adapted from Crous et al. 2017a).
Type genus – Pseudosporidesmium K.D. Hyde & McKenzie
Notes – Pseudosporidesmiaceae, introduced by Crous et al. (2017a), included two species of
Pseudosporidesmium and a taxon tentatively identified as Repetophragma inflatum (GenBank
number DQ408576), of which the characters could not be confirmed. Pseudosporidesmiaceae is
assigned to Xylariales (Xylariomycetidae, Sordariomycetes) based on phylogenetic analysis (Su et
al. 2016b, Crous et al. 2017a).
Ecological and economic significance of Pseudosporidesmiaceae
Saprobic taxa having the ability to decompose lignocellulose in woody litter, softening the
wood and releasing nutrients as simple molecules that can be reused by plants and all other
organisms (Yuen et al. 1998, Bucher et al. 2004). Thus, they play an important role in nutrient and
carbon recycling, biological diversity and ecosystem functioning (Palmer et al. 1997, Wong et al.
1998a).
Genus included in Pseudosporidesmiaceae
Pseudosporidesmium K.D. Hyde & McKenzie, Fungal Divers. 80: 404 (2016)
Index Fungorum number: IF551835; 2 species with sequence data.
Type species – Pseudosporidesmium knawiae (Crous) K.D. Hyde & McKenzie
Notes – Pseudosporidesmium was established by Su et al. (2016b) to accommodate
Sporidesmium knawiae, which was introduced by Crous (2008). Presently, two species are accepted
in Sporidesmium and both were isolated from leaves of living plants (Crous 2008, Crous et al.
2017a).
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Figure 214 – Diluviicola aquatica (Material examined – THAILAND, Prachuap Khiri Khan, on
submerged bamboo in a small river, 30 July 2015, W. Dong 55A, MFLU 15-2701, holotype). a
Herbarium material. b Appearance of black ascomata semi-immersed or superficial on host. c
Vertical section of ascoma. d Structure of peridium. e Paraphyses. f, g Asci. h Asci apicalring, j, l,
m Ascospores mounted in water. k Ascospore mounted in cotton blue. j, k Appendages marked by
arrows. Scale bars: c = 50 μm, d-h, j-m = 20 μm, i = 5 μm.
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Figure 215 – Pseudosporidesmium knawiae (redrawn from Crous 2008). a, b Conidiophores with
conidia. c Conidiogenous cells with conidium. d, e Conidia. Scale bars: 10 μm.
Pseudotruncatellaceae Crous, Persoonia 42: 309 (2019)
Index Fungorum number: IF830823; Facesoffungi number: FoF06878; 2 species.
Saprobic on dead plant stems or associated with leaves. Asexual morph: coelomycetous.
Conidiomata acervular to pycnidioid, gregarious, immersed to semi-immersed, oval. Peridium of
hyaline to pale brown, thick-walled cells forming textura angularis. Conidiophores arising from
basal and lateral cells in cavity, cylindrical, septate, branched or rarely unbranched, sometimes
reduced to conidiogenous cells, hyaline, smooth-walled. Conidiogenous cells subcylindrical,
proliferating percurrently at apex, hyaline, smooth-walled. Conidia fusoid, straight, 2 or 3septate, with 2–3(–4) tubular apical appendages; two median cells guttulate, fusoid, dark brown,
thick-, smooth-walled; basal cell, narrowly obconic with a truncate base, hyaline, smooth-walled.
Sexual morph: Undetermined (adapted from Crous et al. 2019a).
Type genus – Pseudotruncatella R.H. Perera, Camporesi, Maharachch. & K.D. Hyde
Notes – Pseudotruncatellaceae was introduced by Crous et al. (2019a) to accommodate a
single genus Pseudotruncatella, which was previously placed in Amphisphaeriales genera incertae
sedis (Perera et al. 2018).
Ecological and economic significance of Pseudotruncatellaceae
Pseudotruncatella arezzoensis is a saprobe on woody substrates (Perera et al. 2018). Saprobic
taxa play an important role in nutrient and carbon cycling in ecosystems (Harley 1971).
Pseudotruncatella bolusanthi is associated with plant leaves of Bolusanthus speciosus (Crous et al.
2019a). However, no study has confirmed pathogenicity of Pseudotruncatellaceae species.
Genus included in Pseudotruncatellaceae
Pseudotruncatella R.H. Perera, Camporesi, Maharachch. & K.D. Hyde, Phytotaxa 338(2): 181
(2018)
Index Fungorum number: IF553932; 2 species with sequence data.
Type species – Pseudotruncatella arezzoensis .H. Perera, Camporesi, Maharachch. & K.D.
Hyde
Notes – Perera et al. (2018) established asexual morphic genus Pseudotruncatella to
accommodate P. arezzoensis. Pseudotruncatella bolusanthi was added to the genus and family
834
Pseudotruncatellaceae was introduced to accommodate them (Crous et al. 2019a). The genus is
characterized by acervular to pycnidioid conidiomata, holoblastic, annellidic, conidiogenous cells,
2 or 3-septate, fusiform to clavate conidia with up to 2–3(–4) apical appendages and dark brown
median cells (Perera et al. 2018, Crous et al. 2019a).
Figure 216 – Pseudotruncatella arezzoensis (Material examined – ITALY, Arezzo Province, Croce
di Pratomagno, aerial stem of Helichrysum italicum (Roth) G. Don fil. (Asteraceae), 18 July 2013,
Erio Camporesi, IT1322E, MFLU 17-1776, paratype). a Herbarium material. b, c Conidiomata on
host. d Vertical section of conidioma. e, f Conidiogenous cells with mature and immature conidia.
g-k Conidia. Scale bars: c = 500 µm, d = 200 µm, e-k = 10 µm.
Pyriculariaceae Klaubauf, M.-H. Lebrun & Crous, Stud. Mycol. 79: 104 (2014)
Index Fungorum number: IF810202; Facesoffungi number: FoF05054; 82 species.
Pathogenic or saprobic on various plant hosts, commonly on monocotyledons. Sexual morph:
Ascomata perithecial, immersed, solitary, scattered or gregarious, black, coriaceous, with long
cylindrical necks, covered in setae. Peridium brown to dark brown, comprising several layers of
textura angularis. Paraphyses septate, intermingled among asci. Asci 8-spored, unitunicate,
subcylindrical, short pedicellate, with a large, J+, apical ring. Ascospores overlapping 2–3-seriate,
ellipsoid to obclavate, fusiform, with 3 septa, often with median cells pigmented, pale brown.
Asexual morph: Hyphomycetous. Conidiophores solitary or in fascicles, subcylindrical, erect,
olivaceous, pale brown or brown, branched or unbranched. Conidiogenous cells holoblastic,
835
polyblastic, integrated, pigmented, denticulate. Conidia hyaline to brown, pyriform to elliptical, 1–
5-transversely septate, in some genera with apical mucoid appendages (adapted from Klaubauf et
al. 2014).
Type genus – Pyricularia Sacc.
Notes – Klaubauf et al. (2014) introduced Pyriculariaceae as a sister family to the
Ophioceraceae, and accepted two genera Deightoniella and Pyricularia in Pyriculariaceae, and
introduced seven genera (viz. Bambusicularia, Barretomyces, Macgarvieomyces, Neopyricularia,
Proxipyricularia, Pseudopyricularia, Xenopyricularia) based on morphology and phylogenetic
analyses. DNA replication licensing factor (MCM7) and calmodulin (calM) genes were used as a
phylogenetic marker for some genera by Pordel et al. (2015, 2017). Hernández-Restrepo et al.
(2015b) established Neocordana, a new genus in Pyriculariaceae. Wijayawardene et al. (2017a,
2018a) accepted ten genera in Pyriculariaceae. It is difficult to identify some of these genera in this
family without molecular data. We illustrate both morphs of Pyricularia with line diagrams and
describe and illustrate a new collection of Deightoniella africana.
Ecological and economic significance of Pyriculariaceae
Pyriculariaceae species cause serious crop diseases on a wide range of hosts, including major
diseases on monocotyledons (Klaubauf et al. 2014). Species of Pyricularia are important pathogens
in a species-rich genus, and sampling leaf spot diseases in Poaceae, which revealed many taxa
(Klaubauf et al. 2014).
Rice blast disease caused by P. oryzae has been found in more than 85 countries (Kato 2001).
Pyricularia grisea is responsible for a foliar disease of Digitaria (Klaubauf et al. 2014), and it also
causes disease on about 50 species of cereals, other grasses and sedges, including Oryza sativa,
Triticum aestivum, Hordeum vulgare, Zea mays, Avena sativa, Secale cereal, Eleusine corocana,
Lolium perenne and weeds and ornamental grasses (Ou 1985). Both P. grisea and P. oryzae cause
rice blast disease and infect all aerial parts of rice, leading to leaf-blast, neck and panicle rot, collar
rot and node blast (Skamnioti & Gurr 2009).
Genera included in Pyriculariaceae
Bambusicularia Klaubauf, Lebrun & Crous, Stud. Mycol. 79: 104 (2014)
Index Fungorum number: IF810203; 1 species with sequence data.
Type species – Bambusicularia brunnea Klaubauf, M.-H. Lebrun & Crous
Notes – The genus is a pathogenic hyphomycetous asexual morph (Klaubauf et al. 2014).
Barretomyces Klaubauf, Lebrun & Crous, Stud. Mycol. 79: 104 (2014)
Index Fungorum number: IF810205; 1 species with sequence data.
Type species – Barretomyces calatheae (D.J. Soares, F.B. Rocha & R.W. Barreto) Klaubauf,
Lebrun & Crous
Notes – The monotypic genus was introduced by Klaubauf et al. (2014) with B. calatheae, a
hyphomycetous species. It was recorded as a pathogen on leaves of Calathea longifolia in Brazil
(Maharachchikumbura et al. 2015b, Wijayawardene et al. 2017a, 2018a).
Deightoniella S. Hughes, Mycol. Pap. 48: 27 (1952)
Index Fungorum number: IF7934; 14 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Deightoniella africana S. Hughes
Notes – Deightoniella was introduced by Hughes (1952), who described D. africana on living
leaves of Imperata cylindrica var. africana in West Africa. The genus, which is worldwide in
distribution, is pathogenic and only a hyphomycetous asexual stage is known (see Seifert et al.
2011 for morphology). The phylogeny was introduced in Klaubauf et al. (2014) and an outline of
species, and nomenclature has been resolved for this genus (Maharachchikumbura et al. 2015b,
2016b, Zhang et al. 2016, Wijayawardene et al. 2017a, 2018a).
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Figure 217 – Deightoniella africana (Material examined – GHANA, Togoland, on leaves of
Imperata cylindrica var. africana, 28 May 1949, S.J. Hughes 913, IMI 39675(a), type). a-c
Herbarium packets and specimens. d Conidiophores on substrate. e Conidiophores. f, g
Conidiophores with conidia. h-l Conidia (h-k were using the same scale bar as l). Scale bars: c =
1000 µm, d = 100 µm, e-l = 20 µm.
Macgarvieomyces Klaubauf, Lebrun & Crous, Stud. Mycol. 79: 106 (2014)
Index Fungorum number: IF810207; 3 species with sequence data.
Type species – Macgarvieomyces borealis (de Hoog & Oorschot) Klaubauf, Lebrun & Crous
Notes – Macgarvieomyces was introduced to accommodate two species, M. borealis and M.
juncicola occurring on leaf spots and stem bases of Juncus, respectively. The species were based on
host plant and taxa cannot clearly be distinguished from other genera in the family (Klaubauf et al.
2014). However, molecular analyses distinguished a distinct clade sister to Deightoniella.
Pyricularia luzulae was synonymised as Macgarvieomyces luzulae (Marin-Felix et al. 2018).
837
Neocordana Hern.-Restr. &Crous, Phytotaxa 205(4): 233 (2015)
Index Fungorum number: IF811105; 7 morphological species (Species Fungorum 2020), 5
species with sequence data.
Type species – Neocordana musae (Zimm.) M. Hern.-Rest. & Crous
Notes – Neocordana was introduced by Hernández-Restrepo et al. (2015b) to accommodate
N. musae (type species), N. johnstonii, N. versicolor and N. musicola, based on morphology and
phylogenetic studies. The genus occurs on leaves of Musa spp. (Musaceae) and Canna denudata
(Cannaceae) (Hernández-Restrepo et al. 2015b). Neocordana musarum and N. musigena were
introduced with evidence from phylogenetic analysis and morphology (Crous et al. 2016a, 2017b).
Neopyricularia Klaubauf, Lebrun & Crous, Stud. Mycol. 79: 108 (2014)
Index Fungorum number: IF810210; 1 species with sequence data.
Type species – Neopyricularia commelinicola (M.J. Park & H.D. Shin) Klaubauf, Lebrun &
Crous
Notes – Klaubauf et al. (2014) introduced the monotypic genus Neopyricularia from leaves
of Commelina communis in South Korea (Maharachchikumbura et al. 2015b, 2016b,
Wijayawardene et al. 2017a, 2018a).
Proxipyricularia Klaubauf, Lebrun & Crous, Stud. Mycol. 79: 109 (2014)
Index Fungorum number: IF810211; 1 species with sequence data.
Type species – Proxipyricularia zingiberis (Y. Nisik.) Klaubauf, Lebrun & Crous
Notes – Klaubauf et al. (2014) introduced Proxipyricularia to accommodate the type species,
P. zingiberis, which is a pathogen on Zingiber mioga and Z. officinale in Japan. The sexual morph
is undetermined (Klaubauf et al. 2014).
Pseudopyricularia Klaubauf, Lebrun & Crous, Stud. Mycol. 79: 109 (2014)
Index Fungorum number: IF810213; 8 species with sequence data.
Type species – Pseudopyricularia kyllingae Klaubauf, M.-H. Lebrun & Crous
Notes – The genus was introduced by Klaubauf et al. (2014), along with four other species.
The type species, Pseudopyricularia kyllingae is a pathogen on Kyllinga brevifolia (Cyperaceae) in
Japan. Additional species were introduced by Crous et al. (2015d, 2018b), Marin-Felix et al. (2017)
and Pordel et al. (2017).
Pyricularia Sacc., Mycelia 2(no. 6): 20 (1880)
Index Fungorum number: IF9670; 44 morphological species(Species Fungorum 2020), 18
species with sequence data.
Type species – Pyricularia grisea Sacc.
Notes – The name Pyricularia refers to the pyriform shape of the conidia, which is a specific
character of this genus (Bussaban et al. 2005). Pyricularia species can infect various
monocotyledons including Cenchrus, Echinochloa, Lolium, Oryza, Pennisetum or Zingiber.
Pyricularia oryzae is the cause of rice blast disease, one of the most important diseases of rice.
Klaubauf et al. (2014) synonymised Magnaporthe grisea under Pyricularia grisea, by clarifying
the taxonomic relationships among species that are magnaporthe- or pyricularia-like using
molecular phylogeny (LSU, ITS, rpb1, act and calM).
Pyriculariomyces Y. Marín, M.J. Wingf. & Crous, Stud. Mycol. 92: 118 (2019)
Index Fungorum number: IF823760; 1 species with sequence data.
Type species – Pyriculariomyces asari (Crous & M.J. Wingf.) Y. Marín, M.J. Wingf. &
Crous
Notes – Pyriculariomyces was introduced to accommodate P. asari, collected from leaves
and stems of Asarum sp. in Malaysia. Pyriculariomyces is similar to Pyricularia, except the
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character of integrated terminal conidiogenous cells, and papillate necks with the ostioles of
ascomata (Marín-Felix et al. 2018).
Figure 218 – Pyricularia oryzae (a-d, g-h), Pyricularia grisea (e, f). a-d Perithecia, asci and
ascospores (redrawn from Yaegashi & Udagawa 1978). e, f Conidiophores and conidia (redrawn
from Klaubauf et al. 2014). g Pyriform conidia (redrawn from Zhang et al. 2014). h Microconidia
produced from phialides (redrawn from Zhang et al. 2014). Scale bars: a, d = 200 µm, b, c = 20 µm,
e, f = 10 µm, g, h = 5 µm.
Xenopyricularia Klaubauf, Lebrun & Crous, Stud. Mycol. 79: 116 (2014)
Index Fungorum number: IF810223; 1 species with sequence data.
Type species – Xenopyricularia zizaniicola (Hashioka) Klaubauf, Lebrun & Crous
Notes – Xenopyricularia is a monotypic genus, introduced to accommodate X. zizaniicola
which is pathogenic on Zizania latifolia in Japan. Xenopyricularia is similar to Pyricularia, except
the conidia are very wide and more obovoid than in Pyricularia, and some conidia appear to be
irregularly pigmented (Klaubauf et al. 2014).
Requienellaceae Boise, Mycologia 78(1): 37 (1986)
Index Fungorum number: IF81336; Facesoffungi number: FoF06879; 13 species.
Saprobic on dead wood or pathogenic on plants. Sexual morph: Ascomata immersed or
erumpent, globose to subglobose base with prominent apex, solitary or aggregated, perithecioid,
sometimes whitish, greyish to black at the margin. Ostiole inconspicuous or strongly erumpent,
flattened or papillate to conical, black. Peridium comprising thick-walled angular cells, thickened
in upper regions. Paraphyses numerous, apically free, aseptate, curved, scarcely branched. Asci 8spored, unitunicate, cylindrical, subfusiform to narrowly clavate, with thick-walled apex, wide
ocular chamber comprising a slightly refractive, inversely funnel-shaped dome, turning slightly
reddish in Congo Red. Ascospores uni - to biseriate, olivaceous or brown, ellipsoid to oblong, with
round or acute ends, with one or several transverse distosepta and large lumina. Asexual morph:
Undetermined (adapted from Jaklitsch et al. 2016b).
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Type genus – Requienella Fabre.
Notes – Requienellaceae has historically been treated in Melanommatales (Barr 1990a) based
on its trabeculate-like paraphyses (Liew et al. 2000). Aptroot (1991) followed this and added more
lichenised taxa to the non-lichenised Requienella. Jaklitsch et al. (2016b) determined
Requienellaceae as a distinct family in Xylariales based on the DNA based sequence analyses of its
generic type. Phylogenetically this family is a strongly supported monophyletic lineage sister to the
taxa in Cainiaceae (Jaklitsch et al. 2016b). Currently the family comprises four genera viz.
Acrocordiella (Jaklitsch et al. 2016b), Lacrymospora, Parapyrenis and Requienella
(Wijayawardene et al. 2018a).
Ecological and economic significance of Requienellaceae
Requienellaceae species have a widespread distribution and inhabit bark (Cannon & Krik
2007). They frequently occur on economic plants or lichens, such as R. seminuda on bark of dead
Olea europaea trees, R. fraxini on bark of living trunks of old Fraxinus excelsior, Acrocordiella
occulta on bark of Ribes spp., Acrocordiella omanensis saprobic on stems of Juniper sp. and
Parapyrenis lichenicola probably on thalli of Pertusaria erythrella (Aptroot 1997, Hawksworth &
Halici 2007, Jaklitsch et al. 2016b, Maharachchikumbura et al. 2018a). Therefore, members of this
family are saprobes, parasites or mutualistic and play an important role in material recycling and
energy conversion in natural ecosystems.
Genera included in Requienellaceae
Acrocordiella O.E. Erikss., Mycotaxon 15: 189 (1982)
Index Fungorum number: IF47; 2 species with sequence data.
Type species – Acrocordiella occulta (Romell) O.E. Erikss.
Notes – Acrocordiella is characterised by solitary or small groups of immersed ascomata with
white or black margins, a peridium comprising brown to hyaline, thick-walled cells of textura
angularis and 3-distoseptate ascospores with rhomboid lumina. This genus has been considered as
a synonym of Requienella (Boise 1986). Wijayawardene et al. (2018a) listed Acrocordiella in
Pyrenulaceae. However, Jaklitsch et al. (2016b) and Maharachchikumbura et al. (2016b) confirmed
Acrocordiella and Requienella as distinct genera of Requienellaceae in Xylariales
(Sordariomycetes) based on phylogenetic analyses.
Lacrymospora Aptroot, Biblthca Lichenol. 44: 95 (1991)
Index Fungorum number: IF25383; 1 morphological species.
Type species – Lacrymospora parasitica Aptroot
Notes – Lacrymospora is distinguished from Parapyrenis by blackening of the thallus around
the ascomata (Aptroot 1991) and asymmetrical ascospores with several rows of granules, indicating
septation (Calatayud et al. 2001). The generic position is still doubtful and fresh collections are
needed to understand the taxonomic status and provide support as a separate genus.
Parapyrenis Aptroot, Biblthca Lichenol. 44: 96 (1991)
Index Fungorum number: IF26301; 8 morphological species (Species Fungorum 2020), 2
species with sequence data.
Type species – Parapyrenis aurora (Zahlbr.) Aptroot
Notes – Parapyrenis was described to accommodate P. aurora, P. conica, P. elongata and P.
guayaci that are treated as non-lichenized, and are associated with plant bark, having cellular
paraphyses and ascospores with a thickened internal wall layer (Aptroot 1991), Aptroot (1995)
introduced P. maclurae, P. maritima and P. tecomatis and provided a key to Parapyrenis species.
A lichenized fungus P. lichenicola was also assigned to this non-lichenized genus by Aptroot et al.
(1997). Hawksworth & Halici (2007) reviewed this genus and provided a brief description for
Parapyrenis without illustrations. Vu et al. (2019) added LSU and ITS sequences to P. conica and
P. maritima.
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Requienella Fabre, Annls Sci. Nat., Bot., sér. 6 15: 55 (1883)
Index Fungorum number: IF4676; 2 species with sequence data.
Type species – Requienella seminuda (Pers.) Boise.
Notes – Requienella was introduced by Fabre (1883), however, it has been shown that the
three species recognised by him in Requienella belonged to other genera. Boise (1986) reinstated
Requienella by typifying the genus with R. seminuda (= Sphaeria seminuda). Based on
morphology, the genus was been classified in Dothideomycetes, however, Jaklitsch et al. (2016b)
showed the genus belongs to the class Sordariomycetes, despite its apparently fissitunicate asci.
Requienella fraxini and R. seminuda are accepted in Requienella based on morphology and
sequence data and R. princeps has been transferred to Decaisnella (Barr 1990a).
Figure 219 – Requienella seminuda (a, b, d, e, h, i, j), Requienella fraxini (c, k, l), Acrocordiella
occulta (j, m, n) redrawn from Jaklitsch et al. (2016b). a Ascomata. b, c Sections of ascomata. d, e
Asci and paraphyses. e Apical region of asci. f-k Ascospores. Scale bars: a = 0.2 mm, b = 0.2 mm,
c = 0.3 mm, d = 25 µm, d, f = 15 µm, e, k, l, n = 10 µm, i, j, m = 7 µm.
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Reticulascaceae Réblová & W. Gams, Stud. Mycol. 68(1): 180 (2011)
Index Fungorum number: IF515435; Facesoffungi number: FoF01311; 37 species.
Saprobic on wood or bark in terrestrial and freshwater habitats. Sexual morph: Stromata if
present minute. Ascomata perithecial, superficial, gregarious to solitary, brown to black,
subglobose to conical, base flattened, slightly verruculose, glabrous with minute papilla, ostiole
periphysate. Peridium 2-layered, fragile, the whole wall heavily sclerotized in the upper part,
poorly developed towards the rim, absent at the base. Paraphyses numerous, filiform, sparsely
septate, hyaline. Asci 8-spored, unitunicate, cylindrical to clavate, short-pedicellate, apex truncate
to broadly rounded, with J-, apical ring. Ascospores uni to bi-seriate, hyaline or dark brown,
ellipsoidal to fusiform, septate, mostly 2-celled, with a delayed formation of the 2 additional septa,
smooth-walled, with or without end pores. Asexual morphs: Setae straight, cylindrical, septate,
dark brown, paler towards the apex when present. Conidiophores macronematous, mononematous,
solitary, erect, straight, sometimes curved, branched or unbranched, thick-walled, brown to dark
brown, becoming paler towards the apex, terminating in a cylindrical to slightly flask-shaped or
funnel-shaped, smooth-walled. Conidiogenous cells mono- or polyphialidic, collarette hyaline to
subhyaline. Conidia hyaline to brown, pyriform to cylindrical, 1- or multi-septate, distal end
bluntly rounded, basal end truncate, smooth-walled (adapted from Réblová et al. 2011).
Type genus – Cylindrotrichum Bonord.
Notes – Reticulascaceae was introduced by Réblová et al. (2011) in Glomerellales based on
analysis of combined ITS, LSU, SSU and rpb2 sequence data. Three genera were included, i.e.
Cylindrotrichum (as Reticulascus), Kylindria and Sporoschismopsis. Presently, Blastophorum is
also accepted in Reticulascaceae by Hyde et al. (2016b) based on the phylogenetic analysis and
morphology. In addition, Hongsanan et al. (2017) provided further evidence for the placement of
Reticulascaceae in Glomerellales by showing good support in both phylogenetic and MCC trees.
Ecological and economic significance of Reticulascaceae
Member of Reticulascaceae are saprobes involved in nutrient cycling. Marsault & Peterson
(2017) reported a new compound cyclic decadepsipeptide isolated from Cylindrotrichum
tulasneorum.
Genera included in Reticulascaceae
Blastophorum Matsush., Microfungi of the Solomon Islands and Papua-New Guinea (Osaka): 8
(1971)
Index Fungorum number: IF7394; 4 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Blastophorum truncatum Matsush.
Notes – This genus is characterized by unbranched, percurrently proliferating, septate and
dark brown conidiophores; enteroblastic to polyblastic and sympodial conidiogenous cells, which
are subhyaline at the base and hyaline above and inconspicuous flattened denticles at
conidiogenous loci; and septate, elongated, cuneiform and slimy conidia (Hyde et al. 2016b). This
genus resembles Kylindria in its polyblastic and sympodial conidiogenous cells, but differs in
having cuneiform and larger conidia (Hyde et al. 2016b). Hyde et al. (2016b) placed Blastophorum
in Reticulascaceae based on the phylogenetic analysis of a combined LSU and ITS sequence data
and morphology.
Cylindrotrichum Bonord., Handb. Allgem. mykol. (Stuttgart): 88 (1851)
Index Fungorum number: IF7881; 15 morphological species (Species Fungorum 2020), 6
species with sequence data
Type species – Cylindrotrichum oligospermum (Corda) Bonord.
Notes – Cylindrotrichum was introduced by Bonorden (1851) with C. oligospermum as the
type species. Reticulascus was introduced by Réblová et al. (2011) for a holomorphic species R.
tulasneorum (asexual morph C. oligospermum, Réblová & Gams 1999), the generic type, and R.
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clavatus (asexual morph Cylindrotrichum clavatum). Réblová et al. (2016b) recommended the use
of Cylindrotrichum over Reticulascus given its widespread use, priority, and greater number of
names. Cylindrotrichum oligospermum is illustrated in this entry.
Figure 220 – Cylindrotrichum oligospermum (Material examined – CZECH REPUBLIC,
Southwestern Bohemia, Javornická, Hornatina Mts, Strašín, near Sušice, on dead branch of
Sambucus nigra L. (Adoxaceae), 21 October 1997, M. Svrček, PRM 842978, holotype). a
Ascomata on the host. b Section through ascoma. c Peridium. d-g Asci. h Interthecial filaments. i-l
Ascospores (j Ascospores stained with Meltzer reagent). m Conidia (redrawn from Réblová &
Gams 1999). Scale bars: a = 100 µm, b, c = 50 µm, d-g = 25 µm, h-l = 10 µm, m = 20 µm.
Kylindria DiCosmo, S.M. Berch & W.B. Kendr., Mycologia 75(6): 970 (1983)
Index Fungorum number: IF11115; 11 morphological species (Species Fungorum 2020), 2
species with sequence data.
Type species – Kylindria triseptata (Matsush.) DiCosmo, S.M. Berch & W.B. Kendr.
Notes – Kylindria was erected by DiCosmo et al. (1983) with the type species K. triseptatum
(≡Cylindrotrichum triseptatum) and another four Cylindrotrichum species. Based on the
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phylogenetic analyses of a combine ITS, LSU, SSU dataset, Réblová et al. (2011) show that
Kylindria should be accommodated in Reticulascaceae instead of Chaetosphaeriaceae.
Maharachchikumbura et al. (2015, 2016b) confirmed Kylindria belongs to Reticulascaceae based
on the phylogenetic analyses of a combine LSU, SSU, tef1 and rpb2 dataset. Two species were
introduced by Maharachchikumbura et al. (2018b). The genus is characterized by setiform,
branched and septate conidiophores, monophialidic conidiogenous cells and septate, oblongellipsoid with hyaline conidia (Maharachchikumbura et al. 2016b, 2018b).
Figure 221 – Cylindrotrichum oligospermum a-d Asexual morph of Reticulascus tulasneorum,
CBS 101319, PR M 842978, redrawn from Réblová & Gams (1999) e Menispora oligosperma,
redrawn from Corda (1838). a Habit sketch of conidiophores and setae. b-d Conidiophores and
hyaline conidia of Reticulascus tulasneorum. e A flake of conidiophore and spores of Menispora
oligosperma. Scale bars: b-d = 10 µm.
Sporoschismopsis Hol.-Jech. & Hennebert, Bull. Jard. Bot. natn. Belg. 42(4): 385 (1972)
Index Fungorum number: IF10043; 7 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Sporoschismopsis moravica Hol.-Jech. & Hennebert
Notes – Sporoschismopsis was described for saprobic, wood-inhabiting dematiaceous
hyphomycetes by Holubová-Jechová & Hennebert (1972). The sexual-asexual relationship between
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Porosphaerellopsis and Sporoschismopsis has been established by Réblová (2014), and the sexual
morph Porosphaerellopsis has been treated as the synonym of Sporoschismopsis given its priority.
The asexual morph of this genus is characterized by robust conidiophores with a funnel-shaped
collarette, brown, septate, clavate, cuneiform, obovate or pyriform conidia, bluntly rounded at the
distal end and truncate at the base, developing in a basipetal succession in fragile chains. The
sexual morph is characterized by superficial to semi-immersed, subglobose, conical or ovate, dark
brown ascomata and with ellipsoidal, fusiform, septate, brown to dark-brown ascospores (Réblová
2014). In addition, Réblová (2014) also provided key to the species accepted in Sporoschismopsis.
Rhamphoriaceae Réblová, Mycologia: 110(4):750–770 (2018)
Index Fungorum number: IF823918; Facesoffungi number: FoF06880; 17 species.
Saprobes on wood. Sexual morph: Ascomata perithecial, immersed or particularly erumpent
becoming superficial, papillate or with a central cylindrical or rostrate neck. Ostiole periphysate.
Peridium leathery to fragile, comprising two layers. Paraphyses consisting of septate, cylindrical
paraphyses. Asci 8-spored, unitunicate, cylindrical or cylindrical-clavate, persistent, with a J-,
apical ring. Ascospores overlapping uniseriate, hyaline or brown, ellipsoidal, obovoid, clavate,
fusiform to fusiform-clavate, dictyoseptate or transversely septate, lacking mucilaginous sheaths or
appendages, in some species ascospores may produce ascoconidia while in the asci (e.g.
Rhamphoria pyriformis). Asexual morph: Hyphomycetous. Conidiophores macronematous or
semi-macronematous, mononematous or loosely fasciculate. Conidiogenous cells producing
conidia holoblastically, conidial secession rhexolytic on sympodially extending polyblastic
conidiogenous cells, or schizolytic on monoblastic conidiogenous cells. Conidia hyaline
(Rhamphoria, Rhamphoriopsis, Rhodoveronaea and Xylolentia) or brown (Linkosia multiseptum),
aseptate or septate (adapted from Réblová & Štěpánek 2018).
Type genus – Rhamphoria Niessl
Notes – Rhamphoriaceae was introduced by Réblová & Štěpánek (2018) with Rhamphoria as
the type genus and included the genera Rhamphoriopsis, Rhodoveronaea, Xylolentia and a
dematiaceous hyphomycete Linkosia multiseptum. They formed a monophyletic clade with high
statistical support based on a combined dataset of SSU, LSU and rpb2 sequences. Munk (1957)
placed Rhamphoria in the subfamily Rhamphorioidae of the Diaporthaceae based on morphology.
However, the position of Rhamphoria in the Sordariomycetes was uncertain (Lumbsch & Huhndorf
2010). A strongly supported monophyletic clade based on LSU sequence data of Rhamphoria
delicatula and Rhodoveronaea varioseptata was recovered and selected as members of the
Sordariomycetidae (Réblová 2009). Despite Rhamphoria and Rhodoveronaea differing in sexual
characters, their asexual morphs are similar dematiaceous hyphomycetes with same mode of
conidiogenesis.
These genera (except for Linkosia) share a few features such as perithecial ascomata, absence
of stromatic tissues or clypeus, similar anatomy of the two-layered peridium, unitunicate,
pedicellate asci with a J-, apical ring, cylindrical paraphyses and dictyoseptate or transversely
septate ascospores (Réblová & Štěpánek 2018). Linkosia multiseptum, on the other hand, lacks
conidiophores, has distoseptate, thick-walled conidia and ampulliform, monoblastic conidiogenous
cells. Based on the morphology, it is unexpected to place L. multiseptum in Rhamphoriaceae.
Shenoy et al. (2006) suggested that Linkosia (Hernández-Gutiérrez & Sutton 1997) is polyphyletic,
based on LSU sequence data. The most similar relatives of Rhamphoriaceae are members of the
Sporidesmiaceae (Su et al. 2016b).
Ecological and economic significance of Rhamphoriaceae
Members of Rhomphoriaceae are saprobes of decaying wood, reported predominantly from
temperate regions of Europe. Members of Rhamphoria are uncommon except for R. pyriformis,
which occurs on various hard wood trees (Munk 1948, 1957, Sivanesan 1976, Müller & Samuels
1982a, Réblová & Štěpánek 2018). Other Rhamphoria species have been collected only a few times
and mostly only once (Réblová & Štěpánek 2018).
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Genera included in Rhamphoriaceae
Rhamphoria Niessl, Verh. nat. Ver. Brünn 14: 204 (1876)
Index Fungorum number: IF4690; 13 morphological species (Species Fungorum 2020), 3
species with sequence data.
Type species – Rhamphoria delicatula Niessl
Notes – Rhamphoria is characterized by black, astromatic, perithecial ascomata with a
distinct papilla or rostrate neck, unitunicate asci with a J-, apical ring and hyaline, dictyospores that
may produce ascoconidia while still in the asci (e.g. Rhamphoria pyriformis). The species in
Rhamphoria are mainly separated by the size of ascospores, number of transverse and longitudinal
septa and to a lesser degree on spore shape (Réblová & Štěpánek 2018).
Figure 222 – Rhamphoria delicatula (redrawn from Réblová & Štěpánek 2018). a Ascomata. b
Peridium. c Asci. d Paraphyses. e Ascal apices with apical ring. f Ascospores. g Idriella-like
conidiophores and conidia in PDA, 4 weeks. h Phaeoisaria-like conidiophores with conidia in PDA
culture, 4 weeks. Scale bars: a = 200 µm, b = 20 µm, c-h = 10 µm.
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Rhamphoriopsis Réblová & Gardiennet, Mycologia: 6 (2018)
Index Fungorum number: IF823919; 1 species with sequence data.
Type species – Rhamphoriopsis muriformis Réblová & Gardiennet
Notes – Rhamphoriopsis muriformis was introduced by Réblová & Gardiennet (2018) from
decaying wood of Buxus sempervivens. Rhamphoriopsis muriformis differs from Rhamphoria buxi
Richon by longer asci and longer, hyaline ascospores and transverse and longitudinal, thin-walled
septa (Réblová & Štěpánek 2018).
Rhodoveronaea Arzanlou, W. Gams & Crous, Stud. Mycol. 58: 89 (2007)
Index Fungorum number: IF504569; 2 species with sequence data.
Type species – Rhodoveronaea varioseptata Arzanlou, W. Gams & Crous
Notes – Rhodoveronaea was introduced by Arzanlou et al. (2007), with the characters similar
to Veronaea and isolated from Bertia moriformis. This monophyletic genus was described as an
asexually reproducing taxon with no known sexual morph (Arzanlou et al. 2007). Réblová (2009)
showed the connection between the sexual and asexual morphs of Rhodoveronaea varioseptata, the
latter which differs from the similar ramichloridium-like taxa by a basal, marginal conidial frill, and
variably septate conidia.
Xylolentia Réblová, Mycologia: 10 (2018)
Index Fungorum number: IF823921; 1 species with sequence data.
Type species – Xylolentia brunneola Réblová
Notes – Xylolentia brunneola was collected on decaying wood of Fagus sylvatica. This genus
is distinct among members of the Rhamphoriaceae in having brown, septate ascospores. It is similar
to Brunneosporella aquatica (Ranghoo et al. 2001), Hyalorostratum brunneisporum (Raja et al.
2010), and Jobellisia spp. (Huhndorf et al. 1999), all of which have 1-septate, brown ascospores
arranged uniseriately in the ascus. However, analysis of a combined SSU, LSU and rpb2 sequence
dataset did not show a close relationship of Xylolentia with these genera (Réblová & Štěpánek
2018).
Sarocladiaceae Lombard, Persoonia 41: 343 (2018)
Index Fungorum number: IF828245; Facesoffungi number: FoF06265; 25 species.
Saprobic on dried stems and branches of monocotyledons and dicotyledons, occasionally
hemibiotrophic or necrotrophic, soil associated fungi, and clinical trial samples, and clinical wash
fluid. Sexual morph: Undetermined. Asexual morph: Colonies effuse on the natural substrate,
scattered, hairy, dark brown, septate. Vegetative hyphae hyaline, branched, partly immersed,
smooth- and thin-walled, branched, composed of pale brown, septate hyphae. Conidiophores
macronematous, synnematous, tree-like, dark brown, parallel and unbranched in the stipe,
gregarious or scattered, erect, stripes straight or slightly flexuous, constricted at septa, tapering
towards the apex, irregularly branched, cylindrical, smooth. Conidiogenous cells phialidic,
integrated, terminal, cylindrical, straight to slightly curved, slightly narrowing at apex, acropetally
proliferating, hyaline to pale brown, verrucose. Conidia forming slimy heads on the phialides or
produced in chains, unicellular, fusiform to broad fusiform, hyaline, aseptate, thick-walled,
verrucose, bud scars or disjunctions present at the site of attachment, guttulate (adapted from Crous
et al. 2018b).
Type genus – Sarocladium Gams & D. Hawksw.
Notes – Sarocladiaceae was introduced by Crous et al. (2018b) to accommodate the
distinctive lineage of Parasarocladium and Sarocladium as the generic type. Sarocladium was
previously placed in genera incertae sedis of Hypocreales as it is formed a phylogenetic affinity
with species of Acremonium (Summerbell et al. 2011, Giraldo et al. 2015, Maharachchikumbura
et al. 2015b, 2016b, Wijayawardene et al. 2017b). The phylogenetic distinction of Acremonium
and Sarocladium is confirmed since the type species of Acremonium (A. alternatum) formed a
well-supported lineage in Bionectriaceae (Crous et al. 2018b; Fig. 15 in this study). The
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phylogenetic placement of Sarocladium and related species was provided, with Parasarocladium, a
second genus for the Acremonium radiatum complex (Summerbell et al. 2018).
A phylogenetic analysis of the LSU sequence alignment showed that Sarocladiaceae shares a
close relationship with Bionectriaceae (Crous et al. 2018b). In our multilocus phylogenetic
analysis of the combined LSU, ITS, cmdA, rpb2, tef1 and tub2 sequence data of Hypocreales,
Sarocladiaceae formed a close relationship with Flammocladiellaceae as basal lineage of
Hypocreales. Sarocladiaceae comprise two genera, Parasarocladium and Sarocladium.
Ecological and economic significance of Sarocladiaceae
Sarocladiaceae species have been recorded as plant pathogens causing sheath-rot of rice and
bamboo (Sawada 1922, Gams & Hawksworth 1976, Sakthivel et al. 2002, Ayyadurai et al. 2005).
They are also saprobes on monocotyledons and dicotyledons, and soils-associated fungi (Giraldo et
al. 2015, Liu et al. 2017, Crous et al. 2018b, Phukhamsakda et al. 2020). The genus was also
reported as human pathogens from calf tissue of legs, sputum and clinical wash fluid (Giraldo et al.
2015). Sarocladium oryzae produced antibiotic cerulenin and helvolic acid (Bills et al. 2004).
Genera included in Sarocladiaceae
Parasarocladium Summerb., J.A. Scott, Guarro & Crous, Microorganisms 6(88): 17 (2018)
Index Fungorum number: IF826815; 4 species with sequence data.
Type species – Parasarocladium radiatum (Sukapure & Thirum.) Summerb., J.A. Scott,
Guarro & Crous
Notes – Parasarocladium was introduced for Acremonium species that formed a sister clade
with Sarocladium sensu stricto (Crous et al. 2018b, Summerbell et al. 2018). The genus is
distinguished from Sarocladium by branched tufts of up to four conidiophores and the homologues
characteristic of CAACTTT motif at 5’ end of 5.8S rDNA (Summerbell et al. 2018).
Sarocladium Gams & D. Hawksw., Kavaka 3: 57 (1976)
Index Fungorum number: IF9790; 21 species with sequence data.
Type species – Sarocladium oryzae (Sawada) W. Gams & D. Hawksw.
Notes – Sarocladium oryzae was originally described as Acrocylindrium oryzae for a sheathrot associated fungus on rice (Sawada 1922, Sakthivel et al. 2002). Gams & Hawksworth (1976)
introduced Sarocladium for Acrocylindrium oryzae based on Sawada's holotype collection. Giraldo
et al. (2012) re-examined cultures of Acremonium, and included seven species in the Sarocladium
clade. Thereafter, the phylogenetic relationship of Acremonium and Sarocladium species has been
clarified, and 18 species are included in Sarocladium (Giraldo et al. 2015). Liu et al. (2017), Crous
et al. (2018b) and Phukhamsakda et al. (2020) described Sarocladium brachiariae, S. clematidicola
and S. dejongiae, respectively.
Sarocladium can be distinguished by its elongated phialides rising solitary on vegetative
hyphae or on conidiophores that are sparsely or repeatedly branched, and the production of
cylindrical conidia (Fig. 223). Below we illustrate Sarocladium clematidis.
Savoryellaceae Jaklitsch & Réblová, Index Fungorum 209: 1 (2015)
Index Fungorum number: IF551026; Facesoffungi number: FoF01283; 42 species.
Saprobic on submerged wood in freshwater, marine and brackish habitats. Sexual morph:
Ascomata perithecial, brown to black, immersed or superficial, globose to pyriform, coriaceous,
periphysate, ostiolate, papillate. Papilla central or eccentric when lying horizontally to the host.
Peridium membranous, comprising several layers of brown, thick-walled cells of textura angularis,
inner layers hyaline. Paraphyses hypha-like, numerous or sparse, septate. Asci 2–8-spored,
unitunicate, clavate to cylindrical, pedicellate, persistent, with a small or relatively large, J-, apical
ring. Ascospores uniseriate or overlapping biseriate, versicolorous with brown middle cells and
hyaline end cells, ellipsoid, fusiform, 3 to many septate, smooth, thick-walled, with or without-
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Figure 223 – Sarocladium clematidis (Material examined – BELGIUM, Flemish Brabant, Botanic
Garden, Bouchout Domain, on dead stems of Clematis patens (Ranunculaceae), 13 June 2017, D.
Ertz BRCP3, MFLU17 –1507, holotype). a Appearance of synnemata on Clematis viticella. b, c
Synnemata. d Conidiogenous cells and conidia. e Asexual morph produced in culture. f, g
Conidiogenous cells and developing conidia. h Conidia. i Culture on PDA. Scale bars: a = 500 μm,
b, c = 100 μm, d, e = 50 μm, f = 10 μm, g, h = 5 μm.
-polar mucilaginous pads or appendages. Asexual morph: Hyphae septate, branched, hyaline to pale
brown. Conidiophores semi-macronematous (conidiophores that are only slightly different from the
vegetative hyphae). Conidiogenous cells absent or if present erect, smooth and thick-walled,
holoblastic, cylindrical, flask-shaped. Conidia solitary, dry, smooth, applanate or rounded, globose
to subglobose or obovate to oval, with longitudinal septa and transverse septa or only transversely
septate, slightly constricted at the septa, with a pale brown small basal cell, olive green to brown or
blackish brown to black, with prominent scar at the point of secession from the conidiogenous cell
(adapted from Luo et al. 2019).
Type genus – Savoryella E.B.G. Jones & R.A. Eaton
Notes – Savoryellaceae was introduced by Jaklitsch & Réblová (2015) to accommodate
Savoryella. Savoryellaceae was previously placed in Sordariales genera incertae sedis by Jones et
al. (2009), in Savoryellales by Boonyuen et al. (2011). Hyde et al. (2017a) suggested that
Savoryellales be raised to subclass rank based on the estimated stem age (Savoryellales; 267 MYA)
and Hongsanan et al. (2017) supported this by phylogenetic and molecular clock analyses and,
hence, introduced the subclass Savoryellomycetidae. Four other genera, i.e. Ascotaiwania,
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Canalisporium, Savoryella and Neoascotaiwania were included in the family along with the
species Helicoön farinosum and Monotosporella species (Jones et al. 2015, Hernández-Restrepo et
al. 2017, Wijayawardene et al. 2018a). Several types of asexual morphs, bactrodesmium-like,
dictyosporium-like, monodictys-like, monotosporella-like, trichocladium-like are linked under
Savoryellaceae (Sivichai et al. 1998, Ranghoo & Hyde 1998, Chang 2001, Sri-indrasutdhi et al.
2010, Réblová et al. 2015, Hernández-Restrepo et al. 2017). The distant placement of Helicoon
farinosum, the asexual morph of Ascotaiwania hughesii (Fallah et al. 1999), from species of
Savoryellales was revealed by rDNA data (Boonyuen et al. 2011, Réblová et al. 2012) and, hence,
Ascotaiwania was considered polyphyletic (Hernández-Restrepo et al. 2017). Dayarathne et al.
(2019a) revised the family with combined LSU, SSU, tef1 and rpb2 data along with molecular
clock analyses and synonymized Neoascotaiwania under Ascotaiwania and, further, excluded the
Monotosporella species, Helicoön farinosum and Ascotaiwania hughesii from the family.
Dematisporium was introduced by Luo et al. (2019) and therefore, currently Savoryellaceae
comprises four genera, Ascotaiwania, Canalisporium, Dermatrosporium and Savoryella.
Ecological and economic significance of Savoryellaceae
Many Savoryellaceae species cause soft rot decay of wood (Eaton & Jones 1971, Leightly &
Eaton 1979, Mouzouras 1986, Leightly 1980). This type of decay is prevalent where wood is
exposed to wet conditions, such as submerged wood in marine and freshwater habitats (Jones 1972,
Bucher et al. 2004), water cooling towers (Findlay & Savory 1950, Savory 1954a, b), archeological
timbers (Jones & Jones 1993), and terrestrial environments (Duncan 1960).
Genera included in Savoryellaceae
Ascotaiwania Sivan. & H.S. Chang, Mycol. Res. 96(6): 481 (1992)
Index Fungorum number: IF25163; 13 morphological species (Species Fungorum 2020,
Dayarathne et al. 2019a), 7 species with sequence data.
Type species – Ascotaiwania lignicola Sivan. & H.S. Chang
Notes – Ascotaiwania was established by Sivanesan & Chang (1992) for an ascomycete
comprising dark-brown to black ascomata with a short to long, lateral neck, asci with a relatively
massive, J-, apical ring and 5 or 3 or 7-septate pigmented ascospores with hyaline end cells, with A.
lignicola as the type species. This genus currently comprises 12 species (Dayarathne et al. 2019a).
Divergence estimates are a crown age of 146 (200–196) MYA. (Dayarathne et al. 2019a).
Canalisporium Nawawi & Kuthubutheen, Mycotaxon 34(2): 477 (1989)
Index Fungorum number: IF11041; 15 morphological species (Zhao et al. 2012), 11 species
with sequence data.
Type species – Canalisporium caribense (Hol.-Jech. & Mercado) Nawawi & Kuthub.
Notes – Canalisporium was introduced to accommodate Berkleasmium caribense, B.
pulchrum (Holubová-Jechová & Mercado Sierra 1984) and Canalisporium elegans (Nawawi &
Kuthubutheen 1989). At present, 15 species are recognized in this genus (Zhao et al. 2012). Sriindrasutdhi et al. (2010) described a sexual morph for C. grenadoideum and introduced
Ascothailandia, typified by A. grenadoidea. Dayarathne et al. (2019a) showed that this genus
emerged at the crown age of 56 (23–94) MYA.
Dematiosporium Z.L. Luo, K.D. Hyde & H.Y. Su, Fungal Divers.: 99: 573 (2019)
Index Fungorum number: IF555672; 1 species with sequence data.
Type species – Dematiosporium aquaticum Z.L. Luo, K.D. Hyde & H.Y. Su
Notes – Dematisporium which is typified by D. aquaticum which mostly resembles
Conioscypha aquatica in having micronematous conidiophores and dry, dark brown to black,
globose to subglobose conidia (Luo et al. 2019). However, Dematisporium aquaticum is
distinguished from Conioscypha aquatica in having larger conidia and C. aquatica belongs to
Conioscyphaceae.
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Figure 224 – Ascomatal, asci and ascospore morphology of the genera of Savoryellaceae: S.
grandispora (Material examined – MALAYSIA, State Negara, Lipur Lentang Nature Reserve, on
submerged wood, October 1991, K.D. Hyde, BRIP 20918, holotype), Savoryella lignicola
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(Material examined – UK, Flintshire, Connah’s Quay, on Scots pine test-blocks placed for 168 days
amongst the packing timber of a water -cooling tower at Connah's Quay, 16 June 1966 and 1
December 1966, IMI 129784, IMI 129785 holotype), S. longispora (Material examined – UK,
Flintshire, Connah’s Quay, on Scots pine test-blocks placed for 168 days amongst the packing
timber of a water -cooling tower at Connah's Quay, 16 June1966 and 1 December 1966, IMI
129784, IMI 129785, holotype), S. yunnanensis (Material examined – CHINA, Yunnan Province,
Dehong, on submerged wood in a stream, 25 November 2017, W. Dong, H40C, MFLU 18-1203,
holotype), Canalisporium caribense (Material examined – THAILAND, Chiang Rai Province,
stream flowing in Tham Luang Nang Non Cave, on submerged wood, 25 November 2014, J. Yang,
MFLU15-3581, holotype). a, b Ascomata of S. lignicola. c, d Peridium and paraphyses of S.
yunnanensis. e-g Asci of S. lignicola, S. grandispora, S. yunnanensis respectively. h Apical ring of
S. grandispora. i-l Ascospores of S. grandispora, S. lignicola, S. longispora and S. yunnanensis
respectively. m Conidia attached to the conidiophores of C. caribense. n Conidia attached to the
conidiophores of A. fusiformis. o-q Conidiogenous cells and conidia of A. lignicola (Chang 2001).
c, d A. sawadae (Ranghoo & Hyde 1998) A. uniseptata (Kirk 1983) respectively. Scale bars: b, c =
100 μm, d, e = 50 μm, f-i = 10 μm.
Savoryella E.B.G. Jones & R.A. Eaton, Trans. Br. mycol. Soc. 52(1): 161 (1969)
Index Fungorum number: IF4870; 13 morphological species (Species Fungorum 2020), 9
species with sequence data.
Type species – Savoryella lignicola E.B.G. Jones & R.A. Eaton
Notes – Savoryella (Jones & Eaton 1969) currently comprises eleven species (HernandezRestrepo et al. 2017, Dayarathne et al. 2019a). Savoryella limnetica was combined under
Ascotaiwania limnetica, re-described with a new asexual morph by Réblová et al. (2015c) and
synonymized under Neoascotaiwania limnetica by Hernandez-Restrepo et al. (2017). However,
Dayarathne et al. (2019a) synonymized Neoascotaiwania under Ascotaiwania. Savoryella has been
placed in several orders; Sphaeriales family incertae sedis by Kohlmeyer & Kohlmeyer (1979),
ascomycetes incertae sedis by Kohlmeyer (1986), Amphisphaeriaceae (Eriksson & Hawksworth
1986b) and Sordariales (Jones & Hyde 1992). Barr (1990a) transferred this genus to
Halosphaeriales (now Microascales) based on morphology (the catenophyses-like paraphyses) and
ultrastructural observations. Vijaykrishna et al. (2006) and Cai et al. (2006a) referred Savoryella
elongata and S. longispora to Hypocreales, subclass Hypocreomycetidae, based on a phylogenetic
analysis of LSU data, although with weak statistical support. Jones et al. (2009) referred the genus
to the Sordariales genera incertae sedis based on morphology. Boonyuen et al. (2011) introduced
the new order Savoryellales to accommodate the genera Ascotaiwania, Ascothailandia (and its
asexual morph, Canalisporium) and Savoryella based on multigene dataset. Dayarathne et al.
(2019a) showed that divergence of this genus was 108 (72–146) MYA.
Schizoparmaceae Rossman, D.F. Farr & Castl., Mycoscience 48(3): 137 (2007)
Index Fungorum number: IF82150; Facesoffungi number: FoF04613; 57 species.
Saprobic, endophytic, parasitic or pathogenic on fruits, leaves, stems, and roots of wide
variety of woody and herbaceous plants, and in soil. Sexual morph: Ascomata perithecial, solitary,
when collapsed collabent, scattered, subepidermal, erumpent to superficial, globose, coriaceous,
brown to black, papillate, with periphysate ostiole in the center. Peridium of thick-walled, brown
cells of textura angularis. Paraphyses lacking. Asci 8-spored, unitunicate, clavate to subcylindrical,
sessile, with a J-, apical ring, floating free at maturity. Ascospores biseriate, hyaline, becoming pale
brown at maturity, ellipsoidal, aseptate, with or without mucoid caps. Asexual morph:
Coelomycetous. Conidiomata pycnidial, subepidermal, immersed to semi-immersed, erumpent,
unilocular, globose, slightly depressed globose to subglobose, glabrous, brown to dark brown or
black, ostiolate. Peridium of irregularly thickened, with plate-like orientation, comprising brown
cells of textura angularis. Conidiophores densely aggregated, simple or branched at base,
occasionally septate, smooth, hyaline, subulate, invested in mucus, developing from basal pad.
852
Conidiogenous cells phialidic, often annellidic, subcylindrical, obclavate or lageniform, discrete,
hyaline, smooth, proliferating percurrently, or with prominent periclinical thickening. Conidia
hyaline, becoming olivaceous brown to brown at maturity, unicellular, smooth, thin- or thickwalled, ellipsoid, globose, napiform, fusiform or naviculate with a truncate base and an obtuse to
apiculate apex, at times with a longitudinal germ slit, with or without a mucoid appendage (adapted
from Rossman et al. 2007).
Type genus – Coniella Höhn.
Notes – Schizoparmaceae was introduced by Rossman et al. (2007) to accommodate
Schizoparme and the asexual genera Coniella and Pilidiella (Rossman et al. 2007, Alvarez et al.
2016). Species of Schizoparmaceae are common in both tropical and temperate regions, worldwide
(Rossman et al. 2007). Initially, the species in this family were placed in Melanconidaceae.
Following Castlebury et al. (2002), who established the name ‘Schizoparme complex’ and their
distinct lineage in Diaporthales, many comprehensive studies were conducted on the family (van
Niekerk et al. 2004, Rossman et al. 2007, Alvarez et al. 2016). Even though, the family was
established with three genera (Coniella, Pilidiella and Schizoparme), and after several taxonomic
refinements, the family currently comprises a single genus (van Niekerk et al. 2004, Rossman et al.
2007, Alvarez et al. 2016). Based on the International Code of Nomenclature for algae, fungi and
plants (McNeil et al. 2012), Alvarez et al. (2016) synonymized Pilidiella and Schizoparme under
the accepted generic name Coniella. Schizoparmaceae has a stem age at 70 MYA (Hongsanan et al.
2017). However, authors have mentioned that the data set used in the study was not sufficient to
represent this morphologically diverse family. Most of the previous studies of this genus used
single gene phylogenies of ITS and LSU for species delimitation (Castlebury et al. 2002, van
Niekerk et al. 2004, Miranda et al. 2012). This approach is accepted for the preliminary
identification. Confusions and inconsistencies in this method were addressed by multigene
phylogenetic approach used in Alvarez et al. (2016) and Chethana et al. (2017). Therefore, it is
recommended to use ITS, LSU, tef1, rpb2 and histone (selection of 4 genes) in combined analyses
(Fig. 227) to resolve Coniella species (Alvarez et al. 2016, Chethana et al. 2017).
Ecological and economic significance of Schizoparmaceae
Coniella species are economically important due to the negative role they play by causing
foliar, fruit, leaf, stem and root diseases on a range of economically important plants. Coniella
diplodiella and C. vitis cause grape white rot of grapes (Chethana et al. 2017); Coniella
castaneicola causes fruit and leaf diseases on strawberry (Mass 1998). Coniella granti causes
cankers, crown rots, die backs, fruit rots, leaf spots, shoot blights and twig blights on pomegranates
(Chen et al. 2014, Mirabolfathy et al. 2012, KC & Vallad 2016). Many of these hosts are fruit
plants. Therefore, these diseases can lead to production losses and also reduce the marketability of
the fruits.
Yu et al. (2018) reported seven Azaphilone derivatives known as coniellins A-G from
Coniella fragariae. These Coniellins exhibited moderate cytotoxicity, significant inhibition of
NFκB activation in the triple negative breast cancer cell lines and reduced the tumour cell
migration. This is the only report on natural products from this genus. Therefore, further studies
might discover natural products from Coniella species.
Genus included in Schizoparmaceae
Coniella Höhn., Ber. Dt. Bot. Ges. 36(7): 316 (1918)
Index Fungorum number: IF7753; 57 morphological species (Species Fungorum 2020), 38
species with sequence data.
Type species – Coniella fragariae (Oudem.) B. Sutton
Notes – The cosmopolitan genus, Coniella was established by Höhnel (1918a) and is typified
by Coniella pulchella (= Coniella fragariae). Species of this genus are plant pathogens (van
Niekerk et al. 2004, Mirabolfathy et al. 2012, Chethana et al. 2017). In addition, they have
saprobic, endophytic, and parasitic lifestyles, as well as being secondary invaders of plant tissues
853
infected by other organisms or injured by other causes (Samuels et al. 1993, Ferreira et al. 1997,
Alvarez et al. 2016). Several Coniella species are host-specific, contradictory to their wide host
range (Alvarez et al. 2016, Chethana et al. 2017). This genus has been subjected to many revisions.
Initially, Coniella and Pilidiella were given distinct identities due to their conidial pigmentation
(von Arx 1973, 1981b). Conidial pigmentation was rejected as a distinguishing character and
Pilidiella was synonymized under Coniella (Sutton 1980, Nag Raj 1993). With molecular data used
in species delimitation, Coniella and Pilidiella were considered as distinct genera (Castlebury et al.
2002, van Niekerk et al. 2004, Wijayawardene et al. 2016b). In an attempt to resolve many species
complexes residing in the genus, Alvarez et al. (2016) accepted Coniella as the only genus in
Schizoparmaceae and synonymized Pilidiella and Schizoparme. The most recent phylogenetic
status of this genus is by Chethana et al. (2017). Coniella destruens and Coniella vitis are illustrated
in this study (Figs. 225, 226).
Figure 225 – Coniella destruens (redrawn from van Niekerk et al. 2014). a Ascus and ascospores
of Coniella destruens. b Conidiophores and conidia of C. destruens. Scale bars: a, b = 10 µm.
Scortechiniaceae Huhndorf, A.N. Mill. & F.A. Fernández, Mycol Res 108(12): 1387 (2004)
Index Fungorum number: IF82146; Facesoffungi number: FoF01123; 33 species.
Saprobic on woody substrates in terrestrial habitats. Sexual morph: Ascomata perithecial,
scattered or gregarious, superficial, semi-immersed or immersed in host, sitting on or in a
subiculum or subiculum absent, globose, obpyriform or ovoid, dark brown to black, coriaceous to
membranous, turbinate or tuberculate or smooth, with or without brown branched setae, collabent
or not collapsing. Subiculum thin or thick, brown to dark brown, septate, crisp when dry, branched
or unbranched hyphae, with spines or smooth. Peridium thick, Munk pores present or absent, outer
layer composed of black to brown cells of textura angularis, thick-walled; inner layer composed of
hyaline cells of textura prismatica, thin-walled. Hamathecium Quellkörper present, conical, wide at
the base, paraphyses absent or indistinct, filiform, hyaline. Asci 8- to multi-spored, unitunicate,
cylindrical to clavate, long or short pedicellate, apical ring J-, indistinct or absent, evanescent.
Ascospores 2-seriate to irregularly arranged, hyaline or light brown, allantoid, ellipsoid to oval,
straight to slightly curved, aseptate, with or without appendages. Asexual morph: Conidiophores
erect, solitary, straight to flexuous. Conidiogenous cells hyaline, smooth, subcylindrical. Conidia
aseptate, hyaline, smooth, ellipsoid to ovoid, granular (adapted from Huhndorf et al. 2004a).
854
Type genus – Scortechinia Sacc.
Notes – Scortechiniaceae was introduced by Huhndorf et al. (2004a) based on superficial,
black, turbinate ascomata with a quellkörper and clavate asci with hyaline ascospores, as well as on
LSU sequence analyses. The Quellkörper is believed to play a role to rupture the ascoma and
probably is involved in ascospore discharge (Mugambi & Huhndorf 2010). This structure was
accepted as a main character to define this family when other morphs were not significant
(Huhndorf et al. 2004a, Mugambi & Huhndorf 2010, Carneiro de Almeida et al. 2016). Huhndorf et
al. (2004a) accepted Euacanthe, Neofracchiae and Scortechinia as the members in the family based
on LSU sequence data. Mugambi & Huhndorf (2010) added Biciliospora, Coronophorella,
Cryptosphaerella, Scortechiniella, Scortechiniellopsis, and Tympanopsis in Scortechiniaceae based
on multi-gene phylogenetic analyses. Crous et al. (2013) introduced the monotypic genus,
Pseudocatenomycopsis, which closely related to Neofracchiaea, Cryptosphaerella and
Scortechiniellopsis based on LSU sequence analyses. Members of Scortechiniaceae constitute a
strongly supported monophyletic group in Fig. 11. In this entry we illustrate Neofracchiaea callista.
Most members of this family have molecular support for their inclusion, but several of the genera
are monotypic.
Figure 226 – Coniella vitis (Material examined – RUSSIA, Rostov Region, Shakhty City, on dead
and dying branch of Salix alba L. (Salicaceae), 5 March 2016, T.S. Bulgakov, T1278 (MFLU 16–
1572). a Host tissue. b Submerged pycnidia on PDA. c Close view of the pycnidia and the spore
mass. d Conidiogenous cells. e Hyaline to brown conidia. f, g Upper view (f) and the reverse view
(g) of the colony on the PDA. Scale bars: a, b = 1 mm, c = 500 µm, d = 5 µm, e = 10 µm.
Ecological and economic significance of Scortechiniaceae
Most Scortechiniaceae species are saprobic on wood and widespread worldwide (Saccardo
1882, Saccardo & Berlese 1885, Höhnel 1909a, Petrak 1952a).
Genera included in Scortechiniaceae
Biciliospora Petr., Sydowia 6(5-6): 429 (1952)
Index Fungorum number: IF567; 1 species with sequence data.
Type species – Biciliospora velutina Petr.
855
Notes – Biciliospora velutina has tuberculate ascomata and cylindrical asci with ellipsoid
ascospores with filiform appendage at each ends (Petrak 1952a). Biciliospora was accepted as a
member of Coronophoraceae or synonym of Nitschkia in Nitschkiaceae (Müller & von Arx 1973,
Nannfeldt 1975, Subramanian & Sekar 1990). Mugambi & Huhndorf (2010) however, reported it
was a member of Scortechiniaceae based on multi-gene analysis.
Figure 227 – Phylogenetic tree generated by maximum likelihood analysis of combined ITS, LSU,
histone and tef1 sequence data of Coniella species. Related sequences were obtained from
GenBank. Thirty-three strains are included in the analyses, which comprise 2876 characters
including gaps. The tree was rooted with Melanconiella sp. (CBS 110385). Tree topology of the
ML analysis was similar to the MP and BI. The best scoring RAxML tree with a final likelihood
value of -15134.164466 is presented. The matrix had 807 distinct alignment patterns, with 23.87%
of undetermined characters or gaps. Estimated base frequencies were as follows; A = 0.249426, C =
0.245786, G = 0.256290, T = 0.248498; substitution rates AC = 0.996078, AG = 2.077237, AT =
1.158367, CG = 0.708887, CT = 4.169021, GT = 1.000000; gamma distribution shape parameter α
856
= 0.145978. The maximum parsimonious dataset consisted of constant 2083, 512 parsimonyinformative and 282 parsimony-uninformative characters. The parsimony analysis of the data
matrix resulted in the maximum of five equally most parsimonious trees with a length of 2495 steps
(CI = 0.532, RI = 0.583, RC = 0.310, HI = 0.468) in the first tree. RAxML and maximum
parsimony bootstrap support values ≥ 60% (black) and Bayesian posterior probabilities ≥ 0.90 (red)
are shown near the nodes. The scale bar indicates 0.1 changes. The ex-type strains are in bold.
Coronophorella Höhn., Sber. Akad. Wiss. Wien, Math.-naturw. Kl., Abt. 1 118: 1507 (1909)
Index Fungorum number: IF1255; 1 species with sequence data.
Type species – Coronophorella chaetomioides (Penz. & Sacc.) Höhn.
Notes – Coronophorella chaetomioides is characterised by reniform ascospores and is
considered as a member of Coronophoraceae (Nannfeldt 1975). It was accepted as a member of
Scortechiniaceae based on multi-gene analysis by Mugambi & Huhndorf (2010).
Cryptosphaerella Sacc., Syll. fung. (Abellini) 1: 186 (1882)
Index Fungorum number: IF1329; 18 morphological species (Species Fungorum 2020), 6
species with sequence data.
Type species – Cryptosphaerella nitschkei (Auersw.) Sacc.
Notes – Cryptosphaerella has multi-spored asci with allantoid ascospores (Saccardo 1882).
This genus was considered as a synonym of Coronophora (Nannfeldt 1975) or a member of
Coronophoraceae (Subramanian & Sekar 1990). It was clarified as an independent genus of
Scortechiniaceae based on multi-gene analysis (Mugambi & Huhndorf 2010). The monophyletic
Cryptosphaerella is closely related to Biciliospora, Scortechiniella and Scortechiniellopsis (Fig.
11).
Euacanthe Theiss., Annls mycol. 15(3/4): 272 (1917)
Index Fungorum number: IF1906; 1 species with sequence data.
Type species – Euacanthe usambarensis (Rehm) Theiss.
Notes – Euacanthe was established and considered as a synonym of Acanthonitschkea,
Nitschkiaceae or a member of Coronophoraceae (Theissen 1917, Nannfeldt 1975, Subramanian &
Sekar 1990). A second species, E. foveolata (≡ Sphaeria foveolata) was accepted as a member of
Scortechiniaceae based on multi-gene analysis (Subramanian & Sekar 1990, Mugambi & Huhndorf
2010).
Neofracchiaea Teng, Sinensia, Shanghai 9: 255 (1938)
Index Fungorum number: IF3454; 1 species with sequence data.
Type species – Neofracchiaea callista (Berk. & M.A. Curtis) Teng
Notes – Neofracchiaea callista has multi-spored asci with ellipsoid ascospores (Teng 1938)
and was treated as synonym of Nitschkia (Nannfeldt 1975). Mugambi & Huhndorf (2010) accepted
it as a member of Scortechiniaceae based on multi-gene analysis.
Pseudocatenomycopsis Crous & L.A. Shuttlew., Persoonia 31: 221 (2013)
Index Fungorum number: IF805836; 1 species with sequence data.
Type species – Pseudocatenomycopsis rothmanniae Crous & L.A. Shuttlew.
Notes – Pseudocatenomycopsis rothmanniae has hyaline, ovoid, granular conidia.
Pseudocatenomycopsis, the only asexual genus of Scortechiniaceae nests in clade of
Scortechiniaceae (Fig. 11).
Scortechinia Sacc., Atti Inst. Veneto Sci. lett., ed Arti, Sér. 6 3: 713 (1885)
Index Fungorum number: IF4968; 3 morphological species (Species Fungorum 2020), 2
species with sequence data.
Type species – Scortechinia acanthostroma (Mont.) Sacc. & Berl.
857
Notes – Scortechinia is characterized by tuberculate ascomata with a quellkörper, clavate asci
and obovate ascospores (Huhndorf et al. 2004a). It nested in the clade of Scortechiniaceae and is
related to Neofracchiaea callista (Fig. 11).
Figure 228 – Neofracchiaea callista (Material examined – USA, Maryland, Beltsville; on wood,
October 1950, F. Petrak, PDD 54147). a Material label. b Specimen c Ascomata subimmersed in
subiculum on substrate. d Collapsing ascoma. e Section through ascoma with quellkörper
(arrowhead). f Peridium. g, h Asci. i Ascospores. Notes: figs. e, g-h are stained in Congo red
reagent. Scale bars: c = 1 cm, d-e = 100 µm, f = 50 µm, g-i = 20 µm.
Scortechiniella Arx & E. Müll., Beitr. Kryptfl. Schweiz 11(no. 1): 382 (1954)
Index Fungorum number: IF4969; 1 species with sequence data.
Type species – Scortechiniella similis (Bres.) Arx & E. Müll.
Notes – Scortechiniella similis has multi-spored asci and obovate ascospores with filiform
appendages at each end (von Arx & Müller 1954) and was accepted as a member of
Coronophoraceae (Müller & von Arx 1973, Nannfeldt 1975). Mugambi & Huhndorf (2010)
however, accepted it as a member of Scortechiniaceae based on multi-gene analysis.
858
Scortechiniellopsis Sivan., Trans. Br. mycol. Soc. 62(1): 36 (1974)
Index Fungorum number: IF4970; 1 species with sequence data.
Type species – Scortechiniellopsis leonensis Sivan.
Notes – Scortechiniellopsis leonensis has multi-spored asci and reniform ascospores
(Sivanesan 1974) and was accepted as a synonym of Nitschkia (Nannfeldt 1975). Mugambi &
Huhndorf (2010) however, accepted it as a member of Scortechiniaceae based on multi-gene
analysis.
Tympanopsis Starbäck, Bih. K. svenska VetenskAkad. Handl., Afd. 3 19(no. 2): 24 (1894)
Index Fungorum number: IF5651; 5 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Tympanopsis euomphala (Berk. & M.A. Curtis) Starbäck
Notes – Tympanopsis euomphala is characterised by pyriform asci with ellipsoid ascospores
and was considered as a member of Coronophoraceae (Starbäck 1894, Nannfeldt 1975, von Arx
1981a). It was considered as a member of Scortechiniaceae based on multi-gene analysis by
Mugambi & Huhndorf (2010).
Sordariaceae G. Winter, Rabenh. Krypt.-Fl., Edn 2 (Leipzig) 1.2: 162 (1885)
Index Fungorum number: IF81384; Facesoffungi number: FoF01148; 113 species.
Saprobic on wood, rotting vegetation and dung in terrestrial habitats, also associated with
food. Sexual morph: Ascomata perithecial or cleistothecial, brown to black, gregarious or scattered,
solitary, superficial, erumpent or immersed, globose, subglobose to ovoid, carbonaceous,
coriaceous or membranaceous, tuberculate or smooth, with setae or hairs, papillate or papilla
indistinct or absent, ostiolate, with hyaline periphyses or periphyses absent. Peridium relatively
thick, carbonaceous, coriaceous or membranaceous, comprising brown to hyaline cells of textura
angularis or globulosa or prismatica. Paraphyses numerous, septate, un- or branched, filiform or
cylindrical or absent. Asci 8-spored, unitunicate, cylindrical to clavate, pedicellate, J-. Ascospores
uniseriate or biseriate, hyaline, yellowish, brown or black, stellate, oval or ellipsoid, 0–1-septate,
concolorous or versicolorous, with or without guttules, uneven in thickness with striate to reticulate
patterns or with a sheath, with or without germ pores, appendages present or absent. Asexual
morph: Undetermined (adapted from Maharachchikumbura et al. 2016b).
Type genus – Sordaria Ces. & De Not.
Notes – Sordariaceae was introduced by Winter (1885b) based on brown to black ascomata
and cylindrical asci with brown to black ascospores, and is typified by Sordaria. Most species in
this family are coprophilous and can be heterothallic, homothallic or pseudo-homothallic (Dutta et
al. 1976, Cai et al. 2006c). Species of Sordariaceae have unique ascospores and Copromyces,
Effetia, Guilliermondia and Stellatospora are coprophilous taxa but without sequence data.
Ascospores with glutinous sheaths or/and special ornamentations on the wall were considered to be
similar characters as found in Sordariaceae and Lasiosphaeriaceae (Lundqvist 1972, Huhndorf et al.
2004b, Cai et al. 2006c). Multi-gene analysis placed Neurospora, Pseudoneurospora and Sordaria
in Sordariaceae and related to Lasiosphaeriaceae (Huhndorf et al. 2004b, Cai et al. 2006b, c,
Maharachchikumbura et al. 2015b, 2016b, Hyde et al. 2017a). In this entry we illustrate
Guilliermondia saccoboloides Boud.
Ecological and economic significance of Sordariaceae
Most species of Sordariaceae were isolated from dung of herbivores and are saprobes
important in nutrient cycling. Some of the homothallic species has a long-standing history as model
organisms, viz. Neurospora crassa, Sordaria fimicola and Sordaria macrospora (Kück et al. 2009,
Thatoi et al. 2013, Froyd et al. 2014, Kuo et al. 2014, Newcombe et al. 2016).
Genera included in Sordariaceae
Copromyces N. Lundq., Ark. Bot., Ser. 2 6: 327 (1967)
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Index Fungorum number: IF1230; 2 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Copromyces bisporus N. Lundq.
Notes – Copromyces has 2-spored asci and globose ascospores (Lundqvist 1967). Huhndorf
et al. (2004b) and Vu et al. (2019) used LSU and ITS sequence data from Copromyces octosporus
(CBS 386.78) which was provided by J.C. Krug (TRTC 51747).
Effetia Bartoli, Maggi & Persiani, Mycotaxon 19: 517 (1984)
Index Fungorum number: IF25588; 1 morphological species.
Type species – Effetia craspedoconidica Bartoli, Maggi & Persiani
Notes – The monotypic genus Effetia is characterized by aseptate, oval conidia with sheaths
and has irregular, brown chlamydospores (Bartoli et al. 1984).
Guilliermondia Boud., Bull. Soc. mycol. Fr. 20(1): 19 (1904)
Index Fungorum number: IF2150; 1 morphological species.
Type species – Guilliermondia saccoboloides Boud.
Notes – This monotypic genus Guilliermondia has sphaerical asci and ellipsoid ascospores
surrounded by a brown sheath (Boudier 1904).
Figure 229 – Guilliermondia saccoboloides (Material examined – FRANCE, Lyon, 69 Rhôneon,
on horse dung, April 1903, MAA Guilliermond, PC 0167691, holotype). a Label and material. b
Ascomata. c-e Asci. f-i Ascospores. Scale bars: b = 500 µm, c-e = 5 µm, f-i = 2 µm.
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Neurospora Shear & B.O. Dodge, J. Agric. Res., Washington 34: 1025 (1927)
Index Fungorum number: IF3502; 56 morphological species (Species Fungorum 2020), 35
species with sequence data.
Type species – Neurospora sitophila Shear & B.O. Dodge
Notes – Neurospora is the largest genus in Sordariaceae. It is characterized by cylindrical asci
and brown ascospores with striate to reticulate patterns on the wall. Gelasinospora was accepted as
a synonym of Neurospora based on LSU analysis (García et al. 2004).
Pseudoneurospora Dania García, Stchigel & Guarro, Mycol. Res. 108(10): 1139 (2004)
Index Fungorum number: IF28887; 2 species with sequence data.
Type species – Pseudoneurospora amorphoporcata (Udagawa) Dania García, Stchigel &
Guarro
Notes – Gelasinospora amorphoporcata was transferred to Pseudoneurospora as the type
species (García et al. 2004) and a second species P. canariensis was introduced by Crous et al.
(2014a). Species have brown ascomata with cylindrical asci and ellipsoid ascospores with an
epispore with an irregular pattern (García et al. 2004, Crous et al. 2014a).
Sordaria Ces. & De Not., Comm. Soc. crittog. Ital. 1(4): 225 (1863)
Index Fungorum number: IF5061; 50 morphological species (Species Fungorum 2020), 15
species with sequence data.
Type species – Sordaria fimicola (Roberge) Ces. & De Not.
Notes – This mainly coprophilous genus has perithecial ascomata, cylindrical asci and brown
ascospores, some with appendages, sheaths and/or pores and may also be saprobic on wood (Barr
1990b, Maharachchikumbura et al. 2016b). Species such as S. fimicola occur on herbaceous plants,
wood and soil (Lundqvist 1972, Chambers & De Wet 1987, Alma et al. 2000, Kavak 2012, Ivanová
2015). A new species, S. clematidis, will be introduced from the UK on Clematis by Phukhamsakda
et al. (2020).
Stellatospora Tad. Ito & Nakagiri, Mycoscience 35(4): 413 (1994)
Index Fungorum number: IF27456; 1 morphological species.
Type species – Stellatospora terricola Tad. Ito & Nakagiri
Notes – The monotypic genus Stellatospora was isolated from soil and is characterized by
obpyriform asci with stellate ascospores (Ito & Nakagiri 1994).
Spathulosporaceae Kohlm., Mycologia 65(3): 615 (1973)
Index Fungorum number: IF81388; Facesoffungi number: FoF01798; 6 species.
Parasitic on marine algae. Sexual morph: Ascomata subglobose, ovoid, pyriform, subiculate,
coriaceous or leathery, dark brown, sterile hairs enclosing ascoma, ostiolate, ostiolar canal
surrounded by tube-like projection extending into the ascomatal cavity, papillate or epapillate.
Paraphyses lacking. Asci 8-spored, clavate to subglobose, thin-walled, unitunicate, without apical
ring, deliquescing early. Ascospores overlapping multi-seriate, hyaline, fusiform, cylindrical or
ellipsoidal, aseptate to 3-septate, with apical appendages. Reproductive structures: Antheridial.
Spermatia ellipsoidal to fusiform, without appendages. Trichogynes simple or branched, septate,
arising from the margin of young ascoma (adaped from Maharachchikumbura et al. 2016b).
Type genus – Spathulospora A.R. Caval. & T.W. Johnson
Notes – Spathulosporaceae is characterized by crustose, dark thalli, peg-like individual
penetrating cells, intracellular crusts and stromata, antheridia with spermatia and trichogynes,
ostiolate ascomata without paraphyses, deliquescing asci, and 1-celled, appendaged ascospores.
The genus was introduced by Cavaliere & Johnson (1965) and currently comprises Spathulospora
and Retrostium with six species (Nakagiri & Ito 1997, Index Fungorum 2020, Jones et al. 2019).
However, sequence data are available only for S. antarctica and S. adelpha. This family was first
placed in Spathulosporales then so as to read and then in Lulworthia (Kohlmeyer 1973, Inderbitzin
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et al. 2004, Campbell et al. 2005, Jones et al. 2009). However, Maharachchikumbura et al. (2015,
2016b) confirmed the placement of Spathulosporaceae in Spathulosporales by molecular analysis
using available sequence data. Spathulospora was introduced by Cavaliere & Johnson (1965) and
referred to the Spathulosporomycetes, Spathulosporomycetidae (Locquin 1984), Spathulosporales
(Kohlmeyer 1973), Spathulosporaceae (Kohlmeyer 1973). It is a marine genus on the seaweed
Ballia spp. (Rhodophyta), and is typified by Spathulospora phycophila. However, two
Spathulospora species (S. antarctica, S. adelpha), have shown a relationship to Lulworthiales, but
with low support (Inderbitzin et al. 2004, Campbell et al. 2005, Jones et al. 2009, 2019). There are
no sequences for the type of this genus, hence, new collections, isolations and sequencing are
required to determine its phylogenetic placement.
Ecological and economic significance of Spathulosporaceae
Spathulospora species are obligate parasites on some marine algae (Ballia spp.) and may
cause malformations. They proliferate in the host cells without penetrating the plasma membrane,
and only afterwards form the stroma and sexual structures on the surface, presumably using
nutrients acquired from the host. In some cases, the interaction of species of Spathulospora and
host triggers a reaction analogous to witches’ broom in vascular plants: the alga produces long
hairs, which surround the stromata and ascomata of the fungus (Inderbitzin et al. 2004). Retrostium
species inhabit marine Rhodophytes in coastal habitats. This fungus forms mononematous
spermatiophores with penicillate phialides and trichogynes at the edge of ascomata on the surface
of the host alga and consume nutrients (Nakagiri & Ito 1997).
Genera included in Spathulosporaceae
Retrostium Nakagiri & Tad. Ito, Mycologia 89(3): 485 (1997)
Index Fungorum number: IF27834; 1 morphological species.
Type species – Retrostium amphiroae Nakagiri & Tad. Ito
Notes – This genus was introduced by Nakagiri & Ito (1997) to accommodate R. amphiroae,
a taxon inhabiting Amphiroa zonata (marine Rhodophyta) in coastal habitats in Japan (Nakagiri &
Ito1997). Molecular data are unavailable for the species of this genus. Ascomata of R. amphiroae
are conical to hemisphaerical, superficial, carbonaceous, epapillate and ostiolate (Nakagiri & Ito
1997). The ostiolar canal is surrounded by a tube-like projections extending into the ascomatal
cavity and the two-layered peridial structure is similar to that of Spathulospora (Nakagiri & Ito
1997). This taxon differs from Spathulospora by thalli, ascomatal structure, ascospores and host
(Kohlmeyer 1973, Nakagiri & Ito 1997). Retrostium amphiroae has true hyphae that grow between
host cells, unlike Spathulospora, which have crustose thalli with "intramural hyphae" that invade
the host cells (Kohlmeyer 1973). Paraphyses are absent, but pseudoparenchyma cells are present,
similar to S. lanata (Kohlmeyer & Kohlmeyer 1975). Ascospores are 1-celled, oblong and
furnished with a mucilaginous appendage at each end, while some Spathulospora species have
spathulate ascospores (Kohlmeyer & Kohlmeyer 1975). Antheridium of R. amphiroae is a
mononematous spermatiophore rather than a spermodochium as in Hispidicarpomyces and
spermatia lack appendages as in Hispidicarpomyces, but can be distinguished by shape, as the latter
forms a clavate spermatium with a cupulate base (Nakagiri & Ito 1997).
Spathulospora A.R. Caval. & T.W. Johnson, Mycologia 57(6): 927 (1965)
Index Fungorum number: IF5071; 5 morphological species (Species Fungorum 2020), 2
species with sequence data.
Type species – Spathulospora phycophila A.R. Caval. & T.W. Johnson
Notes – Spathulospora phycophila was illustrated in Maharachchikumbura et al. (2016b) and
thus is not illustrated here.
Sporidesmiaceae Fr., Summa veg. Scand., Section Post. (Stockholm): 504 (1849)
Index Fungorum number: IF81405; Facesoffungi number: FoF01830; 188 species.
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Figure 230 – Retrostium amphiroae (Material examined – JAPAN, Honshu: Shizuoka Pref.,
Shimoda, on living thalli of Amphiroa zonata, 30 June 1992, A. Nakagiri, IFO H-12210, holotype)
a Herbarium material. b Host. c Ascomata on host surface. d Section through ascoma (arrow
indicates tube-like projection extending into the ascoma cavity). e Peridium. f Ascus. g-i
Ascospores. j Broken pieces of outgrowths attached to the ascomata outer wall. k Spermatiophores
arising from the margin of young ascoma. Scale bars: d = 100 μm. e = 50 μm, f, i = 20 μm, g-i = 10
μm, j = 5 μm.
Saprobic on woody debris in terrestrial and aquatic habitats or mycoparasites. Sexual morph:
Ascomata perithecial, scattered or solitary, immersed with neck erumpent through host surface,
uniloculate, subglobose or ellipsoidal. Neck cylindrical, central or lateral. Ostiolum periphysate.
Peridium three-layered. Paraphyses numerous, hypha-like, hyaline, septate, unbranched, persistent,
embedded in a gelatinous matrix. Asci 8-spored, unitunicate, long cylindrical, apically rounded,
with a distinct, relatively small, refractive, J-, wedge-shaped, apical ring. Ascospores obliquely uniseriate, hyaline, fusiform, septate, with a thin sheath. Asexual morph: Colonies black, effuse.
Mycelium comprising hyaline to pale brown, branched, septate, thin-walled hyphae. Conidiophores
solitary or in clusters, single, or sometimes aggregated in groups, brown, cylindrical, straight or
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curved, erect or somewhat repent, septate, smooth-walled. Conidiogenous cells monoblastic,
holoblastic, terminal, cylindrical, brown, elongating once or twice percurrently at the apex. Conidia
dry, solitary, medium brown, smooth, obclavate to cylindrical or fusoid, straight to flexuous, apex
obtuse, base obconically truncate, smooth-walled.
Type genus – Sporidesmium Link
Notes – Sporidesmiaceae was introduced by Fries (1849) and has received very little usage.
Presently, there is only one genus accepted in this family. Shenoy et al. (2006) found that
Sporidesmium and morphologically similar genera are not monophyletic, as they are distributed
among different families and orders in Dothideomycetes and Sordariomycetes. Su et al. (2016b)
reported similar results. A monophyletic clade with species resembling the type species of
Sporidesmium was designated as Sporidesmiaceae sensu stricto, and a new family
Distoseptisporaceae was introduced to accommodate a new sporidesmium-like genus
Distoseptispora (Su et al. 2016b).
Ecological and economic significance of Sporidesmiaceae
Saprobes have the ability to decompose lignocellulose in woody litter, resulting in the
softening of the wood and release of nutrients in the form of simple molecules that are reused by
plants and other organisms (Yuen et al. 1998, Bucher et al. 2004). Thus, they play an important role
in nutrient and carbon cycling, biological diversity and ecosystem functioning (Palmer et al. 1997,
Wong et al. 1998a).
Genus included in Sporidesmiaceae
Sporidesmium Link, Mag. Gesell. naturf. Freunde, Berlin 3(1–2): 41 (1809)
Index Fungorum number: IF10024; 188 morphological species (Species Fungorum 2020),
17 species with sequence data.
Type species – Sporidesmium atrum Link
Notes – Sporidesmium is a large and heterogeneous genus. To date, 383 epithets are referred
to the genus (Index Fungorum 2020). However, many previously described species were revised
and transferred to over 30 genera (Iturriaga et al. 2008). Studies based on phylogenetic analysis
have been carried out to further re-examine the classification of sporidesmium-like taxa, given that
the generic delimitations based on morphology data appear to be questionable. Sporidesmium and
morphologically similar genera are clearly not monophyletic, as they are distributed among
different families and orders in Dothideomycetes and Sordariomycetes (Shenoy et al. 2006, Su et
al. 2016b, Yang et al. 2017). Sporidesmium was an asexual genus until Zhang et al. (2017a)
introduced a sexual morph species Sporidesmium thailandense which is collected from freshwater
habitat in Thailand. In this entry, we introduce a new species Sporidesmium dulongense based on
morphology and phylogeny. It was collected from a freshwater river in northwestern Yunnan
Province, China.
Sporidesmium dulongense Z.L. Luo, K.D. Hyde & H.Y. Su, sp. nov.
Fig. 231
Index Fungorum number: IF555377; Facesoffungi number: FoF04924
Etymology – Referring to the collect location of the fungus.
Holotype – HKAS 92659.
Saprobic on submerged decaying wood. Sexual morph: Undetermined. Asexual morph:
Colonies effuse on natural substrate, scattered, hairy, dark brown to black. Mycelium immersed,
composed of septate, branched, brown, smooth hyphae. Conidiophores 88–124 μm (x̅ = 106 μm,
SD = 18, n = 25), 5.5–7.5 μm (x̅ = 6.5 μm, SD = 1, n = 25), macronematous, mononematous,
unbranched, erect, straight or flexuous, dark brown, septate, smooth. Conidiogenous cells
monoblastic, holoblastic, terminal, determinate, dark brown, cylindrical. Conidia 50–58 μm ( x =
54 μm, SD = 4, n = 30), 13–15 μm ( x = 14 μm, SD = 1, n = 30), acrogenous, solitary, dry,
lageniform, straight or slight curved, tapering and hyaline at apex, truncate at base, dark brown,
septate, with sphaerical guttules in almost all cells, smooth-walled.
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Figure 231 – Sporidesmium dulongense (HKAS 92659, holotype). a Colonies on wood. b-d
Conidiophore with conidia. e, f Conidiogenous cells. g-i Conidia. j Germinating conidium. Scale
bars: b = 55 μm, c, d = 45 μm, e, h, j = 25 μm, f-g, i= 30 μm.
Material examined – CHINA, Yunnan Province, saprobic on decaying wood submerged in
Dulong River, May 2015, Z.L. Luo S-436 (HKAS 92659, holotype), ex-type living culture
MFLUCC 17-0116.
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GenBank numbers – ITS: MH795812, LSU: MH795817, rpb2: MH801190, tef1: MH801191.
Notes – Sporidesmium dulongense is most similar to S. submersum in having unbranched,
dark brown, septate conidiophores and acrogenous, solitary, pyriform, septate conidia with
sphaerical guttules in almost all cells. However, S. dulongense differs from S. submersum by its
longer conidiophores (88–124 μm vs. 59–72 μm) and lageniform conidia with a long hyaline apex.
Phylogenetic analysis based on multi-genes also showed that they are distinct species in
Sporidesmium.
Sporocadaceae Corda, Icon. Fungorum (Prague) 5: 34 (1842)
Index Fungorum number: IF81408; Facesoffungi number: FoF06111; 750 species.
Saprobic or pathogenic on leaves, twigs, branches, fruits of flowering plants and
gymnosperms, as endophytes or parasitic on humans and animals. Sexual morph: Ascomata
scattered or confluent, perithecial, immersed to erumpent, black to dark brown. Ostiole circular,
papillate. Peridium comprising several layers of cells of textura prismatica, brown at the base, dark
brown outwardly. Paraphyses numerous, hypha-like, flexuose, filiform. Asci 8-spored, unitunicate,
obclavate to cylindrical, with or without a discoid, J+ subapical ring. Ascospores overlapping
uniseriate or biseriate, obovoid to ellipsoid, transversally septate, longitudinal septum in mid cells,
with or without vertical septa, hyaline, pale brown to brown. Asexual morph: Coelomycetous.
Conidiomata scattered to gregarious, subepidermal or subepidermal in origin, immersed to
erumpent, pycnidial, acervular, stromatic or synnematous in Synnemapestaloides. Conidiomata
wall composed of at least 3–4 layers of cells of textura angularis to textura prismatica, outer 1–2layers brown and inner 1–2 layers hyaline 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, hyaline, smooth. Conidiogenous cells holoblastic, annellidic, ampulliform,
cylindrical to subcylindrical, or lageniform, mostly hyaline. Conidia cylindrical to clavate, or
fusiform, straight or curved, 2–4-euseptate, smooth or verruculose, median cells hyaline, pale
olivaceous or pale brown to dark brown, bearing cellular, terminal cells mostly hyaline, appendages
absent or if present, tubular, filiform, straight or flexuous, branched or unbranched (adapted from
Maharachchikumbura et al. 2016b, Liu et al. 2019a).
Type genus – Sporocadus Corda
Notes – Sporocadaceae includes genera that are typically appendaged bearing coelomycetes
and have been subjected to several taxonomic re-classifications (Nag Raj 1993,
Maharachchikumbura et al. 2016b, Liu et al. 2019a). The family was re-validated by Jaklitsch et al.
(2016b) and Bartaliniaceae, Discosiaceae, Pestalotiopsidaceae and Robillardaceae (Crous et al.
2015a, Senanayake et al. 2015) were treated as synonyms of Sporocadaceae. In addition to the
phylogeny, morphology of the asexual morph genera having acervular conidiomata that produce
hyaline, pale or dark brown, septate conidia were taken into the consideration when they were
assigned to the family. Jaklitsch et al. (2016b) and Wijayawardene et al. (2018a) accepted 22
genera under Sporocadaceae.
Studies on Sporocadaceae were mostly based on ITS and LSU sequence data and these
regions were not informative in resolving generic boundaries within the family (Jaklitsch et al.
2016b, Liu et al. 2019a). The most recent study by Liu et al. (2019a) provided a revision of this
family complete with morphology and multi-gene phylogeny based on the LSU, ITS and rpb2
sequence data and further analysis using protein coding genes (tef1 or tub2) for each genus. As a
result, seven new genera were introduced and the placement of 23 known genera were confirmed.
Here, we follow the treatments and updated classification as presented by Liu et al. (2019a). The
accepted genera are listed below with illustrations and a description with two new host records of
Sporocadaceae.
Ecological and economic significance of Sporocadaceae
Spodocadaceae taxa are ecologically important as they occur as saprobes, endophytes or
pathogens, on leaves, twigs, branches and fruits on a wide range of host plants (Nag Raj 1993,
866
Hyde et al. 2017b, Liu et al. 2019a). A number of species, especially related to Pestalotiopsis,
Neopestalotiopsis, Pseudopestalotiopsis and Truncatella are considered to be destructive pathogens
diseases in several economically important crops (Maharachchikumbura et al. 2016b). Furthermore,
commonly isolated endophytes of Spodocadaceae produce a wide range of chemically novel
diverse metabolites (Maharachchikumbura et al. 2014a, b, Liu et al. 2019a)
Genera included in Sporocadaceae
Allelochaeta Petr., Sydowia 9(1-6): 464 (1955)
Index Fungorum number: IF7095; 41 morphological species (Species Fungorum 2020), 37
species with sequence data.
Type species – Allelochaeta gaubae Petr.
Notes – Allelochaeta was considered as a synonym of Diploceras, one of the five groups of
fungi into which the Seimatosporium complex was split by Nag Raj (1993). Based on a multi-gene
phylogenetic study using the type species, Crous et al. (2018a) resurrected Allelochaeta. Members
of this genus are characterized by mostly 3-septate, hyaline or pigmented, concolourous or
versicolourous conidia, having branched or solitary, cellular or continuous appendages (Crous et al.
2018a). Crous et al. (2018a) discussed the conidial septation, appendage types and pigmentation
loss and gain during the evolution of Allelochaeta species.
Annellolacinia B. Sutton, Mycol. Pap. 97: 31 (1964)
Index Fungorum number: IF7161; 2 morphological species (Species Fungorum 2020).
Type species – Annellolacinia dinemasporioides B. Sutton
Notes – The coelomycete genus Annellolacinia was introduced by Sutton (1964) and is
characterized by conidia that are fusiform, aseptate, bearing a single, cellular, unbranched
appendage at each end (Nag Raj 1993). Wijayawardene et al. (2016b) tentatively placed the genus
in Discosiaceae (now Sporocadaceae) based on its morphological similarities, and we follow the
same. However, there are no cultures or sequence data to confirm the placement of Annellolacinia.
Bartalinia Tassi, Bulletin Labor. Orto Bot. de R. Univ. Siena 3: 4 (1900)
Index Fungorum number: IF7327; 21 morphological species (Species Fungorum 2020), 7
species with sequence data.
Type species – Bartalinia robillardoides Tassi
Notes – Bartalinia was established by Tassi (1990) and is characterized by 3–4-septate
conidia (Nag Raj 1993). They have a worldwide distribution, occurring as saprobes on various
shrubs and trees and are often associated with leaf spots (Wong et al. 2003, Maharachchikumbura
et al. 2016b, Wijayawardene et al. 2017a, Phookamsak et al. 2019, Farr & Rossman 2019). They
have also been recorded in freshwater (Nguyen et al. 2019). The genus was previously placed in
Bartaliniaceae by Senanayake et al. (2015) which was later synonymized under Sporocadaceae by
Jaklitsch et al. (2016b).
Broomella Sacc., Syll. fung. (Abellini) 2: 557 (1883)
Index Fungorum number: IF658; 14 morphological species (Species Fungorum 2020), 2
species with sequence data
Type species – Broomella vitalbae (Berk. & Broome) Sacc.
Notes – Broomella species have been mostly reported from Asia and Europe (Farr &
Rossman 2019), as saprobes from terrestrial habitats (Li et al. 2015d, Wijayawardene et al. 2018a).
The sexual morph is characterized by cylindric-elongate, unitunicate asci, with J-, discoid apical
ring, bearing 2-3-septate, ellipsoid-fusiform ascospores with brown, median and hyaline, terminal
cells bearing centric appendages at each end (Shoemaker & Müller 1963, Li et al. 2015d,
Wanasinghe et al. 2018, Liu et al. 2019a). The asexual morph is pestalotia-like and this is yet to be
confirmed by phylogeny (Wanasinghe et al. 2018, Liu et al. 2019a). Li et al. (2015d) provided an
epitype for Broomella vitalbe illustrating both its sexual and asexual morphs.
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Ciliochorella Syd., Annls mycol. 33(1/2): 62 (1935)
Index Fungorum number: IF7657; 7 morphological species (Species Fungorum 2020), 2
species with sequence data.
Type species – Ciliochorella mangiferae Syd.
Notes – Ciliochorella, introduced by Sydow & Mitter (1935) consists of saprobic species
reported from India, Japan, South America and Thailand (Tangthirasunun et al. 2015, Hyde et al.
2016b, Liu et al. 2019a, Farr & Rossman 2019), from terrestrial habitats. Sutton (1980) mentioned
Ciliochorella mangiferae as a common species found in both tropical and subtropical leaf litter.
Diploceras (Sacc.) Died., Mykol. Untersuch. Ber.: 342 (1915)
Index Fungorum number: IF8043; 2 morphological species (Species Fungorum 2020).
Type species – Diploceras hypericinum (Ces.) Died.
Notes – Diploceras was introduced as a subgenus of Hyaloceras by Saccardo (1892) and
subsequently raised to generic rank by Diedicke (1915). After re-examining the type, Sutton (1975)
re-classified this genus as a synonym of Seimatosporium. Nag Raj (1993) resurrected Diploceras as
a genus to accommodate D. hypericinum. Most other species previously classified as Diploceras
were later included in Allelochaeta during a study based on multi-gene phylogeny by Crous et al.
(2018a). An emended description for this genus was provided by Liu et al. (2019a).
Disaeta Bonar, Mycologia 20: 299 (1928)
Index Fungorum number: IF8066; 1 species with sequence data.
Type species – Disaeta arbuti Bonar
Notes – Disaeta was introduced by Bonar (1928) without mentioning the type species. The
genus is characteristic in producing 4-septate conidia, with bristle-like appendages (Bonar 1928).
Liu et al. (2019a) provided a representative strain of D. arbuti isolated from Acacia pycnantha,
Australia, along with an emended description and further citing the need for a taxonomic revision
of this genus.
Discosia Lib. ex Durieu & Mont., Fl. d'Algérie, Cryptog. 1: 587 (1849)
Index Fungorum number: IF8072; 50 morphological species (Species Fungorum 2020), 11
species with sequence data.
Type species – Discosia artocreas (Tode) Fr.
Notes – Species of Discosia are mostly saprobes and have been recorded worldwide from
terrestrial habitats on a various range of host plants (Farr & Rossman 2019). Based on phylogenetic
evidence, Liu et al. (2019a) synonymised Adiscio under Discosia, the former which was introduced
by Tanaka et al. (2011) to accommodate the sexual morph species of this genus. An epitype for the
type species was designated by Liu et al. (2019a) as the type specimen was destroyed.
Distononappendiculata F. Liu, L. Cai & Crous (2019)
Index Fungorum number: IF828323; 3 species with sequence data.
Type species – Distononappendiculata banksiae (Crous & Summerell) F. Liu, L. Cai &
Crous.
Notes – Distononappendiculata species are characteristic in producing distoseptate conidia
that lack appendages (Liu et al. 2019a). They occur on leaves of Banksia sp. in Australia (Liu et al.
2019a).
Diversimediispora F. Liu, L. Cai & Crous (2019)
Index Fungorum number: IF828326; 1 species with sequence data.
Type species – Diversimediispora humicola F. Liu, L. Cai & Crous.
Notes – Diversimediispora species produce conidia with versicolourous median cells where
the second and third cells (from the apex) are darker, making it characteristically different from
other genera in Sporocadaceae (Liu et al. 2019a).
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Doliomyces Steyaert, Darwiniana 12(2): 169 (1961)
Index Fungorum number: IF11007; 3 morphological species (Species Fungorum 2020).
Type species – Doliomyces senegalensis (Speg.) Steyaert
Notes – Doliomyces was initially placed in Amphisphaeriaceae (Kirk et al. 2008) but due to
the lack of sequence data or cultures, this placement was considered uncertain. Senanayake et al.
(2015) listed Doliomyces in Amphisphaeriales, genera incertae sedis. However, Wijayawardene et
al. (2016b) classified this genus under Bartaliniaceae (now Sporocadaceae), based on its
morphology. In this study we tentatively place Doliomyces in Sporocadaceae. Species of
Doliomyces are characterized by cylindrical to navicular, fusiform, 3–5-septate, conidia with dark
brown median cells, bearing a single, unbranched, basal appendage and several branched, apical
appendages (Nag Raj 1993, Wijayawardene et al. 2016b).
Heterotruncatella F. Liu, L. Cai & Crous (2019)
Index Fungorum number: IF828340; 17 species with sequence data.
Type species – Heterotruncatella lutea (H.J. Swart & D.A. Griffiths) F. Liu, L. Cai & Crous
Notes – Heterotruncatella was introduced by Liu et al. (2019a) to accommodate several new
and previously described Truncatella species. Although morphologically similar, phylogenetically
they were distinct from Truncatella. Most species have been recorded from Australia and Europe
(Senanayake et al. 2015, Liu et al. 2019a, Farr & Rossman 2019).
Hyalotiella Papendorf, Trans. Br. mycol. Soc. 50(1): 69 (1967)
Index Fungorum number: IF8588; 6 morphological species (Species Fungorum 2020), 3
species with sequence data.
Type species – Hyalotiella transvalensis Papendorf
Notes – Hyalotiella was introduced by Papendorf & Du-Toit (1967). Hyalotiella species are
characterized by vase-shaped pycnidia, producing 3-septate, cylindrical conidia bearing more than
one branched apical appendage but without a basal appendage (Nag Raj 1993, Li et al. 2015d,
Senanayake et al. 2016). They occur as saprobes, and have been recorded from India, Italy, South
Africa and South America (Nag Raj 1993, Li et al. 2015d, Farr & Rossman 2019).
Hymenopleella Munk, Dansk bot. Ark. 15(no. 2): 89 (1953)
Index Fungorum number: IF2416; 7 species with sequence data.
Type species – Hymenopleella hippophaëicola Jaklitsch & Voglmayr
Notes – Hymenopleella is a sexual morph genus recorded from Africa, China and India
(Jeewon et al. 2003a, Liu et al. 2019a). In the study by Liu et al. (2019a), the type species of
Dyrithiopsis and Neotruncatella clustered together with Hymenopleella and therefore both genera
were synonymized under the latter genus (Liu et al. 2019a).
Immersidiscosia Kaz. Tanaka, Okane & Hosoya, Persoonia 26: 94 (2011)
Index Fungorum number: IF519746; 1 species with sequence data.
Type species – Immersidiscosia eucalypti (Pat.) Kaz. Tanaka, Okane & Hosoya
Notes – The genus was introduced by Tanaka et al. (2011) to accommodate Immersidiscosia
eucalypti which has been recorded from France, Japan and Tunisia (Tanaka et al. 2011,
Wijeyawardene et al. 2017a, Farr & Rossman 2019).
Monochaetia (Sacc.) Allesch., Rabenh. Krypt.-Fl., Edn 2 (Leipzig) 1(7): 665 (1902)
Index Fungorum number: IF8971; 42 morphological species (Species Fungorum 2020), 11
species with sequence data.
Type species – Monochaetia monochaeta (Desm.) Allesch.
Notes – Members of this genus have been reported worldwide, as saprobes, endophytes or
pathogens, occurring on various hosts, especially on Quercus spp. (Maharachchikumbura et al.
2016b, De Silva et al. 2017, Wijayawardene et al. 2017a, Liu et al. 2019a, Farr & Rossman 2019).
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Morinia Berl. & Bres., Annuario Soc. Alpinisti Trident., 1887-88: 82 (1889)
Index Fungorum number: IF8997; 4 species with sequence data.
Type species – Morinia pestalozzioides Berl. & Bres.
Notes – Morinia species are characterised by transversely or muriformly septate conidia
(Collado et al. 2006). They are recorded from terrestrial habitats (Wijayawardene et al. 2017a).
Zetiasplozna was combined under Morinia based on phylogeny and similarity (Liu et al. 2019a).
Neopestalotiopsis Maharachch., K.D. Hyde & Crous, Stud. Mycol. 79: 135 (2014)
Index Fungorum number: IF809759; 43 species with sequence data.
Type species – Neopestalotiopsis protearum (Crous & L. Swart) Maharachch., K.D. Hyde &
Crous
Notes – Neopestalotiopsis, established by Maharachchikumbura et al. (2014b) is widespread,
occurring as saprobes or pathogens on various host plants (Maharachchikumbura et al. 2014b, Farr
& Rossman 2019). The genus is distinct from Pestalotiopsis in having versicolourous median cells.
Neopestalotiopsis rhizophorae is illustrated in this entry (Fig. 232).
Nonappendiculata F. Liu, L. Cai & Crous, Stud. Mycol. 92: 358 (2018)
Index Fungorum number: IF828374; 1 species with sequence data.
Type species – Nonappendiculata quercina F. Liu, L. Cai & Crous
Notes – Nonappendiculata, introduced by Liu et al. (2019a) produces 3-septate, fusoid
conidia without appendages. It occurs on Quercus from Italy (Liu et al. 2019a).
Parabartalinia F. Liu, L. Cai & Crous, Stud. Mycol. 92: 358 (2018)
Index Fungorum number: IF828376; 1 species with sequence data.
Type species – Parabartalinia lateralis F. Liu, L. Cai & Crous
Notes – This genus was introduced by Liu et al. (2019a) and is similar to Bartalinia, yet
phylogenetically distinct. It is recorded from South Africa on Acacia sp. (Liu et al. 2019a).
Pestalotiopsis Steyaert, Bull. Jard. bot. État Brux. 19: 300 (1949)
Index Fungorum number: IF9272; 225 morphological species (Species Fungorum 2020),
138 species with sequence data.
Type species – Pestalotiopsis maculans (Corda) Nag Raj
Notes – Pestalotiopsis was erected by Steyaert (1949). Pestalotiopsis species occur
worldwide, as saprobes, endophytes or opportunistic pathogens on economically important plants
as well as ornamental crops (Maharachchikumbura et al. 2014b, 2016b). The diversity and
phylogenetic affinities based on ITS, tub2 and tef1 gene regions for this genus has been discussed
comprehensively in several studies (Jeewon et al. 2003a, Maharachchikumbura et al. 2011, 2012,
2013, 2014a, Liu et al. 2019a).
Pseudopestalotiopsis Maharachch., K.D. Hyde & Crous, Stud. Mycol. 79: 180 (2014)
Index Fungorum number: IF809753; 15 species with sequence data.
Type species – Pseudopestalotiopsis theae (Sawada) Maharachch., K.D. Hyde & Crous
Notes – Pseudopestalotiopsis species are widely distributed as saprobes or pathogens,
occurring on leaves (Maharachchikumbura et al. 2014b, 2016a). They have conidia with
concolourous, brown to dark brown or olivaceous median pigmented cells (Maharachchikumbura et
al. 2014b).
Pseudosarcostroma F. Liu, L. Cai & Crous, Stud. Mycol. 92: 374 (2018)
Index Fungorum number: IF828385; 1 morphological species.
Type species – Pseudosarcostroma osyridicola F. Liu, L. Cai & Crous
Notes – Pseudosarcostroma, introduced by Liu et al. (2019a) is phylogenetically related to
Broomella, Bartalinia, Truncatella, Parabartalinia, and Diversimediispora. It shares similar
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characters with Sarcostroma, such as having conidia bearing a single appendage at each end, with
an undulate or verruculose conidial wall (Liu et al. 2019a). The genus was recorded from France on
Osyris alba (Liu et al. 2019a).
Figure 232 – Neopestalotiopsis rhizophorae (Material examined – THAILAND, Kor Chang, Trat
Province, leaf spots of Rhizophora mucronata L., 27 April 2017, Norphanphoun C. KC23-1,
KC23-2 (MFLU); living cultures, MFLUCC 17-1728. THAILAND, Kor Chang, Trat Province, leaf
spots of Rhizophora mucronata L., 27 April 2017, Norphanphoun C. KC23-2 (MFLU); living
cultures, MFLUCC 17-1729). a Collecting place. b, c Leaf spots of Rhizophora mucronata. d, e
Culture on PDA (d-above, e-below). f-h Colony sporulating on PDA. i Conidiogenous cells giving
rise to conidia. j-p Conidia. Scale bars: f = 200 µm, g = 1000 µm, h = 500 µm, i-p = 20 µm.
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Robillarda Sacc., Michelia 2(no. 6): 8 (1880)
Index Fungorum number: IF22470; 17 morphological species (Species Fungorum 2020), 6
species with sequence data.
Type species – Robillarda sessilis (Sacc.) Sacc.
Notes – The genus was introduced by Saccardo (1880a) and have flexuous, narrow tubular,
aseptate appendages and holoblastic conidiogenous cells, proliferating sympodially or percurrently
near the apex (Crous et al. 2015a, Wijayawardene et al. 2016a). Robillarda species have been
reported worldwide, as saprobes (Wijayawardene et al. 2017a, Farr & Rossman 2019). Most
recently, Robillarda mangiferae was introduced from leaf blight on mango in Yunnan, China
(Phookamsak et al. 2019).
Sarcostroma Cooke, Journal of the Quekett microsc. Club 2: 267 (1871)
Index Fungorum number: IF9789; 17 morphological species (Species Fungorum 2020), 11
species with sequence data.
Type species – Sarcostroma berkeleyi Cooke
Notes – Species of this genus are saprobes, endophytes, pathogens on leaves in both tropical
and temperate regions (Maharachchikumbura et al. 2016b, Norphanphoun et al. 2015, Farr &
Rossman 2019). Sarcostroma was resurrected to accommodate several species characterised by
fusoid conidia with four or more cells, having pigmented median cells and paler, thin-walled end
cells, bearing an attenuated tubular apical appendage and a similar excentric basal appendage (Nag
Raj 1993). The holotype of the genus has been reported lost and epitypification is needed (Liu et al.
2019a).
Seimatosporium Corda, Deutschl. Fl., 3 Abt. (Pilze Deutschl.) 3(13): 79 (1837)
Index Fungorum number: IF9865; 71 morphological species (Species Fungorum 2020), 24
species with sequence data.
Type species – Seimatosporium rosae Corda
Notes – Seimatosporium species are widely distributed, occurring as saprobes or pathogens
(Norphanphoun et al. 2015, Maharachchikumbura et al. 2016b, Wijayawardene et al. 2017a,
Wanasinghe et al. 2018). It was introduced by Corda (1833). Phylogenetically, they have been
linked to Discostroma sexual morphs (Tanaka et al. 2011). However, there are no sequence data for
the type species, this requires further study.
Seiridium Nees, Syst. Pilze (Würzburg): 22 (1816)
Index Fungorum number: IF9868; 39 morphological species (Jaklitsch et al. 2016b, Species
Fungorum 2020), 20 species with sequence data.
Type species – Seiridium marginatum Nees
Notes – Seiridium are characterized by 6-celled conidia (Jeewon 2003a,
Maharachchikumbura et al. 2014b) and are widely distributed as saprobes and plant pathogens
(Tsopelas et al. 2007). The genus was established by Nees (1816). Blogiascospora and Lepteutypa
was identified as the sexual morph (Senanayake et al. 2015). Jaklitsch et al. (2016b) epitypified
Seiridium marginatum and confirmed its sexual morph as Blogiascospora marginata based on
molecular data. Seiridium spyridiicola is illustrated in this entry (Fig. 233).
Sporocadus Corda, Icon. fung. (Prague) 3: 23 (1839)
Index Fungorum number: IF10026; 49 morphological species (Species Fungorum), 13
species with sequence data.
Type species – Sporocadus lichenicola Corda
Notes – Sporocadus was established by Corda (1839b) to include four species, but with no
mention of the type. Hughes (1958) lectotypified Sporocadus based on S. lichenicola. The genus
was once synonymised under Seimatosporium by Sutton (1975), but later classified as a distinct
872
genus by Brockman (1976) and Nag Raj (1993). Liu et al. (2019a) showed that both Sporocadus
and Seimatosporium are phylogenetically distinct and species in Sporocadus lack appendages.
Figure 233 – Seiridium spyridiicola (Material examined – ITALY, Forli-Cesena Province, near
Camposonaldo – Santa Sofia, on dead and land cone of Picea excelsa, 15 April 2015, Erio
Camporesi IT2444, MFLU 15-0813; living culture KUMCC 16-0115). a Appearance on host. b
Close up of several conidiomata. c Section of conidioma. d Wall of conidioma. e, f Conidiogenous
cells giving rise to conidia. f, g Conidiophores with conidia. h-k Conidia (upside down). Scale bars:
a = 500 µm, c = 1 mm, d = 100 µm, e-l = 20 µm.
Strickeria Körb., Parerga lichenol. (Breslau) 5: 400 (1865)
Index Fungorum number: IF5283; 33 morphological species (Species Fungorum 2020), 1
species with sequence data.
873
Type species – Strickeria kochii Körb.
Notes – Strickeria was introduced and described as a lichenised fungus by Körber (1865). It
was often classified in Dothideomycetes and upon re-examination of the type material, it was
placed under Sordariomycetes genera incertae sedis (Eriksson & Hawksworth 1991, Lumbsh &
Huhndorf 2010). Subsequently this genus was classified in Xylariales by Jaklitsch et al. (2016b).
Synnemapestaloides T. Handa & Y. Harada, Mycoscience 45:138 (2004)
Index Fungorum number: IF28803; 2 species with sequence data.
Type species – Synnemapestaloides rhododendri T. Handa & Y.Harada.
Notes – Synnemapestaloides species are distinct in producing spores on synnemata
(Watanabe et al. 2016). They occur on Juniperus and Rhododendron (Liu et al. 2019a).
Truncatella Steyaert, Bull. Jard. bot. État Brux. 19: 293 (1949)
Index Fungorum number: IF10328; 13 morphological species (Species Fungorum 2020), 7
species with sequence data.
Type species – Truncatella truncata (Lév.) Steyaert
Notes – Truncatella was established by Steyaert (1949) to accommodate distinct species
having 3-septate, verruculose, pigmented conidia (Steyaert 1949, Jeewon et al. 2002,
Maharachchikumbura et al. 2016b, Tibpromma et al. 2017b). They occur worldwide as saprobes on
various dicotyledons and monocotyledons (Farr & Rossman 2019).
Xenoseimatosporium F. Liu, L. Cai & Crous , Stud. Mycol. 92: 411 (2018)
Index Fungorum number: IF828424; 1 species with sequence data.
Type species – Xenoseimatosporium quercinum (Goonas., R.K. Schumach. & K.D. Hyde) F.
Liu, L. Cai & Crous
Notes – This genus was introduced by Liu et al. (2019a) to accommodate the previously
described species Seimatosporium quercinum on Quercus from Germany (Goonasekara et al.
2016). It is phylogenetically close to the genera Allelochaeta and Sarcostroma.
Xenoseimatosporium is distinct from Seimatosporium in producing allantoid and subcylindrical
conidia with ragged appendages (Liu et al. 2019a).
Stachybotryaceae L. Lombard & Crous, Persoonia 32: 283 (2014)
Index Fungorum number: IF90922; Facesoffungi number: FoF05320; 304 species.
Saprobic or pathogenic on plants and commonly isolated from soil. Sexual morph: Ascomata
superficial or immersed, solitary or scattered, sometimes fused, globose to subglobose, bright to
dark yellow or black, sometimes covered with intertwined hyphae. Papilla central, short, black,
without periphyses. Ostiole conical, thin, black. Peridium composed of several layers of dark
brown, thin-walled cells of textura angularis (Stachybotrys), or textura intricata (Scopinella), or
outwardly composed of thick-walled textura angularis and inwardly with textura prismatica
(Peethambara). Paraphyses moniliform or filiform, hyaline, septate or aseptate, intermingled with
asci. Asci 4–8-spored, unitunicate, clavate to cylindrical, apex rounded to nearly truncate,
deliquescent at maturity, lacking an apical ring. Ascospores 2–3-seriate, hyaline or brown to dark
brown, ellipsoidal to fusiform or broadly reniform, 0–1-septate, with or lacking a mucilaginous
sheath. Asexual morph: Hyphomycetous. Conidiophores single, sporodochial or synnematous, if
conidiophore single, macronematous, mononematous, solitary or gregarious, simple or irregularly
branched, hyaline or pale brown, smooth-walled; if conidiophores sporodochial or synnematous,
pulvinate, gelatinous, hyaline to dark green, with or without setae, with irregularly penicillate, bior triverticillately branched conidiogenous ring. Conidiogenous cells enteroblastic, monophialidic
to polyphialidic, discrete, cylindrical, ellipsoidal or clavate to broadly reniform, with conspicuous
collarettes, determinate or proliferating percurrently, initially hyaline becoming pale brown at
maturity. Conidia arising from the apices of the phialides, elongate, cylindrical, fusiform or
ellipsoidal, 0–3-septate, smooth-walled, smooth, striate or roughened, hyaline, brown to dark
874
brown, aggregated in slimy, dark green to black masses (adapted from Maharachchikumbura et al.
2016b).
Type genus – Stachybotrys Corda
Notes – Stachybotryaceae was established by Crous et al. (2014a) to accommodate
Myrothecium, Peethamabra and Stachybotrys. This family is characterized by asexual morphs
having mononematous, sporodochial or synnematous conidiophores and phialidic conidiogenous
cells that produce conidia in chains or in slimy or dry masses (Seifert et al. 2011, Crous et al.
2014a, Wang et al. 2015b, Lombard et al. 2016). Lombard et al. (2016) monographed the family
based on morphology and LSU, ITS, rpb2, cmdA, tef1 and tub2 sequence data and 33 genera were
accepted. Wijayawardene et al. (2018a) accepted 36 genera, however the family seems to be split
into rather numerous genera based on few distinguishing characters and poor phylogenetic
resolution.
Ecological and economic significance of Stachybotryaceae
Some species in Stachybotryaceae are serious plant, animal and human pathogens. The most
serious plant pathogenic species are Myrothecium roridum and M. verrucaria that cause dieback
and leaf spots of various plant hosts (Zhao et al. 2010, Zhang et al. 2011, Hong et al. 2013, Ben et
al. 2015, Fujinawa et al. 2016). Stachybotrys chartarum (often cited as S. atrus Corda) is one of the
most feared of fungi in the world, which can produce mycotoxins and result in serious buildingrelated illness and agricultural damage (Drobotko 1945, Dankó 1975, Page & Trout 2001, Seifert et
al. 2011, Hyde et al. 2018a). It was reported that mycotoxins produced by Stachybotrys can cause a
variety of symptoms in humans, including cold- and flu-like symptoms, sore throats, diarrhea,
headaches, dermatitis, patches of hair loss, and fatigue, and also can cause the death of animals
(Schneider 1979, Croft et al. 1986, Johanning et al. 1996, Page and Trout 2001).
Genera included in Stachybotryaceae
Achroiostachys L. Lombard & Crous, Persoonia 36: 172 (2016)
Index Fungorum number: IF815916; 6 species with sequence data.
Type species – Achroiostachys humicola L. Lombard & Crous
Notes – Achroiostachys was introduced by Lombard et al. (2016) to accommodate A.
aurantispora, A. betulicola, A. humicola, A. levigata, A. phyllophila and A. saccharicola. The genus
is characterised by hyaline, mostly smooth, thin-walled conidiophores, phialidic conidiogenous
cells and hyaline, aseptate, smooth-walled, ellipsoidal to limoniform conidia (Lombard et al. 2016).
Albifimbria L. Lombard & Crous, Persoonia 36: 177 (2016)
Index Fungorum number: IF815924; 6 species with sequence data.
Type species – Albifimbria verrucaria (Alb. & Schwein.) L. Lombard & Crous
Notes – Albifimbria was established by Lombard et al. (2016) to accommodate A. lateralis, A.
terrestris, A. verrucaria and A. viridis. No sexual morph has been determined. The genus is
characterised by the formation of verrucose setae surrounding the sporodochia and conidia
sometimes bearing a funnel-shaped mucoid appendage (Lombard et al. 2016).
Albosynnema E.F. Morris, Mycopath. Mycol. appl. 33: 179 (1967)
Index Fungorum number: IF7086; 2 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Albosynnema elegans E.F. Morris
Notes – Lombard et al. (2016) accepted this genus in Stachybotryaceae with support from
phylogenetic analysis. The genus is characterised by synnematous, branched, hyaline
conidiophores, phialidic conidiogenous cells and dark, septate, slimy conidia (Seifert et al. 2011).
Alfaria Crous, Montaño-Mata & García-Jim., Persoonia 32: 239 (2014)
Index Fungorum number: IF808923; 15 species with sequence data.
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Type species – Alfaria cyperi-esculenti Crous, Montaño-Mata & García-Jim.
Notes – Alfaria was established to accommodate A. cyperi-esculenti, which is a plant
pathogen. The genus is characterised by solitary or sporodochial, verticillately or penicillately
branched, hyaline conidiophores, phialidic conidiogenous cells and cylindrical to ellipsoidal to
ossiform, hyaline to lightly pigmented, aseptate conidia (Lombard et al. 2016). When setae are
present, they are septate and unbranched (Lombard et al. 2016).
Alfariacladiella Crous & R.K. Schumach., Beihefte zur Sydowia 68: 202 (2016)
Index Fungorum number: IF817207; 1 species with sequence data.
Type species – Alfariacladiella spartii Crous & R.K. Schumach.
Notes – Alfariacladiella is characterised by sporodochial, cupulate conidiomata surrounded
by hyaline and septate marginal hyphae (Hernández-Restrepo et al. 2016b). Conidiophores are
hyaline, smooth, septate and monoverticillate or not. Conidiogenous cells are phialidic. Conidia are
single, hyaline, smooth-walled, guttulate and fusiform. The genus is similar to Alfaria, but differs in
lacking brown setae surrounding the conidiomata, and having fusiform conidia with apical mucoid
caps (Hernández-Restrepo et al. 2016c).
Brevistachys L. Lombard & Crous, Persoonia 36: 182 (2016)
Index Fungorum number: IF815934; 5 species with sequence data.
Type species – Brevistachys variabilis L. Lombard & Crous
Notes – Brevistachys was introduced by Lombard et al. (2016) to accommodate stachybotryslike species having distinctly short conidiophores and conidiogenous cells borne on conidiophores
or directly from vegetative hyphae.
Capitofimbria L. Lombard & Crous, Persoonia 36: 185 (2016)
Index Fungorum number: IF815940; 1 species with sequence data.
Type species – Capitofimbria compacta (R.F. Castañeda, Gusmão, Stchigel & M. Stadler) L.
Lombard & Crous
Notes – Lombard et al. (2016) transferred Myrothecium compactum to Capitofimbria. No
sexual morph was identified. The genus is characterised by the marginal hyphae terminating in a
capitate to clavate thick-walled cells surrounding the sporodochia (Lombard et al. 2016).
Cymostachys L. Lombard & Crous, Persoonia 36: 186 (2016)
Index Fungorum number: IF815942; 4 species with sequence data.
Type species – Cymostachys coffeicola L. Lombard & Crous
Notes – Two species, C. coffeicola and C. fabispora, were introduced by Lombard et al.
(2016). The third species, C. garethjonesii, was introduced by Lin et al. (2016). The genus is
characterised by irregularly cymosely branched conidiophores and olivaceous brown to dark
brown, fabiform conidia.
Didymostilbe Henn., Hedwigia 41: 148 (1902)
Index Fungorum number: IF8021; 10 morphological species (Species Fungorum 2020), 2
species with sequence data.
Type species – Didymostilbe coffeae Henn.
Notes – The genus has large, thick-walled conidia, having prominent apical and/or basal
mammiform protuberances. Lombard et al. (2016) placed the genus in Stachybotryaceae with the
support of phylogenetic analysis.
Digitiseta Gordillo & Decock, Mycol. Progr. 17: 179–190 (2017)
Index Fungorum number: IF820512; 4 species with sequence data.
Type species – Digitiseta setiramosa (R.F. Castañeda) Gordillo & Decock
876
Notes – The genus is characterised by sporodochial conidiomata with a green mucoid mass of
conidia on the top and setoid hypha-like extensions with short apical branches, biverticillate
conidiophores, phialidic conidiogenous cells and aseptate, cylindrical, hyaline to pale greenish
conidia (Gordillo & Decock 2018).
Dimorphiseta L. Lombard & Crous, Persoonia 36: 188 (2016)
Index Fungorum number: IF815956; 3 species with sequence data.
Type species – Dimorphiseta terrestris L. Lombard & Crous
Notes – The genus is similar to Smaragdiniseta, but differs by the setae morphology. Type I
setae of Dimorphiseta are hyaline, and type II setae taper to sharp apices, however, type I setae of
Smaragdiniseta are emerald green, and type II setae narrow to an obtuse apex (Lombard et al.
2016).
Globobotrys L. Lombard & Crous, Persoonia 36: 189 (2016)
Index Fungorum number: IF815990; 1 species with sequence data.
Type species – Globobotrys sansevieriicola (Crous & M.J. Wingf.) L. Lombard & Crous
Notes – The stachybotrys-like genus Globobotrys was introduced by Lombard et al. (2016) to
accommodate G. sansevieriicola (≡ Stachybotrys sansevieriicola). Only one species is accepted in
the genus, which is characterised by mononematous, mostly unbranched, hyaline conidiophores,
phialidic conidiogenous cells and aseptate, hyaline to olivaceous brown, smooth-walled, thickwalled, globose to broadly ellipsoidal conidia (Lombard et al. 2016).
Grandibotrys L. Lombard & Crous, Persoonia 36: 189 (2016)
Index Fungorum number: IF815992; 3 species with sequence data.
Type species – Grandibotrys pseudotheobromae L. Lombard & Crous
Notes – Two species, G. pseudotheobromae and G. xylophilus, were introduced by Lombard
et al. (2016) in Grandibotrys. Hyde et al. (2017b) added G. hyalinus. The genus is characterised by
mononematous, unbranched or branched, hyaline conidiophores, phialidic conidiogenous cells and
olivaceous green to dark brown, limoniform to ellipsoidal conidia, having a mammiform apical
and/or basal protrudance (Lombard et al. 2016).
Gregatothecium L. Lombard & Crous, Persoonia 36: 191 (2016)
Index Fungorum number: IF815995; 1 species with sequence data.
Type species – Gregatothecium humicola L. Lombard & Crous
Notes – The genus differs from the similar genera by the slimy olivaceous green conidial
masses produced on the conidiophores and sporodochia (Lombard et al. 2016). Unbranched and
hyaline setae arise from the basal stroma, (Lombard et al. 2016). Conidiophores are penicillately
branched, conidiogenous cells are phialidic and hyaline and conidia are aseptate and cylindrical to
subcylindrical (Lombard et al. 2016).
Hyalinostachys C.G. Lin & K.D. Hyde, gen. nov.
Index Fungorum number: IF555606; Facesoffungi number: FoF03883; 1 species with
sequence data.
Etymology: Name reflects the hyaline stachybotrys-like conidiophores and conidia.
Saprobic on plant host. Sexual morph: Undetermined. Asexual morph: Hyphomycetous.
Colonies effuse, hairy, pale to pale brown. Mycelium partly superficial and partly immersed.
Conidiophores macronematous, mononematous, single or in small groups, simple or branched,
erect, straight, septate, thick-walled, smooth, hyaline, narrower at the apex. Conidiogenous cells
monophialidic, discrete, determinate, terminal, elongate ampulliform, clavate or subcylindrical, the
outer ones somewhat curved and ventricose, hyaline, smooth; conidial mass slimy, pale orange.
Conidia acrogenous, aseptate, smooth, hyaline, cylindrical, ellipsoidal.
Type species – Hyalinostachys cylindrospora C.G. Lin & K.D. Hyde
877
Notes – The combined LSU and rpb2 phylogenetic analyses indicate that Hyalinostachys
forms a separate clade and sister taxon to Achroiostachys, Globobotrys, Melanopsamma and
Sirastachys in Stachybotryaceae (Fig. 15). Hyalinostachys is distinguished from these genera by
simple or branched, thick-walled, smooth and hyaline conidiophores and hyaline, cylindrical or
ellipsoidal conidia.
Hyalinostachys cylindrospora C.G. Lin & K.D. Hyde, sp. nov.
Fig. 235
Index Fungorum number: IF555605; Facesoffungi number: FoF03884
Etymology – Name reflects the cylindrical conidia.
Holotype – MFLU 17-2650.
Saprobic on plant host. Sexual morph: Undetermined. Asexual morph: Hyphomycetous.
Colonies effuse, hairy, pale to pale brown. Mycelium partly superficial and partly immersed.
Conidiophores macronematous, mononematous, single or in small groups, simple or sometimes
branched, erect, straight, septate, thick-walled, smooth, hyaline, up to 310 μm, 4–12 μm, narrower
at the apex, bearing a whorl of 4–5 conidiogenous cells. Conidiogenous cells monophialidic,
discrete, determinate, terminal, elongate ampulliform, clavate or subcylindrical, the outer ones
somewhat curved and ventricose, hyaline, smooth, 13–19 × 3.5–6 μm; conidial mass slimy, pale
orange. Conidia acrogenous, aseptate, smooth, hyaline, cylindrical or ellipsoidal, 14–16.5 × 5.5–7
μm.
Culture characteristics: Conidia germinating on PDA in 12 h. Colonies on PDA, reaching 3
mm in 2 weeks at 25°C, flat, dark from above and the reverse. Mycelium immersed in media,
composed of branched, septate, smooth, hyaline hyphae.
Material examined – THAILAND, Chiang Mai, Mae Taeng District, Sop Poeng, on decaying
wood, 25 September 2016, Chuan-Gen Lin, LCG 11-5 (MFLU 17-2650, holotype), ex-type living
culture, MFLUCC 17-2583.
Genbank numbers – ITS: MG717501, LSU: MG717503, rpb2: MK133023.
Notes – This species belongs to family Stachybotryaceae, and its placement is supported by
morphology and phylogenetic analysis. Hyalinostachys cylindrospora can be distinguished from
other stachybotrys-like species by its hyaline conidiophores and hyaline, cylindrical or ellipsoidal
conidia.
Inaequalispora L. Lombard & Crous, Persoonia 36: 191 (2016)
Index fungorum number: IF815997; 3 species with sequence data.
Type species – Inaequalispora prestonii (M.C. Tulloch) L. Lombard & Crous
Notes – Inaequalispora was established by Lombard et al. (2016) to accommodate I. prestonii
(≡ Myrothecium prestonii). Gordillo & Decock (2018) introduced Inaequalispora cylindrospora
and I. longiseta to this genus. The genus is characterised by sporodochial, stromatic conidiomata,
penicillately branched, hyaline conidiophores, phialidic, hyaline conidiogenous cells and aseptate,
fusiform to ellipsoidal to asymmetrically ellipsoidal, hyaline conidia, with a funnel-shaped mucoid
apical appendage (Lombard et al. 2016).
Kastanostachys L. Lombard & Crous, Persoonia 36: 192 (2016)
Index Fungorum number: IF815999; 1 species with sequence data.
Type species – Kastanostachys aterrima (Fuckel) L. Lombard & Crous
Notes – The asexual morph of Kastanostachys is characterised by unbranched, percurrent
conidiophores, hyaline, phialidic conidiogenous cells and hyaline, smooth, ellipsoidal conidia
(Lombard et al. 2016). An epitype was designated for K. aterrima (≡ Melanomma aterrima Fuckel)
by Lombard et al. (2016).
Koorchalomella Chona, Munjal & J.N. Kapoor, Indian Phytopath. 11: 130 (1958)
Index Fungorum number: IF8681; 2 morphological species (Species Fungorum 2020), 1
species with sequence data.
878
Figure 234 – Phylogenetic tree generated from maximum likelihood (ML) analysis based on
combined LSU and rpb2 sequence data of genera of Stachybotryaceae. Bootstrap support values for
maximum likelihood (ML, first value) and maximum parsimony (MP, second value) and greater
than 50% and Bayesian posterior probabilities (BYPP, third value) greater than 0.90 are indicated
above or below the nodes. Hyphen (‘‘-’’) indicates a value lower than 50% for ML and MP and a
posterior probability lower than 0.90 for BYPP. New isolates are in blue bold. The tree is rooted
with Calonectria ilicicola (CBS 190.50), Fusarium sambucinum (CBS 146.95), and Nectria
cinnabarina (CBS 125165).
879
Type species – Koorchalomella oryzae Chona, Munjal & J.N. Kapoor
Notes – Hyde et al. (2017b) introduced the second species for this genus and provided DNA
sequence data for it, which placed the genus in Stachybotryaceae. The genus is characterised by
sporodochial conidiomata, unbranched, hyaline setae, branched, hyaline conidiophores, phialidic,
hyaline conidiogenous cells and aseptate, hyaline, slimy conidia with funnel-shaped mucilaginous
polar appendages (Seifert et al. 2011).
Figure 235 – Hyalinostachys cylindrospora (MFLU 17-2650, holotype). a Host material. b
Conidiophores on the host surface. c, d Conidiophores and conidiogenous cells. e, f Conidiogenous
cells. g Conidia. Scale bars: b = 200 μm, c, d = 50 μm, e, f = 20 μm, g = 10 μm.
Melanopsamma Niessl, Verh. nat. Ver. Brünn 14: 200 (1876)
Index Fungorum number: IF3079; 42 morphological species (Species Fungorum 2020), 2
species with sequence data.
Type species – Melanopsamma pomiformis (Pers.) Sacc.
880
Notes – Wang et al. (2015b) synonymised this genus under Stachybotrys, however, Lombard
et al. (2016) resurrected it based on multigene phylogenetic analysis. The genus is characterised by
mononematous, unbranched, hyaline conidiophores, phialidic, hyaline conidiogenous cells, and
aseptate, hyaline to olivaceous brown to dark brown, smooth to verrucose, limoniform to obovoid
to globose to ellipsoidal conidia (Lombard et al. 2016).
Memnoniella Höhn., Centbl. Bakt. ParasitKde, Abt. II 60: 16 (1923)
Index Fungorum number: IF8900; 9 species with sequence data.
Type species – Memnoniella aterrima Höhn.
Notes – Wang et al. (2015b) synonymised this genus under Stachybotrys, however, Lombard
et al. (2016) resurrected it based on multigene phylogenetic analysis. The genus is characterised by
mononematous, unbranched, septate conidiophores, phialidic conidiogenous cells and aseptate,
ellipsoidal to globose to reniform conidia (Lombard et al. 2016). The conidia of this genus
sometimes are in dry or slimy chains (Lombard et al. 2016).
Myrothecium Tode, Fung. mecklenb. sel. (Lüneburg) 1: 25 (1790)
Index Fungorum number: IF9049; 35 morphological species (Species Fungorum 2020), 11
species with sequence data.
Type species – Myrothecium inundatum Tode
Notes – Lombard et al. (2016) narrowed the concept of Myrothecium to include only species
with conidia that are less than 5 μm long, borne in olivaceous to dark green slimy masses
surrounded by a setose fringe; only two species are accepted.
Myxospora L. Lombard & Crous, Persoonia 36: 202 (2016)
Index Fungorum number: IF816011; 5 species with sequence data.
Type species – Myxospora masonii (M.C. Tulloch) L. Lombard & Crous
Notes – This genus is characterised by the formation of mostly fusiform conidia, bearing an
apical hilum without funnel-shaped mucoid appendages; some species form synnematal
conidiomata (Lombard et al. 2016).
Neomyrothecium L. Lombard & Crous, Persoonia 36: 205 (2016)
Index Fungorum number: IF816017; 1 species with sequence data.
Type species – Neomyrothecium humicola L. Lombard & Crous
Notes – The genus is characterised by sporodochial, stromatic conidiomata, penicillately
branched, hyaline conidiophores, phialidic conidiogenous cells and aseptate, hyaline, cylindrical
conidia. The genus is similar to Paramyrothecium but differs by multiseptate and unbranched setae
and pulvinate sporodochia that lack a white setosee fringe (Lombard et al. 2016).
Paramyrothecium L. Lombard & Crous, Persoonia 36: 206 (2016)
Index Fungorum number: IF815988; 15 species with sequence data.
Type species – Paramyrothecium roridum (Tode) L. Lombard & Crous
Notes – Paramyrothecium was introduced by Lombard et al. (2016) to accommodate 12
myrothecium-like species. The genus differs from Myrothecium sensu stricto and other
myrothecium-like genera by having 1–3-septate, thin-walled setae surrounding the sporodochia.
Parasarcopodium Melnik, S.J. Lee & Crous, Mycol. Progr. 3(1): 22 (2004)
Index Fungorum number: IF28802; 3 species with sequence data.
Type species – Parasarcopodium ceratocaryi Melnik, S.J. Lee & Crous
Notes – Lombard et al. (2016) placed this genus in Stachybotryaceae with the support of
phylogenetic analysis. The genus is characterised by sporodochial conidiomata, branched, setiform
conidiophores, phialidic, hyaline conidiogenous cells and aseptate, hyaline conidia with polar
mucilaginous caps (Seifert et al. 2011).
881
Parvothecium L. Lombard & Crous, Persoonia 36: 214 (2016)
Index Fungorum number: IF816019; 2 species with sequence data.
Type species – Parvothecium terrestre L. Lombard & Crous
Notes – Gordillo & Decock (2018) described the second species, P. amazonense in this
genus. The genus is characterised by sporodochial, stromatic conidiomata, penicillately branched,
hyaline conidiophores, phialidic conidiogenous cells and aseptate, ellipsoidal to asymmetrically
ellipsoidal, hyaline to pigmented conidia (Lombard et al. 2016).
Peethambara Subram. & Bhat, Revue Mycol., Paris 42(1): 52 (1978)
Index Fungorum number: IF3786; 1 species with sequence data.
Type species – Peethambara sundara Subram. & Bhat
Notes – Lombard et al. (2016) placed this genus in Stachybotryaceae with the support of
multigene phylogenetic analysis. The genus is characterised by white to yellow, synnematous
conidiomata and ellipsoidal to limoniform, hyaline to subhyaline, 1-septate conidia with a
mammiform basal and/or apical protrudance (Lombard et al. 2016).
Pseudoornatispora Tibpromma & K.D. Hyde, Fungal Divers. 92: 115 (2018)
Index Fungorum number: IF555336; 1 species with sequence data.
Type species – Pseudoornatispora krabiense Tibpromma & K.D. Hyde
Notes – The genus is characterised by black ascomata with seta and ostioles, 6–8-spored,
unitunicate asci, hyaline, 1-septate, verrucose, fusiform ascospores, mononematous conidiophores,
phialidic, hyaline conidiogenous cells, ellipsoid, hyaline to subhyaline, aseptate, guttulate conidia,
without a mucilaginous sheath (Tibpromma et al. 2018). This genus is similar to Koorchaloma and
the species of these two genera grouped together in the phylogenetic analysis generated from
combined ITS, LSU and rpb2 sequence data (Li et al. unpublished data). We maintain these two
genera as distinct. Pseudoornatispora krabiense is illustrated in this entry.
Septomyrothecium Matsush., Bull. natn. Sci. Mus., Tokyo, N.S. 14: 469 (1971)
Index Fungorum number: IF9885; 2 species with sequence data.
Type species – Septomyrothecium uniseptatum Matsush.
Notes – Decock et al. (2008) introduced one new species and one new combination to this
genus. The genus is characterised by sporodochial conidiomata surrounded by white to grey fringe,
long, hyaline dichotomously branched hyphoid extensions extending beyond the olivaceous green
to black conidial masses, and 0–1-septate, cylindrical, straight to curved conidia (Lombard et al.
2016).
Sirastachys L. Lombard & Crous, Persoonia 36: 215 (2016)
Index Fungorum number: IF816021; 9 species with sequence data.
Type species – Sirastachys phaeospora L. Lombard & Crous
Notes – Sirastachys was introduced by Lombard et al. (2016) to accommodate seven
stachybotrys-like species which are characterised by the formation of synnemata in culture from
which the conidiophores arise laterally. The genus is characterised by mononematous, unbranched
or branched conidiophores arising laterally from synnemata, phialidic conidiogenous cells and
aseptate, hyaline to pale olivaceous brown to dark brown, ellipsoidal to obovoid to cylindrical
conidia (Lombard et al. 2016).
Smaragdiniseta L. Lombard & Crous, Persoonia 36: 218 (2016)
Index Fungorum number: IF816029; 1 species with sequence data.
Type species – Smaragdiniseta bisetosa (V.G. Rao & de Hoog) L. Lombard & Crous
Notes – Smaragdiniseta was introduced by Lombard et al. (2016) to accommodate S. bisetosa
(≡ Myrothecium bisetosum). The genus is characterised by sporodochial, stromatic conidiomata
surrounded by two types of setae, verticillately or penicillately branched, hyaline conidiophores,
882
phialidic, hyaline conidiogenous cells and aseptate, obclavate to narrowly ellipsoidal, hyaline
conidia.
Figure 236 – Pseudoornatispora krabiense (Material examined – THAILAND, Krabi Province,
Mueang Krabi District, on Pandanus sp., 15 December 2015, S. Tibpromma KB014, MFLU 161892, holotype, HKAS 96242, isotype; ex-type living culture, MFLUCC 16-0317). a Appearance
of taxon on dead leaves of Pandanus sp. b Paraphyses. c Section of ascoma. d Section of peridium.
e Papilla. f-h Ascospores. i Ascospore stained with India ink. j, k Asci. Scale bars: a = 200 μm, b =
5 μm, c = 50 μm, d, e, i = 10 μm, f, g = 5 μm, h = 30 μm, j, k = 20 μm.
Stachybotrys Corda, Icon. fung. (Prague) 1: 21 (1837)
Index Fungorum number: IF10052; 88 morphological species (Species Fungorum 2020), 30
species with sequence data.
Type species – Stachybotrys atrus Corda
Notes – Lombard et al. (2016) revised this genus and transferred many previously accepted
species to other genera, e.g. Brevistachys, Globobotrys, Sirastachys, and Striatibotrys.
883
Striatibotrys L. Lombard & Crous, Persoonia 36: 224 (2016)
Index Fungorum number: IF816035; 7 species with sequence data.
Type species – Striatibotrys eucylindrosporus (D.W. Li) L. Lombard & Crous
Notes – Striatibotrys was introduced by Lombard et al. (2016) to accommodate six
stachybotrys-like species which are characterised by mononematous conidiophores, phialidic
conidiogenous cells and aseptate, pale olivaceous to dark brown, smooth-walled, longitudinally
striate, ellipsoidal to subcylindrical to fusiform conidia (Lombard et al. 2016).
Striaticonidium L. Lombard & Crous, Persoonia 36: 227 (2016)
Index Fungorum number: IF816042; 5 species with sequence data.
Type species – Striaticonidium cinctum (Corda) L. Lombard & Crous
Notes – Striaticonidium was introduced by Lombard et al. (2016) to accommodate four
myrothecium-like species which are characterised by synnematous or sporodochial or
mononematous, verticillately or penicillately branched conidiophores, phialidic, hyaline
conidiogenous cells and aseptate, fusiform to ellipsoidal, olivaceous green to brown, longitudinally
striate conidia (Lombard et al. 2016).
Tangerinosporium L. Lombard & Crous, Persoonia 36: 231 (2016)
Index Fungorum number: IF816048; 1 species with sequence data.
Type species – Tangerinosporium thalictricola L. Lombard & Crous
Notes – Tangerinosporium was introduced by Lombard et al. (2016) to accommodate T.
thalictricola which is characterised by sporodochial, stromatic conidiomata covered by an apricot
to orange slimy or dry mass of conidia, hyaline, penicillately branched conidiophores, phialidic
conidiogenous cells and aseptate, ellipsoidal to ossiform, hyaline conidia (Lombard et al. 2016).
Virgatospora Finley, Mycologia 59(3): 538 (1967)
Index Fungorum number: IF10411; 2 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Virgatospora echinofibrosa Finley
Notes – This genus was synonymised under Didymostilbe (Rossman 1983, Rossman et al.
1999). Lombard et al. (2016) resurrected the genus based on multigene phylogenetic analysis and
accepted it in Stachybotryaceae. The genus is characterised by white to dark brown or black to
olivaceous grey, synnematous conidiomata, branched conidiophores, 3-septate, fusiform,
olivaceous grey, coarsely striate conidia with papillate and truncate ends (Finley 1967, Lombard et
al. 2016).
Xenomyrothecium L. Lombard & Crous, Persoonia 36: 232 (2016)
Index Fungorum number: IF816050; 1 species with sequence data.
Type species – Xenomyrothecium tongaense (W.B. Kendr., DiCosmo & Michaelides) L.
Lombard & Crous
Notes – Xenomyrothecium was introduced by Lombard et al. (2016) to accommodate X.
tongaense (≡ Myrothecium tongaense). The genus is characterised by sporodochial, stromatic
conidiomata, hyaline conidiophores, phialidic, pale green conidiogenous cells and aseptate, oblongellipsoidal, pale green conidia (Lombard et al. 2016). The genus differs from other genera by its
oblong-ellipsoidal conidia and lack of setae formed in the sporodochia.
Xepicula Nag Raj, Coelomycetous Anamorphs with Appendage-bearing Conidia (Ontario): 979
(1993)
Index Fungorum number: IF26431; 3 species with sequence data.
Type species – Xepicula leucotricha (Peck) Nag Raj
Notes – The genus is characterised by cupulate conidiomata with septate conidiomatal setae
and lack of exipular elements (Rungjindamai et al. 2012).
884
Xepiculopsis Nag Raj, Coelomycetous Anamorphs with Appendage-bearing Conidia (Ontario):
9823 (1993)
Index Fungorum number: IF26432; 2 morphological species (Species Fungorum 2020).
Type species – Xepiculopsis perpulchra Nag Raj
Notes – Xepiculopsis was introduced by Nag Raj (1993) to accommodate X. graminea and X.
perpulchra. The placement of this genus in Stachybotryaceae needs to be confirmed with DNA
sequence data. The genus is characterised by pulvinate to cupulate, stromatic conidiomata, hyaline
conidiophores, phialidic, hyaline conidiogenous cells and fusiform to ellipsoid, aseptate, pale
olivaceous conidia with an obtuse apex (Nag Raj 1993).
Stilbosporaceae Link, Abh. dt. Akad. Wiss. Berlin: 180 (1826)
Index Fungorum number: IF81439; Facesoffungi number: FoF01411; 58 species.
Saprobic on bark of trees and shrubs. Sexual morph: Pseudostromata inconspicuous and
immersed. Ectostromatic disc absent or present, if present inconspicuous, light brown to rarely dark
brown. Entostroma prosenchymatous, pale coloured, slightly differentiated from surrounding bark.
Ascomata loosely arranged as valsoid groups, in a single layer, immersed, aggregated, globose to
subglobose, coriaceous, black, ostiolate, papillate. Ostiole not obvious and convergent in groups.
Paraphyses filiform, aseptate, hyaline. Asci 8-spored, unitunicate, cylindrical, initially attached to
the base, later floating in centrum, with a J-, refractive, apical ring. Ascospores overlapping
uniseriate to biseriate, brown, ellipsoid to oblong, distoseptate. Asexual morph: Coelomycetous.
Conidiomata acervular, with paraphyses. Conidiophores cylindrical, hyaline. Conidiogenous cells
annellidic. Conidia brown, cylindrical, clavate to pyriform, eu- or distoseptate, oblique or
longitudinal septa present or absent (adapted from Voglmayr & Jaklitsch2014, Senanayake et al.
2017a).
Type genus – Stilbospora Pers.
Notes – Stilbosporaceae was introduced by Link (1826) with Prosthecium as the asexual
morph. This family was not phylogenetically well-resolved and synonymized in several families.
Voglmayr & Jaklitsch (2014) accommodated Stilbosporaceae in Diaporthales based on LSU
sequence data. Stegonsporium and Stilbospora were included in the family and Prosthecium was
synonymized under Stilbospora. The type species of Stilbospora is S. macrosperma and it was
connected with its asexual morph Prosthecium ellipsosporum (Voglmayr & Jaklitsch 2008).
Prosthecium ellipsosporum is the generic type of Prosthecium. Maharachchikumbura et al. (2015)
accommodated Natarajania in the family using multigene phylogenetic analysis. Crinitospora was
added to the family by Senanayake et al. (2017a). At present the family comprises four genera
including Crinitospora, Natarajania, Stegonsporium and Stilbospora.
Ecological and economic significance of Stilbosporaceae
Voglmayr & Jaklitsch (2014) reported most taxa in the family as discovered from Europe and
North America. Asexual morphs of several species are opportunistic and moderate phytopathogens.
Those species cause branch dieback or twig blight on several plants in the region (Voglmayr &
Jaklitsch 2008). Some species have been found in dead branches of overwintered plants (Voglmayr
& Jaklitsch 2014).
Genera included in Stilbosporaceae
Crinitospora B. Sutton & Alcorn, Trans. Br. mycol. Soc. 84(3): 437 (1985)
Index Fungorum number: IF11057; 1 species with sequence data.
Type species – Crinitospora pulchra B. Sutton & Alcorn
Notes – The asexual caulicolous Crinitospora is monotypic and the sexual morph has not yet
been determined (Crous et al. 2014b). The genus was recorded from the leaves of Mangifera indica
in Queensland. There are no published data on the ecology and pathology of the genus (Senanayake
et al. 2018). Previously the genus was placed in Melanconidaceae (Diaporthales).
Maharachchikumbura et al. (2015), Senanayake et al. (2017a, 2018) and Voglmayr et al. (2017)
885
have confirmed the taxonomic placement in Stilbosporaceae. The genus is characterised by
coelomycetous conidiomata and hyaline conidia, having several apical appendages which are
tubular or filiform, and unbranched.
Natarajania Pratibha & Bhat, Kavaka 33: 129 (2005)
Index Fungorum number: IF522499; 1 species with sequence data.
Type species – Natarajania indica J. Pratibha & Bhat
Notes – This asexual hyphomycetous genus was recorded as a saprobe on leaf litter. The
sexual morph of the genus has not been identified. Natarajania was accommodated in Diaporthales
genera incertae sedis by Senanayake et al. (2017a). Based on molecular data, Senanayake et al.
(2018) has accommodated Natarajania in Stilbosporaceae.
Stegonsporium Corda, Naturalientausch 11: 458 (1827)
Index Fungorum number: IF10071; 40 morphological species (Species Fungorum 2020), 5
species with sequence data.
Type species – Stegonsporium pyriforme (Hoffm.) Corda
Notes – Members of the genus are saprobes or mild pathogens in overwintered plants such as
Acer sp. (Senanayake et al. 2018). The genus has both sexual and asexual morphs and
Stegonsporium shows a high similarity with Stilbospora (Voglmayr & Jaklitsch 2008).
Stegonsporium has pyriform conidia which bear 2–7 transverse and 1–3 longitudinal distosepta. In
addition the ascospores of the genus are distoseptate. These two significant characters separate
Stegonsporium from Stilbospora (Voglmayr & Jaklitsch 2008).
Stilbospora Pers., Neues Mag. Bot. 1: 93 (1794)
Index Fungorum number: IF10116; 16 morphological species (Species Fungorum 2020), 3
species with sequence data.
Type species – Stilbospora macrosperma Pers.
Notes – The type species Stilbospora macrosperma was epitypified by Voglmayr & Jaklitsch
(2014). Prosthecium (the asexual morph) includes opportunistic and moderately phytopathogenic
taxa, which are responsible for the branch dieback or twig blight in several overwintered plants
(Voglmayr & Jaklitsch 2008). Some species of Prosthecium were also recorded from corticated and
dead branches of the same plants (Voglmayr & Jaklitsch 2014). In this entry, we illustrated the
sexual morph of Stilbospora macrosperma.
Sydowiellaceae Lar.N. Vassiljeva, Pirenomits. Lokuloaskomits. Severa Dal'nego Vostoka
(Leningrad): 210 (1987)
Index Fungorum number: IF81867; Facesoffungi number: FoF06882; 47 species.
Saprobic or parasitic on plant matter. Sexual morph: Stromata well- or poorly developed,
prosenchymatous, scattered, immersed to erumpent, appearing as an aggregation of ostioles,
rounded or elliptic, dark brown to black, composed of compact pseudoparenchymatous tissues,
several ascoma in a stroma, some species turning umber in 5% KOH. Ascomata solitary or
aggregated, immersed or erumpent, globose to subglobose, sometimes circinate, coriaceous, with
central or asymmetrically located ostiolar canal opening through an individual or converged ostiole,
internally covered by filamentous, hyaline periphyses, sometime ostiolar opening wider than canal,
black to brown. Peridium comprising a few layers of brown, thick-walled cells of textura
angularis. Paraphyses cellular, septate, branched, hyaline. Asci 8-spored, unitunicate, cylindrical to
subglobose, short pedicellate, apex blunt, with J-, apical ring. Ascospores uni- to multi-seriate,
hyaline, pale brown to dark brown, filamentous, ellipsoid or long fusoid-cylindrical, 1–11-septate,
sometimes with apical and basal appendages, wall smooth or verruculose ornamentation. Asexual
morph: Coelomycetous. Conidiomata sometimes stromatic, pycnidia, uniloculate, superficial,
aggregated 3–5 in one group, globose, orange to brown. Conidiomatal wall comprising thickwalled, orange cells of textura angularis. Conidiophores elongate, branched, hyaline, few
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conidiogenous cells arising from one conidiophore, attached to conidiomatal wall. Conidiogenous
cells cylindrical, hyaline, ampulliform, septate, ends pointed, phialidic. Conidia ovoid to ellipsoid,
unicellular, hyaline, smooth-walled.
Type genus – Sydowiella Petr.
Notes – Vasilyeva (1987) introduced this family to accommodate an assortment of diversified
taxa. Vasilyeva (2001) and Kruys & Castlebury (2012) introduced several genera to this family.
Senanayake et al. (2017b) revised this family and introduced several genera and species following a
phylogenetic analysis. Voglmayr & Mehrabi (2018) accommodated Caudospora in this family
introducing C. iranica as new species. Currently this family comprises 16 genera.
Figure 237 – Stilbospora macrosperma (Material examined – AUSTRIA, Niederösterreich,
Rekawinkel, grid square 7862/1, on a trunk of Carpinus betulus L. (Betulaceae), 20 October 2001,
W. Jaklitsch, W.J. 1840, D25, WU 24708, epitype). a Herbarium specimen. b Ascomata on
substrate. c Cut showing ascomata in stroma. d Vertical section of stroma. e-g Ascospores. h-j Asci.
Scale bars: b, c = 200 μm, d = 20 μm, e-g = 5 μm, h-j = 20 μm.
Ecological and economic significance of Sydowiellaceae
Species of Sydowiellaceae are saprobes, endophytes or pathogens on herbaceous,
dicotyledonous and hardwood trees. Sydowiella species was reported on leaves and stem cankers
(Nordskog et al. 2003). Cainiella johansonii occurs on both living and dead leaves and petioles of
Dryas species, while Cainiella borealis occurs as a saprobe on dead branches of Cassiope
tetragona (Kruys & Castlebury 2012). Paragnomonia fragariae causes a dieback disease of
strawberry (Klebahn 1918). Hapalocystis berkeleyi forms stromatic fruiting bodies on dead woody
plants. Most of other members in this family are saprobes and help in litter decomposition.
Genera included in Sydowiellaceae
Alborbis Senan. & K.D. Hyde, Mycosphere 8(1): 183 (2017)
Index Fungorum number: IF552717; 1 species with sequence data.
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Type species – Alborbis galericulata (Tul. & C. Tul.) Senan. & K.D. Hyde
Notes – Alborbis is a monotypic genus introduced based on Valsa galericulata. This species
is a mild pathogen causing canker of Fagus species and saprobic on branches of Fagus sylvatica
(Mejía et al. 2011). Mejía et al. (2011) suggested that Valsa galericulata has an affiliation with
Sydowiellaceae based on the molecular data and Senanayake et al. (2017b) introduced Alborbis to
accommodate this taxon. This genus is characterised by inconspicuous or astromatic ascomata and
ellipsoid to cylindrical, 1-septate, 4-guttulate ascospores with short appendages.
Breviappendix Senan. & K.D. Hyde, Mycosphere 8(1): 186 (2017)
Index Fungorum number: IF552719; 3 species with sequence data.
Type species – Breviappendix rubi (Rehm) Senan., Maharachch. & K.D. Hyde
Notes – Breviappendix comprises three species, B. rubi, B. rosae and B. rostellata
(Senanayake et al. 2017b). Breviappendix rubi was reported as a severe stem canker and dieback
forming pathogen of various cultivated and wild Rubus species in both Europe and America
(Nordskog et al. 2003).
Cainiella E. Müll., Sydowia 10(1-6): 120 (1957)
Index Fungorum number: IF720; 2 species with sequence data.
Type species – Cainiella johansonii (Rehm) E. Müll.
Notes – Cainiella was introduced by Muller (1957) based on Lizonia johansonii. Cainiella
comprises C. borealis in addition to the type species. This genus is characterized by dark, immersed
ascomata, with long-necks, deliquescing paraphyses, as well as asci with a J-, refractive ring and 1septate ascospores. Cainiella species are saprobic or very mild pathogens on arctic dwarf shrubs
and prostrate evergreen subshrubs.
Calosporella J. Schröt., Krypt.-Fl. Schlesien (Breslau) 3.2(4): 442 (1897)
Index Fungorum number: IF757; 1 species with sequence data.
Type species – Calosporella innesii (Curr.) J. Schröt.
Notes – Calosporella was typified with C. innesii using a specimen collected from Austria
associated with dead corticated branches of Acer pseudoplatanus (Voglmayr & Jaklitsch 2014).
Calosporella innesii has hyaline, mostly 3-septate ascospores, with small apical appendages.
Caudospora Starbäck, Bih. K. svenskaVetensk Akad. Handl.,Afd. 3 15(no. 2): 11 (1889)
Index Fungorum number: IF854; 2 species with sequence data.
Type species – Caudospora taleola (Fr.) Starbäck
Notes – Caudospora is characterized by immersed perithecia in corticolous pseudostromata
delimited by a distinct blackish zone, a whitish ectostromatic disc with few converging ostioles,
cylindrical asci and hyaline, 2-celled ascospores with or without an appendage at each end and 2–3
median appendages. Caudospora taleola is a saprobe or weak pathogen which can cause canker
diseases (Phillips & Burdekin 1992).
Chapeckia M.E. Barr, Mycol. Mem. 7: 164 (1978)
Index Fungorum number: IF983; 2 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Chapeckia nigrospora (Peck) M.E. Barr
Notes – Chapeckia was introduced based on Diatrype nigrospora (Barr 1978). This genus
comprises Chapeckia ribesia as the second species. Chapeckia was placed in Sydowiellaceae based
on molecular data (De Silva et al. 2009, Kruys & Castlebury 2012, Senanayake et al. 2017b).
Hapalocystis Auersw., Fungi rhenaniexsic., fasc. 6: no. 585 (1863)
Index Fungorum number: IF2222; 7 morphological species (Species Fungorum 2020), 3
species with sequence data.
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Type species – Hapalocystis berkeleyi Auersw.
Notes – Hapalocystis was established to accommodate Sphaeria hapalocystis and Sphaeria
inquinans var. ulmi as Hapalocystis berkeleyi and H. bicaudata respectively (Fuckel 1863).
Clements & Shear (1931) designated H. berkeleyi as the type species. The asexual morph of
Hapalocystis was reported as a phoma-like asexual morph for H. berkeleyi from culture (Glawe
1985, Liu et al. 2015).
Figure 238 – Hapalocystis berkeleyi (Material examined – GERMANY, Rhineland, Hattenheim
(near), Gruenau, L. Fuckel (Fungi rhenani exs. 585; ex herb. C.E. Brome), isotype, K (M) 195743;
ITALY, Province of Forlı`-Cesena, Modigliana, Montebello, on branch of Platanus hybrid, 14
April 2013, E. Camporesi, MFLU 14–0798). a Conidiomata on MEA. b Conidiophores,
conidiogenous cells and conidia. c Conidia. Scale bars: a = 500 μm, b = 10 μm, c = 5 μm.
Italiomyces Senan., Camporesi & K.D. Hyde, Mycosphere 8(1): 196 (2017)
Index Fungorum number: IF552721; 1 species with sequence data.
Type species – Italiomyces centaureae Senan., Camporesi & K.D. Hyde
Notes – Italiomyces is distinct in its bi-guttulate or rarely 4-guttulate ascospores with minute,
spine-like apical appendages. Phylogenetically, Italiomyces forms a distinct clade in
Sydowiellaceae.
Lambro Racib., Parasit. Alg. Pilze Java's (Jakarta) 2: 13 (1900)
Index Fungorum number: IF552801; 3 morphological species (Species Fungorum 2020).
Type species – Lambro insignis Racib.
Notes – Lambro insignis produces necrotic spots in leaves of Sterculia subpeltata (Raciborski
1900). The asexual morph of Lambro is reported as a cylindrosporella-like.
Paragnomonia Senan. & K.D. Hyde, Mycosphere 8(1): 198 (2017)
Index Fungorum number: IF552723; 1 species with sequence data.
Type species – Paragnomonia fragariae (Kleb.) Senan. & K.D. Hyde
Notes – Paragnomonia fragariae was isolated from dead petioles and leaves of strawberry
and causes strawberry decline.
Ranulospora Senan., Camporesi & K.D. Hyde, Mycosphere 8(1): 202 (2017)
Index Fungorum number: IF552725; 1 species with sequence data.
Type species – Ranulospora alnea Senan., Camporesi & K.D. Hyde
Notes – Ranulospora alnea is a saprobe collected from bark of grey alder. This genus
comprises grouped ascomata combined by brown, hyphae.
Rossmania Lar.N. Vassiljeva, Mycoscience 42(4): 401 (2001)
Index Fungorum number: IF28578; 2 morphological species (Species Fungorum 2020), 1
species with sequence data.
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Type species – Rossmania ukurunduensis Lar.N. Vassiljeva
Notes – Rossmania was introduced to accommodate taxa having clustered ascomata with
long beaks, a non-united ectostromatic disc, and septate, elongate ascospores (Vasilyeva 2001).
Rossmania comprises R. ukurunduensis and R. aculeata (Vasilyeva 2001). Rossmania
ukurunduensis was collected in Russia associated with Acer ukurunduense, while R. aculeata was
collected from Sri Lanka associated with Theaceae (Vasilyeva 2001).
Sillia P. Karst., Bidr. Känn. Finl. Nat. Folk 23: 159, 251 (1873)
Index Fungorum number: IF5029; 8 morphological species (Species Fungorum 2020), 3
species with sequence data.
Type species – Sillia ferruginea (Pers.) P. Karst.
Notes – Ascospores of Sillia are filiform or acicular, scolecosporous and multi-septate.
Rossman et al. (2007) was accommodated Sillia in Sydowiellaceae based on its morphology. De
Silva et al. (2009), Kruys & Castlebury (2012) and Senanayake et al. (2017b) placed Sillia in
Sydowiellaceae based on phylogeny.
Sydowiella Petr., Annls mycol. 21(1/2): 30 (1923)
Index Fungorum number: IF5312; 9 morphological species (Species Fungorum 2020), 5
species with sequence data.
Type species – Sydowiella fenestrans (Duby) Petr.
Notes – Sydowiella are mild pathogens or saprobes characterized by solitarily, erumpent
ascomata without stromatic tissues and 2-celled ascospores (Kobayashi 1970). Sydowiella species
are mostly distributed in temperate regions (Farr & Rossman 2018). Betula, Centaurea, Dryas,
Epilobium, Juncus, Rubus, and Urtica are the common host genera from which Sydowiella species
have been collected. In this entry we illustrate the sexual morph of Sydowiella fenestrans. The
asexual morph of Sydowiella has not been reported.
Tenuiappendicula Senan., Camporesi & K.D. Hyde, Mycosphere 8(1): 207 (2017)
Index Fungorum number: IF552728; 1 species with sequence data.
Type species – Tenuiappendicula alnicola Senan., Camporesi & K.D. Hyde
Notes – Tenuiappendicula is characterized by 1-septate ascospores with long, thin
appendages and asci without an apical ring.
Tortilispora Senan. & K.D. Hyde, Mycosphere 8(1): 208 (2017)
Index Fungorum number: IF552802; 3 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Tortilispora aurantiaca (Wehm.) Senan. & K.D. Hyde
Notes – Tortilispora was introduced to accommodate some Winterella species and currently
this genus comprises T. aurantiaca, T. albofusca and T. cinctula. Tortilispora species are
characterized by multi-septate elongated ascospores, and valsoid ascomata.
Synnemasporellaceae X.L. Fan & J.D.P. Bezerra, Persoonia 40: 130 (2018)
Index Fungorum number: IF823994; Facesoffungi number: FoF05208; 2 species.
Saprobic, pathogenic or endophytic on several hosts. Sexual morph: Pseudostromata
appearing upon the bark surface as pustules containing a small cluster of ostioles developing
through the adherent periderm, covered by a whitish pulverulence. Stromatic zones absent.
Ascomata perithecial, sphaerical or slightly flattened, with long necks, thickly clustered underneath
the ectostromatic disks. Asci clavate. Ascospores biseriate, fusoid-ellipsoid, 2-celled, hyaline,
usually with a short, hyaline, bristle-like appendage at both ends. Asexual morph: Coelomycetous
or hyphomycetous. Conidiomata pycnidial or synnematal. Synnemata long and determinate,
parallel, comprised of slender, cylindrical black stalks emerging from host tissue and a sphaerical,
capitate, shiny black mass of conidia, conidiogenous cells zone concave, dark. Pycnidia centrally
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ostiolate, hemisphaerical, immersed, slightly erumpent. Conidiophores aggregated, straight to
curved. Conidiogenous cells aggregated, hyaline, straight to curved, cylindrical. Conidia cylindrical
to clavate, with a discrete hilum, smooth, pale brown (adapted from Fan et al. 2018).
Figure 239 – Sydowiella fenestrans (Material examined – GERMANY, Hessen, Wiesenwald pr.
Oestrich (Nassau), on Epilobii angustifolii, Fuckel, 1894, F109251, holotype). a Herbarium
specimen. b, c Ascomata on host surface. d Cross section of ascoma. e Peridium. f-h Asci. i-k
Ascospores. Scale bars: b, c = 200 µm, d = 100 µm, e-h = 20 µm, i-k = 10 µm.
Type genus – Synnemasporella X.L. Fan & J.D.P. Bezerra
Notes – Synnemasporellaceae was established by Fan et al. (2018) to accommodate taxa with
synnematous conidiomata and which lack the usual characters of any of the hyaline,
didymosporous, stromatic genera in Diaporthales. Based on phylogenetic analyses,
Synnemasporellaceae forms a well-supported clade between the families Apiosporopsidaceae and
Juglanconidaceae. Species in the family have distinct synnemata, ascomatal and/or conidiomatal
characters, ascospore shape, conidiogenous cells and conidia as well as different fungi-host
associations and disease symptoms from species in Apiosporopsidaceae and Juglanconidaceae
(Senanayake et al. 2017a, Voglmayr et al. 2017). Synnemasporellaceae has also been placed in a
sister clade with Melanconidaceae (Mejía et al. 2011), the latter family having different conidia
colour and hilum to the species in Synnemasporellaceae (Wehmeyer 1933b).
Ecological and economic significance of Synnemasporellaceae
Species belonging to Synnemasporellaceae are saprobic, endophytic or pathogenic.
Synnemasporella toxicodendri has been reported to grow solely on diseased wood of
Toxicodendron sylvestre in China, while S. aculeans affects several species of Rhus, namely, Rhus
copallina, R. diversiloba, R. glabra, R. javanica, R. typhina and R. vernix mainly in Japan and the
USA (Wehmeyer 1933b, Kobayashi 1970, Mejía et al. 2011), and R. chinensis in China (Fan et al.
2018).
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Genus included in Synnemasporellaceae
Synnemasporella X.L. Fan & J.D.P. Bezerra, Persoonia 40: 130 (2018)
Index Fungorum number: IF823995; 2 species with sequence data.
Type species – Synnemasporella toxicodendri X.L. Fan & J.D.P. Bezerra
Notes – Synnemasporella accommodates two species, S. aculeans and S. toxicodendri (Fan et
al. 2018). Cryptodiaporthe aculeans, which has perithecial ascomata as well as sporodochial and/or
pycnidial conidiomata (Fan et al. 2018), was transferred to Synnemasporella. Cryptodiaporthe
aculeans was introduced by Wehmeyer (1933b) who noted that Cryptodiaporthe contained a
heterogeneous group of species which would probably be separated into several genera when their
relationships were fully understood. Initially, Cryptodiaporthe was considered a synonym of
Plagiostoma (Sogonov et al. 2008, Mejía et al. 2011, Voglmayr et al. 2017). However,
phylogenetic analyses (Senanayake et al. 2017a) indicated that they are two separate genera.
Synnemasporella toxicodendri differs from Cryptodiaporthe aculeans in having longer synnemata,
a flat to slightly concave conidiogenous cell on the top of the synnemata and smaller conidia.
Synnemasporella toxicodendri is illustrated for this entry.
Telimenaceae Mardones, T. Trampe & M, Piepenbr., Persoonia 39: 83 (2017)
Index Fungorum number: IF818222; Facesoffungi number: FoF06883; 39 species.
Mostly biotrophic on monocotyledons and dicotyledons (except Poaceae). Sexual morph:
Stroma of various shapes, covered by a cuticular or epidermal shiny blackened clypeus, with
limited development around the ostiole or extensively above the ascomata and sometimes below
the ascomata. Pseudostroma strongly developed, interfusing and conspicuously expanding into the
host tissue. Ascomata perithecial, amphigenous, epiphyllous or hyphophyllous, uni- to
multiloculate, sometimes confluent, frequently surrounded by a bright yellow to reddish coloured
zone, subcuticular, epidermal, subepidermal or immersed in the host tissue, pyriform, globose,
lenticular, or deformed by vascular bundles, with a periphysate ostiole. Peridium hyaline to
pigmented, composed of cells of textura intricata. Paraphyses hyaline, thin-walled, slightly longer
than the asci, septate, often dissolving during maturation. Asci 8-spored, rarely 4-spored,
unitunicate, clavate or cylindrical, usually, apical ring often J-. Ascospores 1–2-seriate, usually
hyaline, rarely pale brown, globose to filiform, mostly cylindrical, thin and smooth-walled, aseptate
to 3-septate, sometimes surrounded mucilaginous sheath. Spermatogonia infrequently found,
pycnidial, spermatiogenous cells cylindrical, tapering towards the tip, proliferating percurrently,
with filiform, hyaline, aseptate scolecospores, probably spermatial in function. Asexual morph:
Undetermined (adapted from Mardones et al. 2017).
Type genus – Telimena Racib.
Notes – Mardones et al. (2017) redefined Phyllachoraceae and proposed the new family
Telimenaceae with Telimena erythrinae as the type species, resulting in three families in
Phyllachorales. Telimena species occur on monocotyledons and eudicotyledons, except Poaceae,
and generally have enlarged black pseudostroma around the perithecia (Mardones et al. 2017,
2018). Phylogenetic analyses of combined LSU, SSU, ITS, rpb2 and tef1 sequence data showed
that Telimena spp. formed a well-separated monophyletic clade in Phyllachorales (Mardones et al.
2017) and this was also found in Dayarathne et al. (2017).
Ecological and economic significance of Telimenaceae
Telimena species occur as parasites on several monocotyledons and eudicotyledons
(Mardones et al. 2017). The hyphae of Telimena species, such as Telimena zanthoxylicola,
penetrate intracellularly through the host tissue by appressoria-like structures against the host cell
walls, which are then perforated by fine canals of the hyphae to form haustoria-like swellings on
the other side, which can serve in nutrient absorption (Speer 1980a). Hyphae can continually grow
and infect new cells, which are stimulated to divide spontaneously, and this is clearly visible in the
palisade cells (Speer 1980a). On the leaves, infected areas do not spread far beyond the radius of
the clypeus, however, on twigs hyphal growth is less limited, seeming to cause die-back in the
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youngest shoots (Speer 1980a). Telimena erythrinae, is a parasite of the dicotyledonous plant host
Erythrina variegata (Fabaceae) forming necrotic leaf spots (Mardones et al. 2017).
Figure 240 – Synnemasporella toxicodendri (Material examined – CHINA, Zhejiang Province,
Hangzhou City, Linan, Xijing Mountain, N30°15'32.84" E119°43'31.21", 54 m asl, on twigs and
branches of Toxicodendron sylvestre, 22 April 2017, Q. Yang & Z. Du, CF 2017481, holotype). a,
b Habit of synnemata on branches. c Conidiophores and conidiogenous cells and developing
conidia. d-g Conidia. Scale bars: a, b = 1 mm, c-g = 10 µm.
Genus included in Telimenaceae
Telimena Racib., Parasit. Alg. Pilze Java’s (Jakarta) 1: 18. 1900.
Index Fungorum number: IF5364; 39 morphological species (Species Fungorum 2020), 18
species with sequence data.
Type species – Telimena erythrinae Racib.
Notes – Raciborski (1900) introduced Telimena for phyllachora-like species with 3-septate
ascospores. Currently, there are 14 species in this genus (Mardones et al. 2017). Close relationships
of Telimena with the genera Telimenopsis and Telimenella and Telimenochora have been reported
(Petrak 1931, Müller 1975, Barr 1977). These genera are segregated by the shape and septation of
the ascospores (Sivanesan 1987). However, there are no molecular sequence data from the type
material of most Telimena species (Mardones et al. 2017).
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Figure 241 – Telimena picramniae (Material examined – COSTA RICA, Aserri, (as P. antidesma),
on Picramnia antidesma (= P. bonplandiana), 26 June 1923, F.L. Stevens, ILL00005686, holotype,
ILL00005686, ILL00005687, paratypes). a Herbarium material. b Close up of a leaf spot. c Section
through pseudostroma. d Section through ostiolar region. e Peridium. f Paraphyses. g, h Asci. i-l
Ascospores. Scale bars: b, c = 1000 µm, d = 100 µm, g, h 50 µm, f, i-l = 20 µm.
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Thyridiaceae J.Z. Yue & O.E. Erikss., Syst. Ascom. 6(2): 233 (1987)
Index Fungorum number: IF82030; Facesoffungi number: FoF01913; 35 species.
Hemibiotrophic or saprobic on woody substrates. Sexual morph: Ascomata stromatic.
Stromatal tissue immersed, becoming erumpent to superficial, soft-textured, reddish brown to
brightly pigmented. Ascomata immersed in stromata, globose, ostiolate. Ostioles periphysate, with
short or long papilla or necks, sometimes convergent necks merging into one ostiole. Peridium
composed of compressed rows of cells, externally brown, internally hyaline. Paraphyses
filamentous, septate. Asci 8-spored, unitunicate, oblong cylindric, apical ring J-. Ascospores
uniseriate, shades of brown, ellipsoid or biconoid, symmetric, 1-septate or muriform. Asexual
morph: Coelomycetous or hyphomycetous, often developing on the edge of sexual morph stroma.
Conidiomata immersed in stromata, irregular, multilocular and folded inside. Conidiogenous cells
emerging from the interior of the cavity, bearing one or more conidiogenous cells. Conidiogenous
cells enteroblastic, phialidic, hyaline. Conidia ellipsoidal, aseptate, small, hyaline. Holoblastic
sympodial conidia also produced from hyphae. Sometimes ascospores produce primary conidia
(adapted from Barr 1990b, Leuchtmann & Müller 1986, Maharachchikumbura et al. 2016b).
Type genus – Thyridium Nitschke
Notes – In a phylogenetic and molecular clock analyses, Thyridiaceae showed a close
relationship to Annulatascales and Myrmecridiales (Hyde et al. 2017a, Hongsanan et al. 2017,
Senanayake et al. 2017a). However, only a single strain of the type genus has sequence data
(Thyridium vestitum strain AFTOL-ID 172), and the ordinal placement remains uncertain in
Diaporthomycetidae (Maharachchikumbura et al. 2015b, 2016b, Hongsanan et al. 2017).
Maharachchikumbura et al. (2015, 2016b) and Wijayawardene et al. (2018a) accepted three genera
in Thyridiaceae (Mattirolia, Pleurocytospora, and Thyridium). No sequence data are available for
Pleurocytospora species. Mattirolia has been synonymised under Thyronectria (Nectriaceae) by
Jaklitsch & Voglmayr (2014) based on an ex-epitype sequence of the generic type of Mattirolia, M.
roseovirens. Here we follow the treatment of Jaklitsch & Voglmayr (2014).
Ecological and economic significance of Thyridiaceae
Species of Thyridium are saprobic on woody substrates (Checa et al. 2013, Zhou & Hyde
2002). Pleurocytospora species are saprobic or hemibiotrophic in woody substrates (Barr 1990b,
Zhu et al. 2005) and therefore, involved in nutrient recycling. Pleurocytospora taxi was recorded
from Taxus cuspidata, while P. lycii was isolated from Arabidopsis thaliana as an endophyte (Sun
et al. 2003, Wang et al. 2015a, Junker et al. 2012) and there are no records of pathogenic species in
Thyridiaceae.
Genera included in Thyridiaceae
Pleurocytospora Petr., Annls mycol. 21(3/4): 256 (1923)
Index Fungorum number: IF9459; 3 morphological species (Species Fungorum 2020).
Type species – Pleurocytospora vestita Petr.
Notes – Pleurocytospora was introduced to accommodate P. vestita and P. lycii (Petrak
1923). Pleurocytospora taxi was introduced from the host Taxus cuspidata (Sun et al. 2003).
Thyridium Nitschke, Pyrenomyc. Germ. 1: 110 (1867)
Index Fungorum number: IF5465; 32 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Thyridium vestitum (Fr.) Fuckel
Notes – Thyridium was introduced by Nitschke (1867) to accommodate species with
uniseriate, muriform, dark-coloured ascospores, 8-spored asci and filiform paraphyses (Nitschke
1867). Furthermore, species of Thyridium are characterized by KOH+ stromata, cylindrical or
clavate asci, and very pale brown to dark brown muriform ascospores (Checa et al. 2013,
Maharachchikumbura et al. 2016b). Asexual morphs are coelomycetous, or produce holoblastic
sympodial conidia from hyphae (Leuchtmann & Müller 1986).
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Figure 242 – Pleurocytospora vestita (Material examined – SWITZERLAND, Zurich, Zollikon, in
garden, on Ribes rubrum L., 15 May 1986, E. Müller, ZT MYC 58916). a Herbarium material. b
Appearance of conidiomata on the host substrate. c Transverse section of conidiomata. d Surface of
the stromatic tissue. e, f Conidiophores with conidiogenous cells. g, h Conidiogenous cells with
conidia. i Conidia. Scale bars: b = 1 mm, c = 200 µm, d-f = 50 µm, g-i = 10 µm.
Tilachlidiaceae L. Lombard & Crous, Stud. Mycol. 80: 237 (2015)
Index Fungorum number: IF810273; Facesoffungi number: FoF01280; 20 species.
Saprobic or parasitic on other fungi, entomogenous on lepidopterous insects, associated
with bryophytes, or isolated from soil. Sexual morph: Ascomata arising directly from the
mycelium or formed on loosely arranged stromata, perithecial, KOH-. Ostiole apapillate. Asci 6–
8-spored, clavate or cylindrical, short pedicellate. Ascospores hyaline to yellow, irregularly ovoid
or ellipsoidal, muriform, 2–13 celled, with 1–5 transverse septa. Asexual morph: Hyphomycetous.
Conidiophores synnematous or acremonium-like. Synnemata when present simple to branched,
Conidiophores synnematous or acremonium-like. Synnemata when present simple to branched,
cylindrical, narrowing towards the apex, consisting of bundles of parallel, longitudinal, closely
compacted hyphae, with 1–4 phialides. Phialides scattered, cymbiform to cylindrical or allantoid,
aseptate or septate, sometimes with obvious collarettes, narrowing towards the apex, sometimes
integrated in septate branches, hyaline to yellow, smooth or becoming verrucose. Conidia
fusiform to ellipsoid to subcylindrical or oblong, 0–7-septate, with distinct hilum at both ends,
hyaline to yellow, smooth to finely ornamented, with or without mucoid sheath, formed in chains
or agglutinating into large sphaerical or irregular white masses (adapted from Lombard et al.
2015).
Type genus – Tilachlidium Preuss
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Figure 243 – Thyridium lasiacidis (Material examined – FRENCH GUIANA, Saül, vicinity of the
town, elev. ca. 200 m, on dead culms of Lasiacis ligulata, 3–16 February 1986, G.J. Samuels
3784A, NSF BSR 8500236, holotype, NY 01293357). a, b Herbarium material. c Appearance of
stromata on host substrate. d Stomatal tissue in 5% KOH. e Surface view of peridium. f Transverse
section through ascomata. g. Transverse section through the peridium. h Paraphyses in 3% KOH im Asci. n-p Ascospores. Scale bars: c = 500 µm, e = 50 µm, f = 100 µm, g = 50 µm, h-m = 20 µm,
n-p = 10 µm.
897
Notes – Tilachlidiaceae was introduced to accommodate two synnematous genera,
Septofusidium and Tilachlidium, supported by phylogenetic analysis (Lombard et al. 2015). Gams
(1971) had classified Tilachlidium as a member of Hypocreales family incertae sedis, whereas
Septofusidium was placed in Nectriaceae based on morphology. These two genera share similar
asexual morph characters and are saprobic or parasitic on other fungi (Petch 1931b, Mains 1951,
Gams 1971, Samson 1974, Sun et al. 2019). They produce synnematous asexual morphs with
hyaline conidia.
Ecological and economic significance of Tilachlidiaceae
Species of Tilachlidiaceae are saprobes on other fungi such as dried Basidiomycetes. For
example, Tilachlidium brachiatum was recorded from Hypholoma fasciculare and Agaricus
galericulatus (Mains 1951, Lombard et al. 2015, Preuss 1851). They also grow on dead
lepidopterous insects, eg. Tilachlidium ramosum was isolated from a dried lepidopterous larva.
Some are pathogenic on plants (Mains 1951, Kidd & Beaumont 1924). Septofusidium sp. was
reported as a pathogen of guava (Psidium guajava), causing rapid wilt of trees (Grech 1985). Evans
(1974) discussed the potential use of Tilachlidium spp. for biocontrol of ants, which might have an
important agricultural application in the future.
Members of this family have potential medical applications. Some species of Tilachlidium
and Septofusidium produce antibiotics (Gottshall et al. 1951, Roberts 1952), as well as novel
compounds with antineoplastic properties or cytotoxicity to leukemia cells (Feng et al. 2004, Vann
et al. 2016). Ekiz et al. (2016) revealed the broad-spectrum antibacterial activity of a secondary
metabolite from Septofusidium berolinense. Bioactive secondary metabolites from S. berolinense,
with cytotoxic activities were also screened (Ekiz et al. 2016, Vann et al. 2016). Tilachlidium
humicola had been used for the production of L-glutaminases, an important industrial enzyme
(Shindia et al. 2007, Nandakumar et al. 2003).
Genera included in Tilachlidiaceae
Psychronectria J. Pawłowska, Istel, Wrzosek & D. Hawksw., Mycologia 109(4): 604 (2017)
Index Fungorum number: IF821597; 1 species with sequence data.
Type species – Psychronectria hyperantarctica (D. Hawksw.) J. Pawłowska, Istel, Wrzosek
& D. Hawksw.
Notes – The monotypic genus Psychronectria was introduced to accommodate Thyronectria
hyperantarctica (Pawłowska et al. 2017). Psychronectria species are associated with bryophytes
(Pawłowska et al. 2017). The genus is characterised by pink, orange or pale red ascomata, arising
directly from the mycelium or formed on loose stromata and, hyaline to yellow, muriform
ascospores (Pawłowska et al. 2017).
Septofusidium W. Gams, Cephalosporium-artige Schimmelpilze: 147 (1971)
Index Fungorum number: IF9882; 4 morphological species (Species Fungorum 2020), 2
species with sequence data.
Type species – Septofusidium elegantulum (Pidopl.) W. Gams
Notes – Septofusidium was introduced by Gams (1971). Sequence data or cultures are not
available for the generic type S. elegantulum, although S. berolinense and S. herbarum have
molecular data. Septofusidium differs as conidia are arranged in chains, without a mucoid sheath.
Tilachlidium Preuss, Linnaea 24: 126 (1851)
Index Fungorum number: IF10236; 15 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Tilachlidium brachiatum (Batsch) Petch
Notes – Tilachlidium pinnatum was synonymised under T. brachiatum by Lombard et al.
(2015). There are no records for the sexual morphs of Tilachlidium. Species of Tilachlidium are
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saprobic growing on dried fungi or entomogenous on lepidopterous insects (Mains 1951). Sequence
data are available only for the type species T. brachiatum in GenBank.
Figure 244 – Tilachlidium brachiatum (Material examined – CANADA, Nova Scotia, Salmon
River, fragments of lepidopterous larva on a mossy log, 7 September 1931, L.E. Wehmeyer 1474,
MICH 15719, holotype of Tilachlidium ramosum). a Herbarium material. b, c Synnemata on the
host. d-f Synnemata with phialides. g Conidial mass attached to a phialide. h, i Phialides. j, k
Conidia. Scale bars: b = 1 mm, c = 500 µm, d = 200 µm, e = 100 µm, f = 200 µm, g = 5 µm, h, i =
20 µm, j, k = 5 µm.
Tirisporellaceae Suetrong, E.B.G. Jones & K.L. Pang, Cryptog. Mycol. 36(3): 323 (2015)
Index Fungorum number: IF812184; Facesoffungi number: FoF01413; 3 species.
Saprobic on submerged substrates in freshwater or brackish habitats. Sexual morph:
Ascomata solitary or gregarious, partially immersed to superficial, globose to subglobose, black,
and coriaceous to carbonaceous, papillate, ostiolate, with a neck and periphyses. Peridium thickwalled, brown to black. Paraphyses early deliquescent, irregularly wide, septate, hyaline, tapering
towards the apices and embedded in a mucilaginous matrix. Asci 8-spored, unitunicate, cylindrical
to clavate, indistinctly pedicellate, apex with a J-, subapical ring. Ascospores uniseriate or bi-seriate
to tri-seriate, hyaline to brown, 1–7-septate, fusoid, falcate to lunate, straight or curved, cell wall
smooth or verrucose, with or without appendages. Asexual morph: Hyphomycetous. Conidiophores
macronematous, mononematous, erect, pale brown to brown, paler toward the apex, straight or
flexuous, smooth. Conidiogenous cells enteroblastic, monophialidic, integrated on hyphae,
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terminal, sometimes erect on hyphae, pale brown, subcylindrical, ampulliform. Conidia unicellular,
hyaline, fusoid-ellipsoid, obovoid or ellipsoid, apex obtuse or broadly rounded, tapering to a
truncate base, smooth-walled (adapted from Suetrong et al. 2015).
Type genus – Tirisporella E.B.G. Jones, K.D. Hyde & Alias
Notes – Tirisporellaceae was established by Suetrong et al. (2015) to accommodate the
genera, Tirisporella and Thailandiomyces, based on morphology and molecular phylogeny.
Subsequently, Abdel-Wahab et al. (2017) introduced Bacusphaeria (type species: B. nypae), which
formed a distinct clade in this family. Tirisporellaceae retains its uniqueness in having large, black
perithecial ascomata with an ostiolate neck, a thick-walled peridium, cylindrical asci and 2–3seriate, 1–7-septate, fusoid ascospores, with or without appendages. Sequence data are available for
the three genera of this family and phylogenetic analyses confirmed their placement in
Tirisporellales (Jones et al. 2015). Hyde et al. (2017a) carried out divergence estimates of
Sordariomycetes and estimated the stem age of Tirisporellaceae as 112 MYA.
Ecological and economic significance of Tirisporellaceae
Tirisporellaceae species are saprobes on wood or palm fronds found in freshwater to brackish
water habitats (Jones et al. 1996, Pinruan et al. 2008, Abdel-Wahab et al. 2017). Tirisporella
beccariana and Bacusphaeria nypae grow on the basal petioles of Nypa fruticans, which is
perennially submerged in brackish water. Those species may have salt resistance to survive in this
environment.
Genera included in Tirisporellaceae
Bacusphaeria Norlailatul, Alias et Suetrong, Botanica Marina 60 (4): 11 (2017)
Index Fungorum number: IF812182; 1 species with sequence data.
Type species – Bacusphaeria nypae Norlailatul, Alias & Suetrong
Notes – The monotypic genus Bacusphaeria is similar to Tirisporella, because they have
large, thick-walled ascomata, cylindrical asci and both occur on the basal petioles of Nypa fruticans
in brackish water. However, Bacusphaeria is distinct from Tirisporella because it; i) lacks complex
ascospore appendages; ii) has fewer septa in the ascospores (Bacusphaeria: 1–5-septate vs.
Tirisporella: 5–7-septate); iii) lack of paraphyses; iv) has a prominent apical ring in contrast to faint
apical ring in Tirisporella; v) having generally uniseriate ascospores in contrast to biseriate
ascospores in Tirisporella. Base pair differences of Bacusphaeria nypae (MFLU 13-0617) and
Tirisporella beccariana (BCC38300) are 97 bp different out of 859 bp (11.29%) without gaps in
the LSU gene locus. Furthermore, in phylogeny, Bacusphaeria nypae is distinct from the other two
genera and formed a separate lineage in Tirisporellaceae (Fig. 7).
Thailandiomyces Pinruan, Sakay., K.D. Hyde & E.B.G. Jones, Fungal Divers. 29: 91 (2008)
Index Fungorum number: IF511583; 1 species with sequence data.
Type species – Thailandiomyces bisetulosus Pinruan, Sakay., K.D. Hyde & E.B.G. Jones
Notes – Pinruan et al. (2008) introduced Thailandiomyces to accommodate a single species T.
bisetulosus and described its asexual morph in Craspedodidymum. Their phylogenetic analysis
based on LSU and SSU sequence data showed that the genus is well-placed in Diaporthales, but
differs from Diaporthe species in morphology. Suetrong et al. (2015) conducted phylogenetic
analysis of Tirisporella beccariana and revealed that Thailandiomyces formed a sister clade to
Tirisporella and therefore included Thailandiomyces in Tirisporellaceae. Thailandiomyces
bisetulosus is distinct from other members in Tirisporellaceae by its hyaline ascospores with
bipolar appendages.
Pinruan et al. (2008) showed that Thailandiomyces bisetulosus is well-positioned in
Diaporthales (Sordariomycetes, Sordariomycetidae) using SSU and LSU sequence analyses (Zhang
et al. 2006, Hibbett et al. 2007), but was not referred to any family. Suetrong et al. (2015)
introduced Tirisporellaceae to accommodate this taxon. Thailandiomyces bisetulosus resemble the-
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Figure 245 – Thailandiomyces bisetulosus (Material examined – THAILAND, Narathiwat,
Sirindhorn Peat Swamp Forest, on submerged trunk of Licuala longicalycata, 12 May 2001, U.
Pinruan, BBH Wah 110, holotype). a Herbarium material. b, c Long necked ascomata. d Vertical
section through ascoma. e Peridium. f Periphysate ostiolar region. g Paraphyses. h, i Asci. j, k
Ascospores. Scale bars: c = 500 μm, d, h, i = 50 μm, e-g = 20 μm, j, k = 10 μm.
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-morphologies of Diaporthales, including its saprobic habitat on decaying plant material, partiallyimmersed ascomata, long periphysate necks, unbranched paraphyses that deliquesce early in
development, unitunicate asci that float free in the centrum and asci with a refractive, J-, apical ring
(Barr 1991, Samuels & Blackwell 2001).
Tirisporella E.B.G. Jones, K.D. Hyde & Alias, Can. J. Bot. 74(9): 1489 (1996)
Index Fungorum number: IF27659; 1 species with sequence data.
Type species – Tirisporella beccariana (Ces.) E.B.G. Jones, K.D. Hyde & Alias
Notes – The monotypic genus Tirisporella was introduced by Jones et al. (1996) to
accommodate the type species T. beccariana (≡ Sphaeria beccariana), which was frequently
encountered on intertidal petioles of Nypa fruticans (mangrove palm). This genus has historically
been classified in the Loculoascomycetes incertae sedis, and Pleosporales incertae sedis because of
the bitunicate-like asci (Jones et al. 2009), while the familial placement was confirmed and
assigned into Sordariomycetes based on phylogenetic analysis (Suetrong et al. 2015). The most
obvious characters of Tirisporella are the first basal septum delimiting a hyaline to light-coloured
basal cell, the remaining cells brown and verrucose ascospores with apical appendages. During
examination of intertidal fungi from Nypa fruticans, fresh collections of T. beccariana were made
and enabled a re-description and illustration of the fungus, and herein we provide an updated
phylogenetic tree with all strains of this order. Tirisporella beccariana is illustrated in this entry.
Togniniaceae Réblová, L. Mostert, W. Gams & Crous, Stud. Mycol. 50: 540 (2004)
Index Fungorum number: IF500154; Facesoffungi number: FoF01414; 64 species.
Saprobic on dead wood or pathogenic on human and various woody plants, frequently
associated with Petri and Esca disease of grapevines. Sexual morph: Ascomata perithecial, black,
superficial to subimmersed, astromatic, with 1–3 necks. globose to subglobose base, with straight
or flexuous, periphysate ostiole. Peridium fragile to leathery, two-layered, the outer layer
comprising thin-walled, brown cells of textura prismatica to textura angularis, the inner layer
comprising non-pigmented flattened cells. Paraphyses abundant, hyaline, branching, septate,
attenuate towards the apex, slightly constricted at septa. Ascogenous hyphae hyaline, elongate,
branched, smooth-walled, sympodial, bearing remnant of base from which a single ascus is
produced. Asci 8-spored, unitunicate, clavate, with obtuse, sessile bases and thickened ascal apex
without a discharge mechanism, aggregating in spicate arrangement. Ascospores 3–4-seriate,
hyaline, aseptate, allantoid, oblong-ellipsoidal, slightly curved, with rounded ends. Asexual morph:
Hyphomycetous. Mycelium mid-brown, dense, branched, septate, some species produce wart-like
droplets. Conidiophores arising from bundled or single mycelium, branched at base or unbranched,
septate, pale brown, paler towards the apex, with small warts or verruculose ornamentations at the
base, percurrent rejuvenation. Conidiogenous cells usually monophialidic, integrated or discrete,
hyaline to pale brown, with or without basal septum. Conidia hyaline, septate, smooth-walled,
cylindrical, oblong-ellipsoidal or allantoid, straight or slightly curved, generally adhering in
globose head at the tip of phialides (adapted from Réblová et al. 2004).
Type genus – Phaeoacremonium Berl.
Notes – Togninia was previously placed in the Calosphaeriales (Mostert et al. 2003).
However, phylogenetic analysis based on a combined LSU and SSU dataset showed that Togninia
spp. formed a distinct clade in Diaporthales, for which Togniniaceae was proposed (Réblová et al.
2004) and this was followed by later researchers (Gryzenhout et al. 2006b, Damm et al. 2008,
Réblová 2008). Subsequently, Togniniaceae was relocated to a new order Togniniales based on its
phylogenetic and morphological distinctiveness in the class Sordariomycetes and this family was
considered to include three genera Conidiotheca, Togninia and Phaeoacremonium
(Maharachchikumbura et al. 2015b). Gramaje et al. (2015) reduced Togninia under its asexual
genus Phaeoacremonium and this was accepted by Dai et al. (2016) and Maharachchikumbura et
al. (2016b). To date, Togniniales comprised a single family Togniniaceae, but Conidiotheca was
treated as a member of Calosphaeriaceae, order Calosphaeriales (Wijayawardene et al. 2018a).
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Figure 246 – Tirisporella beccariana (Material examined – THAILAND, Ranong, Ngao (Ranong)
Mangrove Forest Research Center, on intertidal petiole of Nypa fruticans Wurmb., 7 December,
2016, S.N. Zhang, SNT82, living culture MFLUCC 18-1572, specimen voucher MFLU 18-1582,
HKAS 97482; THAILAND, Prachuap Khiri Khan, Pak Nam Pran, on intertidal petiole of Nypa
fruticans, 2 December, 2016, S.N. Zhang, SNT102, living culture MFLUCC 18-1571, specimen
voucher MFLU 18-1584; THAILAND, Krabi, Pali, on intertidal petiole of Nypa fruticans, 30
August, 2017, S.N. Zhang, SNT203, specimen voucher MFLU 18-1585, HKAS 97483). a, b
Appearance of ascomata on host surface with ostioles. c Vertical section through the ascoma. d-g
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Asci. h Ostiole with periphyses. i Structure of peridium. j Paraphyses. k Apex of ascus in Lugol’s
iodine, with a J-, apical ring. l-p Ascospores. q Germinating spore. r Colony on PDA. s-u Asexual
morph structure in culture. Scale bars: a = 500 μm, b, c = 200 μm, d-g = 50 μm, h-j, l-q = 20 μm, k,
s, u = 10 μm, t = 5 μm.
Ecological and economic significance of Togniniaceae
Phaeoacremonium species are found on wide range of hosts, including animals, plants and
humans. They can be saprobic, parasitic, or hyperparasitic and play an important role in agriculture,
forestry and medicine (Mostert et al. 2006, Reátegui et al. 2006). Many species of
Phaeoacremonium causes trunk diseases on forest and ornamental trees (e.g. willow and poplar)
resulting in severe and widespread tree decline and mortality (Hashemi & Mohammadi 2016,
Kazemzadeh et al. 2017). Phaeoacremonium minimum is a pathogenic species associated with petri
disease and esca symptoms in young grapevines worldwide (Crous & Gams 2000, Graniti et al.
2000, Fourie & Halleen 2002, 2004, Halleen et al. 2003, Edwards & Pascoe 2004, Feliciano et al.
2004, Essakhi et al. 2008, Gramaje et al. 2009). Phaeoacremonium minimum was also reported as a
pathogen causing subcutaneous lesions on human fingers (Choi et al. 2011). In addition, novel
bioactive compounds from Phaeoacremonium species have been investigated. Compounds with
antifungal bioactivity against Aspergillus flavus and Fusarium verticillioides (Reátegui et al. 2006),
lactone derivatives with activity against the phytopathogenic fungi (Silva et al. 2017),
naphthalenone which is involved in the symptom expression of esca-effected grapevines (Evidente
et al. 2000, Abou-Mansour et al. 2004) and phytotoxins p-hydoxybenzaldehyde (Tabacchi et al.
2000) have been recovered from Phaeoacremonium species.
Genera included in Togniniaceae
Conidiotheca Réblová & L. Mostert, Mycol. Res.111(3): 305 (2007)
Index Fungorum number: IF510227; 1 morphological species.
Type species – Conidiotheca tympanoides (M.E. Barr) Réblová & L. Mostert
Notes – Conidiotheca was introduced by Réblová & Mostert (2007) as a monotypic genus to
accommodate Conidiotheca tympanoides (≡ Romellia tympanoides). Lack of molecular data and
absence of characteristics of its asexual morph resulted in placing this genus as incertae sedis in
perithecial ascomycetes (Réblová & Mostert 2007). The taxonomic placement of Conidiotheca is
still unresolved. All studies of this genus were based on those descriptions provided by Réblová &
Mostert (2007). Lumbsch & Huhndorf (2010) listed Conidiotheca under Calosphaeriaceae
(Calosphaeriales), this treatment was followed by Wijayawardene et al. (2017a, 2018a). However,
Maharachchikumbura et al. (2015, 2016b) considered Conidiotheca as a member of Togniniaceae.
The exact familial position should be established with further fresh collections and based on both
morpho-phylogenetic examinations.
Phaeoacremonium W. Gams, Crous & M.J. Wingf., Mycologia 88(5): 789 (1996)
Index Fungorum number: IF27679; 63 morphological species (Species Fungorum 2020), 62
species with sequence data.
Type species – Phaeoacremonium parasiticum (Ajello, Georg & C.J.K. Wang) W. Gams,
Crous & M.J. Wingf.
Notes – Phaeoacremonium was introduced by Crous et al. (1996) for five new species with P.
parasiticum as the type species. They mentioned this genus is similar to Phialophora and
Acremonium. Mostert et al. (2003) confirmed Phaeoacremonium as the asexual morph of Togninia
based on their morphological comparison, sexual compatibility and DNA phylogeny. Togninia was
introduced by Berlese (1990) with Togninia minima as the type species. Togninia has previously
classified in Calosphaeriaceae, Calosphaeriales (Berlese 1900, Barr 1985, Mostert et al. 2003).
Réblová et al. (2004) erected Togniniaceae in Diaporthales to accommodate Phaeoacremonium and
Togninia based on phylogenetic analysis of LSU and SSU sequence data. Subsequently, Mostert et
al. (2006) added several species to both Phaeoacremonium and Togninia, coupled with the generic
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descriptions. Currently, 67 epithets of Phaeoacremonium are listed in Index Fungorum (2020).
Among them, Phaeoacremonium chlamydosporum was transferred to Phaeomoniella (Gams &
Crous 2000). Phaeoacremonium aleophilum, P. mortoniae and P. novae-zealandiae were
synonymized under P. minimum, P. fraxinopennsylvanicum and P. leptorrhynchum, respectively
(Réblova 2011, Gramaje et al. 2015). To date, 63 species are accepted in this genus, of which P.
aquaticum only have ITS gene sequence available. For P. inconspicuum, no ex-type culture or
DNA exists (Gramaje et al. 2015). The remaining 62 species have been confirmed with molecular
data.
Herewith, we illustrated Phaeoacremonium inconspicuum (S F6209, holotype), an old
collection without any available ex-type culture or DNA sequence data. The previous description
was provided with hand-drawn illustrations (Eriksson & Yue 1990). But the morphology of this
species are poorly described. Gramaje et al. (2015) treated it as a new combined species of P.
inconspicuum based on the original description derived from Eriksson & Yue (1990). Here we
refine it to clarify its classification with the microscopic character of ascomata, asci and ascospores
and make sure that this species belonging to this genus.
Torpedosporaceae E.B.G. Jones & K.L. Pang, Cryptog. Mycol. 35(2): 135 (2014)
Index Fungorum number: IF91154; Facesoffungi number: FoF01099; 3 species.
Saprobic on intertidal mangrove wood and roots, bark leaves, and sand in marine habitats.
Sexual morph: Ascomata perithecial, hyaline, immersed or superficial, subglobose, ostiolate,
papillate, subcarbonaceous to coriaceous. Paraphyses narrow, irregular, persistent or early
deliquescing. Asci 8-spored, unitunicate, thin-walled, clavate to ellipsoidal, short pedicellate,
lacking an apical ring, early deliquescing. Ascospores fasciculate, hyaline, cylindrical to ellipsoidal,
3–5-septate, with several radiating appendages at one or both ends. Asexual morph:
Hyphomycetous. Hyphae septate, branched, hyaline. Conidiophores present or obsolete,
cylindrical, clavate, septate or aseptate, acrogenous or laterally on the hyphae, hyaline to light
brown. Conidia holoblastic, irregularly helicoid, muriform, cells of the conidia tightly fused, more
or less similar in size and colour, acrogenous, solitary, constricted at the septa, yellow to brown.
Conidial cells up to 50 in number (adapted from Maharachchikumbura et al. 2016b).
Type genus – Torpedospora Meyers
Notes – Torpedosporaceae was introduced with two Torpedospora species and
Glomerulispora mangrovis (asexual morph), based on molecular analysis of partial sequences of
the nuclear SSU and LSU ribosomal DNA (Jones et al. 2014). Hence, the family comprised two
genera, Torpedospora and Glomerulispora (Jones et al. 2014). However, G. mangrovei, grouped
with the two Torpedospora species such that these generic names are considered synonyms (AbdelWahab et al. 2010, Jones et al. 2015). Based on the widespread utility of Torpedospora and its
priority, the use of Torpedospora over Glomerulispora was recommended by Réblová et al.
(2016b). The family groups in the Hypocreomycetidae, order incertae sedis (Jones et al. 2014,
Maharachchikumbura et al. 2015b). Subsequently, Jones et al. (2015) had referred the family to a
new order Torpedosporales and this was followed by Maharachchikumbura et al. (2016b) and
Wijayawardene et al. (2018a).
Ecological and economic significance of Torpedosporaceae
Torpedosporaceae species are saprobic on intertidal wood or mangrove wood and roots, bark
and leaves, in marine habitats. Weber et al. (2015) reported that T. radiata has an abundance of
recognizable secondary metabolite gene clusters and are under investigation (Pang et al. 2016).
Genus included in Torpedosporaceae
Torpedospora Meyers, Mycologia 49: 496 (1957)
Index Fungorum number: IF5501; 3 morphological species (Species Fungorum 2020), 2
species with sequence data.
Type species – Torpedospora radiata Meyers
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Figure 247 – Phaeoacremonium inconspicuum (Material examined – PHILIPPINES, Luzon,
Laguna Province, Los Baños, on Gigantochloa schrebneriana, 10 September 1913, M.B.
Raimundo, S-F6209, holotype). a, b Envelope with original information. c-e Aggregated perithecia
on bamboo column. f, h Perithecial necks (arrows). g Vertical section of ascomata. i Peridium. j-l
Ascogenous hyphae with asci attached. m Asci containing ascospores. n-q Ascospores. r
Paraphyses. Scale Bars: d, e = 200 µm, f = 1 mm, g = 100 µm, h = 150 µm, i, k = 25 µm, j = 50
µm, l, m, r = 15 µm, n-q = 5 µm.
906
Notes – Torpedospora is characterised by torpedo-like, cylindrical to elongate-ellipsoidal
ascospores, together with the radiating appendages at one or both ends. This genus comprises three
species, T. ambispinosa, T. mangrovei, and T. radiata (type). Molecular studies by Sakayaroj et al.
(2005), Schoch et al. (2007), Abdel Wahab et al. (2010) and Jones et al. (2015) showed that they
group in the same clade with high bootstrap support, which suggest that they are congeneric. A
study by Abdel-Wahab et al. (2018) also supported this.
Figure 248 – Phaeoacremonium krajdenii (CBS 109479) (reproduced from Mostert et al. 2006) (a,
f, h), P. aleophilum (CBS 246.91) (b), P. austroafricanum (CBS 112949) (c), P. parasiticum (CBS
860.73) (d, e) P. inflatipes (CBS 391.71) (g). a, b Conidiophores. c Conidia. d Mycelium with
mucus and phialides. e-h Phialides. Scale bars: a-h = 10 µm.
Tracyllaceae Crous, Persoonia 40: 365 (2018)
Index Fungorum number: IF825423; Facesoffungi number: FoF06884; 3 species.
Endophytic, saprobic or pathogenic on leaves of grasses and trees. Sexual morph:
Undetermined. Asexual morph: Coelomycetous. Conidiomata pycnothyrial, superficial on leaves,
round, brown, with central column of cells; ostiole lacking, margin of catenate, darker brown cells.
Conidiophores reduced to conidiogenous cells arising from a central columella, doliiform to
ellipsoid, hyaline, smooth, with a single conidiogenous locus, phialidic. Conidia solitary, hyaline,
aseptate, smooth, guttulate, falcate to naviculate or ellipsoid, apex subobtusely rounded, base
truncate; with or without unbranched polar appendages, not delimited by septa (adapted from Crous
et al. 2018d).
Type genus – Tracylla (Sacc.) Tassi
Notes – Tracyllaceae was introduced by Crous et al. (2018d) to accommodate the monotypic
genus Tracylla. This was well-supported by the multigene analysis (ITS, LSU, SSU) of T. aristata
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and T. eucalypti that form a monophyletic lineage which is distinct from other groups in
Sordariomycetes.
Figure 249 – Torpedospora radiata (Material examined – Taiwan, Keelung City, Chaojing Park,
rocky shore, on driftwood, 3 October 2014, K.L. Pang). a Section of immersed ascoma. b Clavate
ascus. c Cylindrical ascospore with unipolar appendages and three septa. d Scanning electron
micrograph of the ascospore. Scale bars: a = 20 µm, b, c = 5 µm.
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Ecological and economic significance of Tracyllaceae
Both Trachylla spartinae and T. aristata were reported as leaf litter saprobes, while T.
eucalypti was observed from symptomatic leaves of Eucalyptus urophylla.
Figure 250 – Torpedospora mangrovei. (Material examined – JAPAN, Okinawa, Naha, Gesashi
Bay Mangrove, 26°12′N, 127°37 E, on decayed driftwood in the intertidal zone, 17 July 2008,
M.A. Abdel-Wahab, IMI 397963, holotype; ex-type strain, MF1019, NBRC 105264). a-i Conidia at
different stages of development. Scale bars: a-i = 5 µm.
Genus included in Tracyllaceae
Tracylla (Sacc.) Tassi, Bollettino del Laboratorio de Orto Botanico Reale Universita Siena 6: 62
(1904)
Index Fungorum number: IF10260; 3 morphological species (Species Fungorum 2020), 2
species with sequence data.
Type species – Tracylla spartinae (Peck) Tassi
Notes – Tracylla was introduced as a subgenus by Saccardo (1895), and raised to generic
rank by Tassi (1904). The genus is typified by Tracylla spartinae observed on Spartina patens and
several other grasses in Brazil, Jamaica, Philippines, Sri Lanka, Thailand and USA (Nag Raj 1993).
Two other species include T. aristata and T. eucalypti. Tracylla aristata was described from
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Eucalyptus leaf litter collected in Australia (Cooke 1891), New Zealand and South Africa (Crous &
Van der Linde 1993), and on Hakea dactyloides (Nag Raj 1993). Tracylla eucalypti was observed
from leaves of Eucalyptus urophylla in Colombia (Crous et al. 2018d). The three species differ in
their conidial morphology with the presence (T. spartinae:14–18 × 6–8 µm, with apical and basal
appendages 11–30 µm; T. aristata: 13-17 × 2-3 µm, apical appendage 6.5-14 µm) or absence of
appendages [T. eucalypti 1–1.5 µm diam, (12–)17–19(–20) × (2.5–)3 µm].
Triadelphiaceae Y.Z. Lu, J.K. Liu, Z.L. Luo & K.D. Hyde, Fungal Divers. 99: 555 (2019)
Index Fungorum number: IF555668; Facesoffungi number: FoF05449; 22 species.
Saprobic on decaying wood, or pathogenic on human, or found from the gut of red palm
weevils. Sexual morph Undetermined. Asexual morph Hyphomycetous. Conidiophores lacking.
Conidiogenous cells holoblastic, monoblastic, integrated, flask-shaped, fusiform, cylindrical or
clavate, arising from undifferentiated hyphae, hyaline to pale brown, smooth-walled. Conidia
acrogenous, solitary, develop one to five forms in distinct species (adapted from Luo et al. 2019).
Type genus – Triadelphia Shearer & J.L. Crane
Notes – Triadelphia is placed in Microascales as genus incertae sedis (Wijayawardene et al.
2017a, 2018a, Lu et al. 2018). Luo et al. (2019) showed that the Triadelphia phylogenetically
shares a sister relationship to the Graphiaceae clade with good bootstrap support (98% ML), but
Triadelphia taxa are obviously distinct from the species of Graphiaceae in conidial and
conidiophore characters. They introduced a new family, Triadelphiaceae, to accommodate
Triadelphia based on morphology and phylogeny.
Figure 251 – Tracylla spartinae (redrawn from Nag Raj 1993). a Cross section of conidioma. b
Conidiogenous cells and conidia. c Conidia. Scale bars: a = 50 µm, b, c = 20 µm.
Ecological and economic significance of Triadelphiaceae
Saprobic taxa have the ability to decompose lignocellulosic matter in woody litter, resulting
in softening of the wood and releasing nutrients in the form of simple molecules (Yuen et al. 1998,
Bucher et al. 2004). Thus, they play an important role in nutrient and carbon cycling, biological
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diversity and ecosystem functioning (Palmer et al. 1997, Wong et al. 1998a). Most Triadelphia
species are found from woody substrates, however, two Triadelphia species were found on humans.
One is T. pulvinata, which has been reported from infections in humans in Saudi Arabia (AlHedaithy 2001), including a fatal disseminated infection in a woman with leukaemia (Edathodu et
al. 2013). The second is T. disseminata, which was reported from a disseminated infection of an
immunocompromised patient in Saudi Arabia (Crous et al. 2015d).
Genera included in Triadelphiaceae
Synnematotriadelphia Chuaseehar., Somrith., Nuankaew & Boonyuen, Mycological Progress
19:137 (2020)
MycoBank number: MB833278; 2 species with sequence data.
Type species – Synnematotriadelphia stilboidea (Mercado & R.F. Castañeda) Chuaseehar.,
Somrith., Nuankaew & Boonyuen
Notes – Triadelphia was erected by Shearer & Crane (1971) to accommodate taxa with
sporodochia and characterized by Triadelphia heterospora with short, subhyaline to dark coloured
conidiogenous cells with dark, septate conidia with two different pleomorphic forms.
Constantinescu & Samson (1982) re-examined herbarium specimens and emended the generic
concept to agglomerated sporodochium-like structures, conidiogenous cells arising from hyphae,
and blastic conidia with at least two pleomorph forms produced on synthetic media. Mercado &
Castañeda (1983) and Matsushima (1995) added two synnematous species to Triadelphia, T.
stilboidea and T. synnematofera. However, they did not emend the generic descriptions of
Triadelphia to include those with synnematous conidiophores. Therefore, Chuaseeharonnachai et
al. (2020) introduced Synnematotriadelphia for species with synnematous conidiomata that produce
pleomorphic spores on natural substrata or on culture media. This is supported by a phylogenetic
study with the new genus forming a well-supported separate clade to Triadelphia.
Triadelphia Shearer & J.L. Crane, Mycologia 63(2): 247 (1971)
Index Fungorum number: IF10270; 20 morphological species (Species Fungorum 2020), 10
species with sequence data.
Type species – Triadelphia heterospora Shearer & J.L. Crane
Notes – The type species character is sporodochial hyphomycete, which was isolated from
wood blocks submerged in the Patuxent River, USA and is characterized by two different forms of
conidia. Li & Ye (2017) accepted 18 species in Triadelphia and provided a key to species. Lu et al.
(2018) introduced a new species of Triadelphia and accepted 21 species in this genus. Triadelphia
fusiformis is illustrated in this entry.
Trichosphaeriaceae G. Winter, Rabenh. Krypt.-Fl., Edn 2 (Leipzig) 1.2: 191 (1885)
Index Fungorum number: IF81492; Facesoffungi number: FoF01809; 125 species.
Saprobic and pathogenic on plants, commonly isolated from herbivore dung. Sexual morph:
Ascomata superficial, semi-immersed or immersed, ostiolate, globose to pyriform, dark brown to
black, sometimes setose. Ostiole situated in a small papilla, with bristles. Setae absent or present;
brown, septate, smooth when present. Peridium carbonaceous, coriaceous or membranaceous,
brown to dark brown, comprising cells of textura angularis. Paraphyses simple or branched,
septate, sometimes guttulate. Asci 4–8-spored, unitunicate, cylindrical to clavate, pedicellate, with
rounded apex, sometimes curved, rounded above, most genera have a J-, apical ring. Ascospores
uniseriate to biseriate, hyaline or brown, ellipsoidal to fusiform, aseptate or 1–3-sepate, sometimes
guttulate. Asexual morph: Coelomycetous (Koorchaloma) or hyphomycetous (Brachysporium).
When coelomycetous: Conidiomata sporodochioid to acervular, superficial, scattered to gregarious,
gelatinous or not, bright coloured, setose. Conidiophores hyaline, branched, septate, often reduced
to conidiogenous cells. Conidiogenous cells phialidic, ampulliform to lageniform or clavate,
hyaline. Conidia blastic-phialidic, fusiform to naviculate, aseptate, hyaline, bearing a mucoid,
funnel-shaped appendage at only apex or both ends. When hyphomycetous: Colonies effuse, brown
911
to dark brown, hairy. Mycelium mostly immersed. Conidiophores macronematous, mononematous,
erect, straight or flexuous, cylindrical, unbranched, often swollen at the base, brown to dark brown.
Conidiogenous cells polyblastic, terminal, proliferating sympodially, cylindrical, denticulate.
Conidia usually pendulous, clavate, ellipsoidal, fusiform, limoniform, obovoid or pyriform, septate,
brown, often with polar cells paler than middle cells (adapted from Maharachchikumbura et al.
2016b).
Figure 252 – Triadelphia fusiformis (Material examined – THAILAND, Krabi, Plai Praya, Khao
To, Ban Bang Thao Mae, on decaying wood in a freshwater stream, 17 December 2015,
Saranyaphat Boonmee BTM05–2, MFLU 18–1436, holotype; HKAS 102206). a Colony on
decaying wood. b-m Conidia. Scale bars: b-m = 20 μm.
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Type genus – Trichosphaeria Fuckel
Notes – Winter (1887) introduced Trichosphaeriaceae with Trichosphaeria as the type genus
and seven other astromatic genera. These seven genera were excluded from Trichosphaeriaceae by
molecular evidence. In multi-gene phylogenetic analyses of LSU, SSU, tef1 and rpb2 sequence data
by Maharachchikumbura et al. (2015), Trichosphaeriaceae had affinities with Papulosaceae and
Thyridiaceae, but they maintained Trichosphaeriaceae as a separate family until further sequence
data become available. However, due to lacking molecular recognition of T. pilosa, the use of
Trichosphaeriales in phylogenetic studies was not recommended by Réblová & Gams (2016).
Hongsanan et al. (2017) recognized Trichosphaeriaceae as family incertae sedis in
Diaporthomycetidae based on phylogenetic and molecular clock evidence, and this treatment was
followed by Wijayawardene et al. (2018a). Certain species in this family are coprophilic, while
other members are saprobic or pathogenic on plants, including Chrysopogon zizanioides, Arenga
engleri and Ulmus minor (Hudson 1963, Yanna et al. 1998, Calatayud & Aguirre-Hudson 2001).
Barr (1990b) accepted four genera in Trichosphaeriaceae, i.e. Acanthostigma, Eriosphaeria,
Rhamphoria, and Trichosphaeria. Acanthostigma was transferred to Tubeufiaceae (Réblová & Barr
2000, Boonmee et al. 2011, 2014), while Rhamphoria was placed in Annulatascaceae
(Maharachchikumbura et al. 2016b) and more recently Rhamphoriaceae. Collematospora was
introduced by Jeng & Cain (1976) who assigned it to Trichosphaeriaceae based on the similar
morphology with previously described genera, Eriosphaeria and Trichosphaeria in
Trichosphaeriaceae. Réblová (1999b) introduced Coniobrevicolla and placed it in
Trichosphaeriaceae based on the characters of peridium, ascal and hamathecium anatomy. Réblová
& Seifert (2004b) found some sexual morphs which produced Brachysporium asexual morphs in
culture. On the basis of morphology of perithecia, asci, ascospores and conidiogenesis,
Brachysporium was placed in Trichosphaeriaceae. Pinnoi et al. (2003) described Unisetosphaeria
in Trichosphaeriaceae rather than Chaetosphaeriaceae based on the morphology. Réblová & Gams
(2016) studied the type material of Acanthosphaeria and relegated this genus to a synonymy of
Chaetosphaeria. Voglmayr et al. (2019a) transferred all Cresporhaphis species including the type
to Leptosillia (Leptosilliaceae) and Rhaphidicyrtis (Pyrenulales) except C. rhoina, and they did not
give a clear classification for C. rhoina. Réblová et al. (2016b) recommended using the name
Stromatographium rather than Fluviostroma because of its greater use and priority, and accepted
Stromatographium in Sordariales. Trichosphaeriaceae is in need of further phylogenetic studies,
including studies of types, and integration of DNA sequence data.
Ecological and economic significance of Trichosphaeriaceae
Trichosphaeriaceae species often occur as saprobes in both terrestrial and aquatic habitats
(Pinnoi et al. 2003, Maharachchikumbura et al. 2016b) and are important in nutrient cycling. They
can be pathogens of fungi and living plants and lichens (Aguirre-Hudson 1991,
Maharachchikumbura et al. 2016b). Some species are coprophilous (Jeng & Cain 1976).
Genera included in Trichosphaeriaceae
Brachysporium Sacc., Syll. fung. (Abellini) 4: 423 (1886)
Index Fungorum number: IF7444; 25 morphological species (Species Fungorum 2020), 3
species with sequence data.
Type species – Brachysporium obovatum (Berk.) Sacc.
Notes – Brachysporium was introduced by Saccardo (1886). Brachysporium comprise many
saprobic wood-inhabiting species (Ellis 1971, Holubová-Jechová 1972, Réblová & Seifert 2004b).
Réblová & Seifert (2004b) established the sexual morph genus Cryptadelphia and linked it with
Brachysporium asexual morphs. Cryptadelphia was synonymized under Brachysporium due to
nomenclatural priority. The sexual morph of this genus is characterised by superficial, globose to
subglobose ascomata with ostioles, 8-spored, unitunicate, cylindrical to clavate asci with long
pedicels and a distinct, refractive apical ring, and hyaline, fusiform to ellipsoidal to oblong-lemonshaped ascospores with a central septum (Réblová & Seifert 2004b). The asexual morph has
913
macronematous, mononematous, unbranched conidiophores, polyblastic, terminal, denticulate
conidiogenous cells, and clavate, ellipsoidal, fusiform, limoniform, obovoid or pyriform, septate,
brown conidia, which often have one or more cells paler than the others (Ellis 1971, Réblová &
Seifert 2004b, Markovskaja & Treigienė 2007).
Collematospora Jeng & Cain, Can. J. Bot. 54(21): 2429 (1976)
Index Fungorum number: IF1177; 1 morphological species.
Type species – Collematospora venezuelensis Jeng & Cain
Notes – Jeng & Cain (1976) introduced the monotypic genus Collematospora to
accommodate C. venezuelensis from horse dung in Venezuela. The genus is characterized by
subglobose to pyriform, ostiolate, hairy ascomata, 8-spored, unitunicate, cylindrical asci with a
distinct, J-, apical ring, and hyaline, globose, aseptate ascospores containing one to several oil
drops (Jeng & Cain 1976). The asexual morph is undetermined.
Coniobrevicolla Réblová, Mycotaxon 70: 422 (1999)
Index Fungorum number: IF28272; 1 morphological species.
Type species – Coniobrevicolla larsenii Réblová
Notes – Coniobrevicolla is a monotypic genus introduced by Réblová (1999b) to
accommodate C. larsenii, which is saprobic on decayed wood of an unidentified deciduous tree in
Denmark. This genus is characterized by superficial, conical, ostiolate ascomata, 8-spored asci with
an indistinct apical ring, and ellipsoidal, 3-septate, ascospores with two brown central cells and
hyaline end cells (Réblová 1999b). The asexual morph is undetermined.
Eriosphaeria Sacc., Atti Soc. Veneto-Trent. Sci. Nat., Padova, Sér. 4 4: 86 (1875)
Index Fungorum number: IF1892; 23 morphological species (Species Fungorum 2020).
Type species – Eriosphaeria vermicularia (Nees) Sacc.
Notes – Eriosphaeria was introduced by Saccardo (1875a). The genus is characterized by
superficial, setose ascomata and 1-septate, oval ascospores (Réblová 1997). Fresh collections are
required to solve the natural classification of Eriosphaeria.
Koorchaloma Subram., J. Indian bot. Soc. 32: 124 (1953)
Index Fungorum number: IF8680; 11 morphological species (Species Fungorum 2020), 1
species with sequence data.
= Kananascus Nag Raj, Mycotaxon 19: 201 (1984)
Type species – Koorchaloma madreeya Subram. (= Kananascus koorchalomagnatus Nag
Raj)
Notes – Nag Raj (1984) reported two sexual morph species of Koorchaloma, including the
type species, Ko. madreeya, and erected Kananascus. Based on nomenclatural priority, we
synonymize Kananascus with Koorchaloma. The sexual morph of Koorchaloma is characterized
by its superficial, globose to pyriform, unilocular and setose ascomata, hyaline, clavate, 8-spored
asci with J-, apical rings and fusiform, 1-septate, hyaline, guttulate ascospores (Nag Raj 1984,
Treigiene 2006), while the asexual morph is characterized by superficial, sporodochioid to
acervular, setose conidiomata, hyaline, branched and septate conidiophores which are often reduced
to phialidic conidiogenous cells, and fusiform conidia which mucoid appendages at both ends or
only at the apex (Nag Raj 1984, Yanna et al. 1998, Treigiene 2006).
Rizalia Syd. & P. Syd., Annls mycol. 12(6): 546 (1914)
Index Fungorum number: IF4761; 8 morphological species (Hyde et al. 2017b, Niranjan &
Sarma 2018).
Type species – Rizalia fasciculata Syd. & P. Syd.
Notes – Rizalia was introduced by Sydow & Sydow (1914). Hyde et al. (2017b) accepted five
species in Rizalia, viz. R. byrsonimae, R. confusa, R. fasciculata, and R. glaziovii and R. guianensis.
914
The sixth species was described by Niranjan & Sarma (2018) from India. Rizalia is characterized
by globose to pyriform, setose ascomata with long necks, clavate asci and hyaline, fusiform or
falcate ascospores, sometimes with apical mucilaginous frills (Hyde et al. 2017b, Niranjan & Sarma
2018). The asexual morph is undetermined.
Figure 253 – Brachysporium obovatum, redrawn from Seifert et al. (2011)
Schweinitziella Speg., Anal. Soc. cient. argent. 26(1): 45 (1888)
Index Fungorum number: IF4920; 4 morphological species (Species Fungorum 2020).
Type species – Schweinitziella styracum Speg.
Notes – Schweinitziella was introduced by Spegazzini (1888). Schweinitziella is a poorly
known genus since no molecular data are available. The sexual morph of Schweinitziella has
clavate asci (Maharachchikumbura et al. 2016b). The asexual morph of this genus is undetermined.
Setocampanula Sivan. & W.H. Hsieh, Mycological Research 93: 87 (1989)
Index Fungorum number: IF25337; 1 morphological species.
Type species – Setocampanula taiwanensis Sivan. & W.H. Hsieh
Notes – Setocampanula is a monotypic genus introduced by Sivanesan & Hsieh (1989) to
accommodate S. taiwanensis. Sivanesan & Hsieh (1989) assigned Setocampanula in
Trichosphaeriaceae, because Setocampanula resembles other genera of Trichosphaeriaceae in its
superficial setose ascomata and the unitunicate asci without any apical ring. Setocampanula
taiwanensis is a bambusicolous fungus collected from Taiwan.
Trichosphaeria Fuckel, Jb. nassau. Ver. Naturk. 23-24: 144 (1870)
Index Fungorum number: IF5588; 50 morphological species (Species Fungorum 2020), 1
species with sequence data.
915
Figure 254 – Setocampanula taiwanensis (Material examined – Taiwan, Kaohsiung, on Yushania
niitakayensis, 29 January 1988, A. Sivanesan, IMI 322672, holotype). a Herbarium label and
specimen. b, c Ascomata on substrate. d Vertical section of ascoma. e Peridium. f Paraphyses. g-j
Asci. k Ascospores. Scale bars: b = 1000 µm, c = 500 µm, d = 100 µm, e = 40 µm, f-k = 20 µm.
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Type species – Trichosphaeria pilosa (Pers.) Fuckel
Notes – Trichosphaeria was established by Fuckel (1870) with the type species T. pilosa
collected from rotten wood from Germany. Trichosphaeria species are characterized by globose to
subglobose ascomata with setae, unitunicate, 4–8-spored asci with a J-, apical ring, and hyaline,
aseptate, ellipsoidal ascospores. Since most Trichosphaeria species were introduced before 1940,
molecular data are lacking in this genus. The genus needs recollection and study.
Unisetosphaeria Pinnoi, E.B.G. Jones, McKenzie & K.D. Hyde, Mycoscience 44(5): 377 (2003)
Index Fungorum number: IF28746; 1 species without molecular data (Pinnoi et al. 2003).
Type species – Unisetosphaeria penguinoides Pinnoi, E.B.G. Jones, McKenzie & K.D. Hyde
Notes – Pinnoi et al. (2003) introduced the monotypic genus Unisetosphaeria on submerged
petiole of Eleiodoxa conferta from Sirindhorn Peat Swamp Forest in Thailand. Unisetosphaeria is
characterized by pyriform, ostiolate, papillate ascomata, 8-spored, clavate, unitunicate, short
pedicellate asci with a refractive, J-, apical ring, and 2-seriate, hyaline, septate ascospores (Pinnoi et
al. 2003). The asexual morph of this genus is undetermined.
Figure 255 – Unisetosphaeria penguinoides (Material examined – THAILAND, Narathiwat,
Sirindhorn Peat Swamp Forest, on submerged petiole of Eleiodoxa conferta, June 22, 2001, A.
Pinnoi, BBH9843, holotype). a Herbarium packet. b-d Ascomata on substratum. e Section of
ascoma. f Asci. g-i Ascospores (redrawn from Pinnoi et al. 2003). Scale bars: b = 2000 μm, c = 500
μm, d = 200 μm, e = 100 μm, f-i = 20 μm.
Tubakiaceae U. Braun, J.Z. Groenew. & Crous, Fungal Systematics and Evolution 1:62 (2018)
Index Fungorum number: IF823660; Facesoffungi number: FoF06366; 29 species.
Saprobic or endophytic and pathogenic in leaves and twigs causing leaf spots and twig
dieback. Sexual morph: apiognomonia- and dicarpellum-like, diaporthaloid, dark, rostrate, ostiolate
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perithecial ascomata, with dark stromatic layers, polyascal. Asci 8-spored, unitunicate. Ascospores
hyaline, aseptate or with a single septum near the apex. Asexual morph:. Coelomycetous.
Conidiomata pycnothyria, Conidiogenous cells monophialidic, hyaline, having collarettes. Conidia
globose to broad ellipsoid-obovoid, aseptate, hyaline to pigmented, often with basal frill or truncate
peg-like hilum (adapted from Braun et al. 2018).
Type genus – Tubakia B. Sutton
Notes – Species of Tubakiaceae are characterized by conidiomata comprising a convex
scutella with radiating threads of cells connected to the substratum by a central columnella, mostly
surrounded by a sheath of small fertile cells which develop to 1-celled, phialidic conidiogenous
cells (Braun et al. 2018). Most Tubakiaceae species have been recorded as endophytes, nevertheless
some act as mild pathogens forming leaf spots or saprobes on shed or still attached leaves. Based
on taxonomic investigations, Tubakiaceae comprises eight genera, viz. Tubakia, Apiognomonioides,
Involutiscutellula, Oblongisporothyrium, Paratubakia, Racheliella, Saprothyrium and
Sphaerosporithyrium (Braun et al. 2018, Senanayake et al. 2018).
Ecological and economic significance of Tubakiaceae
Tubakiaceae is an important family as the members are also considered as plant pathogens.
The initial stages of leaf spot spread through the host plant until resulting in die back (Braun et al.
2018). Tubakia leaf spot is one of the major diseases diagnosed on oak species. Tubakia dryina is
recorded from chestnuts with its typical symptom of brown necrotic leaf spots (El-Gholl et al.
1996).
Genera included in Tubakiaceae
Apiognomonioides U. Braun, J.Z. Groenew. & Crous, Fungal Systematics and Evolution 1:63
(2018)
Index Fungorum number: IF824479; 1 species with sequence data.
Type species – Apiognomonioides supraseptata (S. Kaneko & Tak. Kobay) U. Braun, J.Z.
Groenew. & Crous
Notes – Apiognomonioides was introduced based on the sexual morph. Kaneko & Kobayashi
(1984) showed Apiognomonia as having two celled ascospores in which the upper cell was larger
than the basal cell, however, A. supraseptata is distinct in having a smaller upper cell.
Subsequently, Harrington & McNew (2018) suggested that A. supraseptata represents the only
clearly demonstrated sexual morph of Tubakia. Owing to its isolated position in the LSU tree in
Tubakiaceae, but distant from all other genera in this family, Apiognomonia supraseptata shows its
phylogenetic distince from all other genera in the LSU tree of family Tubakiaceae (Braun et al.
2018). Apiognomonioides supraseptata and the sexual morph of Tubakia suttoniana have similar
morphology (rostrate perithecia, unitunicate 8-spored asci, colourless conidia), but A. supraseptata
differs in forming uniseptate ascospores with a septum near the apex (Braun et al. 2018).
Involutiscutellula U. Braun & C. Nakash., Fungal Systematics and Evolution 1:64 (2018)
Index Fungorum number: IF824481; 1 species with sequence data.
Type species – Involutiscutellula rubra (T. Yokoy. & Tubaki) U. Braun & C. Nakash.
Notes – Involutiscutellula rubra can be distinguished from all other Tubakia species in
having reddish brown colonies and hyphae with pycnothyria which is more or less undulate with
involute margin, having, small, cylindrical to oblong bacilliform conidia (Braun et al. 2018).
Oblongisporothyrium U. Braun & C. Nakash., Fungal Systematics and Evolution 1:66 (2018)
Index Fungorum number: IF824483; 1 species with sequence data.
Type species – Oblongisporothyrium castanopsidis (T. Yokoy. & Tubaki) U. Braun & C.
Nakash.
Notes – Oblongisporothyrium was introduced by Braun et al. (2018) to accommodate O.
castanopsidis (≡ Actinopelte castanopsidis) as the type species. Oblongisporothyrium castanopsidis
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is similar to Paratubakia subglobosa in having scutellae with inwardly curved margins, hyaline to
pigmented conidiogenous cells, and hyaline conidia, however it differs in having oblong conidia
(vs. globose to subglobose in T. subglobosa) (Braun et al. 2018).
Paratubakia U. Braun, & C. Nakash, Fungal Systematics and Evolution 1:67 (2018)
Index Fungorum number: IF824485; 1 species with sequence data.
Type species – Paratubakia subglobosa (T. Yokoy. & Tubaki) U. Braun, & C. Nakash
Notes – Paratubakia subglobosoides is phylogenetically related to P. subglobosa as
pycnothyria formation is similar. However, they differ in the conidial shape and size and the
formation of distinct leaf spots. Furthermore, the culture characteristics of the two species on MEA
are quite different, and they are genetically distinct (Braun et al. 2018).
Racheliella Crous & U. Braun, Fungal Systematics and Evolution 1:69 (2018)
Index Fungorum number: IF824487; 2 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Racheliella wingfieldiana Crous & U. Braun
Notes – The conidiomata of convex scutella with radiating threads of cells connected to the
substratum of type specimen are similar to other genera of Tubakiaceae. However, the genera
having hyaline conidia arranged in a line are excluded from Tubakia sensu stricto. Racheliella
saprophytica is characterized by crustose conidiomata and it is different from all other genera of
Tubakiaceae in having branched, septate conidiophores with phialidic conidiogenous cells provided
with flared, conspicuous collarettes with serrate margins (Braun et al. 2018).
Saprothyrium U. Braun, Crous & J.Z. Groenew., Fungal Systematics and Evolution 1:72 (2018)
Index Fungorum number: IF824491; 1 species with sequence data.
Type species – Saprothyrium thailandense (Senan., Tangthir. & K.D. Hyde) U. Braun, Crous
& J.Z. Groenew
Notes – Pycnothyria of Saprothyrium differ in having a scutellum with obtuse outer strands
with globose to subglobose hyaline conidia, and is similar with Sphaerosporithyrium. However, it
clusters in a phylogenetically distant clade to clades of Tubakia sensu stricto (Braun et al. 2018).
Saprothyrium can be distinguished from all other species of Tubakia sensu stricto by hyaline,
globose to subglobose conidia formed in pycnothyria (Braun et al. 2018). Saprothyrium
thailandense is illustrated in this entry.
Sphaerosporithyrium U. Braun, Crous, O. Moreno-Rico & Marm., Fungal Systematics and
Evolution 1:73 (2018)
Index Fungorum number: IF824492; 1 species with sequence data.
Type species – Sphaerosporithyrium mexicanum O. Moreno-Rico, U. Braun & Marm.
Notes – Sphaerosporithyrium is similar with Paratubakia (Braun et al. 2018) in conidial
shape and colour. However, the formation of pycnothyria formed by species of the latter genus
differs in pointed tips of hyphal scutellum strands. Sphaerosporithyrium is characterized by hyaline
globose-subglobose conidia (Braun et al. 2018).
Tubakia B. Sutton, Trans. Br. Mycol. Soc. 60(1): 164 (1973)
Index Fungorum number: IF10330; 21 morphological species (Species Fungorum 2020), 15
species with sequence data.
Type species – Tubakia japonica (Sacc.) B. Sutton
Notes – Tubakia species may form different types of asexual morphs. Pycnothyria with
typical scutella are a distinct character of this genus. In addition, sporodochial conidiomata are
formed with clusters of conidiogenous cells e.g. on leaf veins as in T. iowensis. Crustose or
pustulate pycnidioid conidiomata are formed in several Tubakia species, including T. californica, T.
dryina, and T. iowensis. Holdenrieder & Kowalski (1989) introduced these conidiomata as
919
“pycnidial” even though they lacked ostioles with conidiomata dehiscing by irregular rupture.
Harrington et al. (2012) described them as “conidioma” and wrote “pycnothyrium” in brackets.
These conidiomata cannot be described as pycnidia as they lack ostioles. Rather, with stromatic
wall layers and a dehiscence by irregular fissures, these should rather be classified as stromatic
(Braun et al. 2018).
Figure 256 – Saprothyrium thailandense (Material examined – Thailand, Chiang Rai, Doi Mae
Salong, on dead leaf, 2 May 2012, K. Wisitrassameewong, NTCL059, MFLU 13–0260, holotype).
a Herbarium specimen. b Conidiomata on the host surface. c Vertical section of pycnothyrium. d
Top view of radiate scutellum and conidiogenous cells with developing conidia. e-g Conidia. Scale
bars: c = 50 μm, d-g = 10 μm.
Vermiculariopsiellaceae Hern.-Rest., J. Mena, Gené & Crous, Studies in Mycology 86: 91 (2017)
Index Fungorum number: IF820347; Facesoffungi number: FoF05408; 22 species.
Parasitic or saprobic on wood or leaves. Sexual morph: Undetermined. Asexual morph:
Stroma present or absent. Conidiomata sporodochial, scattered, setose. Setae branched or
unbranched, brown, septate, straight to flexuous. Conidiophores macronematous, subcylindrical,
hyaline to pale brown, septate, densely packed in a palisade. Conidiogenous cells monophialidic,
terminal, hyaline to pale brown, cylindrical to lageniform, with collarette. Conidia cylindrical to
oblong, straight or slight curved, hyaline, aseptate or septate, sometimes guttulate (adapted from
Hernández-Restrepo et al. 2017).
Type genus – Vermiculariopsiella Bender
Notes – Vermiculariopsiellaceae was established by Hernández-Restrepo et al. (2017) to
accommodate a well-supported monophyletic clade containing Vermiculariopsiella. The sexual
morph of Vermiculariopsiella was linked with Echinosphaeria based on culture studies (Puja et al.
2006, Dhargalkar & Bhat 2009). However, phylogenetic studies, based on LSU and tub2 sequence
data, showed that E. canescens, the type species of Echinosphaeria, resides in
920
Helminthosphaeriaceae (Miller et al. 2014). Therefore, it is possible that the asexual morphs of
Echinosphaeria were wrongly identified as Vermiculariopsiella. The linkage between
Echinosphaeria and Vermiculariopsiella requires further molecular studies to confirm. We regard
the sexual morph of Vermiculariopsiella as undetermined.
Ecological and economic significance of Vermiculariopsiellaceae
Vermiculariopsiella species are mostly saprobic on leaves. As decomposers and recyclers,
species in this order are involved in nutrient cycling and supply members of other kingdoms with
nutrients. Thus, they are important for ecological balance.
Genus included in Vermiculariopsiellaceae
Vermiculariopsiella Bender, Mycologia 24(4): 412 (1932)
Index Fungorum number: IF10383; 22 morphological species (Species Fungorum 2020), 9
species with sequence data.
Type species – Vermiculariopsiella immersa (Desm.) Bender
Notes – Torrend (1912) and Höhnel (1918a) separately introduced Vermiculariopsis. Bender
(1932) synonymized Vermiculariopsis (1918) and established Vermiculariopsiella.
Vermiculariopsiella endophytica and V. pteridis were linked with the Echinosphaeria sexual morph
in culture (Puja et al. 2006, Dhargalkar & Bhat 2009). Crous et al. (2014a) introduced V.
dichapetali and placed it in Microascales incertae sedis (Hypocreomycetidae).
Maharachchikumbura et al. (2015, 2016b) followed this treatment. Crous et al. (2016b) introduced
another V. acacia and V. eucalypti and assigned the genus to Chaetosphaeriaceae based on LSU
sequence data. Hernández-Restrepo et al. (2017) transferred Vermiculariopsiella to
Vermiculariopsiellaceae based on SSU, LSU and ITS sequence data.
Figure 257 – Vermiculariopsiella immersa (redrawn from Seifert et al. 2011). a Colonies on leaves.
b Sporodochia with setae. c Conidiophores, conidiogenous cells and conidia. Scale bars: a = 100
μm, b, c = 10 μm, based on Nawawi et al. (1990).
921
Vialaeaceae P.F. Cannon, Mycol. Res. 99(3): 368 (1995)
Index Fungorum number: IF81967; Facesoffungi number: FoF00686; 4 species.
Parasitic or saprobic on dead plant matter. Sexual morph: Pseudostroma appears as
ellipsoidal, black dots slightly raised from substrate, thick around the upper part of the ascomata.
Ascomata perithecial, solitary or aggregates, immersed, globose, subglobose to ellipsoidal,
coriaceous, black to brown, papillate, ostiolate with periphyses. Papilla long, sometimes distal end
curving towards the substrate, periphysate. Peridium thick, composed of outer, dark-brown, thickwalled cells of textura angularis and inner, hyaline, thick-walled cells of textura angularis.
Paraphyses filiform, rarely branched, septate, hyaline, sometimes apex slightly swollen. Asci 8spored, unitunicate, cylindrical, sometimes tapering towards the apex or base, short-stalked or
sessile, thin-walled except apex, apex obtuse or truncate, apical ring subconical or subapical, J+.
Ascospores biseriate, triseriate to fasciculate, sometimes weakly helically coiled, hyaline, strongly
isthmoid, ends fusiform to rhombic, apical portion sometimes slightly larger than the basal part, 1–
3-septate, smooth-walled. Asexual morph: Ceolomycetous. Conidiomata pycnidia, superficial,
solitary, scattered, globose, with slimy, shining spore mass and basal mycelium forming thick,
black strands. Conidiophores erect, branched, septate, hyaline. Conidiogenous cells phialidic,
discrete or in small whorls, lageniform to cylindrical, hyaline. Conidia oblong to ellipsoidal, 1celled, hyaline smooth, with truncate abscission scar (adapted from Senanayake et al. 2014).
Type genus – Vialaea Sacc.
Notes – The phylogenetic placement of Vialaeaceae was confirmed in Xylariales (Shoemaker
et al. 2013, Senanayake et al. 2014, Maharachchikumbura et al. 2015b). However, this family was
accommodated in Amphisphaeriales based on phylogeny (Hyde et al. 2017a) and this is confirmed
in this study.
Ecological and economic significance of Vialaeaceae
Members of Vialaeaceae are phytopathogens and endophytes. Vialaea insculpta was reported
from an ornamental plant, Ilex aquifolium as a mild pathogen (Shoemaker et al. 2013). Vialaea
insculpta causes dieback of tips of new twigs and older branches, as a result of cankers. However, it
is not yet clear whether Vialaea insculpta are plant pathogens or colonize infected or necrotic
branches (Redlin 1989). Vialaea minutella and V. mangiferae causes branch dieback or cankers on
mango trees (McTaggart et al. 2013, Senanayake et al. 2014) and is likely an endophyte before it
becomes a mild pathogen (McTaggart et al. 2013).
Genus included in Vialaeaceae
Vialaea Sacc., Bull. Soc. mycol. Fr. 12: 66 (1896)
Index Fungorum number: IF5736; 4 morphological species (Species Fungorum 2020), 3
species with sequence data.
Type species – Vialaea insculpta (Fr.) Sacc.
Notes – Vialaea comprises Vialaea bambusae, V. insculpta, V. minutella and V. mangiferae
(Shoemaker et al. 2013, Senanayake et al. 2014, Index Fungorum 2020). Vialaea insculpta was
reported from twigs and leaf veins of Ilex aquifolium grown commercially as ornamental plants.
This species is reported only from in Europe and North America (Atkinson & Trelawny 1962,
Shaw 1973). In this entry we illustrate Vialaea insculpta.
Woswasiaceae H. Zhang, K.D. Hyde & Maharachch., Fungal Divers. 85: 104 (2017)
Index Fungorum number: IF553769; Facesoffungi number: FoF03348; 3 species.
Saprobic, hypersaprotrophic on stromata of Diaporthe in freshwater or terrestrial habitats.
Sexual morph: Ascomata perithecial, scattered, aggregated or in groups, stromatic or astromatic,
immersed to erumpent, globose, subglobose or ellipsoid, coloured, sometimes seated in a stroma or
basal stroma. Neck or papilla centrally located, cylindrical, upright or slightly horizontal to the
substrate, pale brown, dark brown to black. Ostiolum periphysate. Peridium leathery to fragile, twoor three-layered. Paraphyses abundant, persistent, septate. Asci 8-spored, unitunicate, cylindrical922
Figure 258 – Vialaea insculpta (Material examined – GERMANY, Nordrhein-Westfalen,
Deutschland, Westfalen, Bottrop, an der zerzogstrasse, on twigs of Ilex aquifolium, 2 August 1922,
H. Rupprecht, S-F121733). a Packet of the herbarium. b Herbarium specimens. c Ascostromata on
host. d Vertical section of ascoma. e Peridium. f Periphyses in ostiolar canal. g-k Asci. l Apical ring
bluing in Melzer’s reagent. m-r Ascospores. Scale bars: c = 500 µm, d, f = 100 µm, e = 20 µm, g-k
= 50 µm, l = 10 µm, m-r = 20 µm.
923
Figure 259 – Asexual morph of Vialaea mangiferae (Material examined – THAILAND, Chaing
Rai Province, Muang District, near Bandu, Baan Khuakhae, at 31M. 17, (19° 59’ 52.05” N; 99° 49’
25.15” E), on twigs of Mangifera indica, 15 Nov. 2012, leg. K. D. Hyde, CHUNI001; MFLU13–
0342, HKAS 81790, holotype). a Culture from above. b, c Conidiomata forming on Water Agar. d,
e Conidia forming on phialides attached to conidiophores. f Conidia. Scale bars: b = 1 mm, c = 500
µm, e, f =10 µm.
-or fusoid, with a distinctive, J-, apical ring. Ascospores uniseriate or overlapping uni-seriate,
hyaline, globose, subglobose, ellipsoidal or fusiform, unicellular or septate, thin- or thick-walled,
verruculose or smooth-walled, with or without a gelatinous sheath. Asexual morph:
Coelomycetous, produced in culture with or without sporodochial conidiomata. Conidiophores
aggregated, hyaline to subhyaline, with a clavate or penicillate head or conidiogenous cells
developing from hyphae. Conidiogenous cells monoblastic or polyblastic, sympodially
proliferating, terminal, cylindrical, hyaline. Conidia ellipsoid to obovoid, hyaline, aseptate, smoothwalled (adapted from Raja et al. 2003, Jaklitsch et al. 2013, Réblová et al. 2014, Zhang et al.
2017a).
Type genus – Woswasia Jaklitsch, Réblová & Voglmayr
Notes – Woswasiaceae was introduced by Zhang et al. (2017a) and placed in
Diaporthomycetidae, families incertae sedis (Jaklitsch et al. 2013, Réblová et al. 2014, Senanayake
et al. 2016, Zhang et al. 2017a). The family presently contains three genera, Cyanoannulus,
Woswasia and Xylochrysis. The family status was strongly supported in the MCC tree, which has a
stem age at ca 115 MYA (Hyde et al. 2017a, Zhang et al. 2017a).
Ecological and economic significance of Woswasiaceae
Woswasiaceae currently includes three taxa isolated from decaying wood (Zhang et al.
2017a). They may be important decomposers participating in nutrient cycling, especially on
submerged and dry wood.
Genera included in Woswasiaceae
Cyanoannulus Raja, J. Campb. & Shearer, Mycotaxon 88: 8 (2003)
Index Fungorum number: IF28772; 1 species with sequence data.
924
Type species – Cyanoannulus petersenii Raja, J. Campb. & Shearer
Notes – The monotypic genus Cyanoannulus was isolated from decorticated wood submerged
in a stream in North Carolina. The genus is characterised by pale reddish-brown ascomata which
are slightly horizontal to the substrate surface, fusoid asci and thick-walled ascospores with a
narrow channel at the apices and with a mucilaginous sheath (Raja et al. 2003).
Woswasia Jaklitsch, Réblová & Voglmayr, Mycologia 105(2): 479 (2013)
Index Fungorum number: IF800841; 1 species with sequence data.
Type species – Woswasia atropurpurea Jaklitsch, Réblová & Voglmayr
Notes – Woswasia is monotypic genus (Jaklitsch et al. 2013). The genus, together with
Cyanoannulus and Xylomelasma, was placed in Sordariomycetidae genera incertae sedis
(Maharachchikumbura et al. 2016b). Later, the three genera were transferred to a new family
Woswasiaceae based on phylogenetic analysis and divergence estimates (Zhang et al. 2017a). The
asexual morph of this genus has not yet been reported.
Xylochrysis Réblová, Mycologia 106(3): 567 (2014)
Index Fungorum number: IF805253; 1 species with sequence data.
Type species – Xylochrysis lucida Réblová, V. Štěpánek & R.K. Schumach.
Notes – The monotypic genus Xylochrysis was isolated from decaying bark and wood of
Rhododendron in Germany. The genus is characterised by ascomata often aggregated in small
groups, with astromatic or stromatic base, surrounded by a pigmented layer of cells, cylindrical asci
and hyaline, ellipsoidal ascospores (Réblová et al. 2014).
Xylochrysis lucida was introduced by Réblová et al. (2014) in a newly established genus
Xylochrysis with conidiomata produced in culture. Xylochrysis lucida was not assigned to any other
known families when first reported. Zhang et al. (2017a) established a new family Woswasiaceae to
accommodate three genera including X. lucida.
Figure 260 – Asexual morph of Xylochrysis lucida (redrawn from Réblová et al. 2014). a
Conidiophores bearing conidia. b Conidia. Scale bars: a, b = 10 μm.
925
Figure 261 – Xylochrysis lucida (Material examined – GERMANY, Near Neuendorf, on decaying
wood and bark of a twig of a cultivated shrubby Rhododendron sp., 14 January 2012, R.K.
Schumacher R.S. 044, PRM 922621, holotype). a Herbarium material with label. b Appearance of
superficial ascomata on host. c Vertical section through ascoma. d, e Vertical section through
peridium. f Structure of peridium. g Paraphyses. h-l Unitunicate asci. m-q Ascospores. Scale bars:
d, e = 50 μm, f, g, i-l = 20 μm, h, m-q = 5 μm.
926
Xenodactylariaceae Crous, Persoonia 41: 289 (2018)
Index Fungorum number: IF828248; Facesoffungi number: FoF06885; 1 species.
On leaves of unidentified vine. Sexual morph: Undetermined. Asexual morph: Mycelium
consisting of smooth, hyaline, branched, septate, hyphae. Conidiophores reduced to conidiogenous
cells on hyphae, erect to flexuous, hyaline, smooth, with one to several denticulate apical loci.
Conidia occurring in branched chains, hyaline, smooth, subcylindrical, septate (adapted from Crous
et al. 2018b).
Type genus – Xenodactylaria Crous
Notes – Crous et al. (2018b) introduced Xenodactylariaceae mainly focusing on similarity
with Dactylaria and Cylindrosympodium and being a phylogenetically stable independent clade.
Ecological and economic significance of Xenodactylariaceae
Xenodactylaria thailandica was found on leaves of unidentified vine in Thailand (Crous et al.
2018b).
Genus included in Xenodactylariaceae
Xenodactylaria Crous, Persoonia 41: 289 (2018)
Index Fungorum number: IF828193; 1 species with sequence data.
Type species – Xenodactylaria thailandica Crous
Notes – Dactylaria (de Hoog 1985) is heterogeneous genera (Crous et al. 2016a, 2017a) with
the type species, D. purpurella, clustering in Magnaporthales (Klaubauf et al. 2014) and has
similarities with Dactylaria and Cylindrosympodium. Xenodactylaria is distinct from Dactylaria
and Cylindrosympodium in having hyaline hyphae, conidiophores, conidiogenous cells and hyaline
conidia with occur in short chains. Xenodactylaria phylogenetically clusters on its own. In this
study, X. thailandica is illustrated.
Figure 262 – Xenodactylaria thailandica (redrawn from Crous et al. 2018b). a, b Conidial
attachments to the mycelia. c Conidiogenous cells. d-g conidia. Scale bars: a-g = 10 µm.
Xenospadicoidaceae Hern.-Restr., J. Mena & Gené, Studies in Mycology 86: 91 (2017)
Index Fungorum number: IF820349; Facesoffungi number: FoF05404; 74 species.
= Lentomitellaceae H. Zhang, K.D. Hyde & Maharachch, Fungal Divers. 85: 95 (2017)
Lignicolous in terrestrial and freshwater environments. Sexual morph: Ascomata perithecial,
astromatic, partially superficial, scattered or grouped. Neck cylindrical or rostrate with or without
sulcations, subhyaline to dark brown. Ostiole periphysate. Paraphyses septate, tapering. Peridium
two layered. Asci 8-spored, unitunicate, cylindrical or cylindrical-clavate, pedicellate, with a J-,
apical ring. Ascospores overlapping uniseriate, hyaline or pale brown, aseptate or septate,
ellipsoidal to fusiform, ovoid, suboblong, smooth-walled or ornamented. Asexual morph:
Hyphomycetous. Colonies effuse. Conidiophores macronematous, mononematous, branched or
unbranched, septate, brown near base, subhyaline to hyaline towards the apex. Conidiogenous cells
927
tretic or blastic, terminal or intercalary, hyaline or brown, subcylindrical, sympodially proliferating.
Conidia globose, ellipsoidal, obovoid to clavate, hyaline or brown, aseptate or septate, smoothwalled. Synasexual morph: selenosporella-like asexual morphs are sometimes formed in vitro and
in vivo. Conidiophores macronematous or semi-macronematous, mononematous, branched or
unbranched, often reduced to conidiogenous cells. Conidiogenous cells blastic, discrete, terminal or
intercalary, subcylindrical to flask-shaped, with a short terminal, rachis bearing a few denticles.
Conidia globose, clavate to obovate, or narrow fusiform, hyaline, aseptate, smooth-walled (adapted
from Hernández-Restrepo et al. 2017).
Type genus – Spadicoides S. Hughes
Notes – Xenospadicoidaceae was introduced by Hernández-Restrepo et al. (2017) as a
monotypic family in Xenospadicoidales, which was established at the same time.
Xenospadicoidaceae included two monotypic genera, the type genus Xenospadicoides and
Pseudodiplococcium (Hernández-Restrepo et al. 2017). Hernández-Restrepo et al. (2017)
synonymized six species under Xenospadicoides, including Spadicoides atra, the type species.
Réblová et al. (2018) emended Xenospadicoidaceae based on combined act1, ITS, LSU, SSU, tub2
and rpb2 sequence data. Pseudodiplococcium and Xenospadicoides was reduced to synonymy
under Spadicoides, which was assigned as the type genus (Réblová et al. 2018). Three more genera,
Calyptosphaeria, Lentomitella and Torrentispora, were added to Xenospadicoidaceae. In addition,
Lentomitellaceae was synonymized under Xenospadicoidaceae (Réblová et al. 2018).
Subsequently, the fifth genus, Neospadicoides in Xenospadicoidaceae was introduced by Luo et al.
(2019) from freshwater habitats.
Ecological and economic significance of Xenospadicoidaceae
Species of Xenospadicoidaceae are lignicolous in both terrestrial and freshwater habitats. The
ecological role of freshwater fungi has increasingly been recognized (Hyde et al. 2016a).
Decomposing dead plant material is the primary role. The transfer of nutrients and energy in the
food web occurs through decomposition by freshwater fungi (Wong et al. 1998a).
Genera included in Xenospadicoidaceae
Calyptosphaeria Réblová & A.N. Mill., Studies in Mycology 89: 13 (2017)
Index Fungorum number: IF821760; 4 species with sequence data.
Type species – Calyptosphaeria tenebrosa Réblová & A.N. Mill.
Notes – Calyptosphaeria, introduced by Réblová et al. (2018), was segregated from
Lentomitella to accommodate C. subdenudata, C. tropica, C. tenebrosa and C. collapsa based on
morphology and phylogeny. Calyptosphaeria species are saprobic and have a worldwide
distribution (Huhndorf et al. 2008, Réblová et al. 2018). The genus is characterized by semiimmersed, globose to conical ascomata with ostioles and necks, 8-spored, unitunicate, cylindrical,
short-pedicellate asci with J-, apical rings, and ellipsoidal to fusiform, brown, septate ascospores
(Réblová et al. 2018). The asexual morph is undetermined.
Lentomitella Höhn., Annales Mycologici 3 (6): 552 (1906)
Index Fungorum number: IF2736; 11 morphological species, 9 species with sequence data.
Type species – Lentomitella vestita (Sacc.) Höhn.
Notes – Lentomitella was introduced by von Höhnel (1905) who segregated this genus from
Ceratostomella Sacc. based on the ornamentation of the ascospores of C. vestita as Lentomitella
vestita. Lentomitella can be distinguished from Ceratostomella by asci, ascospores and the centrum
(Réblová 2006). Lentomitella was historically assigned to Lasiosphaeriaceae (Untereiner 1993) and
Clypeosphaeriaceae (Barr 1990b, Eriksson et al. 2003). Réblová (2006) reinstated Lentomitella
with three accepted species, L. cirrhosa, L. crinigera and L. tomentosa, and their phylogenetic
analyses showed Lentomitella resided in Sordariomycetes incertae sedis. Réblová et al. (2018)
divided Ceratostomella species and other similar taxa into four genera, Calyptosphaeria,
Lentomitella, Spadicoides, and Torrentispora, which represent Xenospadicoidaceae.
928
Lentomitella cirrhosa is saprobic on decaying wood. Since the type specimen of L. cirrhosa
is unavailable, Réblová (2006) designated a collection from New Zealand on decayed wood of
Podocarpus totara (PDD 81434) as the epitype. Sphaeria cirrhosa was accepted in Ceratostoma by
Fuckel (1870). However, Saccardo (1882) transferred S. cirrhosa to Ceratostomella as C. cirrhosa.
Réblová (2006) again transferred C. cirrhosa to Lentomitella.
Figure 263 – Lentomitella cirrhosa (Material examined – NEW ZEALAND, Westland, Hokitika,
40 km S of Greymouth, Mananui Point, Lake Mahinapua, Swimmers Beach walks, on decayed
wood of Podocarpus totara, 5 March 2003, M. Réblová, M.R. 2705, PDD 81434, epitype). a
Herbarium packet. b Herbarium material. c Ascoma on the host. d Ascoma in cross section. e
Peridium. f, g Asci. h-j Ascospores. Scale bars: d = 100 μm, e = 20 μm, f, g = 10 μm, h-j = 5 μm.
Neospadicoides Z.L. Luo, K.D. Hyde & H.Y. Su, Fungal Divers. 99: 513 (2019)
Index Fungorum number: IF555652; 3 species with sequence data.
Type species – Neospadicoides lignicola Z.L. Luo, K.D. Hyde & H.Y. Su
Notes – Neospadicoides was introduced by Luo et al. (2019) to accommodate N. aquatica, N.
lignicola and N. yunnanensis based on morphology and phylogeny. All Neospadicoides species are
929
reported from freshwater habitats in Yunnan Province, China. The genus is characterized by
macronematous, mononematous, unbranched, cylindrical, brown, septate conidiophores, polytretic,
terminal conidiogenous cells, and fusiform or obovoid, septate conidia (Luo et al. 2019). The
sexual morph of this genus is undetermined.
Figure 264 – Lentomitella unipretoriae (Material examined – SOUTH AFRICA, Kogelberg Nature
Reserve Western Cape, on dead twigs of Protea lepidocarpodendrom, 3 November 2000, S.
Marincowitz, PREM 59518, holotype). a Herbarium packet and material. b, c Ascoma on the host.
d Peridium. e, f Asci. g-j Ascospores. Scale bars: d = 50 μm, e-g = 10 μm, h-j = 5 μm.
Spadicoides S. Hughes, Can. J. Bot.36: 805 (1958)
Index Fungorum number: IF9961; 46 morphological species, 5 species with sequence data
(Ma et al. 2016a, Réblová et al. 2018, Qiao et al. 2019).
= Xenospadicoides Hern.-Restr., J. Mena & Gené, Studies in Mycology 86: 92 (2017)
= Pseudodiplococcium Hern.-Restr., J. Mena & Gené, Studies in Mycology 86: 92 (2017)
Type species – Spadicoides bina (Corda) S. Hughes
Notes – Spadicoides, based on S. bina, was introduced by Hughes (1958), who regarded
930
unbranched conidiophores as an important character to separate Spadicoides from Diplococcium
(Hughes 1958). Sinclair et al. (1985) considered conidial catenation as the main character in
identification between Diplococcium and Spadicoides. Shenoy et al. (2010) indicated that
Spadicoides is not monophyletic and is unrelated to Diplococcium based on LSU sequence data.
Réblová et al. (2018) established the sexual-asexual connection for S. bina, S. fuscolutea and S.
hyalostoma, and a selenosporella-like synasexual morph was observed for S. bina and S. fuscolutea
in vitro (Réblová et al. 2018). The asexual morph of Spadicoides is characterized by
mononematous, unbranched or sparingly branched conidiophores, polytretic conidiogenous cells
with acropleurogenus, obovoid to ellipsoid, dark brown conidia, formed singly or in a chain and
selenosporella-like synasexual morphs (Goh & Hyde 1996, Ho et al. 2002, Cai et al. 2004, Réblová
et al. 2018), while sexual morph has astromatic ascomata with long necks, 8-spored asci with a
short pedicel and J-, apical ring and hyaline, aseptate or 1-septate ascospores (Réblová et al. 2018).
Figure 265 – Spadicoides bina, redrawn from Seifert et al. (2011). This drawing shows the
conidiophores, mycelium, conidiogenous cells and conidia of S. bina. Scale bar: 10 μm.
Torrentispora K.D. Hyde, W.H. Ho, E.B.G. Jones, K.M. Tsui & S.W. Wong, Mycological
Research 104 (11): 1399 (2000)
Index Fungorum number: IF28453; 10 morphological species (Species Fungorum 2020), 6
species with sequence data.
= Pseudoannulatascus Z.L. Luo, Maharachch. & K.D. Hyde, Phytotaxa 239 (2): 179 (2015)
= Fusoidispora Vijaykr., Jeewon & K.D. Hyde, Sydowia 57 (2): 272 (2005)
Type species – Torrentispora fibrosa K.D. Hyde, Wai H. Ho, E.B.G. Jones, K.M. Tsui &
S.W. Wong
931
Notes – Torrentispora was introduced by Hyde et al. (2000) to accommodate T. fibrosa on
decaying wood from freshwater habitat. Subsequently, the type species was also reported on
Nothofagus sp. and dried driftwood in New Zealand from terrestrial environments (Réblová et al.
2018). In the multigene phylogenetic analysis, Fusoidispora aquatica and Pseudoannulatascus
biatriisporus grouped with T. fibrosa, therefore, the genera Fusoidispora and Pseudoannulatascus
are synonymised with Torrentispora (Réblová et al. 2018). The sexual morph of Torrentispora is
characterized by semi-immersed, globose to conical ascomata with ostioles and necks, 8-spored,
unitunicate, cylindrical, short-pedicellate asci with J-, massive, apical rings, and ellipsoidal to
fusiform, hyaline, aseptate or transverse septate ascospores, sometimes with a fibrillar sheath
(Réblová et al. 2018). The asexual morph is undetermined in this genus.
Xyladictyochaetaceae Crous & Hern.-Restr., Fungal Systematics and Evolution 1: 212 (2018)
Index Fungorum number: IF824802; Facesoffungi number: FoF06787; 1 species.
Saprobic on leaf litter. Sexual morph: Undetermined. Asexual morph: Mycelium medium
brown, smooth, septate, branched hyphae, forming globose, intercalary, brown, smooth,
chlamydospore-like structures. Conidiophores erect, brown, smooth, subcylindrical, flexuous,
multiseptate. Conidiogenous cells terminal and intercalary, polyphialidic, lacking flared collarettes.
Conidia solitary, aggregating in slimy mass, hyaline, smooth, fusoid-ellipsoid, slightly curved, apex
subacute, base truncate, medially 1-septate; each end with flexuous, unbranched appendage
(adapted from Crous et al. 2018c).
Type genus – Xyladictyochaeta Hern.-Restr., R.F. Castañeda & Gené
Notes – Hernández-Restrepo et al. (2017) introduced Xyladictyochaeta typified by X.
lusitanica in Xylariales based on LSU phylogeny. Xyladictyochaetaceae was introduced by Crous
et al. (2018c) as a monotypic family, sister to Phlogicylindriaceae, based on LSU phylogeny. Multigene phylogeny (LSU-ITS-rpb2-tub2) in this study clearly shows that Xyladictyochaetaceae is
sister to Phlogicylindriaceae in Amphisphaeriales with high statistical support (95% ML).
Ecological and economic significance of Xyladictyochaetaceae
Xyladictyochaetaceae species are saprobes on dry leaves of Eucalyptus sp. (HernándezRestrepo et al. 2017).
Figure 266 – Xyladictyochaeta lusitanica (FMR 12177 ex-type, redrawn from Hernández-Restrepo
et al. 2017). a-c Conidiophores. d Conidia. Scale bars: 10 µm.
Genus included in Xyladictyochaetaceae
Xyladictyochaeta Hern.-Restr., R.F. Castañeda & Gené, Stud. Mycol. 86: 94 (2017)
Index Fungorum number: IF820355; 1 species with sequence data.
Type species – Xyladictyochaeta lusitanica Hern.-Restr., R.F. Castañeda & Gené
932
Notes – Xyladictyochaeta is similar to Dictyochaeta in having setiform conidiophores with
integrated, intercalary and terminal phialides and (0–)1-septate conidia with setulae at both ends.
However, the phylogenetic analyses show that those genera in different orders i.e. Dictyochaeta
(Chaetosphaeriales) and Xyladictyochaeta (Amphisphaeriales) (Crous et al. 2018c, Wei et al. 2018,
this study). At present, Xyladictyochaeta is a monotypic genus with Xyladictyochaeta lusitanica
isolated from dry leaves of Eucalyptus sp. in Australia and Portugal (Fig. 266) (HernándezRestrepo et al. 2017, Crous et al. 2018c).
Xylariaceae Tul. & C. Tul., Select. fung. carpol. (Paris) 2: 3 (1863)
Index Fungorum number: IF81528; Facesoffungi number: FoF00070; 1236 species.
Saprobic, pathogenic, or endophytic in wood, leaves and fruits or associated with insect
vectors. Sexual morph: Stromata extremely variable in size, shape and colour, erect or applanate or
effuse-pulvinate, or sometimes rudimentary or lacking, arising singly or aggregated, with one to
several ascomata, ostiolate, mostly without extractable stromal pigments, unipartite or bipartite.
Ascomata variable in size, globose-pyriform, single or multi-layered. Ostioles papillate, umbilicate
or at the same level as the stromal surface. Paraphyses hyaline, filamentous, septate, embedded in a
gelatinous matrix. Asci 4–8-spored, unitunicate, cylindrical to clavate, pedicellate or apedicellate,
apically rounded, with or without a J+, apical ring, or with apical thickenings. Ascospores
uniseriate-biseriate, brown to black, rarely hyaline, 1–2-celled, variously-shaped, mostly
ellipsoidal, subglobose or reniform, mostly with a germ slit, straight, spiral or sigmoid. Perispore
dehiscent or lacking, smooth or with patterns. Asexual morph: Hyphomycetous. Mostly
geniculosporium-like. Conidiophores hyaline to light brown, smooth, branched. Conidia hyaline,
roughened or smooth, ellipsoidal. Several hyphomycetous genera have been linked to Xylariaceae
(adapted from Maharachchikumbura et al. 2016b, Daranagama et al. 2018).
Type genus – Xylaria Hill ex Schrank
Notes – Maharachchikumbura et al. (2016b) accepted 87 genera with a complete family
description of Xylariaceae, which included the subfamilies Xylaroideae and Hypoxyloideae. Wendt
et al. (2018) re-established Hypoxylaceae (Hypoxyloideae) based on morphology, phylogeny and
secondary metabolite analyses. Based on morphology, phylogeny and molecular clock analyses,
Clypeosphaeriaceae was synonymized under Xylariaceae with a stem age of 63 MYA (Jaklitsch et
al. 2016b, Hongsanan et al. 2017). However, in present study, we accept Clypeosphaeriaceae as a
distinct family (see notes under Clypeosphaeriaceae). Daranagama et al. (2018) provided an
updated account of genera of Xylariaceae accepting 37 genera after observing type specimens.
However, Wendt et al. (2018) placed Gigantospora, Nipicola, Sabalicola, Spirodecospora,
Striatodecospora and Xylotumulus in Xylariales genera incertae sedis, due to lack of molecular
data and undetermined asexual morphs. We accept those genera in the Xylariaceae based on study
of herbarium specimens by Daranagama et al. (2018). Lee et al. (2016) and Johnston et al. (2016)
introduced Abieticola and Entalbostroma in Xylariaceae. Wijayawardene et al. (2017a) validly
published the genus Palmaria in Xylariaceae due to invalid nomenclature as Palmomyces (Hyde et
al. 1998a). Following recent updates, we accept 32 genera in Xylariaceae.
Xylariaceae species are important producers of bioactive compounds and secondary
metabolites (Stadler & Hellwig 2005, Helaly et al. 2018). Song et al. (2014) summarized 188
secondary metabolites with bioactive properties produced by Xylaria species. Since the taxa
presently included under Xylaria are not well-resolved, secondary metabolites may be possible
markers for species identification (Lee et al. 2000, Song et al. 2014). Isaka et al. (2012) described
novel secondary metabolites from Poronia gigantea, and suggested that poronitin A (1) and (R)-5methylmellein as useful chemotaxonomic markers for this genus. There are several other
xylariaceous genera such as Coniolariella, Entoleuca, Hypocopra, Kretzschmaria, Rosellinia and
Xylotumulus that have been subjected to secondary metabolites analyses (Stadler et al. 2001,
Stadler & Hellwig 2005, Helaly et al. 2018). Therefore, it is important to extend the morphology,
phylogeny and chemotaxonomy approaches to discover relationships in Xylariaceae (Helaly et al.
2018).
933
Ecological and economic significance of Xylariaceae
There are several important plant pathogenic species reported in this family especially
belonging to the genera Entoleuca and Rosellinia. Entoleuca mammata causes canker diseases
(hypoxylon canker) on Malus sp. (Rosaceae), Populus sp., Salix sp. (Salicaceae), and Sorbus sp.
(Rosaceae) (Kasanen et al. 2004, Ostry & Anderson 2009). Rosellinia necatrix and R. desmazieresii
which are known from temperate regions, and R. bunodes known from the tropics, causes root rot
of fruit trees and vines (Agrios 2005, Ten Hoopen & Krauss 2006). Among the root-diseases
causing Rosellinia species, R. bunodes is responsible for black root rot, R. necatrix for white root
rot and R. pepo for stellate root rot (Castro et al. 2013). Most species of this family are saprobes on
plant material. Cheng et al. (2015) introduced two new saprobic species of Ascotricha on the brown
alga Padina tetrastromatica, in intertidal regions of China. Species of Hypocopra, Podosordaria,
Poronia and Wawelia have been reported as coprophilous worldwide. Most of the species have
been investigated from Poaceae hosts as saprobes worldwide. Currently, extensive studies of
secondary metabolites and bioactive compouds from xylariaceous species have become a key
research area (Stadler & Hellwig 2005, Cheng et al. 2015, Hung et al. 2015, Chang et al. 2018,
Helaly et al. 2018). Therefore, species in Xylariaceae are of great interest due to their economic and
environmental importance as saprobes, endophytes, pathogens and secondary metabolites
producers.
Genera included in Xylariaceae
Abieticola Hyang B. Lee, Mycotaxon 131(4): 755 (2016)
Index Fungorum number: IF811702; 1 species with sequence data.
Type species – Abieticola koreana Hyang B. Lee
Notes – Abieticola koreana was described as an endophyte from the inner bark of a
Manchurian fir (Abies holophylla) in Korea. The genus differs from Poronia in having slightly
curved conidia, and its shorter conidiogenous cells that sometimes bear 3 conidia.
Amphirosellinia Y.M. Ju, J.D. Rogers, H.M. Hsieh & Lar.N. Vassiljeva, Mycologia 96(6): 1393
(2004)
Index Fungorum number: IF28880; 6 morphological species (Species Fungorum 2020), 3
species with sequence data.
Type species – Amphirosellinia nigrospora Y.M. Ju, J.D. Rogers & H.M. Hsieh
Notes – Amphirosellinia has rarely erumpent to mostly immersed perithecioid, bipartite
stromata, and has a close affinity to Rosellinia. The genus has synnematous, geniculosporium-like
asexual morphs that produce lacrymoid conidia (Ju et al. 2004). However, Amphirosellinia differs
from Rosellinia in having stromata that develop beneath the host epidermis with a thick, carbonized
crust. The genus has been identified with a potential antimicrobial agent using A. nigrospora
(Nguyen et al. 2018).
Anthostomella Sacc., Atti Soc. Veneto-Trent. Sci. Nat., Padova, Sér. 4 4: 84 (1875)
Index Fungorum number: IF226; 118 morphological species (Lu & Hyde 2000, Species
Fungorum 2020), 35 species with sequence data.
Type species – Anthostomella tomicoides Sacc.
Notes – Anthostomella is characterized by immersed, dark, clypeate ascomata with
periphysate ostiolar canals, 8-spored, cylindrical, unitunicate asci and mostly dark, unicellular
ascospores, sometimes with dwarf cells or appendages at the ends (Daranagama et al. 2015).
Daranagama et al. (2015, 2016) described the polyphyletic nature of the genus and emphasized the
need for recollection and providing molecular data. Voglmayr et al. (2018) re-evaluated the genus
with suggestions to provide a new family for Anthostomella with the availability of sequence data
of the type species. In this study, Anthostomella does not appear to be phylogenetically related to
Xylariaceae. Daranagama et al. (2018) re-examined several herbarium specimens and tentatively
synonymized Appendixia with Anthostomella based on morphological similarities.
934
Anthostomelloides Tibpromma & K.D. Hyde, Turkish Journal of Botany 41: 107-116 (2017)
Index Fungorum number: IF552117; 5 morphological species (Daranagama et al. 2018), 2
species with sequence data.
Type species – Anthostomelloides krabiensis Tibpromma & K.D. Hyde
Notes – Anthostomelloides was introduced as a saprobe on dead leaves of Pandanus odorifer.
It is characterized by immersed, globose ascomata and a peridium comprising cells of textura
prismatica (Tibpromma et al. 2017a).
Ascotricha Berk., Ann. nat. Hist., Mag. Zool. Bot. Geol. 1: 257 (1838)
Index Fungorum number: IF384; 18 morphological species (Cheng et al. 2015, Li & Zhao
2018, Species Fungorum 2020), 9 species with sequence data.
Type species – Ascotricha chartarum Berk.
Notes – Ascotricha species are characterized by ascomata having long hairs with shorter,
hyaline branches, brown, ellipsoidal ascospores with an equatorial germ slit and geniculosporium
type asexual morphs (Berkeley 1838, Cheng et al. 2015). Cheng et al. (2015) added two novel
species and one combination of Ascotricha from algae and confirmed the taxonomic placement of
the genus in Xylariaceae. Li & Zhao (2018) added one new species and two combinations.
However, BLASTn results of the available sequences and phylogenetic affinities of our multigene
analyses that show Ascotricha is close to Zygosporium. Therefore, revision of Ascotricha is needed
to establish its taxonomic placement.
Astrocystis Berk. & Broome, J. Linn. Soc., Bot. 14(no. 74): 123 (1873)
Index Fungorum number: IF439; 24 morphological species (Species Fungorum 2020), 7
species with sequence data.
Type species – Astrocystis mirabilis Berk. & Broome
Notes – Astrocystis accommodates mostly superficial, uniascomatal stromata bearing species
(Berkeley & Broome 1875). The genus formed a monophyletic clade in analyses of ITS, LSU,
SSU, rpb2 and tub2 sequence data (Hsieh et al. 2010, Li et al. 2015b, c). Most Astrocystis species
occur on bamboo and are widespread in tropical and subtropical regions.
Brunneiperidium Daranag., Camporesi & K.D. Hyde, Fungal Divers. 73:221 (2015)
Index Fungorum number: IF809521; 2 species with sequence data.
Type species – Brunneiperidium gracilentum Daranag., Camporesi & K.D. Hyde
Notes – Brunneiperidium was introduced based on its unique peridium characters, with the
outermost wall of irregularly arranged brown and yellowish-brown cells. Phylogenetic analyses
revealed that the genus clustered between Kretzschmaria and Xylaria (Daranagama et al. 2018).
Collodiscula I. Hino & Katum., Bull. Faculty of Agriculture, Yamaguchi University 6: 55 (1955)
Index Fungorum number: IF1187; 5 species with sequence data.
Type species – Collodiscula japonica I. Hino & Katum.
Notes – Collodiscula was introduced by Hino & Katumoto (1955) to accommodate species
with brown, 1-septate ascospores lacking a germ slit, superficial, stromatic ascomata and large, J+,
wedge-shaped ascal apical ring. There are five species of Collodiscula are mostly associated with
bamboo (Li et al. 2015b, Hyde et al. 2017b).
Coniolariella Dania García, Stchigel & Guarro, Mycol. Res. 110(11): 1285 (2006)
Index Fungorum number: IF29050; 4 species with sequence data.
Type species – Coniolariella gamsii (Asgari & Zare) Dania García, Stchigel & Guarro
Notes – Coniolariella is reminiscent of Obolarina in some morphological characters as both
genera have J-, short pedicellate asci without a distinct apical ring. Coniolariella is also similar to
Rosellinia in asci and ascospore morphology.
935
Engleromyces Henn., Bot. Jb. 28(3): 327 (1900)
Index Fungorum number: IF1819; 2 morphological species (Species Fungorum 2020).
Type species – Engleromyces goetzei Henn.
Notes – Engleromyces is characterized by massive, hard stromata similar to
macroxylariaceous genera such as Sarcoxylon (Whalley et al. 2010). Engleromyces goetzei
possesses medicinal values and is used in Africa and China (Kokwaro 1983). Both, Engleromyces
goetzei and E. sinensis have been analysed for the secondary metabolites and contain
neoengleromycin and cytochalasins (Liu et al. 2002). Daranagama et al. (2018) accepted the genus
in Xylariaceae until molecular data are available.
Entalbostroma J.D. Rogers & P.R. Johnst., Mycotaxon 131(4): 766 (2017)
Index Fungorum number: IF817225; 1 species with sequence data.
Type species – Entalbostroma erumpens J.D. Rogers & P.R. Johnst.
Notes – Entalbostroma is a monotypic genus. Entalbostroma erumpens was described on
dead leaves of Phormium tenax and P. cookianum (Johnston et al. 2016). The genus is
characterized by pulvinate to applanopulvinate, orbicular to elliptical to irregular stromata, dark
brown ascospores with a germ slit on flattened side, a hyaline gelatinous sheath in sexual and
palisade-like layer of short, cylindric conidiophores and hyaline, smooth, ellipsoid conidia with
narrowed flattened bases (Johnston et al. 2016).
Entoleuca Syd., Annls mycol. 20(3/4): 186 (1922)
Index Fungorum number: IF1833; 3 species with sequence data.
Type species – Entoleuca callimorpha Syd.
Notes – Sydow & Petrak (1922) introduced the genus with E. callimorpha as the type species.
Until 1994, Hypoxylon mammatum was considered as similar taxa to E. callimorpha. However,
Læssøe & Spooner (1994) and Læssøe (1994) treated the H. mammatum as separate, but synonym
of Rosellinia. Based on these taxonomic confusions, Rogers & Ju (1996) revised the type, authentic
and other specimens and re-established the genus. Entoleuca consists of saprobic and plant
pathogenic species distributed in Europe. As a pathogen, E. mammata causes canker (hypoxylon
canker) on Malus sp. (Rosaceae), Populus sp., Salix sp. (Salicaceae), and Sorbus sp. (Rosaceae)
(Shaw 1973, Callan 1998, Kasanen et al. 2004, Eriksson 2014) and is a saprobe on decaying tree
trunks.
Euepixylon Füisting, Bot. Ztg. 25(no. 39): 309 (1867)
Index Fungorum number: IF27402; 2 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Euepixylon udum (Pers.) Læssøe & Spooner
Notes – The genus is characterized by semi-immersed, multiascomatal, linear stomata with
conspicuous ascomatal mounds, forming a clypeoid carbonaceous layer and cylindrical, unitunicate
asci with short-pedicel. Daranagama et al. (2018) and Wendt et al. (2018) accepted the genus in
Xylariaceae. However, sequence data of the type species are needed to investigate its potential
synonymy with Nemania.
Halorosellinia Whalley, E.B.G. Jones, K.D. Hyde & Læssøe, Mycol. Res. 104(3): 368 (2000)
Index Fungorum number: IF28368; 3 morphological species (Species Fungorum 2020), 2
species with sequence data.
Type species – Halorosellinia oceanica (S. Schatz) Whalley, E.B.G. Jones, K.D. Hyde &
Læssøe
Notes – Halorosellinia was established to accommodate H. oceanicum, which was earlier
accepted as Hypoxylon oceanicum. The sexual morph of the genus is characterized by immersed
ascomata in a poorly developed pseudostromata, unitunicate, 8-spored asci with a cylindrical, J+,
apical ring and dark brown-opaque, ellipsoidal ascospores with a straight germ slit (Whalley et al.
936
2010). Based on sexual and asexual morphology coupled with phylogenetic analyses, Daranagama
et al. (2018) and Wendt et al. (2018) accepted this genus in Xylariaceae.
Helicogermslita Lodha & D. Hawksw., Trans. Br. mycol. Soc. 81(1): 91 (1983)
Index Fungorum number: IF2254; 9 morphological species (Species Fungorum 2020).
Type species – Helicogermslita celastri (S.B. Kale & S.V.S. Kale) Lodha & D. Hawksw.
Notes – Hawksworth & Lodha (1983) described the genus which is characterized by asci with
a J-, apical ring and ascospores with spiral germ slits. The asexual morph has been described as
geniculosporium-like. There are no sequence data for this genus.
Hypocopra (Fr.) J. Kickx f., Fl. Crypt. Flandres (Paris) 1: 362 (1867)
Index Fungorum number: IF2431; 31 morphological species (Species Fungorum 2020), 2
species with sequence data.
Type species – Hypocopra merdaria (Fr.) J. Kickx f.
Notes – Hypocopra species are characterized by sessile, clypeoid or sometimes even
rudimentary stromata, mainly inhabiting dung (Krug & Cain 1973).
Hypocreodendron Henn., Hedwigia 36(4): 223 (1897)
Index Fungorum number: IF8615; 1 species with sequence data.
Type species – Hypocreodendron sanguineum Henn.
Notes – Hypocreodendron is a monotypic genus introduced by Hennings (1897) which is
characterized by more or less cylindrical, stromata with an apical discoid or shallow cup bearing
conidia. The species is associated with ant nests. Based on sexual asexual morph connections,
Hypocreodendron was validated over Discoxylaria (Stadler et al. 2013, Maharachchikumbura et al.
2015b, Réblová et al. 2016b).
Kretzschmaria Fr., Summa veg. Scand., Sectio Post. (Stockholm): 409 (1849)
Index Fungorum number: IF2589; 34 morphological species (Pourmoghaddam et al. 2018,
Species Fungorum 2020), 11 species with sequence data.
Type species – Kretzschmaria clavus (Fr.) Sacc.
Notes – Kretzschmaria was established by Fries (1849) which is similar to Xylaria. The
genus is characterized by pulvinate or effused-pulvinate stromata attached to the substrate by
narrow connectives, or clavate, turbinate, obconical, peltate-discoid, often aggregated into a crustlike mass. Pourmoghaddam et al. (2018) provided an update for the genus while introducing two
new species K. hedjaroudei and K. iranica from Iran.
Kretzschmariella Viégas, Bragantia 4(1-6): 105 (1944)
Index Fungorum number: IF2590; 1 species with sequence data.
Type species – Kretzschmariella culmorum (Cooke) Y.-M. Ju & J. D. Rogers
Notes – Kretzschmariella is a monotypic genus which was resurrected by Ju & Rogers
(1994). Kretzschmariella culmorum has superficial stromata on bamboo and bamboo-like
substrates and and one to multicellular, ellipsoid to oblong-obovate or cylindrical conidia. There
are confusion associated among Kretzschmariella and some Hypoxylon species.
Leprieuria Læssøe, J.D. Rogers & Whalley, Mycol. Res. 93(2): 152 (1989)
Index Fungorum number: IF25391; 1 morphological species.
Type species – Leprieuria bacillum (Mont.) Læssøe, J.D. Rogers & Whalley
Notes – Leprieuria is characterized by cylindrical to rod-shaped stromata, umbilicate, usually
2–10 ostioles per stroma, cylindrical, short-pedicellate asci lacking an apical ring, reniform
ascospores glabrous in face view and straight germ slit and a geniculosporium-like asexual morph
(see Daranagama et al. 2018). Wendt et al. (2018) accepted Leprieuria in Xylariaceae based on its
asexual morph. Molecular analyses are needed to resolve this genus.
937
Lunatiannulus Daranag., Camporesi & K.D. Hyde, Fungal Divers.: 73: 227 (2015)
Index Fungorum number: IF809519; 1 species with sequence data.
Type species – Lunatiannulus irregularis Daranagama, Camporesi & K.D. Hyde
Notes – This diatrypaceous-like genus was introduced to accommodate species with ascomata
with a well-developed clypeus, cylindrical asci with an apical ring, ellipsoidal ascospores with
germ slits and hyaline appendages or dwarf cells and a libertella-like asexual morph. Daranagama
et al. (2018) accepted this genus in Xylariaceae based on morphology and multigene phylogeny.
Nemania Gray, Nat. Arr. Brit. Pl. (London) 1: 516 (1821)
Index Fungorum number: IF3437; 53 morphological species (Species Fungorum 2020), 14
species with sequence data.
Type species – Nemania serpens (Pers.) Gray
Notes – Nemania is a species rich and well-established genus in Xylariaceae (Daranagama et
al. 2018, Wendt et al. 2018) characterized by stromata not associated with bark rupturing structures,
lack KOH-extractable pigments and finely papillate ostioles. Maharachchikumbura et al. (2016b)
and Réblová et al. (2016b) proposed the use of Nemania over Geniculosporium following Stadler et
al. (2013). Nemania bipapillata is illustrated in this paper (Fig. 267).
Podosordaria Ellis & Holw., Bot. Gaz. 24: 37 (1897)
Index Fungorum number: IF4282; 33 morphological species (Daranagama et al. 2018), 5
species with sequence data.
Type species – Podosordaria mexicana Ellis & Holw.
Notes – Podosordaria was described by Holway (1897) and has erumpent, obconical
stromata with glomerate head and stipe, clavate-slightly pyriform, short-pedicellate asci with a J+,
apical ring and equilateral ellipsoidal, dark brown ascospores with a gelatinous sheath and a
geniculosporium-like asexual morph. Multigene phylogenetic studies revealed that the genus has a
close affinity to Poronia in Xylariaceae (Daranagama et al. 2018, Wendt et al. 2018).
Poronia Willd., Fl. berol. prodr.: 400 (1787)
Index Fungorum number: IF4346; 9 morphological species (Species Fungorum 2020), 3
species with sequence data.
Type species – Poronia gleditschii Willd.
Notes – Poronia has a close affinity to Podosordaria and is distinguished by a lindquistia
conidial state (Rogers & Ju 1998). The sexual morph of the genus is typically stalked stromata in
the form of a flattened disc (Krug & Cain 1973). The genus is accepted in Xylariaceae based on
morphology and multigene phylogeny (Daranagama et al. 2018, Wendt et al. 2018).
Rosellinia De Not., G. bot. ital. 1(1): 334 (1844)
Index Fungorum number: IF4785; 158 morphological species (Jayawardena et al. 2019), 27
species with sequence data.
Type species – Rosellinia aquila (Fr.) Ces. & De Not.
Notes – Rosellinia was introduced to accommodate uniascomatal species (De Notaris 1844).
Petrini (2013) published a monograph accepting 142 species and several species were added
(Ariyawansa et al. 2015, Su et al. 2016a, Tibpromma et al. 2017b, Fournier et al. 2017, Niranjan &
Sarma 2018). We observed a herbarium specimen of Rosellinia bunodes (Fig. 268), the species
occurred on dead wood and was described from Sri Lanka. It has long ascospores with thread-like
ends and short germ slits. Rosellinia bunodes causes black root rot of fruit trees and vines, mainly
in the tropics (Ten Hoopen & Krauss 2006, Castro et al. 2013, dos Santos et al. 2017).
Sarcoxylon Cooke, Grevillea 12(no. 62): 50 (1883)
Index Fungorum number: IF4863; 5 morphological species (Daranagama et al. 2018), 1
species with sequence data.
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Figure 267 – Nemania bipapillata (Material examined – INDIA, Andaman and Nicobar Islands,
South Andaman, Mount Harriet, Gun Point (11˚72’23”N 92˚73’80”E), decaying twig, on 7
December, 2017, collected by M. Niranjan & V.V.Sarma. Herbarium accession AMH-10074,
Ajrekar Mycological Herbarium, Agarkar Research Institute (ARI), Pune, India; living culture
PUFNI 17612. INDIA, South Andaman, Jirkatang (11˚45’5” N 92˚39’7”E), terrestrial forest, on an
unidentified decaying twig 28 March 2017, collected by M. Niranjan & V.V. Sarma (PUFNI
T128F1) Fungal Biotechnology Lab, Department of Biotechnology, Pondicherry University inhouse herbarium code). a, b Stromata on host. c Stromata vertical section. d, e Culture on malt
extract agar plates (d-from above, e-reverse). f Paraphyses. g-j Asci. k J+ apical ring. l-n
Ascospores. Scale bars: i = 50 μm, f, h = 20 μm, g, j-n = 10 μm.
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Figure 268 – Rosellinia bunodes (Material examined – SRI LANKA, Peradeniya, on dead wood,
November 1867, G.H.K. Thwaites, K(M) 62957, holotype). a, b Stromata (b ostiole in white
arrow). c Synnemata. d-g Herbarium details. h Paraphyses. i-m Ascospores (j germ slit in black
arrow, m in 10% KOH). Scale bars: a = 1000 µm, b = 500 µm, c = 200 µm, i-m = 20 µm, h = 5 µm.
Type species – Sarcoxylon compunctum (Jungh.) Cooke
Notes – Sarcoxylon was established by Cooke (1883). Rogers (1981) maintained Sarcoxylon
as a distinct genus apart from Hypoxylon, Entonaema and Xylaria. The genus is characterized by
superficial, pulvinate to hemispherical to more or less upright stromata with a gelatinous zone
beneath ascomatal layer, cylindrical, pedicellate asci with a J+, apical ring and unicellular,
ellipsoid-inequilateral, brown ascospores. The phylogenetic studies of Senanayake et al. (2015),
Daranagama et al. (2018) and Wendt et al. (2018) accepted Sarcoxylon in Xylariaceae.
Squamotubera Henn., Hedwigia 42(Beibl.): (308) (1903)
Index Fungorum number: IF5180; 1 morphological species (Daranagama et al. 2018).
Type species – Squamotubera le-ratii Henn.
Notes – Hennings (1903) erected the monotypic genus Squamotubera which is characterized
by erect, cylindrical to clavate, unbranched stromata with fertile upper part and unicellular,
ellipsoidal ascospores with one flattened side, sometimes bean-shaped, brown, with narrowly
rounded ends and a spiral germ slit. Based on characters, Daranagama et al. (2018) and Wendt et al.
(2018) accepted Squamotubera in Xylariaceae.
940
Stilbohypoxylon Henn., Hedwigia 41: 16 (1902)
Index Fungorum number: IF5264; 11 morphological species (Daranagama et al. 2018), 2
species with sequence data.
Type species – Stilbohypoxylon moelleri Henn.
Notes – Hennings (1903) described the genus Stilbohypoxylon and tentatively placed it in
Xylariaceae. Stilbohypoxylon has superficial, sphaerical stromata with a wrinkled surface (Rogers
& Ju 1997). Daranagama et al. (2018) and Wendt et al. (2018) accepted the genus in Xylariaceae
based on morphology and phylogenetic studies.
Vamsapriya Gawas & Bhat, Mycotaxon 94: 150 (2006)
Index Fungorum number: IF29041; 8 morphological species (Jiang et al. 2018), 6 species
with sequence data.
Type species – Vamsapriya indica Gawas & Bhat
Notes – Vamsapriya was introduced by Gawas & Bhat (2005) based on asexual morph, and
Dai et al. (2017) described the sexual morph for the genus. The sexual morph has solitary,
scattered, immersed ascomata with papillate, ostiole, cylindrical, shortly pedicellate asci with a J+,
apical ring, and fusiform, apiosporous, 1-septate, hyaline ascospores with pointed ends surrounded
by inconspicuous sheath. The asexual morph has dark brown, erect, rigid, synnematous
conidiophores, terminal or intercalary, monotretic, enteroblastic, dark brown, ellipsoidal
conidiogenous cells and cylindrical, fusiform, straight to flexuous, brown to dark brown, 0–20septate, slightly verrucose conidia. Most of the species are described from bamboo (Dai et al.
2014a, 2017, Jiang et al. 2018). Based on phylogeny and morphology, Jiang et al. (2018) and
Wendt et al. (2018) accepted the genus in Xylariaceae.
Virgaria Nees, Syst. Pilze (Würzburg): 54 (1816)
Index Fungorum number: IF10408; 11 morphological species (Species Fungorum 2020), 2
species with sequence data.
Type species – Virgaria nigra (Link) Nees
Notes – Virgaria is accepted over Ascovirgaria with its geniculosporium-like asexual morph
(Rogers & Ju 2002, Stadler et al. 2013, Maharachchikumbura et al. 2015b, Réblová et al. 2016b).
The sexual morph is characterized by rudimentary stromata, globose ascomata, cylindrical, long
pedicellate asci with a J+, apical ring and unicellular, ellipsoid, brown ascospores. The genus is
accepted in Xylariaceae (Daranagama et al. 2018, Wendt et al. 2018).
Wawelia Namysł., Bull. int. Acad. Sci. Lett. Cracovie, Cl. sci. math. nat. Sér. B, sci. nat. 2: 602
(1908)
Index Fungorum number: IF5764; 5 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Wawelia regia Namysł.
Notes – Wawelia was described based on W. regia, and originally placed in Hypocreaceae
(Namyslowski 1908). With the introduction of more Wawelia species by Lundqvist (1992) and
Webster et al. (1999) the genus was accepted in Xylariaceae. Wendt et al. (2018) confirmed the
genus in Xylariaceae based on geniculosporium-like asexual morph.
Xylaria Hill ex Schrank, Baier. Fl. (München) 1: 200 (1789)
Index Fungorum number: IF5832; more than 670 morphological species (Species Fungorum
2020), 132 species with sequence data.
Type species – Xylaria hypoxylon (L.) Grev.
Notes – Mostly, species of Xylaria are characterized by large stromata, cylindrical asci with
long stipes, ellipsoidal, dark ascospores in their sexual and geniculosporium-like asexual morph (Ju
& Rogers 1996, Stadler et al. 2013). Xylaria is a highly diversified genus, most likely exceeds 500
species (Peršoh et al. 2009). Most Xylaria species are saprobes growing on moist, decaying
941
dicotyledonous wood and less often on monocotyledons, fruits, seeds and fallen leaves. The
endophytic species are common in tropical plants. Interestingly, the subgenus Pseudoxylaria is
associated with termite nests, mainly macrotermitine termites (Visser et al. 2009, Hsieh et al.
2010). Based on rDNA sequences data Arthroxylaria and Geniculisynnema were synonymised
under Xylaria (Seifert et al. 2002, Okane & Nakagiri 2007, Réblová et al. 2016b). The significant
variations of the characters of Xylaria species might be the result of many convergent evolution
within the genus (Lee et al. 2000).
Zygosporiaceae J.F. Li, Phookamsak & K.D. Hyde, Mycosphere 8(10): 1860 (2017)
Index Fungorum number: IF553846; Facesoffungi number: FoF0376; 21 species.
Saprobic on various host plants, especially on monocotyledons. Sexual morph:
Undetermined. Asexual morph: Hyphomycetous. Colonies effuse, white to light pink. Mycelium
immersed or partly superficial, composed of smooth, thin-walled, white or light pink hyphae. Setae
or setiform conidiophores with sphaerical apex. Conidiophores macronematous, mononematous,
solitary or in small groups, pale brown, thin-walled, unbranched, septate, smooth, bearing swollen
dark brown, thick-walled vesicles. Conidiogenous cells holoblastic, discrete, hyaline or light
brown, smooth, sphaerical to ellipsoid, borne in groups of 2–4 on the vesicular cell. Conidia
solitary, aseptate, hyaline or pale brown, globose or ellipsoid, thin- or thick-walled (adapted from
Li et al. 2017b).
Type genus – Zygosporium Mont.
Notes – Zygosporiaceae was introduced by Li et al. (2017b) to accommodate a single genus
Zygosporium. It was based on phylogenetic support as a distinct lineage and separated from other
taxa in Xylariales using analysis of LSU and ITS sequence data. Species of Zygosporiaceae are
saprobes on plant litter, especially on monocotyledons such as Pandanaceae and palms (Li et al.
2017b).
Ecological and economic significance of Zygosporiaceae
Species are saprobes on a wide variety of plant substrates. Zygosporiaceae species are also
sources of biologically active secondary metabolites (Hayakawa et al. 1968, Oh et al. 2006, Kanoh
et al. 2008). For example, Zygosporium masonii is known for the production of cytochalasins
(Hayakawa et al. 1968).
Genus included in Zygosporiaceae
Zygosporium Mont., Annls Sci. Nat., Bot., sér. 2 17: 120 (1842)
Index Fungorum number: IF10473; 21 morphological species (Species Fungorum 2020), 8
species with sequence data.
Type species – Zygosporium oscheoides Mont.
Notes – Zygosporium was introduced by Montagne (1842) with Z. oscheoides as the type
species. Species are usually saprobes in temperate to tropical regions (Photita et al. 2001, Whitton
et al. 2003, Manoharachary et al. 2006, McKenzie et al. 2007, Abbas et al. 2011, Pratibha et al.
2012, Taheriyan et al. 2014). Taxa are hyphomycetous (asexual morphs) with darkly pigmented
colonies, incurved vesicular cells and globose or ellipsoid and smooth-walled or variously
ornamented conidia (Mason 1941, Hughes 1951c). Sequence data are available only for Z.
chartarum, Z. echinosporum, Z. gibbum, Z. masonii, Z. minus, Z. mycophilum, Z. oscheoides and Z.
pseudogibbum. In this entry a new collection of Z. masonii from Dracaena is illustrated (Fig. 269).
Placement of genera in orders incertae sedis
Amphisphaeriales genera incertae sedis
Chitonospora E. Bommer, M. Rousseau & Sacc., Bull. Soc. R. Bot. Belg. 29(no. 1): 270 (1890)
Index Fungorum number: IF1003; 1 morphological species.
Type species – Chitonospora ammophila E. Bommer, M. Rousseau & Sacc.
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Figure 269 – Zygosporium masonii (Material examined – THAILAND, Songkhla Province, on
dead leaves of Dracaena sp., 5 May 2018, Napalai Chaiwan NSW5, MFLU 18-0124). a, b
Appearance on host surface. c-f Vesicle, conidiophores and conidiogenous cell. g, h Conidia. Scale
bars: a, b = 1000 µm, c, d = 50 μm, e, f = 20 μm, g, h = 10 μm.
Fasciatispora K.D. Hyde, Trans. Mycol. Soc. Japan 32(2): 265 (1991)
Index Fungorum number: IF14386; 10 morphological species (Species Fungorum 2020), 4
species with sequence data.
Type species – Fasciatispora nypae K.D. Hyde
Annulatascales genera incertae sedis
Clohiesia K.D. Hyde, Nova Hedwigia 61(1-2): 125 (1995)
Index Fungorum number: IF27574; 3 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Clohiesia corticola K.D. Hyde
Calosphaeriales genus incertae sedis
Calosphaeriopsis Petr., Annls mycol. 39(4/6): 272 (1941)
Index Fungorum number: IF754; 1 morphological species.
Type species – Calosphaeriopsis huberiana (Kirschst.) Petr.
Enchnoa Fr., Summa veg. Scand., Sectio Post. (Stockholm): 410 (1849)
Index Fungorum number: IF1778; 19 morphological species (Species Fungorum 2020).
Type species – Enchnoa lanata (Fr.) Fr.
Kacosphaeria Speg., Boln Acad. nac. Cienc. Córdoba 11(2): 214 (1887)
Index Fungorum number: IF2541; 1 morphological species.
Type species – Kacosphaeria antarctica Speg.
Sulcatistroma A.W. Ramaley, Mycotaxon 93: 140 (2005)
943
Index Fungorum number: IF28974; 1 species with sequence data.
Type species – Sulcatistroma nolinae A.W. Ramaley
Chaetosphaeriales genera incertae sedis
Calvolachnella Marinc., T.A. Duong & M.J. Wingf., Sydowia 68: 203 (2016)
Index Fungorum number: IF815619; 1 species with sequence data.
Type species – Calvolachnella guaviyunis (Marinc., T.A. Duong, M.J. Wingf. & C.A. Perez)
Marinc., T.A. Duong, M.J. Wingf.
Caudatispora J. Fröhl. & K.D. Hyde, Sydowia 47(1): 38 (1995)
Index Fungorum number: IF27569; 2 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Caudatispora palmicola J. Fröhl. & K.D. Hyde
Erythromada Huhndorf, A.N. Mill., F.A. Fernández & Lodge, Fungal Divers. 20: 63 (2005)
Index Fungorum number: IF501320; 1 species with sequence data.
Type species – Erythromada lanciospora Huhndorf, A.N. Mill., F.A. Fernández & Lodge
Lasiosphaeriella Sivan., Trans. Br. mycol. Soc. 64(3): 443 (1975)
Index Fungorum number: IF2657; 5 morphological species (Species Fungorum 2020), 3
species with sequence data.
Type species – Lasiosphaeriella dennisii Sivan.
Neoleptosporella Phukhams., Perera & K.D. Hyde, Fungal Divers. (in prep)
Index Fungorum number: IF557060;1 species with sequence data.
Type species – Neoleptosporella clematidis Phukhams., Konta & K.D. Hyde
Rimaconus Huhndorf, F.A. Fernández, Joanne E. Taylor & K.D. Hyde, Mycologia 93(6): 1073
(2001)
Index Fungorum number: IF28573; 2 species with sequence data.
Type species – Rimaconus jamaicensis (Seaver) Huhndorf, F.A. Fernández, Joanne E. Taylor
& K.D. Hyde
Coniochaetales genera incertae sedis
Cannonia Joanne E. Taylor & K.D. Hyde, Mycol. Res. 103(11): 1398 (1999)
Index Fungorum number: IF28356; 1 morphological species.
Type species – Cannonia australis (Speg.) Joanne E. Taylor & K.D. Hyde
Pseudogliomastix W. Gams, Proc. Indian Acad. Sci., Pl. Sci. 94(2-3): 279 (1985)
Index Fungorum number: IF11172; 1 morphological species.
Type species – Pseudogliomastix protea (Sacc.) W. Gams
Coronophorales genera incertae sedis
Papulaspora Preuss, Linnaea 24: 112 (1851)
Index Fungorum number: IF9205; 33 morphological species (Species Fungorum 2020), 7
species with sequence data.
Type species – Papulaspora sepedonioides Preuss
Sphaerodes Clem., Gen. fung. (Minneapolis): 44 (1909)
944
Index Fungorum number: IF 5099; 9 morphological species (Species Fungorum 2020), 4
species with sequence data.
Type species – Sphaerodes episphaerium (W. Phillips & Plowr.) Clem.
Diaporthales genera incertae sedis
Ceratoporthe Petr., Annls mycol. 23(1/2): 14 (1925)
Index Fungorum number: IF894; 1 morphological species.
Type species – Ceratoporthe didymospora Petr.
Cryptoleptosphaeria Petr., Annls mycol. 21(3/4): 196 (1923)
Index Fungorum number: IF1319; 1 morphological species.
Type species – Cryptoleptosphaeria moravica Petr.
Cytomelanconis Naumov, Bot. Mater. Otd. Sporov. Rast. Bot. Inst. Komarova Akad. Nauk
S.S.S.R. 7: 108 (1951)
Index Fungorum number: IF1396; 1 morphological species.
Type species – Cytomelanconis systema-solare Naumov
Diaporthella Petr., Annls mycol. 22(1/2): 30 (1924)
Index Fungorum number: IF1498; 1 species with sequence data.
Type species – Diaporthella aristata (Fr.) Petr.
Diatrypoidiella Manohar., Kunwar & D.K. Agarwal, Indian Phytopath. 58(2): 205 (2005)
Index Fungorum number: IF29064; 1 morphological species.
Type species – Diatrypoidiella lignicola Manohar., Kunwar & D.K. Agarwal
Ditopellina J. Reid & C. Booth, Can. J. Bot. 45: 1481 (1967)
Index Fungorum number: IF1678; 1 morphological species.
Type species – Ditopellina saccardoana (Traverso & Spessa) J. Reid & C. Booth
Durispora K.D. Hyde, Sydowia 46(2): 315 (1994)
Index Fungorum number: IF27442; 2 morphological species (Species Fungorum 2020).
Type species – Durispora elaeidicola K.D. Hyde
Exormatostoma Gray, Nat. Arr. Brit. Pl. (London) 1: 521 (1821)
Index Fungorum number: IF1965; 2 morphological species (Species Fungorum 2020).
Type species – Exormatostoma nebulosa (Pers.) Gray
Fremineavia Nieuwl., Am. Midl. Nat. 4: 501 (1916)
Index Fungorum number: IF2014; 1 morphological species.
Type species – Fremineavia berkeleyi (Berl.) Nieuwl.
Gibellia Sacc., Atti Inst. Veneto Sci. lett., ed Arti, Sér. 6 3: 714 (1885)
Index Fungorum number: IF2063; 1 morphological species.
Type species – Gibellia dothideoides Sacc. & Berl.
Gyrostroma Naumov, Bull. Soc. mycol. Fr. 30(3): 386 (1914)
Index Fungorum number: IF8413; 3 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Gyrostroma sinuosum Naumov
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Hyalorostratum Raja & Shearer, Mycosphere 1(4): 281 (2010)
Index Fungorum number: IF518893; 1 species with sequence data.
Type species – Hyalorostratum brunneisporum Raja & Shearer
Hypophloeda K.D. Hyde & E.B.G. Jones, Trans. Mycol. Soc. Japan 30(1): 61 (1989)
Index Fungorum number: IF25308; 1 morphological species.
Type species – Hypophloeda rhizospora K.D. Hyde & E.B.G. Jones
Hypospilina (Sacc.) Traverso, Fl. ital. crypt., Pyrenomycetae (Florence): 495 (1913)
Index Fungorum number: IF2451; 4 morphological species (Species Fungorum 2020).
Type species – Hypospilina bifrons (DC.) Traverso
Kapooria J. Reid & C. Booth, Can. J. Bot. 67(3): 898 (1989)
Index Fungorum number: IF25312; 1 morphological species.
Type species – Kapooria musarum (J.N. Kapoor) J. Reid & C. Booth
Keinstirschia J. Reid & C. Booth, Can. J. Bot. 67(3): 897 (1989)
Index Fungorum number: IF25314; 1 morphological species.
Type species – Keinstirschia megalosperma (Kirschst.) J. Reid & C. Booth
Lollipopaia Inderb., Can. J. Bot. 79(9): 1100 (2001)
Index Fungorum number: IF28542; 1 species with sequence data.
Type species – Lollipopaia minuta Inderb.
Macrodiaporthe Petr., Annls mycol. 17(2/6): 94 (1920)
Index Fungorum number: IF2963; 1 morphological species.
Type species – Macrodiaporthe occulta (Fuckel) Petr. (≡ Calospora occulta Fuckel)
Melanamphora Lafl., Sydowia 28(1-6): 243 (1976)
Index Fungorum number: IF3055; 1 morphological species.
Type species – Melanamphora spinifera (Wallr.) Lafl.
Phragmodiaporthe Wehm., Mycologia 33(1): 54 (1941)
Index Fungorum number: IF4024; 3 morphological species.
Type species – Phragmodiaporthe caryae (Peck) Wehm. (≡ Cryptospora caryae Peck)
Phruensis Pinruan, Mycologia 96(5): 1165 (2004)
Index Fungorum number: IF28878; 1 species with sequence data.
Type species – Phruensis brunneispora Pinruan
Plagiophiale Petr., Sydowia 9(1-6): 585 (1955)
Index Fungorum number: IF4154; 2 morphological species (Species Fungorum 2020).
Type species – Plagiophiale eucarpa (P. Karst.) Petr.
Plagiostigme Syd., Annls mycol. 23(3/6): 341 (1925)
Index Fungorum number: IF4156; 7 morphological species (Species Fungorum 2020).
Type species – Plagiostigme couraliae Syd.
Prostratus Sivan., W.H. Hsieh & Chi Y. Chen, Mycol. Res. 97(10): 1179 (1993)
Index Fungorum number: IF26496; 1 morphological species.
Type species – Prostratus cyclobalanopsidis Sivan., W.H. Hsieh & Chi Y. Chen
946
Pseudocryptosporella J. Reid & C. Booth, Can. J. Bot. 47: 1058 (1969)
Index Fungorum number: IF4423; 1 morphological species.
Type species – Pseudocryptosporella polylepidis (E. Müll.) J. Reid & C. Booth
Pseudothis Theiss. & Syd., Annls mycol. 12(3): 274 (1914)
Index Fungorum number: IF4494; 11 morphological species.
Type species – Pseudothis machaerii (Rehm) Theiss. & Syd.
Pseudovalsella Höhn., Annls mycol. 16(1/2): 123 (1918)
Index Fungorum number: IF4503; 2 morphological species (Species Fungorum 2020).
Type species – Pseudovalsella thelebola (Fr.) Höhn.
Savulescua Petr., cu Prilejul Implinirii a 70 de Ani (Bucuresti): 591 (1959)
Index Fungorum number: IF4871; 1 morphological species.
Type species – Savulescua insignis Petr.
Sphaerognomoniella Naumov & Kusnezowa, Bot. Mater. Otd. Sporov. Rast. Bot. Inst. Komarova
Akad. Nauk S.S.S.R. 8: 153 (1952)
Index Fungorum number: IF5101; 1 morphological species.
Type species – Sphaerognomoniella cornicola Naumov & Kusnezowa
Stioclettia Dennis, Kew Bull. 30(2): 362 (1975)
Index Fungorum number: IF5267; 1 morphological species.
Type species – Stioclettia luzulina Dennis
Trematovalsa Jacobesco, C. r. hebd. Séanc. Acad. Sci., Paris 142: 290 (1906)
Index Fungorum number: IF5526; 1 morphological species.
Type species – Trematovalsa matruchotii Iacobescu
Wehmeyera J. Reid & C. Booth, Can. J. Bot. 67(3): 895 (1989)
Index Fungorum number: IF25349; 1 morphological species.
Type species – Wehmeyera acerina (Wehm.) J. Reid & C. Booth
Glomerellales genera incertae sedis
Ascocodinaea Samuels, Cand. & Magni, Mycologia 89(1): 156 (1997)
Index Fungorum number: IF27833; 2 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Ascocodinaea stereicola Samuels, Cand. & Magni
Hypocreales genera incertae sedis
Acremoniopsis A. Giraldo, Gené & Guarro, Persoonia 33: 265 (2014)
Index Fungorum number: IF809883; 1 species with sequence data.
Type species – Acremoniopsis suttoniae A. Giraldo, Gené & Guarro
Berkelella (Sacc.) Sacc., Syll. fung. (Abellini) 9: 989 (1891)
Index Fungorum number: IF549; 3 morphological species (Species Fungorum 2020).
Type species – Berkelella caledonica (Pat.) Sacc.
Bulbithecium Udagawa & T. Muroi, Bull. natn. Sci. Mus., Tokyo, B 16(1): 13 (1990)
Index Fungorum number: IF25501; 1 species with sequence data.
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Type species – Bulbithecium hyalosporum Udagawa & T. Muroi
Cephalosporiopsis Peyronel, Mém. R. Accad. Sci. Torino, Ser. 2 66(no. 10): 52 (1916)
Index Fungorum number: IF7524; 7 morphological species (Species Fungorum 2020).
Type species – Cephalosporiopsis alpina (Peyronel) Peyronel
Chondronectria Etayo, Flakus & Kukwa, Opera Lilloana 50: 130 (2017)
Index Fungorum number: IF818310; 1 morphological species.
Type species – Chondronectria eriodermaticola Etayo, Flakus & Kukwa
Cylindronectria Etayo, Opera Lilloana 50: 154 (2017)
Index Fungorum number: IF818315; 1 morphological species
Type species – Cylindronectria cyanobacteriicola Etayo
Diploospora Grove, J. Bot., Lond. 54: 220 (1916)
Index Fungorum number: IF622352; 6 morphological species (Species Fungorum 2020), 2
species with sequence data.
Type species – Diploospora rosea Grove
Gynonectria Döbbeler, Mycol. Progr. 11(2): 474 (2012)
Index Fungorum number: IF582708; 1 morphological species.
Type species – Gynonectria intraspora Döbbeler
Hapsidospora Malloch & Cain, Can. J. Bot. 48(10): 1819 (1970)
Index Fungorum number: IF2243; 2 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Hapsidospora irregularis Malloch & Cain
Haptospora G.L. Barron, Can. J. Bot. 69(3): 503 (1991)
Index Fungorum number: IF25554; 3 morphological species (Species Fungorum 2020).
Type species – Haptospora appendiculata G.L. Barron
Illosporiopsis D. Hawksw., Mycol. Res. 105(4): 457 (2001)
Index Fungorum number: IF28491; 1 morphological species.
Type species – Illosporiopsis christiansenii (B.L. Brady & D. Hawksw.) D. Hawksw.
Illosporium Mart., Fl. crypt. erlang. (Nürnberg): 325 (1817)
Index Fungorum number: IF8626; 49 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Illosporium roseum Mart.
Leptobarya Etayo, Opera Lilloana 50: 214 (2017)
Index Fungorum number: IF818327; 2 morphological species (Species Fungorum 2020).
Type species – Leptobarya nigra Etayo
Lichenopenicillus Etayo, Opera Lilloana 50: 248 (2017)
Index Fungorum number: IF818339; 1 morphological species.
Type species – Lichenopenicillus versicolor Etayo
Metadothella Henn., Hedwigia 43(6): 384 (1904)
Index Fungorum number: IF3140; 1 morphological species.
Type species – Metadothella stellata Henn.
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Munkia Speg., Anal. Soc. cient. argent. 22(4): 202 (1886)
Index Fungorum number: IF9010; 4 morphological species (Species Fungorum 2020).
Type species – Munkia martyris Speg.
Neomunkia Petr., Sydowia 1(4-6): 329 (1947)
Index Fungorum number: IF9105; 1 species with sequence data.
Type species – Neomunkia sydowii Petr.
Peloronectria Möller, Bot. Mitt. Trop. 9: 297 (1901)
Index Fungorum number: IF3788; 3 morphological species (Species Fungorum 2020).
Type species – Peloronectria vinosa Möller
Pseudoacremonium Crous, Persoonia 32: 241 (2014)
Index Fungorum number: IF808925; 1 species with sequence data.
Type species – Pseudoacremonium sacchari Crous
Pseudoidriella Crous & R.G. Shivas .
Index Fungorum number: IF560696; 1 species with sequence data.
Type species – Pseudoidriella syzygii Crous & R.G. Shivas
Pseudomeliola Speg., Boln Acad. nac. Cienc. Córdoba 11(4): 522 (1889)
Index Fungorum number: IF4456; 10 morphological species (Species Fungorum 2020).
Type species – Pseudomeliola brasiliensis Speg.
Rodentomyces Doveri, Pecchia, Sarrocco & Vannacci, Index Fungorum 307: 7 (2016)
Index Fungorum number: IF552580; 1 species with sequence data.
Type species – Rodentomyces reticulatus Doveri, Pecchia, Sarrocco & Vannacci
Roselliniella Vain., Acta Soc. Fauna Flora fenn. 49(no. 2): 77 (1921)
Index Fungorum number: IF4786; 19 morphological species (Species Fungorum 2020), 2
species with sequence data.
Type species – Roselliniella nephromatis (P. Crouan & H. Crouan) Matzer & Hafellner
Saksenamyces A.N. Rai & P.N. Singh, Mycotaxon 133(3): 424 (2018)
Index Fungorum number: IF821284; 1 morphological species.
Type species – Saksenamyces kamalii Surywanshi, A.N. Rai & P.N. Singh
Sedecimiella K.L. Pang, Alias & E.B.G. Jones, Bot. Mar. 53(6): 495 (2010)
Index Fungorum number: IF518530; 1 species with sequence data.
Type species – Sedecimiella taiwanensis K.L. Pang
Stanjemonium W. Gams, O'Donnell, Schroers & M. Chr., Can. J. Bot. 76(9): 1579 (1999)
Index Fungorum number: IF27934; 4 morphological species (Species Fungorum 2020), 4
species with sequence data.
Type species – Stanjemonium grisellum W. Gams, Schroers & M. Chr.
Stilbella Lindau, Nat. Pflanzenfam., Teil. I (Leipzig) 1(1**): 489 (1900)
Index Fungorum number: IF10110; 68 morphological species (Species Fungorum 2020), 3
species with sequence data.
Type species – Stilbella fimetaria (Pers.) Lindau
Ticonectria Döbbeler, Nova Hedwigia 66(3-4): 362 (1998)
949
Index Fungorum number: IF27898; 3 morphological species (Species Fungorum 2020).
Type species – Ticonectria perianthii Döbbeler
Tilakidium Vaidya, C.D. Naik & Rathod, Indian J. Mycol. Plant Path. 16(2): 115 (1986)
Index Fungorum number: IF25210; 1 morphological species.
Type species – Tilakidium indicum Vaidya, C.D. Naik & Rathod
Microascales genera incertae sedis
Bisporostilbella Brandsb. & E.F. Morris, Mycologia 63(5): 1078 (1971)
Index Fungorum number: IF7379; 1 species with sequence data.
Type species – Bisporostilbella fusca Brandsb. & E.F. Morris
Cephalotrichiella Crous, Persoonia 32: 279 (2014)
Index Fungorum number: IF808954; 1 species with sequence data.
Type species – Cephalotrichiella penicillata Crous
Cornuvesica C.D. Viljoen, M.J. Wingf. & K. Jacobs, Mycol. Res. 104(3): 366 (2000)
Index Fungorum number: IF28384; 4 species with sequence data.
Type species – Cornuvesica falcata (E.F. Wright & Cain) C.D. Viljoen, M.J. Wingf. & K.
Jacobs
Gabarnaudia Samson & W. Gams, Stud. Mycol. 6: 88 (1974)
Index Fungorum number: IF8302; 2 species with sequence data.
Type species – Gabarnaudia betae (Delacr.) Samson & W. Gams
Sporendocladia G. Arnaud ex Nag Raj & W.B. Kendr., Monogr. Chalara Allied Genera
(Waterloo): 162 (1975)
Index Fungorum number: IF10020; 6 morphological species (Species Fungorum 2020), 3
species with sequence data.
Type species – Sporendocladia castaneae G. Arnaud ex Nag Raj & W.B. Kendr.
Phyllachorales genus incertae sedis
Marinosphaera K.D. Hyde, Can. J. Bot. 67(10): 3080 (1989)
Index Fungorum number: IF25372; 1 morphological species.
Type species – Marinosphaera mangrovei K.D. Hyde
Sordariales genera incertae sedis
Abyssomyces Kohlm., Ber. dt. bot. Ges. 83(9-10): 505 (1970)
Index Fungorum number: IF6; 1 morphological species.
Type species – Abyssomyces hydrozoicus Kohlm.
Acanthotheciella Höhn., Sber. Akad. Wiss. Wien, Math.-naturw. Kl., Abt. 1 120: 451 (1911)
Index Fungorum number: IF22; 3 morphological species (Species Fungorum 2020).
Type species – Acanthotheciella barbata (Pat.) Höhn.
Ascolacicola Ranghoo & K.D. Hyde, Mycologia 90(6): 1055 (1998)
Index Fungorum number: IF27878; 2 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Ascolacicola aquatica Ranghoo & K.D. Hyde
950
Bombardiella Höhn., Sber. Akad. Wiss. Wien, Math.-naturw. Kl., Abt. 1 118: 1192 (1909)
Index Fungorum number: IF618; 1 morphological species.
Type species – Bombardiella caespitosa Höhn.
Coronatomyces Dania García, Stchigel & Guarro, Stud. Mycol. 50(1): 144 (2004)
Index Fungorum number: IF500037; 1 species with sequence data.
Type species – Coronatomyces cubensis Dania García, Stchigel & Guarro
Cuspidatispora Shearer & Bartolata, Mycoscience 47(4): 220 (2006)
Index Fungorum number: IF29051; 1 species with sequence data.
Type species – Cuspidatispora xiphiago Shearer & Bartolata
Globosphaeria D. Hawksw., Lichenologist 22(3): 301 (1990)
Index Fungorum number: IF25444; 1 morphological species.
Type species – Globosphaeria jamesii D. Hawksw.
Isia D. Hawksw & Manohar., Trans. Br. mycol. Soc. 71(2): 332 (1978)
Index Fungorum number: IF2510; 2 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Isia neocaledoniensis (C. Moreau) D. Hawksw. & Manohar.
Lasiosphaeris Clem., Gen. fung. (Minneapolis): 35 (1909)
Index Fungorum number: IF2659; 3 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Lasiosphaeris hispida (Tode) Clem.
Lunulospora Ingold, Trans. Br. mycol. Soc. 25(4): 408 (1942)
Index Fungorum number: IF8795; 2 species with sequence data.
Type species – Lunulospora curvula Ingold
Lockerbia K.D. Hyde, Sydowia 46(1): 23 (1994)
Index Fungorum number: IF27284; 2 morphological species (Species Fungorum 2020).
Type species – Lockerbia palmicola K.D. Hyde
Nitschkiopsis Nannf. & R. Sant., Svensk bot. Tidskr. 69: 322 (1975)
Index Fungorum number: IF3516; 1 morphological species.
Type species – Nitschkiopsis stictarum Nannf. & R. Sant.
Onygenopsis Henn., Ann. Jard. Bot. Buitenzorg, Treub-Festschr., suppl. III 23: 64 (1910)
Index Fungorum number: IF3575; 1 morphological species.
Type species – Onygenopsis engleriana Henn.
Phaeosporis Clem., Gen. fung. (Minneapolis): 39 (1909)
Index Fungorum number: IF3954; 4 morphological species (Species Fungorum 2020).
Type species – Phaeosporis melasperma (Nyl.) Clem.
Reconditella Matzer & Hafellner, Biblthca Lichenol. 37: 46 (1990)
Index Fungorum number: IF25462; 1 morphological species.
Type species – Reconditella physconiarum Hafellner & Matzer
Rhexodenticula W.A. Baker & Morgan-Jones, Mycotaxon 79: 363 (2001)
951
Index Fungorum number: IF28519; 4 morphological species (Species Fungorum 2020), 2
species with sequence data.
Type species – Rhexodenticula cylindrospora (R.F. Castañeda, Saikawa & Hennebert) W.A.
Baker & Morgan-Jones
Rhexosporium Udagawa & Furuya, Trans. Mycol. Soc. Japan 18: 302 (1977)
Index Fungorum number: IF4698; 1 morphological species.
Type species – Rhexosporium terrestre Udagawa & Furuya
Roselliniomyces Matzer & Hafellner, Biblthca Lichenol. 37: 91 (1990)
Index Fungorum number: IF25464; 2 morphological species (Species Fungorum 2020).
Type species – Roselliniomyces trichotheliorum Matzer & Hafellner
Roselliniopsis Matzer & Hafellner, Biblthca Lichenol. 37: 97 (1990)
Index Fungorum number: IF25465; 7 morphological species (Species Fungorum 2020).
Type species – Roselliniopsis groedensis (Zopf) Matzer & Hafellner
Stromatographium Höhn., Denkschr. Kaiserl. Akad. Wiss., Math.-Naturwiss. Kl. 83: 37 (1907)
Index Fungorum number: IF10129; 2 morphological species (Species Fungorum 2020).
Type species – Stromatographium stromaticum (Berk.) Höhn.
Utriascus Réblová, Mycologia 95(1): 128 (2003)
Index Fungorum number: IF28700; 1 morphological species.
Type species – Utriascus gabretae Réblová
Ypsilonia Lév., Annls Sci. Nat., Bot., sér. 3 5: 284 (1846)
Index Fungorum number: IF10464; 3 morphological species (Species Fungorum 2020).
Type species – Ypsilonia cuspidata Lév.
Xylariales genera incertae sedis
Adomia S. Schatz, Trans. Br. mycol. Soc. 84(3): 555 (1985)
Index Fungorum number: IF82; 1 morphological species.
Type species – Adomia avicenniae S. Schatz
Alloanthostomella Daranag., Camporesi & K.D. Hyde, Cryptog. Mycol. 37(4): 518 (2016)
Index Fungorum number: IF552371; 1 species with sequence data.
Type species – Alloanthostomella rubicola (Speg. ex Sacc. & Trotter) Daranag., Camporesi
& K.D. Hyde
Anungitea B. Sutton, Mycol. Pap. 132: 10 (1973)
Index Fungorum number: IF7181; 20 morphological species (Species Fungorum 2020), 6
species with sequence data.
Type species – Anungitea fragilis B. Sutton
Ascotrichella Valldos. & Guarro, Trans. Br. mycol. Soc. 90(4): 601 (1988)
Index Fungorum number: IF25217; 1 morphological species.
Type species – Ascotrichella hawksworthii Valldos. & Guarro
Basifimbria Subram. & Lodha, Curr. Sci. 37: 247 (1968)
Index Fungorum number: IF7338; 1 morphological species.
Type species – Basifimbria aurea Subram. & Lodha
952
Biporispora J.D. Rogers, Y.M. Ju & Cand., Nova Hedwigia 68(3-4): 421 (1999)
Index Fungorum number: IF28322; 1 morphological species.
Type species – Biporispora europaea J.D. Rogers, Y.M. Ju & Cand.
Castellaniomyces Senan., Camporesi & K.D. Hyde, Fungal Divers. 87: 183 (2017)
Index Fungorum number: IF553639; 1 species with sequence data.
Type species – Castellaniomyces rosae Senan., & K.D. Hyde
Chaenocarpus Rebent., Prodr. fl. neomarch. (Berolini): 350 (1804)
Index Fungorum number: IF930; 3 morphological species (Species Fungorum 2020).
Type species – Chaenocarpus setosus Rebent., Prodr. fl. neomarch. (Berolini): 350, tab.
3:12a-b
Circinotrichum Nees, Syst. Pilze (Würzburg): 19 (1816)
Index Fungorum number: IF7667; 16 morphological species (Species Fungorum 2020), 4
species with sequence data.
Type species – Circinotrichum maculiforme Nees
Cryptostroma P.H. Greg. & S. Waller, Trans. Br. mycol. Soc. 34(4): 593 (1952)
Index Fungorum number: IF7839; 1 species with sequence data.
Type species – Cryptostroma corticale (Ellis & Everh.) P.H. Greg. & S. Waller
Cyanopulvis J. Fröhl. & K.D. Hyde, Fungal Divers. Res. Ser. 3: 308 (2000)
Index Fungorum number: IF28469; 1 morphological species.
Type species – Cyanopulvis australiensis J. Fröhl. & K.D. Hyde
Diamantinia A.N. Mill., Læssøe & Huhndorf, Sydowia 55(1): 94 (2003)
Index Fungorum number: IF28693; 1 species with sequence data.
Type species – Diamantinia citrina A.N. Mill., Læssøe & Huhndorf
Gigantospora B.S. Lu & K.D. Hyde, Nova Hedwigia 76(1-2): 202 (2003)
Index Fungorum number: IF28792; 1 morphological species.
Type species – Gigantospora gigaspora B.S. Lu & K.D. Hyde
Guestia G.J.D. Sm. & K.D. Hyde, Fungal Divers. 7: 107 (2001)
Index Fungorum number: IF28525; 1 morphological species.
Type species – Guestia gonetropospora G.J.D. Sm. & K.D. Hyde
Gyrothrix (Corda) Corda, Anleit. Stud. Mykol., Prag: LXIII, 49 (1842)
Index Fungorum number: IF8414; 22 morphological species (Species Fungorum 2020), 6
species with sequence data.
Type species – Gyrothrix podosperma (Corda) Rabenh
Hadrotrichum Fuckel, Fungi rhenani exsic., suppl., fasc.: no. 1522 (1865)
Index Fungorum number: IF8419; 15 morphological species (Species Fungorum 2020), 2
species with sequence data.
Type species – Hadrotrichum phragmitis Fuckel
Idriellopsis Hern.-Restr. & Crous, Persoonia 36: 76 (2015)
Index Fungorum number: IF811882; 1 species with sequence data.
Type species – Idriellopsis uncinospora (R.F. Castañeda & W.B. Kendr.) Hern.-Restr. &
Crous
953
Kirstenboschia Quaedvl., Verkley & Crous, Stud. Mycol. 75: 385 (2013)
Index Fungorum number: IF804467; 1 species with sequence data.
Type species – Kirstenboschia diospyri Quaedvl., Verkley & Crous
Lanceispora Nakagiri, Okane, Tad. Ito & Katum., Mycoscience 38(2): 208 (1997)
Index Fungorum number: IF27737; 2 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Lanceispora amphibia Nakagiri, Okane, Tad. Ito & Katum.
Lasiobertia Sivan., Trans. Br. mycol. Soc. 70(3): 383 (1978)
Index Fungorum number: IF2647; 2 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Lanceispora amphibia Nakagiri, Okane, Tad. Ito & Katum.
Leptomassaria Petr., Annls mycol. 12(5): 474 (1914)
Index Fungorum number: IF2779; 4 morphological species.
Type species – Leptomassaria simplex (Nitschke ex G.H. Otth) Petr.
Neoanthostomella D.Q. Dai & K.D. Hyde, Fungal Divers. 82(1), 1–105 (2017)
Index Fungorum number: IF552041; 2 species with sequence data.
Type species – Neoanthostomella pseudostromatica D.Q. Dai & K.D. Hyde
Neoidriella Hern.-Restr. & Crous, Persoonia 36: 78 (2015)
Index Fungorum number: IF811884; 1 species with sequence data.
Type species – Neoidriella desertorum (Nicot & Mouch.) M. Hern.-Restr. & Crous
Nipicola K.D. Hyde, Cryptog. Bot. 2(4): 330 (1992)
Index Fungorum number: IF26307; 4 morphological species (Species Fungorum 2020).
Type species – Nipicola carbospora K.D. Hyde
Occultitheca J.D. Rogers & Y.M. Ju, Sydowia 55(2): 359 (2003)
Index Fungorum number: IF26307; 1 morphological species.
Type species – Occultitheca costaricensis J.D. Rogers & Y.M. Ju
Ophiorosellinia J.D. Rogers, A. Hidalgo, F.A. Fernández & Huhndorf, Mycologia 96(1): 172
(2004)
Index Fungorum number: IF28822; 1 morphological species.
Type species – Ophiorosellinia costaricensis J.D. Rogers, A. Hidalgo, F.A. Fernández &
Huhndorf
Palmicola K.D. Hyde, Sydowia 45(1): 15 (1993)
Index Fungorum number: IF22434; 4 morphological species (Species Fungorum 2020).
Type species – Palmicola archontophoenicis K.D. Hyde
Pandanicola K.D. Hyde, Sydowia 46(1): 35 (1994)
Index Fungorum number: IF22419; 2 morphological species (Species Fungorum 2020).
Type species – Pandanicola calocarpa (Syd. & P. Syd.) K.D. Hyde
Paraidriella Hern.-Restr. & Crous, Persoonia 37: 79 (2015)
Index Fungorum number: IF811886; 1 species with sequence data.
Type species – Paraidriella jambosae (R.F. Castañeda & W.B. Kendr.) M. Hern.-Restr. &
Crous
954
Paramphisphaeria F.A. Fernández, J.D. Rogers, Y.M. Ju, Huhndorf & L. Umaña, Mycologia
96(1): 175 (2004)
Index Fungorum number: IF28823; 1 morphological species.
Type species – Paramphisphaeria costaricensis F.A. Fernández, J.D. Rogers, Y.M. Ju,
Huhndorf & Umaña
Paraphysalospora Crous, Persoonia 39: 373 (2017)
Index Fungorum number: IF823380; 1 species with sequence data.
Type species – Paraphysalospora eucalypti Crous
Paucithecium Lloyd, Mycol. Notes (Cincinnati) 7(4): 1200 (1923)
Index Fungorum number: IF3774; 1 morphological species.
Type species – Paucithecium rickii Lloyd
Pidoplitchkoviella Kiril., Mikrobiol. Zh. 37(5): 603 (1975)
Index Fungorum number: IF4097; 1 species with sequence data.
Type species – Pidoplitchkoviella terricola Kiril.
Polyancora Voglmayr & Yule, Mycol. Res. 110(10): 1247 (2006)
Index Fungorum number: IF500689; 1 species with sequence data.
Type species – Polyancora globosa Voglmayr & Yule
Polyscytalum Riess, Bot. Ztg. 11: 138 (1853)
Index Fungorum number: IF9508; 21 morphological species, 4 species with sequence data.
Type species – Polyscytalum fecundissimum Riess
Poroleprieuria M.C. González, Hanlin, Ulloa & Elv. Aguirre, Mycologia 96(3): 676 (2004)
Index Fungorum number: IF28828; 1 morphological species.
Type species – Poroleprieuria rogersii M.C. González, Hanlin, Ulloa & Elv. Aguirre
Pseudoanthostomella Daranag., Camporesi & K.D. Hyde, Cryptog. Mycol. 37(4): 527 (2016)
Index Fungorum number: IF552377; 4 species with sequence data.
Type species – Pseudoanthostomella pini-nigrae Daranag., Camporesi & K.D. Hyde
Pseudophloeospora Crous & R.G. Shivas, Persoonia 25: 141 (2010)
Index Fungorum number: IF517539; 4 species with sequence data.
Type species – Pseudophloeospora eucalypti Crous & R.G. Shivas
Pulmosphaeria Joanne E. Taylor, K.D. Hyde & E.B.G. Jones, Sydowia 48(2): 256 (1996)
Index Fungorum number: IF27728; 1 morphological species.
Type species – Pulmosphaeria archontophoenicis Joanne E. Taylor, K.D. Hyde & E.B.G.
Jones
Pyriformiascoma Daranag., Camporesi & K.D. Hyde, Fungal Divers. 73: 230 (2015)
Index Fungorum number: IF809526; 1 species with sequence data.
Type species – Pyriformiascoma trilobatum Daranag., Camporesi & K.D. Hyde
Roselymyces Fiuza, C.R. Silva, R.F. Castañeda & Gusmão, Sydowia 70: 60 (2018)
Index Fungorum number: IF822003; 1 morphological species.
Type species – Roselymyces brasiliensis Fiuza, C.R. Silva, R.F. Castañeda & Gusmão
Sabalicola K.D. Hyde, Nova Hedwigia 60(3–4): 596 (1995)
955
Index Fungorum number: IF27636; 1 morphological species.
Type species – Sabalicola sabalensioides (Ellis & G. Martin) K.D. Hyde
Spirodecospora B.S. Lu, K.D. Hyde & W.H. Ho, Fungal Divers. Res. Ser. 1: 170 (1998)
Index Fungorum number: IF27893; 2 morphological species (Species Fungorum 2020).
Type species – Spirodecospora bambusicola B.S. Lu, K.D. Hyde & W.H. Ho
Sporidesmina Subram. & Bhat, Kavaka 15(1-2): 69 (1989)
Index Fungorum number: IF11053; 1 species with sequence data.
Type species – Sporidesmina malabarica Subram. & Bhat
Striatodecospora D.Q. Zhou, K.D. Hyde & B.S. Lu, Mycotaxon 76: 142 (2000)
Index Fungorum number: IF28472; 1 morphological species.
Type species – Striatodecospora bambusae D.Q. Zhou, K.D. Hyde & B.S. Lu
Stromatoneurospora S.C. Jong & E.E. Davis, Mycologia 65(2): 459 (1973)
Index Fungorum number: IF5289; 2 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Stromatoneurospora phoenix (Kunze) S.C. Jong & E.E. Davis
Surculiseries Okane, Nakagiri & Tad. Ito, Mycoscience 42(1): 116 (2001)
Index Fungorum number: IF28496; 1 species with sequence data.
Type species – Surculiseries rugispora Okane, Nakagiri & Tad. Ito
Synnemadiella Crous & M.J. Wingf., Persoonia 37: 339 (2016)
Index Fungorum number: IF819088; 1 species with sequence data.
Type species – Synnemadiella eucalypti Crous & M.J. Wingf.
Tristratiperidium Daranag., Camporesi & K.D. Hyde, Mycol. Progr. 15(no. 8): 4 (2015)
Index Fungorum number: IF551386; 1 species with sequence data.
Type species – Tristratiperidium microsporum Daranag., Camporesi & K.D.Hyde
Xylocrea Möller, Bot. Mitt. Trop. 9: 307 (1901)
Index Fungorum number: IF5840; 2 morphological species.
Type species – Xylocrea piriformis Möller
Xylotumulus J.D. Rogers, Y.M. Ju & Hemmes, Sydowia 58(2): 291 (2006)
Index Fungorum number: IF522310; 1 species with sequence data.
Type species – Xylotumulus gibbisporus J.D. Rogers, Y.M. Ju & Hemmes
Yuea O.E. Erikss., Mycotaxon 85: 314 (2003)
Index Fungorum number: IF28721; 1 morphological species.
Type species – Yuea chusqueicola O.E. Erikss.
Placement of genera in Subclasses incertae sedis
Diaporthomycetidae genera incertae sedis
Aquapteridospora Jiao Yang, K.D. Hyde & Maharachch., Cryptog. Mycol. 36(4): 474 (2015)
Index Fungorum number: IF551731; 2 species with sequence data.
Type species – Aquapteridospora lignicola J. Yang, K.D. Hyde & Maharachch.
956
Aquaticola W.H. Ho, C.K.M. Tsui, Hodgkiss & K.D. Hyde, Fungal Divers. Res. Ser. 3: 88 (1999)
Index Fungorum number: IF28389; 4 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Aquaticola hyalomura W.H. Ho, K.M. Tsui, Hodgkiss & K.D. Hyde
Fusoidispora Vijaykr., Jeewon & K.D. Hyde, Sydowia 57(2): 272 (2005)
Index Fungorum number: IF28975; 1 species with sequence data.
Type species – Fusoidispora aquatica Vijaykr., Jeewon & K.D. Hyde
Platytrachelon Réblová, Mycologia 105(2): 466 (2013)
Index Fungorum number: IF800340; 1 species with sequence data.
Type species – Platytrachelon abietis (Réblová) Réblová
Proliferophorum G.N. Wang, H. Zhang & Senan., Fungal Divers. 9: 128 (2019)
Index Fungorum number: IF555401; 1 species with sequence data.
Type species – Proliferophorum thailandicum G.N. Wang, H. Zhang & Senan.
Pseudostanjehughesia J. Yang & K.D. Hyde, Mycol. Progr. 17(5): 609 (2017)
Index Fungorum number: IF821274; 2 species with sequence data.
Type species – Pseudostanjehughesia aquitropica J. Yang & K.D. Hyde
Hypocreomycetidae genera incertae sedis
Campylospora Ranzoni, Farlowia 4(3): 373 (1953)
Index Fungorum number: IF7482; 5 morphological species (Species Fungorum 2020), 5
species with sequence data.
Type species – Campylospora chaetocladia Ranzoni
Dendroclathra Voglmayr & G. Delgado, Can. J. Bot. 79(9): 995 (2001)
Index Fungorum number: IF28541; 2 species with sequence data.
Type species – Dendroclathra caeruleofusca Voglmayr & G. Delgado
Sordariomycetidae genera incertae sedis
Arecacicola Joanne E. Taylor, J. Fröhl. & K.D. Hyde, Mycoscience 42(4): 370 (2001)
Index Fungorum number: IF28577; 1 morphological species.
Type species – Arecacicola calami Joanne E. Taylor, J. Fröhl. & K.D. Hyde
Bullimyces A. Ferrer, A.N. Mill., Sarmiento & Shearer, Mycologia 104(4): 868 (2012)
Index Fungorum number: IF561094; 3 species with sequence data.
Type species – Bullimyces communis A. Ferrer, A.N. Mill., Sarmiento & Shearer
Cancellidium Tubaki, Trans. Mycol. Soc. Japan 16(4): 357 (1975)
Index Fungorum number: IF7483; 2 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Cancellidium applanatum Tubaki
Ceratolenta Réblová, Mycologia 105(2): 466 (2013)
Index Fungorum number: IF800338; 1 species with sequence data.
Type species – Ceratolenta caudata Réblová
Chaetosphaerides Matsush., Matsush. Mycol. Mem. 10: 146 (2003)
957
Index Fungorum number: IF28561; 1 morphological species.
Type species – Chaetosphaerides ramichloridifera Matsush.
Cryptophyllachora L. Kiss, Kovács & R.G. Shivas, Scientific Reports 8(no. 10806): 4 (2018)
Index Fungorum number: IF825649; 2 species with sequence data.
Type species – Cryptophyllachora eurasiatica L. Kiss, Kovács & R.G. Shivas
Hanliniomyces Raja & Shearer, Mycologia 100(3): 471 (2008)
Index Fungorum number: IF507448; 1 morphological species.
Type species – Hanliniomyces hyaloapicalis Raja & Shearer
Hydromelitis A. Ferrer, A.N. Mill., Sarmiento & Shearer, Mycologia 104(4): 876 (2012)
Index Fungorum number: IF561101; 1 species with sequence data.
Type species – Hydromelitis pulchella A. Ferrer, A.N. Mill., Sarmiento & Shearer
Merugia Rogerson & Samuels, Mem. N. Y. bot. Gdn 64: 166 (1990)
Index Fungorum number: IF25540; 1 morphological species.
Type species – Merugia palicoureae Rogerson & Samuels
Mycomedusiospora G.C. Carroll & Munk, Mycologia 56(1): 91 (1964)
Index Fungorum number: IF3325; 1 morphological species.
Type species – Mycomedusiospora flavida (Rick) G.C. Carroll & Munk
Myxocephala G. Weber, Spaaij & Oberw., Sydowia 41: 360 (1989)
Index Fungorum number: IF11231; 1 species with sequence data.
Type species – Myxocephala albida G. Weber, Spaaij & Oberw.
Nigromammilla K.D. Hyde & J. Fröhl., Cryptog. Mycol. 24(1): 17 (2003)
Index Fungorum number: IF28740; 1 morphological species.
Type species – Nigromammilla calami K.D. Hyde & J. Fröhl.
Phaeotrichosphaeria Sivan., Trans. Br. mycol. Soc. 81(2): 313 (1983)
Index Fungorum number: IF25837; 4 morphological species (Species Fungorum 2020).
Type species – Phaeotrichosphaeria indica Sivan. & N.D. Sharma
Phragmodiscus Hansf., Proc. Linn. Soc. London 159: 31 (1947)
Index Fungorum number: IF4026; 2 morphological species (Species Fungorum 2020).
Type species – Phragmodiscus arundinariae Hansf.
Plagiosphaera Petr., Annls mycol. 39(4/6): 289 (1941)
Index Fungorum number: IF4155; 11 morphological species.
Type species – Plagiosphaera moravica (Petr.) Petr.
Pseudobotrytis Krzemien. & Badura, Acta Soc. Bot. Pol. 23: 761 (1954)
Index Fungorum number: IF9556; 2 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Pseudobotrytis fusca Krzemien. & Badura
Xylariomycetidae genera incertae sedis
Calceomyces Udagawa & S. Ueda, Mycotaxon 32: 448 (1988)
Index Fungorum number: IF25219; 1 species with sequence data.
958
Type species – Calceomyces lacunosus Udagawa & S. Ueda
Placement of genera in Sordariomycetes genera incertae sedis
Acerbiella Sacc. & D. Sacc., Syll. fung. (Abellini) 17: 768 (1905)
Index Fungorum number: IF33; 4 morphological species (Species Fungorum 2020)
Type species – Acerbiella macrospora (Rick) Sacc. & D. Sacc.
Acrospermoides J.H. Mill. & G.E. Thomps., Mycologia 32(1): 12 (1940)
Index Fungorum number: IF53; 2 morphological species (Species Fungorum 2020)
Type species – Acrospermoides subulata J.H. Mill. & G.E. Thomps.
Ameromassaria Hara, Journal of Plant Protection, Tokyo 5: 886 (1918)
Index Fungorum number: IF159; 1 morphological species
Type species – Ameromassaria japonica Hara
Amphisphaerellula Gucevič, Bot. Mater. Otd. Sporov. Rast. Bot. Inst. Komarova Akad. Nauk
S.S.S.R. 8: 142 (1952)
Index Fungorum number: IF172; 2 morphological species (Species Fungorum 2020)
Type species – Amphisphaerellula fagi Gucevič
Amphisphaerina Höhn., Sber. Akad. Wiss. Wien, Math.-naturw. Kl., Abt. 1 128(7–8): 581 (1919)
Index Fungorum number: IF174; 1 morphological species
Type species – Amphisphaerina texensis (Cooke) Höhnel
Amphorulopsis Petr., Sydowia 13(1–6): 181 (1959)
Index Fungorum number: IF178; 1 morphological species
Type species – Amphorulopsis polygonacearum Petr.
Amylis Speg., Fung. parag.: 405 (1922)
Index Fungorum number: IF184; 1 morphological species
Type species – Amylis memorabilis Speg.
Anisomycopsis I. Hino & Katum., J. Jap. Bot. 39: 325 (1964)
Index Fungorum number: IF203; 1 morphological species
Type species – Anisomycopsis rosae I. Hino & Katum.
Anthostomaria (Sacc.) Theiss. & Syd., Annls mycol. 16(1/2): 27 (1918)
Index Fungorum number: IF225; 1 morphological species
Type species – Anthostomaria apogyra (Nyl.) Theiss. & Syd.
Anthostomellina L.A. Kantsch., Bolêz. Rast. 17: 82 (1928)
Index Fungorum number: IF227; 2 morphological species (Species Fungorum 2020)
Type species – Anthostomellina carpinea L.A. Kantsch.
Apodothina Petr., Sydowia 23(1–6): 276 (1970)
Index Fungorum number: IF276; 1 morphological species
Type species – Apodothina pringlei (Peck) Petr.
Apogaeumannomyces Matsush., Matsush. Mycol. Mem. 10: 152 (2003)
Index Fungorum number: IF28562; 1 morphological species
Type species – Apogaeumannomyces perplexus Matsush.
959
Aquadulciospora Fallah & Shearer, Mycologia 93(3): 570 (2001)
Index Fungorum number: IF28486; 1 morphological species
Type species – Aquadulciospora rhomboidia Fallah & Shearer
Aropsiclus Kohlm. & Volkm.-Kohlm., Syst. Ascom. 13(1): 24 (1994)
Index Fungorum number: IF6185; 1 morphological species
Type species – Aropsiclus junci (Kohlm. & Volkm.-Kohlm.) Kohlm. & Volkm.-Kohlm.
Ascorhiza Lecht.-Trinka, C. r. hebd. Séanc. Acad. Sci., Paris 192: 499 (1931)
Index Fungorum number: IF372; 1 morphological species
Type species – Ascorhiza leguminosarum Lecht.-Trinka
Ascoyunnania L. Cai & K.D. Hyde, Fungal Divers. 18: 2 (2005)
Index Fungorum number: IF28946; 1 morphological species
Type species – Ascoyunnania aquatica L. Cai & K.D. Hyde
Atrogeniculata J.S. Monteiro, Gusmão & R.F. Castañeda, Mycotaxon 127: 40 (2014)
Index Fungorum number: IF805084; 1 morphological species
Type species – Atrogeniculata submersa J. S. Monteiro, Gusmão & R. F. Castañeda
Aulospora Speg., Anal. Mus. nac. B. Aires, Ser. 3 12: 404 (1909)
Index Fungorum number: IF25578; 1 morphological species
Type species – Aulospora epimyces (Speg.) Speg.
Azbukinia Lar.N. Vassiljeva, Mycotaxon 35(2): 395 (1989)
Index Fungorum number: IF25350; 1 morphological species
Type species – Azbukinia ferruginea (Fuckel) Lar.N. Vassiljeva
Bactrosphaeria Penz. & Sacc., Malpighia 11(9–10): 407 (1897)
Index Fungorum number: IF482; 1 morphological species
Type species – Bactrosphaeria asterostoma Penz. & Sacc.
Basidiobotrys Höhn., Sber. Akad. Wiss. Wien, Math.-naturw. Kl., Abt. 1 118: 420 (1909)
Index Fungorum number: IF7336; 2 morphological species (Species Fungorum 2020)
Type species – Basidiobotrys clautriavii (Pat.) Höhn.
Biciliopsis Diederich, Biblthca Lichenol. 64: 25 (1997)
Index Fungorum number: IF27753; 2 morphological species (Species Fungorum 2020)
Type species – Biciliopsis leptogiicola Diederich
Bombardiastrum Pat., Bull. Soc. mycol. Fr. 9(3): 153 (1893)
Index Fungorum number: IF617; 1 morphological species
Type species – Bombardiastrum andinum Pat.
Botryosporium Corda, Deutschl. Fl., 3 Abt. (Pilze Deutschl.) 3(11): 9 (1831)
Index Fungorum number: IF7429; 10 morphological species (Species Fungorum 2020), 3
species with sequence data.
Type species – Botryosporium diffusum Corda
Brenesiella Syd., Annls mycol. 27(1/2): 16 (1929)
Index Fungorum number: IF649; 1 morphological species
960
Type species – Brenesiella erythroxyli Syd.
Byrsomyces Cavalc., Instituto de Micologia da Universidade de Pernambuco 675: 6 (1972)
Index Fungorum number: IF696; 1 morphological species
Type species – Byrsomyces olivaceus Cavalc.
Byssotheciella Petr., Annls mycol. 21(3/4): 281 (1923)
Index Fungorum number: IF713; 2 morphological species (Species Fungorum 2020)
Type species – Byssotheciella tiliae Petr.
Caleutypa Petr., Annls mycol. 32(5/6): 434 (1934)
Index Fungorum number: IF726; 1 morphological species
Type species – Caleutypa maculans Petr.
Caproniella Berl., Icon. fung. (Abellini) 2(2–3): 62 (1896)
Index Fungorum number: IF817; 1 morphological species
Type species – Caproniella pleiospora (Mouton) Berl.
Chaetoamphisphaeria Hara, Journal of Plant Protection, Tokyo 5: 349 (1918)
Index Fungorum number: IF939; 1 morphological species
Type species – Chaetoamphisphaeria japonica Hara
Ciliofusospora Bat. & J.L. Bezerra, Publicações Inst. Micol. Recife 385: 15 (1963)
Index Fungorum number: IF1055; 1 morphological species
Type species – Ciliofusospora oenocarpi Bat. & J.L. Bezerra
Clypeoceriospora Sousa da Câmara, Agron. lusit. 8: 30 (1946)
Index Fungorum number: IF1113; 1 morphological species
Type species – Clypeoceriospora rubi Sousa da Câmara
Clypeosphaerulina Sousa da Câmara, Agron. lusit. 1: 195 (1939)
Index Fungorum number: IF1126; 1 morphological species
Type species – Clypeosphaerulina vincae (Sousa da Câmara) Sousa da Câmara
Cryptoascus Petri, Atti Reale Accad. Lincei, Rendic., Sér. 5 18(2): 642 (1909)
Index Fungorum number: IF1311; 2 morphological species (Species Fungorum 2020)
Type species – Cryptoascus oligosporus Petri
Cryptomycella Höhn., Mitt. bot. Inst. tech. Hochsch. Wien 2(3): 48 (1925)
Index Fungorum number: IF7831; 2 morphological species (Species Fungorum 2020)
Type species – Cryptomycella pteridis (Kalchbr.) Höhn.
Cryptomycina Höhn., Annls mycol. 15(5): 322 (1917)
Index Fungorum number: IF1321; 2 morphological species (Species Fungorum 2020)
Type species – Cryptomycina pteridis (Rebent.) Höhn.
Cucurbitopsis Bat. & Cif., Publicações Inst. Micol. Recife 95: 3 (1957)
Index Fungorum number: IF1351; 1 morphological species
Type species – Cucurbitopsis ramulorum Bat., Nascim. & Cif.
Curvatispora V.V. Sarma & K.D. Hyde, Nova Hedwigia 72(3–4): 480 (2001)
Index Fungorum number: IF28530; 1 morphological species
961
Type species – Curvatispora singaporensis V.V. Sarma & K.D. Hyde
Dasysphaeria Speg., Anal. Mus. nac. Hist. nat. B. Aires 23: 60 (1912)
Index Fungorum number: IF1421; 1 morphological species
Type species – Dasysphaeria andicola Speg.
Delpinoëlla Sacc., Bull. Soc. R. Bot. Belg. 38(2): 162 (1899)
Index Fungorum number: IF1446; 1 morphological species
Type species – Delpinoëlla insignis Sacc. & Trotter
Diacrochordon Petr., Sydowia 9(1–6): 591 (1955)
Index Fungorum number: IF7973; 1 morphological species
Type species – Diacrochordon rehmii Petr.
Didymobotryum Sacc., Syll. fung. (Abellini) 4: 626 (1886)
Index Fungorum number: IF8009; 6 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Didymobotryum rigidum (Berk. & Broome) Sacc.
Duradens Samuels & Rogerson, Mem. N. Y. bot. Gdn 64: 170 (1990)
Index Fungorum number: IF25541; 1 morphological species
Type species – Duradens lignicola Samuels & Rogerson
Esfandiariomyces Ershad, Iranian Journal of Plant Pathology 21(1–4): 8 (1985)
Index Fungorum number: IF25103; 1 morphological species
Type species – Esfandiariomyces insignis (Petr.) Ershad
Fantasmomyces D.Hyeon Lee, Marinc., Z.W. de Beer & M.J. Wingf., Persoonia 37: 249 (2016)
Index Fungorum number: IF816620; 1 species with sequence data.
Type species – Fantasmomyces hyalinus Dong Hyeon Lee, Marinc., Z.W. de Beer, M.J.
Wingf.
Fassia Dennis, Bull. Jard. bot. État Brux. 34: 240 (1964)
Index Fungorum number: IF1979; 1 morphological species
Type species – Fassia scabrosa Dennis
Flammispora Pinruan, Sakay., K.D. Hyde & E.B.G. Jones, Stud. Mycol. 50(2): 384 (2004)
Index Fungorum number: IF500093; 2 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Flammispora bioteca Pinruan, Sakayaroj, K.D. Hyde & E.B.G. Jones
Frondisphaeria K.D. Hyde, Mycoscience 37(2): 169 (1996)
Index Fungorum number: IF27664; 2 morphological species (Species Fungorum 2020)
Type species – Frondisphaeria palmicola K.D. Hyde
Hapsidascus Kohlm. & Volkm.-Kohlm., Syst. Ascom. 10: 113 (1991)
Index Fungorum number: IF25237; 1 morphological species
Type species – Hapsidascus hadrus Kohlm. & Volkm.-Kohlm.
Heliastrum Petr., Annls mycol. 29(3/4): 190 (1931)
Index Fungorum number: IF2256; 1 morphological species
Type species – Heliastrum concinnum Petr.
962
Hyaloderma Speg., Anal. Soc. cient. argent. 17(3): 131 (1884)
Index Fungorum number: IF2385; 1 morphological species
Type species – Hyaloderma imperspicuum Speg.
Hydronectria Kirschst., Verh. bot. Ver. Prov. Brandenb. 67: 87 (1925)
Index Fungorum number: IF2405; 1 morphological species
Type species – Hydronectria kriegeriana Kirschst.
Immersisphaeria Jaklitsch, Mycotaxon 101: 18 (2007)
Index Fungorum number: IF510734; 1 morphological species
Type species – Immersisphaeria eichleriana (Bres.) Jaklitsch
Iraniella Petr., Sydowia 3(1-6): 135 (1949)
Index Fungorum number: IF2502; 1 morphological species
Type species – Iraniella rechingeri Petr.
Konenia Hara, Bot. Mag., Tokyo 27(no. 317): 250 (1913)
Index Fungorum number: IF2580; 2 morphological species (Species Fungorum 2020)
Type species – Konenia bambusae Hara
Kravtzevia Schwarzman, Trudy Inst. Bot., Alma-Ata 9: 75 (1961)
Index Fungorum number: IF2587; 1 morphological species
Type species – Kravtzevia halimodendri Schwarzman
Kurssanovia Kravtzev, Trudy Inst. Bot., Alma-Ata 2: 145 (1955)
Index Fungorum number: IF2599; 1 morphological species
Type species – Kurssanovia vassiljevskii Kravtzev
Lecythiomyces Doweld, Index Fungorum 30: 1 (2013)
Index Fungorum number: IF550264; 1 morphological species
Type species – Lecythiomyces aerugineus (Zukal) Doweld
Leptosacca Syd., Annls mycol. 26(1/2): 109 (1928)
Index Fungorum number: IF2796; 1 morphological species
Type species – Leptosacca lumae Syd.
Leptosphaerella Speg., Anal. Mus. nac. Hist. nat. B. Aires 23: 56 (1912)
Index Fungorum number: IF92186; 7 morphological species (Species Fungorum 2020)
Type species – Leptosphaerella fagaricola Speg.
Mangrovispora K.D. Hyde & Nakagiri, Syst. Ascom., Reprint of Volumes 1–4 (1982–1985)
10(1): 19 (1991)
Index Fungorum number: IF25564; 2 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Mangrovispora pemphii K.D. Hyde & Nakagiri
Marisolaris Jørg. Koch & E.B.G. Jones, Can. J. Bot. 67(4): 1190 (1989)
Index Fungorum number: IF25320; 1 morphological species
Type species – Marisolaris ansata Jørg. Koch & E.B.G. Jones
Melanographium Sacc., Annls mycol. 11(6): 557 (1913)
963
Index Fungorum number: IF8891; 13 morphological species (Species Fungorum 2020)
Type species – Melanographium spleniosporum Sacc.
Microcyclephaeria Bat., Revta Biol., Lisb. 1(3–4): 301 (1958)
Index Fungorum number: IF3159; 1 morphological species
Type species – Microcyclephaeria palmicola (Syd.) Bat. & H. Maia
Mirannulata Huhndorf, F.A. Fernández, A.N. Mill. & Lodge, Sydowia 55(2): 173 (2003)
Index Fungorum number: IF28775; 2 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Mirannulata samuelsii Huhndorf, F.A. Fern., A.N. Mill. & Lodge
Natantiella Réblová, Mycol. Res. 113(9): 996 (2009)
Index Fungorum number: IF512833; 1 species with sequence data.
Type species – Natantiella ligneola (Berk. & Broome) Réblová
Naumovela Kravtzev, Trudy Inst. Bot., Alma-Ata 2: 153 (1955)
Index Fungorum number: IF3426; 2 morphological species (Species Fungorum 2020)
Type species – undetermined
Neocryptospora Petr., Sydowia 13(1–6): 51 (1959)
Index Fungorum number: IF3448; 1 morphological species
Type species – Neocryptospora rickii Petr.
Neoeriomycopsis Crous & M.J. Wingf., Persoonia 34: 207 (2015)
Index Fungorum number: IF812453; 1 species with sequence data.
Type species – Neoeriomycopsis aristata (B. Sutton & Hodges) Crous & M.J. Wing
Neolamya Theiss. & Syd., Annls mycol. 16(1/2): 29 (1918)
Index Fungorum number: IF3465; 3 morphological species (Species Fungorum 2020).
Type species – Neolamya peltigerae (Mont.) Theiss. & Syd
Neonawawia a J. Yang, K.D. Hyde & J.K. Liu, Mycosphere 9(6): 1132–1150 (2018)
Index Fungorum number: IF555449; 1 species with sequence data.
Type species – Neonawawia malaysiana (Crous & S.S. Lee) J. Yang, K.D. Hyde & J.K. Liu
Neothyridaria Petr., Annls mycol. 32(5/6): 347 (1934)
Index Fungorum number: IF3486; 1 morphological species.
Type species – Neothyridaria moravica Petr
Ophiomassaria Jacz., Bull. Herb. Boissier 2: 685 (1894)
Index Fungorum number: IF3606; 1 morphological species.
Type species – Ophiomassaria selenospora (G.H. Otth) Jacz
Ophiomeliola Starbäck, Bih. K. svenska VetenskAkad. Handl., Afd. 3 25(no. 1): 22 (1899)
Index Fungorum number: IF3607; 4 morphological species (Species Fungorum 2020).
Type species – Ophiomeliola lindmanii Starbäck
Paoayensis Cabanela, Jeewon & K.D. Hyde, Cryptog. Mycol. 28(4): 303 (2007)
Index Fungorum number: IF510824; 2 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Paoayensis lignicola Cabanela, Jeewon & K.D. Hyde
964
Paradiplococcium Hern.-Restr., J. Mena & Gené, Stud. Mycol. 86: 94 (2017)
Index Fungorum number: IF820295; 1 species with sequence data.
Type species – Paradiplococcium singulare (Hern. Restr., J. Mena, Gené & Guarro) Hern.
Restr., J. Mena & Gené
Paramicrodochium Hern.-Restr. & Crous, Persoonia 36: 62 (2015)
Index Fungorum number: IF811869; 1 species with sequence data.
Type species – Paramicrodochium gracile (Mouch. & Samson) M. Hern. Restr. & Crous
Pareutypella Y.M. Ju & J.D. Rogers, Mycologia 87(6): 891 (1996)
Index Fungorum number: IF27622; 2 morphological species (Species Fungorum 2020).
Type species – Pareutypella sulcata Y.M. Ju & J.D. Rogers
Phialemoniopsis Perdomo, Dania García, Gené, Cano & Guarro, Mycologia 105(2): 408 (2013)
Index Fungorum number: IF563874; 6 species with sequence data.
Type species – Phialemoniopsis ocularis (Gené & Guarro) Perdomo
Phragmeriella Hansf., Mycol. Pap. 15: 89 (1946)
Index Fungorum number: IF4019; 1 morphological species.
Type species – Phragmeriella ireninae Hansf
Phyllocelis Syd., Annls mycol. 23(3/6): 353 (1925)
Index Fungorum number: IF4061; 2 morphological species (Species Fungorum 2020).
Type species – Phyllocelis oyedaeae Syd
Pleocryptospora J. Reid & C. Booth, Can. J. Bot. 47: 1055 (1969)
Index Fungorum number: IF4209; 1 morphological species.
Type species – Pleocryptospora bambusae (Speg.) J. Reid & C. Booth
Pleosphaeria Speg., Anal. Soc. cient. argent. 12(4): 181 (1881)
Index Fungorum number: IF4227; 21 morphological species.
Type species – Pleosphaeria lichenothricis Henssen
Pleurophragmium Costantin, Mucéd. Simpl. (Paris): 100 (1888)
Index Fungorum number: IF9470; 21orphological species (Species Fungorum 2020), 5
species with sequence data.
Type species – Pleurophragmium bicolor Costantin
Protocucurbitaria Naumov, Bot. Mater. Otd. Sporov. Rast. Bot. Inst. Komarova Akad. Nauk
S.S.S.R. 7: 110 (1951)
Index Fungorum number: IF4385; 1 morphological species.
Type species – Protocucurbitaria ribicola Naumov
Pulvinaria Bonord., Handb. Allgem. mykol. (Stuttgart): 272 (1851)
Index Fungorum number: IF4552; 4 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Pulvinaria typica Rodway
Pumilus Viala & Marsais, C. r. hebd. Séanc. Agric. France 198: 1557, 1560 (1934)
Index Fungorum number: IF4555; 1 morphological species.
Type species – Pumilus medullae Viala & Marsais
965
Rehmiomycella E. Müll., Beitr. Kryptfl. Schweiz 11(no. 2): 601 (1962)
Index Fungorum number: IF4667; 1 morphological species.
Type species – Rehmiomycella phoradendri (Rehm) E. Müll
Rhamphosphaeria Kirschst., Annls mycol. 34(3): 189 (1936)
Index Fungorum number: IF338249; 1 morphological species.
Type species – Rehmiomycella phoradendri (Rehm) E. Müll
Rhizophila K.D. Hyde & E.B.G. Jones, Mycotaxon 34(2): 527 (1989)
Index Fungorum number: IF25336; 1 morphological species.
Type species – Rhizophila marina K.D. Hyde & E.B.G. Jones
Rhopographella (Henn.) Sacc. & Trotter, Syll. fung. (Abellini) 22: 440 (1913)
Index Fungorum number: IF4723; 2 morphological species (Species Fungorum 2020).
Type species – Rhopographella gaduae (Henn.) Sacc. & Trotter
Rhynchosphaeria (Sacc.) Berl., Icon. fung. (Abellini) 1(1): 40 (1890)
Index Fungorum number: IF4733; 17 morphological species (Species Fungorum 2020).
Type species – Rhynchosphaeria dusenii Henn.
Rivulicola K.D. Hyde, Nova Hedwigia 64(1-2): 186 (1997)
Index Fungorum number: IF27726; 3 morphological species (Species Fungorum 2020).
Type species – Rivulicola incrustata K.D. Hyde
Romellina Petr., Sydowia 9(1-6): 597 (1955)
Index Fungorum number: IF4781; 1 morphological species.
Type species – Romellina variabilis Petr
Saccardoëlla Speg., Michelia 1(no. 5): 461 (1879)
Index Fungorum number: IF4811; 14 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Saccardoella montellica Speg
Sartorya Vuill., C. r. hebd. Séanc. Acad. Sci., Paris 184: 136 (1927)
Index Fungorum number: IF1768; 8 morphological species (Species Fungorum 2020).
Type species – Sartorya fumigata Vuill
Scharifia Petr., Sydowia 9(1-6): 448 (1955)
Index Fungorum number: IF4883; 1 morphological species.
Type species – Scharifia procumbens Petr
Scoliocarpon Nyl., Mém. Soc. Acad. Maine Loire 4: 81 (1858)
Index Fungorum number: IF4960; 1 morphological species.
Type species – Scoliocarpon pupula (Tuck.) Nyl
Scotiosphaeria Sivan., Trans. Br. mycol. Soc. 69(1): 119 (1977)
Index Fungorum number: IF4971; 1 morphological species.
Type species – Scotiosphaeria endoxylinae Sivan
Selenosporella G. Arnaud ex MacGarvie, Scientific Proc. R. Dublin Soc., Ser. B 2(16): 153 (1969)
966
Index Fungorum number: IF9872; 13 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Selenosporella curvispora G. Arnaud ex MacGarvie
Servazziella J. Reid & C. Booth, Can. J. Bot. 65(7): 1334 (1987)
Index Fungorum number: IF25136; 1 morphological species.
Type species – Servazziella longispora (Servazzi) J. Reid & C. Booth
Sporoctomorpha J.V. Almeida & Sousa da Câmara, Revta agron., Lisb. 1: 90 (1903)
Index Fungorum number: IF5162; 1 morphological species.
Type species – Sporoctomorpha magnoliae J.V. Almeida & Sousa da Câmara
Stanjehughesia Subram., Proc. Indian natn Sci. Acad., Part B. Biol. Sci. 58(4): 184 (1992)
Index Fungorum number: IF22443; 17 morphological species (Species Fungorum 2020), 2
species with sequence data.
Type species – Stanjehughesia hormiscioides (Corda) Subram
Stearophora L. Mangin & Viala, C. r. hebd. Séanc. Acad. Sci., Paris 140: 1478 (1905)
Index Fungorum number: IF5188; 1 morphological species.
Type species – Stearophora radicicola L. Mangin & Viala
Steganopycnis Syd. & P. Syd., Annls mycol. 14(5): 370 (1916)
Index Fungorum number: IF5189; 1 morphological species.
Type species – Steganopycnis oncospermatis Syd. & P. Syd
Stegophorella Petr., Sydowia 1(1-3): 15 (1947)
Index Fungorum number: IF5198; 1 morphological species.
Type species – Stegophorella lagerheimii Petr
Stellosetifera Matsush., Matsush. Mycol. Mem. 9: 26 (1996)
Index Fungorum number: IF27672; 1 morphological species.
Type species – Stellosetifera malaysiana Matsush
Stereosphaeria Kirschst., Annls mycol. 37(1/2): 96 (1939)
Index Fungorum number: IF5226; 2 morphological species (Species Fungorum 2020), 1
species with sequence data.
Type species – Stereosphaeria phloeophila Kirschst
Stomatogenella Petr., Sydowia 9(1-6): 507 (1955)
Index Fungorum number: IF5271; 1 morphological species.
Type species – Stomatogenella mirabilis Petr
Sungaiicola Fryar & K.D. Hyde, Cryptog. Mycol. 25(3): 250 (2004)
Index Fungorum number: IF28836; 1 morphological species.
Type species – Sungaiicola bactrodesmiella Fryar & K.D. Hyde
Synsphaeria Bonord., Handb. Allgem. mykol. (Stuttgart): 271 (1851)
Index Fungorum number: IF5340; 2 morphological species (Species Fungorum 2020).
Type species – Sphaeria versatilis Fr
Teracosphaeria Réblová & Seifert, Mycol. Res. 111(3): 291 (2007)
Index Fungorum number: IF510202; 1 species with sequence data.
967
Type species – Teracosphaeria petroica Réblová & Seifert
Thelidiella Fink, Mycologia 25(4): 305 (1933)
Index Fungorum number: IF5423; 1 morphological species.
Type species – Thelidiella blastenicola Fink
Thyridella (Sacc.) Sacc., Syll. fung. (Abellini) 11: 351 (1895)
Index Fungorum number: IF5464; 3 morphological species (Species Fungorum 2020).
Type species – Thyridium vestitum (Fr.) Fuckel
Thyrotheca Kirschst., Hedwigia 81: 215 (1944)
Index Fungorum number: IF5473; 1 morphological species.
Type species – Thyrotheca nyssae (Berk. & M.A. Curtis) Kirschst
Trichospermella Speg., Anal. Mus. nac. Hist. nat. B. Aires 23: 38 (1912)
Index Fungorum number: IF5585; 2 morphological species (Species Fungorum 2020).
Type species – Trichospermella pulchella Speg
Trichosphaeropsis Bat. & Nascim., Atas Inst. Micol. Univ. Recife 1: 299 (1960)
Index Fungorum number: IF5589; 1 morphological species.
Type species – Trichosphaeropsis crescentiae Bat. & Nascim
Tunstallia Agnihothr., Phytopath. Z. 40(3): 280 (1961)
Index Fungorum number: IF5642; 2 morphological species (Species Fungorum 2020).
Type species – Tunstallia aculeata (Petch) Agnihothr
Tulipispora Révay & Gönczöl, Nova Hedwigia 88(1-2): 42 (2009)
Index Fungorum number: IF537065; 1 morphological species.
Type species – Tulipispora ingoldii Révay & J. Gönczöl
Urosporella G.F. Atk., Bulletin of Cornell University, Science 3(no. 1): 9 (1897)
Index Fungorum number: IF5690; 5 morphological species (Species Fungorum 2020).
Type species – Urosporella americana G.F. Atk
Urupe Viégas, Bragantia 4(1-6): 125 (1944)
Index Fungorum number: IF5691; 1 morphological species.
Type species – Urupe guaduae Viégas
Vleugelia J. Reid & C. Booth, Can. J. Bot. 47: 1057 (1969)
Index Fungorum number: IF5749; 1 morphological species.
Type species – Vleugelia betulina (Bubák & Vleugel) J. Reid & C. Booth
Xenodium Syd., Annls mycol. 33(1/2): 95 (1935)
Index Fungorum number: IF5817; 1 morphological species.
Type species – Xenodium petrakii Syd.
Zalerion R.T. Moore & Meyers, Can. J. Microbiol. 8: 408 (1962)
Index Fungorum number: IF10467; 6 morphological species (Species Fungorum 2020), 4
species with sequence data.
Type species – Zalerion nepura R.T. Moore & Meyers
968
Acknowledgments
Kevin D. Hyde thanks the Foreign Experts Bureau of Yunnan Province, Foreign Talents
Program (2018; grant no. YNZ2018002), a Visiting Professorship at Chiang Mai University,
Thailand, the Thailand Research grants entitled Biodiversity, phylogeny and role of fungal
endophytes on above parts of Rhizophora apiculata and Nypa fruticans (grant no: RSA5980068),
the future of specialist fungi in a changing climate: baseline data for generalist and specialist fungi
associated with ants, Rhododendron species and Dracaena species (grant no: DBG6080013),
Impact of climate change on fungal diversity and biogeography in the Greater Mekong Subregion
(grant no: RDG6130001), Mae Fah Luang University for the grant “Taxonomy diversity,
phylogeny and evolution of fungi in Capnodiales” (grant no: 666713), for supporting this study.
This work was partially supported by the National Natural Science Foundation of China (No.
NSFC 31950410558, 31760013). E.B.G. Jones is supported under the Distinguished Scientist
Fellowship Program (DSFP), King Saud University, Kingdom of Saudi Arabia. M. Stadler and
K.D. Hyde thank the Thailand Research Fund (TRF) and the German Academic Exchange Service
(DAAD) for a joint TRF-DAAD [PPP 2017–2018] academic exchange grant. V.V. Sarma would
like to thank SERB, Department of Science and Technology, Government of India, for funding a
project (SERB/SB/SO/PS/18/2014 dt.19.5.2015) and Ministry of Earth Sciences (MOES), Govt. of
India for funding a project (Sanction order: MOES/36/OO1S/Extra/40/ 2014/PC-IV dt.
14.01.2015); the Department of Biotechnology, Pondicherry University for facilities; forest
departments of Andaman and Nicobar Islands and Tamil Nadu, India are thanked for providing
permission to collect samples. M Niranjan thanks SERB, Govt. of India for a fellowship and B.
Devadatha thanks MOES, Govt. of India for a fellowship. Dong-Qin Dai would like to thank
Yunnan Province Universities of the Science and Technology Innovation Team for the exploitation
and utilisation of endophytes and the Thousand Talents Plan, Youth Project of Yunnan Provinces
for support. D. N. Wanasinghe would like to thank the CAS President’s International Fellowship
Initiative (PIFI) for funding his postdoctoral research (number 2019PC0008), the National Science
Foundation of China and the Chinese Academy of Sciences for financial support under the
following grants: 41761144055, 41771063 and Y4ZK111B01. Rajesh Jeewon thanks the
University of Mauritius for support. Saowaluck Tibpromma would like to thanks the International
Postdoctoral Exchange Fellowship Program (number Y9180822S1), CAS President’s International
Fellowship Initiative (PIFI) (number 2020PC0009), and China Postdoctoral Science Foundation for
funding her postdoctoral research and financial support. Shi-Ke Huang, Xiang-Yu Zeng and YuanPin Xiao were supported by the National Natural Science Foundation of China (No. 31760014) and
the Science and Technology Foundation of Guizhou Province (No. [2017]5788). They would like
to thank Ting-Chi Wen for his support and help. LS Dissanayake would like to thank JC Kang for
funding her Research (NSFC Grants nos. 31670027, 31460011, 30870009). Mingkwan Doilom
would like to thank the 5th batch of Postdoctoral Orientation Training Personnel in Yunnan
Province (grant no.: Y934283261) and the 64th batch of China Postdoctoral Science Foundation
(grant no.: Y913082271). We would like to thank Prof. Abdel-Wahab, Prof. Chang-Hsin Kuo, Dr.
Rungtiwa Phookamsak, Dr. Anupama Daranagama and Dr. Lu Zonglong for their support and give
advice. Jing Yang would like to thank Zuoyi Liu for his support and great help with the laboratory
work. M. Réblová is appreciated for commenting on some parts of the manuscript.
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